The Mycota Edited by K. Esser and P.A. Lemke

Springer-Verlag Berlin Heidelberg GmbH

The Mycota

I Growth, Differentiation and Sexuality Ed. by IG.H. Wessels and F. Meinhardt

II Genetics and Biotechnology Ed. by U. Kiick

III Biochemistry and Molecular Biology Ed. by R. Brambl and G. Marzluf

IV Environmental and Microbial Relationships Ed. by D. Wicklow and B. Soderstrom

V Plant Relationships Ed. by G. Carroll and P. Tudzynski

VI Human and Animal Relationships Ed. by D.H. Howard and ID. Miller

VII Systematics and Evolution Ed. by D.I McLaughlin, E.G. McLaughlin, and P.A. Lemke

VIII Biology of the Fungal Cell Ed. by R.I Howard and N.A.R. Gow

IX Fungal Associations Ed. by B. Hock

X Industrial Applications Ed. by H.D. Osiewacz

XI Agricultural Applications Ed. by F. Kempken

XII Human Fungal Pathogens Ed. by lE. Domer and G.S. Kobayashi

The Mycota A Comprehensive Treatise an Fungi as Experimental Systems for Basic and Applied Research

Edited by K. Esser and P.A. Lemke

VII Volume Editors:

Systematics and Evolution PartA

D.I McLaughlin, E.G. McLaughlin, and P.A. Lemke

With 217 Figures and 20 Tables

Springer

Series Editors

Professor Dr. Dr. h.c. mult. KARL ESSER Allgemeine Botanik Ruhr-Universităt

44780 Bochum, Germany Tel.: +49(234)32-22211 Fax: +49(234)32-14211 e-mail: Karl.Esser@ruhr-uni-bochum.de

Professor Dr. PAUL A. LEMKE t, Auburn, USA

Volume Editors

Professor Dr. David 1. McLaughlin Department of Plant Biology University of Minnesota St. Paul, MN 55108-1095, USA Tel.: +1(612)625-5736 Fax: +1(612)625-1738 e-mail: davem@tc.umn.edu

Professor Dr. Esther G. McLaughlin Department of Biology Augsburg College Minneapolis, MN 55454, USA Tel.:+ 1(612)330-1074 Fax: +1(612)330-1649 e-mail: McLaugh@augsburg.edu

Professor Dr. PAUL A. LEMKE t, Auburn, USA

Library of Congress Cataloging-in-Publication Data The Mycota. Includes bibliographical references and index. Contents: 1. Growth, differentiation, and sexuality/editors. J.G.H. Wessels and F. Meinhardt - 2. Genetics and biotechnology. 1. Mycology. 2. Fungi. 3. Mycology - Research. 4. Research.1. Esser, Karl, 1924- . II. Lemke, Paul A., 1937- . QK603.M87 1994 589.2 ISBN 978-3-642-08193-4 ISBN 978-3-662-10376-0 (eBook) DOI 10.1007/978-3-662-10376-0

This work is subject to copyright. AlI rights are reserved, whether the whole or part of the material is concerned, specifi­cally the rights of translation, reprinting, reuse of illustrations, recitation, broadcasting. reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag Berlin Heidelberg GmbH. Violations are liable for prosecution under the German Copyright Law.

© Springer-Verlag Berlin Heidelberg 2001

Originally published by Springer-Verlag Berlin Heidelberg New York in 2001 Softcover reprint of the hardcover 1 st edition 2001

The use of general descriptive names, registered names, trademarks, etc, in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and there­fore free for general use.

Production Editor: PRO EDIT GmbH, Heidelberg, Germany Cover design: Springer-Verlag, E. Kirchner Typesetting: Best-set Typesetter Ltd., Hong Kong Printed on acid-free paper SPIN: 10033665 3113130/So 5 4 3 2 1 O

Series Preface

Mycology, the study of fungi, originated as a subdiscipline of botany and was a des­criptive discipline, largely neglected as an experimental science until the early years of this century. A seminal paper by Blakeslee in 1904 provided evidence for self-incom­patibility, termed "heterothallism", and stimulated interest in studies related to the control of sexual reproduction in fungi by mating-type specificities. Soon to folIow was the demonstration that sexualIy reproducing fungi exhibit Mendelian inheritance and that it was possible to conduct formal genetic analysis with fungi. The names Burgeff, Kniep and Lindegren are alI associated with this early period of fungal genetics research.

These studies and the discovery of penicillin by Fleming, who shared a Nobel Prize in 1945, provided further impetus for experimental research with fungi. Thus began a period of interest in mutation induction and analysis of mutants for biochemical traits. Such fundamental research, conducted largely with Neurospora crassa, led to the one gene: one enzyme hypothesis and to a second Nobel Prize for fungal research awarded to Beadle and Tatum in 1958. Fundamental research in biochemical genetics was extended to other fungi, especialIy to Saccharomyces cerevisiae, and by the mid-1960s fungal systems were much favored for studies in eukaryotic molecular biology and were soon able to corn pete with bacterial systems in the molecular arena.

The experimental achievements in research on the genetics and molecular biology of fungi have benefited more generalIy studies in the re1ated fields of fungal biochemistry, plant pathology, medical mycology, and systematics. Today, there is much interest in the genetic manipulation of fungi for applied research. This current interest in biotechnical genetics has been augmented by the development of DNA­mediated transformation systems in fungi and by an understanding of gene expres sion and regulation at the molecular level. Applied research initiatives involving fungi extend broadly to are as of interest not only to industry but to agricultural and envi­ronmental sciences as welI.

It is this burgeoning interest in fungi as experimental systems for applied as well as basic research that has prompted publication of this series of books under the title The Mycota. This title knowingly relegates fungi into a separate realm, distinct from that of either plants, animals, or protozoa. For consistency throughout this Series of Volumes the names adopted for major groups of fungi (representative genera in paren­theses) are as folIows:

Pseudomycota

Division: Division:

Eumycota

Division: Division: Division:

Oomycota (Achlya, Phytophthora, Pythium) Hyphochytriomycota

Chytridiomycota (Allomyces) Zygomycota (Mucor, Phycomyces, Blakeslea) Dikaryomycota

Volume Preface

This is an exciting time to produce an overview of the systematics and evolution of the fungi. Homoplasy is evident in alllineages, e.g., those based on the gross morphology of the chytrid zoospore, the perithecium and apothecium, the smut teliospore and the agaric fruiting body, and some classifications based on light microscope morphology have been shown to be unsound. Molecular and subcellular characters, aided by new methods of phylogenetic analysis, have allowed us to see through the conflicts between various phenetic classification schemes and have given us some confidence that we are beginning to achieve a true phylogeny of the fungi. Molecular data have both supported ultra structural characters that first began to unravel the homoplasies unrec­ognized at the light microscopic level, and have also revealed the relationships of fungi to other eukaryotes. They continue to enlarge the scope of the fungi, e.g., with the recent addition of the Microsporidia (see Cavalier-Smith, Chap. 1, VoI. VII, Part A), and they have shown the need for more detailed chemical, subcellular, and developmental studies for a fuller understanding of these organisms and their relationships.

This volume is a mixture of phylogenetic and more classical systematics. Progress in knowledge of species and development of taxonomic characters is mixed. Groups with few species have been studied in great detail, while in groups with large numbers of species much effort is still needed to find and determine the taxa. Classical system­atics groups organisms on a phenetic basis, then sets up a classification; phylogeny is a secondary consideration. Phylogenetic systematics first determines organism relation­ships, then constructs a systematic classification that reflects the phylogeny. Molecular characters have made possible the establishment of a monophyletic and hopefully more permanent classification for the fungi. Thus, Volume VII of The Mycota contains both classical and phylogenetic classifications, reflecting the available data and the ori­entation of different authors. The incompleteness of some classifications, e.g., those for the Urediniomycetes (Swann, Frieders and McLaughlin, Chap. 2, VoI. VII, Part B) and Homobasidiomycetes (Hibbett and Thorn, Chap. 5, VoI. VII, Part B), demonstrates that we are in the early stages of a phylogenetic systematics for these groups.

The taxonomic outline used in The Mycota, VoI. VII, differs somewhat from that of other volumes in the series (Table 1), reflecting current mycological systematics. There is a lack of agreement on the naming of higher taxa, and the rules of nomen­clature permit more than one name for these taxa. Cavalier-Smith (Chap. 1, VoI. VII, Part A) presents an alternative view to the taxonomie outline used for the remainder of the volume (Table 2). Some of the nomenclatural problems stern from a lack of res­olution of deep branches in molecular evolutionary trees, a problem that appears likely to be resolved only with additional data from multiple genes and the addition of missing taxa to the analysis. Problems also arise from a differenee of opinion among authors. The term fungi has assumed an ecological meaning for all organisms with a similar nutritional mode, and, therefore, Eumycota, rather than Fungi, is less confus­ing for the members of the phylum that encompasses a monophyletic group of these organisms. Pseudofungi (Cavalier-Smith, Chap. 1, VoI. VII, Part A) implies that organ­isms that lie outside the Eumycota but possess the fungal lifestyle are not fungi, but

Series Preface VII

insect-associated fungi and the medically important pathogens afflicting humans. Yes, fungi are ubiquitous and important.

There are many fungi, conservative estimates are in the order of 100000 species, and there are many ways to study them, from descriptive accounts of organisms found in nature to laboratory experimentation at the cellular and molecular level. AU such studies expand our knowledge of fungi and of fungal processes and improve our ability to utilize and to control fungi for the benefit of humankind.

We have invited leading research specialists in the field of mycology to contribute to this Series. We are especially indebted and grateful for the initiative and leadership shown by the Volume Editors in selecting topics and assembling the experts. We have aU been a bit ambitious in producing these Volumes on a timely basis and therein lies the possibility of mistakes and oversights in this first edition. We encourage the read­ership to draw our attention to any error, omission or inconsistency in this Series in order that improvements can be made in any subsequent edition.

Finally, we wish to acknowledge the willingness of Springer-Verlag to host this project, which is envisioned to require more than 5 years of effort and the publication of at least nine Volumes.

Bochum, Germany Auburn, AL, USA April1994

KARL ESSER

PAUL A. LEMKE

Series Editors

Addendum to the Series Preface

In early 1989, encouraged by Dieter Czeschlik, Springer-Verlag, Paul A. Lemke and 1 began to plan The Mycota. The first volume was released in 1994, five other volumes followed in the subsequent years. Also on behalf of Paul A. Lemke, 1 would like to take this opportunity to thank Dieter Czeschlik, his colleague Andrea Schlitzberger, and Springer-Verlag for their help in realizing the enterprise and for their excellent cooperation for many years.

Unfortunately, after a long and serious illness, Paul A. Lemke died in November 1995. Without his expertise, his talent for organization and his capability to grasp the essentials, we would not have been able to work out a concept for the volumes of the series and to acquire the current team of competent volume editors. He also knew how to cope with unexpected problems which occurred after the completion of the manu­scripts. His particular concern was directed at Volume VII; in this volume, a posthu­mous publication of his is included.

Paul A. Lemke was an outstanding scientist interested in many fields. He was extremely wise, dedicated to his profession and a preeminent teacher and researcher. Together with the volume editors, authors, and Springer-Verlag, 1 mourn the loss of a very good and reliable friend and colleague.

Bochum, Germany April2000

KARL ESSER

Volume Preface

This is an exciting time to produce an overview of the systematics and evolution of the fungi. Homoplasy is evident in alllineages, e.g., those based on the gross morphology of the chytrid zoospore, the perithecium and apothecium, the smut teliospore and the agaric fruiting body, and some classifications based on light microscope morphology have been shown to be unsound. Molecular and subcellular characters, aided by new methods of phylogenetic analysis, have allowed us to see through the conflicts between various phenetic classification schemes and have given us some confidence that we are beginning to achieve a true phylogeny of the fungi. Molecular data have both supported ultra structural characters that first began to unravel the homoplasies unrec­ognized at the light microscopic level, and have also revealed the relationships of fungi to other eukaryotes. They continue to enlarge the scope of the fungi, e.g., with the recent addition of the Microsporidia (see Cavalier-Smith, Chap. 1, VoI. VII, Part A), and they have shown the need for more detailed chemical, subcellular, and developmental studies for a fuller understanding of these organisms and their relationships.

This volume is a mixture of phylogenetic and more classical systematics. Progress in knowledge of species and development of taxonomic characters is mixed. Groups with few species have been studied in great detail, while in groups with large numbers of species much effort is still needed to find and determine the taxa. Classical system­atics groups organisms on a phenetic basis, then sets up a classification; phylogeny is a secondary consideration. Phylogenetic systematics first determines organism relation­ships, then constructs a systematic classification that reflects the phylogeny. Molecular characters have made possible the establishment of a monophyletic and hopefully more permanent classification for the fungi. Thus, Volume VII of The Mycota contains both classical and phylogenetic classifications, reflecting the available data and the ori­entation of different authors. The incompleteness of some classifications, e.g., those for the Urediniomycetes (Swann, Frieders and McLaughlin, Chap. 2, VoI. VII, Part B) and Homobasidiomycetes (Hibbett and Thorn, Chap. 5, VoI. VII, Part B), demonstrates that we are in the early stages of a phylogenetic systematics for these groups.

The taxonomic outline used in The Mycota, VoI. VII, differs somewhat from that of other volumes in the series (Table 1), reflecting current mycological systematics. There is a lack of agreement on the naming of higher taxa, and the rules of nomen­clature permit more than one name for these taxa. Cavalier-Smith (Chap. 1, VoI. VII, Part A) presents an alternative view to the taxonomie outline used for the remainder of the volume (Table 2). Some of the nomenclatural problems stern from a lack of res­olution of deep branches in molecular evolutionary trees, a problem that appears likely to be resolved only with additional data from multiple genes and the addition of missing taxa to the analysis. Problems also arise from a differenee of opinion among authors. The term fungi has assumed an ecological meaning for all organisms with a similar nutritional mode, and, therefore, Eumycota, rather than Fungi, is less confus­ing for the members of the phylum that encompasses a monophyletic group of these organisms. Pseudofungi (Cavalier-Smith, Chap. 1, VoI. VII, Part A) implies that organ­isms that lie outside the Eumycota but possess the fungal lifestyle are not fungi, but

XII Volume Preface

Table 1. Taxonomie out1ine at the kingdom, phylum, and class levels as used in other volumes in the series and in this Volume. The clas­sification in this volume is necessarily confusing at this time because authors are using their own classifications, rather than an imposed classification

Mycota, VoI. I

PSEUDOMYCOTA Oomycota

Hyphochytriomycota

EUMYCOTA Chytridiomycota

Zygomycota

Dikaryomycota Ascomycotina

Saccharomycetes Ascomycetes

Basidiomycotina Heterobasidiomycetes

Homobasidiomycetes

• Artificial taxon.

Mycota, VoI. VII

PSEUDOMYCOTA··b

OomycotaC

Peronosporomycetes Hyphochytriomycota

Hyphochytriomycetes Plasmodiophoromycota

Plasmodiophoromycetes

EUMYCOTA Chytridiomycotad

Chytridiomycetes Zygomycotad

Zygomycetes Trichomycetes

Ascomycotae

Saccharomycetes Plectomycetes Hymenoascomycetes' Loculoascomycetes'

Basidiomycota Urediniomycetes Ustilaginomycetes Heterobasidiomycetes'.f Homobasidiomycetes··f

b For a natural classification for Oomycota and Hyphochytrio­mycota, kingdom Stramenopila (Stramenipila, Dick, Chap. 2, VoI. VII, Part A) or Chromista have been proposed, and for Plasmodio­phoromycota, kingdom Protozoa (see Cavalier-Smith, Chap. 1, VoI. VII, Part A). C Or Heterokonta (see Cavalier-Smith, Chap. 1, and Dick, Chap. 2, VoI. VII, Part A). d Probably paraphyletic (see Cavalier-Smith, Chap. 1, VoI. VII, Part A, and Berbee and Taylor, Chap. 10, VoI. VII, Part B). e A phylogenetic classification for Ascomycota is not available. Current thinking among ascomycete scholars is that three classes should be recognized, as follows: "Archiascomycetes", which may not be monophyletic, Hemiascomycetes (see Kurtzman and Sugiyama, Chap. 9, VoI. VII, Part A), and a filamentous group, Euascomycetes, that eventually will be subdividable, perhaps at the subclass level [M.E. Berbee and J.w. Taylor, 1995, Can J Bot 73 (SuppI.1):S677, and Chap. 10, VoI. VII, Part B; J.w. Spatafora, 1995, Can J Bot 73 (Suppl. 1):S811]. Saccharomycetes as used here (see Barr, Chap. 8, VoI. VII, Part A) includes "Archiascomycetes" and Hemiascomycetes. See the relevant chapters for further speculation on the ultimate disposition of these groups. f Heterobasidiomycetes as used in VoI. VIIB cannot be separated from Homobasidiomycetes. Hymenomycetes [E.C. Swann and J.W. Taylor, 1995, Can J Bot 73 (Suppl. 1):S862] has been proposed as a class for these groups (see Berbee and Taylor, Chap. 10, VoI. VII, Part B).

Volume Preface

Table 2. Taxonomie outline at the kingdom, phylum, and class levels as used in the rest of this volume compared with that of Cavalier­Smith, Chap. 1, VoI. VII, Part A

Mycota, VoI. VII

PSEUDOMYCOTN Oomycota

Peronosporomycetes Hyphochytriomycota

Hyphochytriomycetes

Plasmodiophoromycota Plasmodiophoromycetes

EU MYCOTA Chytridiomycota

Chytridiomycetes

Zygomycota Zygomycetes

Trichomycetes

Ascomycota Saccharomycetes

Plectomycetes H ymenoascomycetes

Loculoascomycetes Basidiomycota

Urediniomycetes Ustilaginomycetes Heterobasidiomycetes Homobasidiomycetes

a Artificial taxon. b Probably paraphyletic.

Chapter 1, VoI. VII, Part A

CHROMISTA Bigyra

Oomycetes

Hyphochytrea

PROTOZOA Cercozoa

Phytomyxea

FUNGI Archemycota

Chytridiomycetes Allomycetes

Zygomycetes Bolomycetes Glomomycetesb

Enteromycetes Zoomycetes'

Microsporidia Minisporea Microsporea

Ascomycota Taphrinomycetes Geomycetes Endomycetes Plectomycetes Discomycetes Pyrenomycetes Loculomycetes

Basidiomycota Septomycetes Ustomycetes Gelimycetesb

Homobasidiomycetes

, lncludes Zygomycetes, Ascomycetes, and Trichomycetes.

XIII

in an ecological sense they are fungi. Pseudomycota is therefore used in this series for these fungal organisms that lie outside the Eumycota.

The Mycota, VoI. VII, includes treatments of the systematics and related topics of the Eumycota and Pseudomycota as well as specialized chapters on nomenclature, techniques, and evolution. Certain groups are not treated in this volume: the Labyrinthulomycetes (Pseudomycota) and the slime molds. The evolutionary position of the slime molds has been controversial. Recent evidence suggests that most slime molds are more closely related to the Eumycota than previously believed (S.L. Baldauf and w.F. Doolittle, 1997, Proc Natl Acad Sci USA 94:12007) and they should continue to be of interest to those who study fungi for both ecological and phylogenetic reasons.

Chapters 2 to 4, VoI. VII, Part A, cover the Pseudomycota and Chapters 5-14, VoI. VII, Part A, and Chapters 1-5, VoI. VII, Part B, the Eumycota. The Pseudomycota con­tains distantly related groups of fungi (Table 1). The Chytridiomycota and Zygomy­cota are treated in one and two chapters, respectively, while the Ascomycota and Basidiomycota are treated in five or six chapters each, with separate chapters for yeasts

XIV Volume Preface

in each phylum, although the yeasts are not monophyletic groups. Chapter 14, VoI. VII, Part A, discusses the special problems of anamorphic genera and their relationships to the teleomorphic genera, and describes the attempts being made to incorporate anamorphs into modern phylogenetic systematics. In Chapter 6, VoI. VII, Part B, Hawksworth discusses the development of a unified system of biological nomencla­ture. Chapters 7 and 8, VoI. VII, Part B, deal with techniques for cultivation and data analysis, respectively. The final two chapters in VoI. VII, Part B, consider speciation and molecular evolution.

The Mycota, VoI. VII, was originally intended to have been Volume I in the series. Several changes in editors and the unfortunate death of Paul Lemke delayed its pro­duction. Added to these difficulties was the fact that these are tumultuous times in sys­tematics because of the rapid development of molecular and phylogenetic analysis techniques and the explosive accumulation of data. As these techniques and new data are more broadly incorporated into systematics, a more stable and useful classification of the fungi will re suIt.

We thank Heather 1. Olson for her substantial efforts in compiling the indices.

St. Paul, Minnesota, USA Minneapolis, Minnesota, USA April2000

DAVID 1. McLAUGHLIN

ESTHER G. McLAUGHLIN

Volume Editors

Contents Part A

The Fungal Hierarchy

1 What are Fungi? T. CAVALIER-SMITH (With 6 Figures) 3

2 The Peronosporomycetes M.W. DICK (With 25 Figures). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39

3 Hyphochytriomycota M.S. FULLER (With 1 Figure) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 73

4 Plasmodiophoromycota IP. BRASELTON (With 8 Figures) 81

5 Chytridiomycota D.IS. BARR (With 11 Figures) ....................................... 93

6 Zygomycota: Zygomycetes G.L. BENNY, R.A. HUMBER, and lE. MORTON (With 37 Figures)

7 Zygomycota: Trichomycetes G.L. BENNY (With 8 Figures)

8 Ascomycota

113

147

M.E. BARR (With 2 Figures) ........................................ 161

9 Ascomycetous Yeasts and Yeastlike Taxa c.P. KURTZMAN and l SUGlYAMA (With 10 Figures) ..................... 179

10 The Monophyletic Plectomycetes: Ascosphaeriales, Onygenales, Eurotiales D.M. GEISER and K.F. LoBuGLIO (With 19 Figures) .................... 201

11 Pyrenomycetes - Fungi with Perithecia G.I SAMUELS and M. BLACKWELL (With 9 Figures) ..................... 221

12 Discomycetes D.H. PFISTER and IW. KIMBROUGH (With 3 Figures) 257

13 Loculoascomycetes M.E. BARR and S.M. HUHNDORF (With 63 Figures) 283

14 The Taxonomy of Anamorphic Fungi K.A. SEIFERT and W. GAMS (With 15 Figures) 307

Subject Index ....................................................... 349

Biosystematic Index .................................................. 353

Contents Part B

The Fungal Hierarchy

1 Basidiomycetous Yeasts J.w. FELL, T. BOEKHOUT, A. FONSECA, and J.P. SAMPAIO (With 24 Figures)

2 Urediniomycetes E.C. SWANN, E.M. FRIEDERS, and D.J. McLAUGHLIN (With 20 Figures)

3 Ustilaginomycetes R BAUER, D. BEGEROW, F. OBERWINKLER, M. PIEPENBRING, and M.L. BERBEE (With 34 Figures)

4 Heterobasidiomycetes K. WELLS and RJ. BANDONI (With 16 Figures)

5 Homobasidiomycetes D.S. HIBBEIT and RG. THORN (With 14 Figures)

Nomenclature and Documentation

6 The N aming of Fungi D.L. HAWKSWORTH (With 4 Figures)

7 Cultivation and Preservation of Fungi in Culture S.-c. JONG and J.M. BIRMINGHAM

8 Computer-Assisted Taxonomy and Documentation o. PETRINI and T.N. SIEBER (With 2 Figures)

Evolution and Speciation

9 Speciation Phenomena P.A. LEMKE (With 1 Figure)

10 Fungal Molecular Evolution: Gene Trees and Geologic Time M.L. BERBEE and J.w. TAYLOR (With 5 Figures)

Subject Index

Biosystematic Index

List of Contributors

BARR, D.lS., Centre for Land and Biological Resources Research, Research Branch, Agriculture Canada, Ottawa, Ontario, Canada K1A OC6. (present address: 18 St. Osmund Close, Yetminster, Sherborne, Dorset DT9 6LU, UK)

BARR, M.E., 9475 Inverness Road, Sidney, British Columbia, Canada V8L 5G8

BENNY, G.L., Department of Plant Pathology, 1453 Fifield HaU, University of Florida, P.O. Box 110680, Gainesville, Florida 32611-0680, USA

BLACKWELL, M., Department of Biological Sciences, Louisiana State University, Baton Rouge, Louisiana 70803, USA

BRASELTON, lP., Department of Environmental and Plant Biology, Ohio University, Athens, Ohio 45701-2979, USA

CAVALIER-SMITH, T., Department of Zoology, University of Oxford, South Parks Road, Oxford, OX1 3PS, UK

DICK, M.W., Centre for Plant Diversity and Systematics, Department of Botany, School of Plant Sciences, University of Reading, Reading RG6 6AU, UK

FULLER, M.S., Darling Marine Center, University of Maine, Walpole, Maine 04573, USA

GAMS, W., Centraalbureau voor Schimmelcultures, P.0. Box 273, 3740 AG Baarn, The Netherlands

GEISER, D.M., Department of Plant Pathology, 204 Buckhout Laboratory, The Pennsylvania State University, University Park, Pennsylvania 16802, USA

HUHNDoRF, S.M., Department of Botany, The Field Museum, Roosevelt Road at Lake Shore Drive, Chicago, Illinois 60605-2496, USA

HUMBER, R.A., USDA-ARS Plant Protection Research Unit, US Plant, Soil, and Nutrition Laboratory, Tower Road, Ithaca, New York 14853-2901, USA

KIMBROUGH, lW., Plant Pathology Department, P.0. Box 110680, University of Florida, Gainesville, Florida 32611, USA

KURTZMAN, c.P., Microbial Properties Research Unit, USDA/ARS, National Center for Agricultural Utilization Research, 1815 N. University St., Peoria, Illinois 61604-3999. USA

xx List of Contributors

LoBUGLIO, K.F., Department of Plant and Microbial Biology, 111 Koshland HaU, University of California, Berkeley, California 94720, USA

MORTON, lE., Department of Plant & Soil Sciences, West Virginia University, 401 Brooks HaU, P.o. Box 6057, Morgantown, West Virginia 26506-6057, USA

PFISTER, D.H., Department of Organismic and Evolutionary Biology, Harvard University, 22 Divinity Ave., Cambridge, Massachusetts 02138, USA

SAMUELS, G.I, United States Department of Agriculture, Agricultural Research Service, Systematic Botany and Mycology Laboratory, Rm. 304, B-011A, BARC-W, BeltsviUe, Maryland 20705, USA

SEIFERT, K.A., Eastern CereaI and Oilseed Research Centre, Agriculture and Agri-Food Canada, Research Branch, Ottawa, Ontario KIA OC6, Canada

SUGlYAMA,I, Department of Botany, The University Museum, The University of Tokyo, 3-1, Hongo 7 chome, Bunkyko-ku, Tokyo 113-0033, Japan