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Systematics of Calonectria: a Genus of Root, Shoot and Foliar Pathogens

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Systematics of Calonectria: a Genus of Root, Shoot and Foliar Pathogens Studies in Mycology 66 (2010) Systematics of Calonectria: a genus of root, shoot and foliar pathogens Lorenzo Lombard, Pedro W. Crous, Brenda D. Wingfield and Michael J. Wingfield CBS-KNAW Fungal Biodiversity Centre, Utrecht, The Netherlands An institute of the Royal Netherlands Academy of Arts and Sciences Studies in Mycology The Studies in Mycology is an international journal which publishes systematic monographs of filamentous fungi and yeasts, and in rare occasions the proceedings of special meetings related to all fields of mycology, biotechnology, ecology, molecular biology, pathology and systematics. For instructions for authors see www.cbs.knaw.nl. ExEcutivE Editor Prof. dr dr hc Robert A. Samson, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] Layout Editor Manon van den Hoeven-Verweij, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] SciEntific EditorS Prof. dr Dominik Begerow, Lehrstuhl für Evolution und Biodiversität der Pflanzen, Ruhr-Universität Bochum, Universitätsstr. 150, Gebäude ND 44780, Bochum, Germany. E-mail: [email protected] Prof. dr Uwe Braun, Martin-Luther-Universität, Institut für Biologie, Geobotanik und Botanischer Garten, Herbarium, Neuwerk 21, D-06099 Halle, Germany. E-mail: [email protected] Dr Paul Cannon, CABI and Royal Botanic Gardens, Kew, Jodrell Laboratory, Royal Botanic Gardens, Kew, Richmond, Surrey TW9 3AB, U.K. E-mail: [email protected] Prof. dr Lori Carris, Associate Professor, Department of Plant Pathology, Washington State University, Pullman, WA 99164-6340, U.S.A. E-mail: [email protected] Prof. dr Pedro W. Crous, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] Prof. dr David M. Geiser, Department of Plant Pathology, 121 Buckhout Laboratory, Pennsylvania State University, University Park, PA, U.S.A. 16802. E-mail: [email protected] Dr Johannes Z. Groenewald, CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. E-mail: [email protected] Prof. dr David S. Hibbett, Department of Biology, Clark University, 950 Main Street, Worcester, Massachusetts, 01610-1477, U.S.A. E-mail: [email protected] Prof. dr Hyeon-Dong Shin, Division of Environmental Science & Ecological Engineering, Korea University, Seoul 136-701, Korea. E-mail: [email protected] Dr Lorelei L. Norvell, Pacific Northwest Mycology Service, 6720 NW Skyline Blvd, Portland, OR, U.S.A. 97229-1309. E-mail: [email protected] Dr Erast Parmasto, Institute of Zoology & Botany, 181 Riia Street, Tartu, Estonia EE-51014. E-mail: [email protected] Prof. dr Alan J.L. Phillips, Faculdade de Ciências e Tecnologia, Universidade Nova de Lisboa, Quinta de Torre, 2829-516 Caparica, Portugal. E-mail: [email protected] Dr Amy Y. Rossman, Rm 304, Bldg 011A, Systematic Botany & Mycology Laboratory, Beltsville, MD, U.S.A. 20705. E-mail: [email protected] Dr Keith A. Seifert, Research Scientist / Biodiversity (Mycology and Botany), Agriculture & Agri-Food Canada, KW Neatby Bldg, 960 Carling Avenue, Ottawa, ON, Canada K1A OC6. E-mail: [email protected] Dr Roger Shivas, Plant Pathology Herbarium, Agri-Science Queensland, Indooroopilly, Qld 4068, Australia. E-mail: [email protected] Prof. dr Jeffrey K. Stone, Department of Botany & Plant Pathology, Cordley 2082, Oregon State University, Corvallis, OR, U.S.A. 97331-2902. E-mail: [email protected] Dr Richard C. Summerbell, 27 Hillcrest Park, Toronto, Ont. M4X 1E8, Canada. E-mail: [email protected] Prof. dr Brett Summerell, Royal Botanic Gardens and Domain Trust, Mrs. Macquaries Road, Sydney, NSW 2000, Australia. E-mail: [email protected] Prof. dr Ulf Thrane, Department of Systems Biology, Center for Microbial Biotechnology, Technical University of Denmark, Søltofts Plads 221, DK-2800 Kgs. Lyngby, Denmark. E-mail: [email protected] Copyright 2010 CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. 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. Publication date: 15 June 2010 Published and distributed by CBS-KNAW Fungal Biodiversity Centre, P.O. Box 85167, 3508 AD Utrecht, The Netherlands. Internet: www.cbs.knaw.nl. E-mail: [email protected]. ISBN/EAN : 978-90-70351-81-6 Online ISSN : 1872-9797 Print ISSN : 0166-0616 Cover: Top from left to right: Cylindrocladium leaf blight of a Eucalyptus sp. Calonectria polizzii conidiophore branches. Yellow perithecium of Ca. colhounii. Bottom from left to right: Calonectria colombiana, macroconidiophore. Calonectria polizzii, conidiogenous apparatus with conidiophore branches and doliiform to reniform phialides. Calonectria eucalypti, perithecium. Asci containing eight ascospores. CONTENTS L. Lombard, P.W. Crous, B.D. Wingfield and M.J. Wingfield: Species concepts in Calonectria (Cylindrocladium) ......................................... 1 L. Lombard, P.W. Crous, B.D. Wingfield and M.J. Wingfield: Multigene phylogeny and mating tests reveal three cryptic species related to Calonectria pauciramosa ............................................................................................................................................................................... 15 L. Lombard, P.W. Crous, B.D. Wingfield and M.J. Wingfield: Phylogeny and systematics of the genus Calonectria ................................... 31 INDEX ............................................................................................................................................................................................................ 70 available online at www.studiesinmycology.org StudieS in Mycology 66: 1–14. 2010. doi:10.3114/sim.2010.66.01 species concepts in Calonectria (Cylindrocladium) L. Lombard1*, P.W. Crous2, B.D. Wingfield3 and M.J. Wingfield1 1Department of Microbiology and Plant Pathology, Tree Protection Co-operative Programme, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0002, South Africa; 2CBS-KNAW Fungal Biodiversity Centre, Uppsalalaan 8, 3584 CT Utrecht, The Netherlands; 3Department of Genetics, Forestry and Agricultural Biotechnology Institute, University of Pretoria, Pretoria 0002, South Africa *Correspondence: Lorenzo Lombard, [email protected] Abstract: Species of Calonectria and their Cylindrocladium anamorphs are important plant pathogens worldwide. At present 52 Cylindrocladium spp. and 37 Calonectria spp. are recognised based on sexual compatibility, morphology and phylogenetic inference. The polyphasic approach of integrating Biological, Morphological and Phylogenetic Species Concepts has revolutionised the taxonomy of fungi. This review aims to present an overview of published research on the genera Calonectria and Cylindrocladium as they pertain to their taxonomic history. The nomenclature as well as future research necessary for this group of fungi are also briefly discussed. Key words: Calonectria, Cylindrocladium, species concepts, nomenclature, pathogenicity. INtroductIoN topic is not treated other than in the manner in which it applies to Calonectria. The genus Calonectria (Ca.) was erected in 1867 by De Notaris, based on Ca. daldiniana collected on leaves of Magnolia grandiflora (Magnoliaceae), in Daldini, Italy (Rossman 1979a). Rossman TaxoNomIc hIstory (1979a) later reduced Ca. daldiniana to synonymy under Ca. pyrochroa, and defined this nectrioid fungus as having an ascocarp Calonectria resides in the Nectriaceae, one of three families wall structure that is brightly coloured, changing to blood-red in in Hypocreales, an order that has been reviewed extensively 3 % KOH solution, warty to scaly and with a Cylindrocladium (Cy.) (Rogerson 1970, Rossman 1983, Rossman et al. 1996, 1999). The anamorph (Rossman 1993, Rossman et al. 1999). However, due Nectriaceae is circumscribed as having uniloculate ascomata that to the restricted morphological characteristics of the teleomorph are orange to purple and not immersed in well-developed stromata (Rossman 1979b, 1983), specimens can in many cases only be (Rossman et al. 1999). The family includes approximately 20 identified to species level if the anamorph is present (Schoch et al. genera of socio-economic importance and of these, Calonectria is 2000b, Crous 2002). most clearly distinguished from the others by its Cylindrocladium The anamorph genus Cylindrocladium, which is based on Cy. anamorphs and relevance as plant pathogens. scoparium, was first described by Morgan (1892) in the U.S.A., The first monograph of Cylindrocladium by Boedijn & Reitsma where it was found growing as saprobe on a pod of Gleditsia (1950), introduced seven Cylindrocladium species with one triacanthos. Although Morgan (1892) failed to mention the
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