Diseases and Pathogens of EUCALYPTS Dedication This book is dedicated to the memory of Dr Geoffrey Charles Marks (1932–90) who, after 27 years of experience as a Forest Pathologist with the Victorian Government and as a teacher of Forest Pathology at the University of Melbourne, saw the need for such a book. He initiated its writing and undertook the initial planning of its structure and authorship. Geoff was born in Sri Lanka (formerly Ceylon) and was an outstanding athlete and scholar, representing his country as a swimmer at the Helsinki Olympics and completing an honours degree in science at the University of Ceylon, followed by an MSc (1961) and PhD (1963) while on a Rockefeller Foundation Scholarship at the University of Wisconsin. He took up his appointment with the then Forests Commission of Victoria in Melbourne in 1963 and thereafter was involved in the study of eucalypt diseases, being a central participant in the initial studies of dieback caused by Phytophthora cinnamomi in Victorian eucalypt forests. He also initiated early studies on the eucalypt leaf pathogen, Mycosphaerella, in southern Australia. His keen intellect, enlightened attitudes, great enthusiasm and curiosity, waspish good humour and propensity to provoke critical discussion inspired colleagues and led to fruitful collaborations with many people. He developed a deep appreciation of the eucalypt forests of his adopted country, and from this grew his keen understanding of their ecology and pathology. He conducted wide ranging field studies on many aspects of forest pathology and greatly enjoyed field trips to investigate disease problems in the diverse forests of Victoria, tirelessly speculating about the complex causes of the diseases and planning experimental studies with whomever was fortunate enough to accompany him. While returning home from one of these long trips to the forests of eastern Victoria he was seriously injured in a car accident. He showed great courage in recovery, and never lost his enthusiasm, expressed at the time by his delight in initiating and planning this book. He was not destined to see this work to fruition as he died suddenly in August 1990. Vale, Geoff. We hope this book has not fallen too far short of your vision. Title Diseases and Pathogens of E U CA LY P T S P. J. Keane, G. A. Kile, F. D. Podger and B. N. Brown (Editors) © CSIRO 2000 All rights reserved. Except under the conditions described in the Australian Copyright Act 1968 and subsequent amendments, no part of this publication may be reproduced, stored in a retrieval system or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, duplicating or otherwise, without the prior permission of the copyright owner. Contact CSIRO PUBLISHING for all permission requests. National Library of Australia Cataloguing-in-Publication entry Diseases and pathogens of eucalypts. Bibliography. Includes index. ISBN 0 643 06523 7 (paperback). ISBN 0 643 09012 6 (eBook). 1. Eucalyptus – Diseases and pests – Australia. I. Keane, Philip J. 634.9737660994 This book is available from: CSIRO PUBLISHING PO Box 1139 (150 Oxford Street) Collingwood VIC 3066 Australia Tel: (03) 9662 7666Int: +(613) 9662 7666 Fax: (03) 9662 7555Int: +(613) 9662 7555 Email: [email protected] http://www.publish.csiro.au Typeset by Desktop Concepts P/L, Melbourne Printed in Australia by Brown Prior Anderson Cover photographs Upper: Crown decline typical of that observed in trees of Eucalyptus obliqua and E. regnans affected by regrowth dieback and Armillaria. Lower: Crinkle leaf caused by Mycosphaerella cryptica on tip leaves of a seedling of Eucalyptus obliqua. Publication of this book was supported by a grant from the Standing Committee on Forestry of the Australian Ministerial Council on Forestry, Fisheries and Aquaculture. C O N T E N T S Preface xiii Contributors xvi Section I: The Eucalypts—Their Importance, Diversity and Biology 1 Economic and Social Importance of Eucalypts 1 J.W. Turnbull Summary 1 1.1 Introduction 2 1.2 Eucalypts in native forests 2 1.3 Eucalypts as exotics 3 1.4 Industrial eucalypt plantations 4 1.5 Eucalypts in the rural landscape 5 1.6 Conclusion 7 1.7 References 7 2 Morphology, Phylogeny, Origin, Distribution and Genetic Diversity of the Eucalypts 11 B.M. Potts and L.A. Pederick Summary 11 2.1 Introduction 12 2.2 Morphology 12 2.3 Phylogeny 14 2.4 Origins 16 2.5 Distribution 18 2.6 Hybridisation 21 2.7 Genetic variation 23 2.8 Genetic variation in susceptibility to disease 24 2.9 Factors affecting disease risk in plantations 26 2.10 Acknowledgments 27 2.11 References 27 v D I S E A S E S A N D P A T H O G E N S O F E U C A L Y P T S 3 Growth Habits and Silviculture of Eucalypts 35 R.G. Florence Summary 35 3.1 Introduction 36 3.2 Growth habits of the eucalypts 36 3.3 Sivicultural practice in natural forests 38 3.4 Ecologically sustainable silviculture 40 3.5 New directions in silviculture 42 3.6 Eucalypt plantations 43 3.7 Conclusion 44 3.8 References 44 4 Ecology of Eucalypt Regeneration 47 D.H. Ashton Summary 47 4.1 Introduction 48 4.2 Characteristics of eucalypts in relation to their regeneration 50 4.3 Conditions required for regeneration 51 4.4 Major modes of regeneration 51 4.5 In synthesis—the strategies of survival 58 4.6 References 59 5 Physiology of Eucalypts in Relation to Disease 61 C.L. Beadle Summary 61 5.1 Introduction 62 5.2 Productivity and light interception 62 5.3 Biomass production, partitioning of dry mass and foliage 63 5.4 Gas exchange and stomatal conductance 64 5.5 Water relations 66 5.6 Conclusion 68 5.7 References 68 6 Mycorrhizas of Eucalypts 71 G.A. Chilvers Summary 71 6.1 Introduction 72 6.2 Structure of eucalypt mycorrhizas 73 6.3 Functioning of mycorrhizas 74 6.4 The fungal partners of eucalypts 83 6.5 Mycorrhizal infection cycles 86 6.6 Ecology of mycorrhizas 87 6.7 Manipulating eucalypt mycorrhizas 90 6.8 References 94 vi C O N T E N T S Section II: The Diseases of Eucalypts—Their Causes and Biology 7 Diseases and Fungi of the Reproductive Structures of Eucalypts 103 B.N. Brown Summary 103 7.1 Introduction 104 7.2 Fungi of flowers and capsules and their pathogenicity 104 7.3 Seed fungi of eucalypts 106 7.4 Control of seed fungi 114 7.5 Conclusion 115 7.6 References 115 8 Disease during Propagation of Eucalypts 119 B.N. Brown and F.A. Ferreira Summary 119 8.1 Introduction 120 8.2 Diseases with abiotic causes 120 8.3 Fungal diseases 121 8.4 Principal diseases of eucalypt cuttings 143 8.5 Conclusion 143 8.6 References 143 9 Fungal Diseases of Eucalypt Foliage 153 R.F. Park, P.J. Keane, M.J. Wingfield and P.W. Crous Summary 153 9.1 Introduction 154 9.2 Target spot (Aulographina eucalypti) 155 9.3 Leaf spot, leaf blotch and crinkle leaf blight (Mycosphaerella species) 163 9.4 Biotrophic infections 175 9.5 Powdery mildews (Oidium species) 191 9.6 Eucalypt rust (Puccinia psidii) 191 9.7 Angular, vein-limited leaf spots 193 9.8 White leaf and shoot blight (Sporothrix pitereka ) 206 9.9 Winter leaf spot (Piggotia substellata and Ceuthospora innumera) 206 9.10 Leaf spots and speckles of minor importance 210 9.11 Leaf spots and blights of stressed plants 218 9.12 Conclusion 227 9.13 Acknowledgments 230 9.14 References 230 10 Canker Diseases of Eucalypts 241 K.M. Old and E.M. Davison Summary 241 10.1 Introduction 242 10.2 Fungal invasion and host responses 242 10.3 Effect of plant stress on the development of cankers 244 10.4 Major canker diseases of eucalypts 245 vii D I S E A S E S A N D P A T H O G E N S O F E U C A L Y P T S 10.5 Opportunistic pathogens associated with cankers in eucalypts 248 10.6 Conclusion 252 10.7 Acknowledgments 252 10.8 References 253 11 Diseases of Eucalypts Caused by Soilborne Species of Phytophthora and Pythium 259 B.L. Shearer and I.W. Smith Summary 259 11.1 Introduction 260 11.2 Origin of the pathogens 261 11.3 Host range 261 11.4 Distribution patterns and effect of disease 262 11.5 Host–pathogen interactions 264 11.6 Pathogenicity of Pythiaceae on eucalypts 267 11.7 Effects of disease on wood production and conservation values 269 11.8 Pathogen dynamics 272 11.9 Resistance mechanisms 280 11.10 Effects of environment on disease development in established infections 282 11.11 Conclusion 284 11.12 Acknowledgment 284 11.13 References 284 12 Woody Root Rots of Eucalypts 293 G.A. Kile Summary 293 12.1 Introduction 294 12.2 Armillaria root disease 294 12.3 Pseudophaeolus root disease 300 12.4 Ganoderma root rot 301 12.5 Other woody root diseases 301 12.6 Conclusion 302 12.7 References 303 13 Stem and Butt Rot of Eucalypts 307 G.A. Kile and G.C. Johnson Summary 307 13.1 Introduction 308 13.2 Causal organisms and hosts 308 13.3 The decay process 323 13.4 Factors affecting decay development 327 13.5 Particular heart rots and stem conditions 331 13.6 Conclusion 332 13.7 Acknowledgments 333 13.8 References 333 viii C O N T E N T S 14 Diseases of Eucalypts Associated with Viruses, Phytoplasmas, Bacteria and Nematodes 339 T.J. Wardlaw, G.A. Kile and J.C. Dianese Summary 339 14.1 Introduction 340 14.2 Virus-like diseases 340 14.3 Diseases associated with phytoplasmas 341 14.4 Diseases caused by bacteria 343 14.5 Diseases associated with nematodes 346 14.6 Conclusion 348 14.7 References 349 15 Mistletoes and other Phanerogams Parasitic on Eucalypts 353 N. Reid and Z. Yan Summary 353 15.1 Introduction 354 15.2 Mistletoes parasitic on eucalypts 354 15.3 Native cherries parasitic on eucalypts 375 15.4 Dodder-laurels parasitic on eucalypts 377 15.5 Acknowledgments 378 15.6 References 378 16 Nutritional Disorders and other Abiotic Stresses of Eucalypts 385 P. Snowdon Summary 385 16.1 Introduction 386 16.2 Diagnosis 386 16.3 Nutrient deficiencies 386 16.4 Toxicities 395 16.5 Water as an abiotic factor 398 16.6 Frost 400 16.7 Artificial environments 401 16.8 Miscellaneous abnormalities 402 16.9 Conclusion 402 16.10 References 403 17 Eucalypt Diseases of Complex Etiology 411 K.M. Old Summary 411 17.1 Introduction 412 17.2 Etiology of diebacks and declines 412 17.3 Diebacks of native forests and woodlands 414 17.4 Forest diebacks associated with drought 414 17.5 Forest diebacks associated with successional changes 416 17.6 Forest and woodland diebacks associated with chronic insect herbivory 416 17.7 Plantation diseases of complex etiology 421 ix