TREES AND FORESTS A Colour Guide Biology, Pathology, Propagation, Silviculture, Surgery, Biomes, Ecology, Conservation Edited by BRYAN G BOWES Formerly Senior Lecturer at the Department of Botany University of Glasgow, UK MANSON PUBLISHING General note All figures are supplied by the author(s) of each chapter unless otherwise acknowledged. Copyright © 2010 Manson Publishing Ltd ISBN 978-1-84076-085-9 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means without the written permission of the copyright holder or in accordance with the provisions of the Copyright Act 1956 (as amended), or under the terms of any licence permitting limited copying issued by the Copyright Licensing Agency, 33–34 Alfred Place, London WC1E7DP. Any person who does any unauthorized act in relation to this publication may be liable to criminal prosecution and civil claims for damages. 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For full details of all Manson Publishing titles, please write to: Manson Publishing Ltd 73 Corringham Road London NW117DL, UK Tel: +44 (0)20 8905 5150 Fax: +44 (0)20 8201 9233 Website: www.mansonpublishing.com Commissioning editor: Jill Northcott Project manager: Paul Bennett Copy editor: Janet Tosh Layout: DiacriTech, Chennai, India Colour reproduction: Tenon & Polert Colour Scanning Ltd, Hong Kong Printed by: Grafos SA, Barcelona, Spain Contents Preface 5 Temperate evergreen forests of Contributors 6 South Africa 77 Abbreviations 7 Temperate evergreen forests of northern Dedications 8 and central Chile 80 Temperate mixed evergreen forests of the SECTION1 INTRODUCTIONTOWOODYHABITAT Mediterranean 82 Chapter 1 Survey of trees: their global significance, architecture, and early Chapter 5Tropical and sub-tropical evolution 9 rain and dry forests 87 Bryan G Bowes Ghillean T Prance Introduction 9 Introduction 87 Types of trees 11 The forests of tropical America 90 Importance of trees for civilization 17 The forests of southeast Asia and Australia 97 Tree architecture 24 The forests of Africa 99 The evolution of land plants and trees 32 Old world dry semi-deciduous forests 101 Montane forests 101 SECTION2 WORLDDISTRIBUTIONOFFORESTS Secondary forests 102 Chapter 2 Northern boreal and montane coniferous forests 40 SECTION3 TREEMORPHOLOGY,ANATOMY, Aljos Farjon ANDHISTOLOGY Introduction 40 Chapter 6 Woody thickening in trees Conifers of the boreal forest belt 41 and shrubs 104 Montane conifers 44 Bryan G Bowes Coastal conifers 46 Introduction 104 Concluding remarks 50 Vascular activity in broadleaved and coniferous trees 107 Chapter 3 Temperate deciduous and Structure and function of bark 120 temperate rain forests 51 Hugh Angus SECTION4 TREEPATHOLOGY Temperate deciduous forests (TDFs) 51 Chapter 7 The role of cell-wall polymers Temperate rain forests (TRFs) 61 in disease resistance in woody plants 126 Christopher T Brett Chapter 4 Temperate mixed evergreen Introduction 126 forests 68 The molecular structure of cell-wall Stephen D Hopper, Erika Pignatti Wikus, polymers 128 and Sandro Pignatti The cell wall as a physical barrier to Introduction 68 infection 133 Temperate evergreen forests of Australia 70 Active defence mechanisms involving Temperate evergreen forests of Tasmania the cell wall 135 and New Zealand 76 Future developments 137 4 Chapter 8 Microbial and viral pathogens, Classification of systems 212 and plant parasites of plantation and Even-aged systems 212 forest trees 139 Uneven-aged systems 218 Stephen Woodward Silvicultural systems in different parts Introduction 139 of the tropics 220 Tree root diseases 140 Concluding remarks 224 Diseases of stems 149 Diseases of foliage 155 Chapter 12 Tree pruning and surgery in Decay 160 arboriculture 225 Rust diseases 165 David Thorman Wilt diseases 169 Introduction 225 Virus diseases 173 Rationale and problems of tree surgery 229 Parasitic plants 174 Tree pruning techniques 230 Acknowledgements 177 Phenology 234 Pollarding 234 Chapter 9 Insect pests of some important Artificial support systems 235 forest trees 178 Tree felling 236 Claire Ozanne Concluding remarks 236 Introduction 178 Sap feeders 179 Chapter 13 Tree propagation for forestry Bark and stem borers 183 and arboriculture 238 Shoot borers 188 Brent McCown and Thomas Beuchel Bark chewers 191 Introduction 238 Defoliators 192 Overview of concepts 238 Termites 195 Propagation by seed 239 Concluding remarks 197 Vegetative propagation 242 Handling of young propagules 248 SECTION5 FORESTECOLOGYAND Ethics in tree propagation 251 MANAGEMENT,TREESURGERY, PROPAGATION,ANDCONSERVATION Chapter 14 Forest and woodland Chapter 10 General forest ecological conservation 252 processes 198 Ghillean T Prance Peter A Thomas Introduction 252 Introduction 198 Reserve size 253 Size and growth 198 Population size 255 Light 199 Conservation genetics 257 Water 203 The human component 257 Nutrients 203 Maintaining biological interactions 261 Coarse woody debris 206 Reserve management 262 Evergreen and deciduous leaves 207 Education and capacity building 262 Concluding remarks 209 Concluding remarks 262 Chapter 11 Silvicultural systems 210 References and general reading 264 Peter Savill and Nick Brown Introduction 210 Index 279 Ecological principles in forest regeneration 210 5 Preface A grove of sequoias [Sequoiodendron Buddha is believed to have attained enlightenment giganteum]–occupy an area of perhaps less while sitting under the bodhi tree (Ficus religiosa), than a hundred acres–The perfect specimens while ginkgo (Ginkgo biloba) is almost extinct in the not burned or broken are singularly regular and wild of China but has survived as a sacred tree in symmetrical–showing infinite variety in general temple gardens. unity and harmony; the noble shafts with rich Sadly, for the last century or more, such respect purplish brown fluted bark, free of limbs for one for trees has generally been forsaken. Today, hundred and fifty feet or so, ornamented here environmentally detrimental logging and forest and there with leafy rosettes; main branches of clearances still continue, either illegally or by official the oldest trees very large, crooked and rugged, connivance, driven by world population growth, zigzagging stiffly outward seemingly lawless, yet industrialization, market globalization, and the unexpectedly stooping at the right distance from pursuit of maximal (but patently unsustainable) the trunk and dissolving in dense bossy masses commercial profits. of branchlets, thus making a regular though The principal aim of this present volume is to greatly varied outline,–a cylinder of leafy, bring together and review a number of features of outbulging spray masses, terminating in a noble forests, trees, their development, natural environ- dome, that may be recognised while yet far off ments, and ecology, which are usually disparately upheaved against the sky–the king of all considered. These topics are expertly treated by conifers, not only in size but in sublime majesty scientists from various countries and presented in a of behaviour and port. format in which, as an integral part of each chapter, (John Muir, My First Summer in the numerous high-quality colour illustrations comple - Sierra,September 17th1869) ment the concise but clearly written text. This Guide will be of importance and interest to A reverence and respect for trees, such as are anyone studying plant science, forestry, or ecology exhibited in Muir’s scientific but beautiful prose and of practical and theoretical use to conserva- describing his experiences in the Californian Sierra tionists, foresters, tree propagators, and tree surgeons Nevada is deeply embedded in the human psyche. A working in the field. The 16 contributing authors, sacred oak (Quercus) was at the centre of the Greek based in Britain, the USA, Australia, and Italy, reflect oracle at Dodona founded in about 1800 BP; while a wealth of expertise, which will help ensure that this Artemis, the goddess of woodlands, and Apollo (her volumeis of value and use for students and interested twin brother) were said to be born under a date amateurs in countries throughout the world. palm. For the Pehuenche Amerindians of Chile, the monkey puzzle tree (Araucaria araucana) is holy, Bryan G Bowes 6 Contributors Chapters 1 and 6 Chapter 8 BRYANG BOWES STEPHENWOODWARD Formerly at the Department of Botany, School of Biological Sciences, University of Glasgow, Glasgow, UK University of Aberdeen, Aberdeen, UK Chapter 2 Chapter 9 ALJOSFARJON CLAIREOZANNE Royal Botanic Gardens Kew, Richmond, School of Human and Life Sciences, Surrey, UK Roehampton University, London, UK Chapter 3 Chapter 10 HUGHANGUS PETERA THOMAS Head of Tree Collections, Westonbirt Arboretum, School of Life Sciences, Keele University, Keele, UK near Tetbury, Gloucestershire, UK Chapter 11 Chapter 4 PETERSAVILL STEPHEND HOPPER Oxford Forestry Institute, Department of Plant Director, Royal Botanic Gardens Kew, Richmond, Sciences, University of Oxford, Oxford, UK Surrey, UK NICKBROWN ERIKAPIGNATTIWIKUS Oxford Forestry Institute, Department of Plant Dipartimento di Biologia,Universita di Trieste, Sciences, University of Oxford, Oxford, UK Trieste, Italy Chapter 12 SANDROPIGNATTI DAVIDTHORMAN Dipartimento di Biologia Vegetale, Rostrevor, Station Road, Whittington, Oswestry, Universita di Roma ‘La Sapienza’, Roma, Italy Shropshire, UK Chapters 5 and 14 Chapter 13 GHILLEANT PRANCE BRENTH MCCOWN School of Plant Sciences, University of Reading, Professor, Department of Horticulture, Whiteknights, Reading, UK University of Wisconsin-Madison, Madison, Wisconsin, USA Chapter 7 CHRISTOPHERT BRETT THOMASBEUCHEL Institute of Biomedical and Life Sciences, McKay Nursery, Waterloo, Wisconsin, USA University of Glasgow, Glasgow, UK 7 Abbreviations BP before present PGA polygalacturonic acid B.t. Bacillus thuringiensis PME pectin methyl esterase B.t.k Bacillus thuringiensisvar. kurstaki PRPs pathogenesis-related proteins CWD coarse woody debris REGUA Reserva Ecologica Guapi-açú RG-I rhamnogalacturonan-I DED Dutch elm disease RG-II rhamnogalacturonan-II DON dissolved organic nitrogen SMS selective management system E east W west HRGPs hydroxyproline-rich glycoproteins TDF temperate deciduous forest IBA indolebutyric acid TEF temperate evergreen forest IUCN International Union for Conservation of TSS tropical shelterwood system Nature IPM integrated pest management Note: all sectioned specimens were photographed under the light microscope except where MUS Malayan Uniform System designated TEM (transmission electron MVP minimum viable population micrograph). Abbreviations:LS, longitudinal mya million years ago section; RLS, radial longitudinal section; TS, transverse section; TLS, tangential NEP net ecosystem productivity longitudinal section. NPP net primary productivity 8 Dedications To the Editor’s family With great love to my dear, supportive, and forbearing wife Diane; in warm memory of my late wife Ruth; with all love and affection for our children Tanya and Adrian; and to my six bright, creative, and affectionate grandchildren–Sean, Aidan, Declan, Cian, Myles, and Marcus. To William Shakespeare The quality of mercy is not strain’d; It droppeth as the gentle rain from heaven Upon the place beneath. It is twice blest: It blesseth him that gives and him that takes. ’Tis mightiest in the mightiest; it becomes The throned monarch better than his crown; His sceptre shows the force of temporal power, The attribute to awe and majesty, Wherein doth sit the dread and fear of kings; But mercy is above this sceptred sway, It is enthroned in the hearts of kings, It is an attribute to God himself (The Merchant of Venice) 9 S 1 I ECTION NTRODUCTION TO WOODY HABITAT CHAPTER 1 Survey of trees: their global significance, architecture, and early evolution Bryan G Bowes INTRODUCTION role of trees in the complex web of life. Most trees – apart from those growing in According to Jane (1970), there are some 20,000 plantations, horticulture, arboreta, parks, or urban species of woody plants but, more recently, Oldfield settings–are distributed in various biomes, or major et al.(1998) estimated that there are possibly up to regions, of the terrestrial world (1; Packham et al., 100,000 tree (arborescent) species: these were 1992). Each biome is characterized by its own range defined as single-stemmed species growing to at least of animal and plant life. Chapters 2–5 of this Guide 2 m in height at maturity. However, this definition consider the various natural regions populated by seems unduly restrictive, since it would apparently trees, while Chapters 10 and 14 emphasize the vital exclude some natural multi-stemmed species of 1 1Overview of the Blue Mountains Eucalyptus forest near Sydney, Australia. This forest is dominated by eucalypts, with some 90 arborescent species of Eucalyptusand other members of the family Myrtaceae occurring here.