Major clades of Agaricales: a multilocus phylogenetic overview

P. Brandon Matheny*, Judd M. Curtisa, Valérie Hofstetterb, M. Catherine Aimec, Jean-Marc Moncalvod, Zai-Wei Ge, Zhu-Liang Yange, Jason C. Slotf, Joseph F. Ammiratig, Timothy J. Baronih, Neale L. Bougheri, Karen W. Hughesj, D. Jean Lodgek, Richard W. Kerriganl, Michelle T. Seidlm, Duur K. Aanenn, Matthew DeNitiso, Graciela M. Danielep, Dennis E. Desjardinq, Bradley R. Kroppr, Lorelei L. Norvells, Andrew Parkert, Else C. Vellingau, Rytas Vilgalysb & David S. Hibbettaa Biology Department, Clark University, 950 Main Street, Worcester, Massachusetts, 01610b Department of Biology, Box 90338, Duke University, Durham, North Carolina 27708c USDA-ARS, Systematic Botany and Mycology Laboratory, Room 304, Building 011A, 10300 Baltimore Avenue, Beltsville, Maryland 20705-2350d Centre for Biodiversity and Conservation Biology, Royal Ontario Museum and Department of Botany, University of Toronto, Toronto, Ontario, M5S 2C6 Canadae Kunming Institute of Botany, Chinese Academy of Sciences, Kunming 650204, P.R. Chinaf Biology Department, Clark University, 950 Main Street, Worcester, Massachusetts, 01609g University of Washington, Biology Department, Box 355325, Seattle, Washington 98195h Department of Biological Sciences, SUNY Cortland, Box 2000, Cortland, New York 13045-0900i Department of Environment and Conservation, Locked Bag 104, Bentley Delivery Centre, WA 6983, Australiaj Botany Department, 437 Hesler Biology Building, University of Tennessee, Knoxville, Tennessee 37996-1100k International Institute of Tropical Forestry, USDA Forest Service – FPL, PO Box 1377 Luqillo, PR 00773-1377l Sylvan Research, 198 Nolte Drive, Kittanning, Pennsylvania 16201m Environmental Microbiology Laboratory Inc., 1400 12th Avenue SE, Bellevue, Washington 98004n Laboratory of Genetics, Arboretumlaan 4, 6703 BD, Wageningen, The Netherlandso 127 Harrington Way, Worcester, Massachusetts 01604p Instituto Multidisciplinario de Biología Vegetal, CONICET-Universidad Nacional de Córdoba, Casilla de Correo 495, 5000 Córdoba, Argentinaq Department of Biology, San Francisco State University, San Francisco, California 94132r Department of Biology, Utah State University, Logan, Utah 84322s Pacific Northwest Mycology Service, 6720 NW Skyline Boulevard, Portland, Oregon 97229-1309t 127 Raven Way, Metaline Falls, Washington 99153-9720u Department of Plant and Microbial Biology, 111 Koshland Hall, University of California, Berkeley, California 94720-3102 1 Corresponding author. E-mail: pmatheny@clarku.eduAn overview of the phylogeny of the Agaricales is presented based on a multilocus analysis of a six-gene region supermatrix. Bayesian analyses of 5611 nucleotide characters of rpb1, rpb1-intron 2, rpb2 and 18S, 25S, and 5.8S ribosomal RNA genes recovered six major clades, which are recognized informally and labeled the Agaricoid, Tricholomatoid, Marasmioid, Pluteoid, Hygrophoroid and Plicaturopsidoid clades. Each clade is discussed in terms of key morphological and ecological traits. At least 11 origins of the ectomycorrhizal habit appear to have evolved in the Agaricales, with possibly as many as nine origins in the Agaricoid plus Tricholomatoid clade alone. A family-based phylogenetic classification is sketched for the Agaricales, in which 30 families, four unplaced tribes and two informally named clades are recognized.