Contents

Preface page xi

Acknowledgements xiii

Metric equivalents xiv

1 Introduction: Forest basics 1

1.1 Characteristics of woodlands and forests 1

1.2 The value of woodlands and forests 5

1.3 Tree biology and how it influences woodland ecology 6

1.4 Spatial structure 13

1.5 The woodland ecosystem: food chains, food webs and the plant, animal and decomposition subsystems 19

1.6 Forest types and classification 25

1.7 Regional classifications of forests and woodlands 35

2 Forest soils, climate and zonation 39

2.1 Soils and trees 39

2.2 Features of forest soils 41

2.3 Roots, foraging and competition 58

2.4 Forest zonation and site quality 66

2.5 Rain forests: climate, soils and variation 76

3 Primary production and forest development 84

3.1 Plant life forms and biological spectra 84

3.2 Light and shade 91

3.3 Water 110

3.4 Temperature and pollutant influences on tree growth 115

3.5 Altitudinal zonation and timberlines 117

3.6 Evergreen and deciduous strategies: aspects of competitive advantage 127

3.7 Contrasts between three widespread tree genera: the pines, beeches and oaks 131

3.8 Ecology and significance of ageing trees 139

4 Reproductive strategies of forest plants 144

4.1 Plant strategies 144

4.2 Regenerative strategies and vegetative spread 152

4.3 Reproduction and fruiting 162

4.4 Masting 165

4.5 Roles and influences of animals 178

4.6 Time constraints 181

5 Biotic interactions 187

5.1 Producers and consumers 187

5.2 The interdependence of producers and consumers 188

5.3 Insect defoliation and damage 191

5.4 Forest fungi 201

5.5 Specialized heterotrophs: epiphytes, parasites and saprotrophs 214

5.6 Exotic plants 218

5.7 Herbivorous mammals and birds 219

5.8 The impact ofwoodland carnivores and omnivores 230

5.9 Herbivores and the Holocene: did the lowland

European forest have a closed canopy? 235

6 Biodiversity in woodlands 241

6.1 Genetic variation in populations and its implications 241

6.2 Selection pressures and biodiversity 242

6.3 Biodiversity at organism, population and habitat levels 247

6.4 Changes in species diversity over time 257

6.5 What allows species to co-exist in a woodland? 266

6.6 Conservation, biodiversity, population integrity and uniqueness 273

7 Decomposition and renewal 276

7.1 The vital key to a working forest 276

7.2 Decomposition 277

7.3 Degradative stages 285

7.4 How much dead material is there? 288

7.5 What controls the rate of decomposition? 291

7.6 Rates of decomposition 298

7.7 Woody material 302

8 Energy and nutrients 318

8.1 Growth of forests 318

8.2 Energy flow through forest ecosystems 326

8.3 Nutrient cycling 328

8.4 Nitrogen 331

8.5 Nutrient dynamics in different forests 341

8.6 Human influences 345 9 Forest change and disturbance 350

9.1 Ecology of past forests 350

9.2 Ecological processes that govern change 366

9.3 Disturbance, patch dynamics and scales of change 374

9.4 Examples of forest change 387

9.5 Stability and diversity 395

10 Working forests 397

10.1 Forest resources and products 397

10.2 Single-and multi-use forests 409

10.3 Silviculture and the replacement of trees 410

10.4 Improving the forest: choice of species and provenance 420

10.5 Forest practices 424

10.6 Sustainable forest management 427

10.7 Landscape ecology and forests 429

11 The future - how will our forests change? 441

11.1 Threats to forests and the increasing demand for timber 441

11.2 Desertification 445

11.3 Climate change 447

11.4 Other causes of forest decline 461

11.5 Problems in urban forests - the social interface 471

11.6 Agroforestry and new forests 474

11.7 The final challenge 481 References 483 Index 514

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