FOREST ECOLOGYFourth Edition

BURTON V. BARNESUniversity of Michigan

DONALD R. ZAKUniversity of Michigan

SHIRLEY R. DENTONBiological Research Associates

Tampa, Florida

STEPHEN H. SPURRUniversity of Texas at Austin

John Wiley & Sons, Inc.NEW YORK • CHICHESTER • WEINHEIM • BRISBANE • SINGAPORE • TORONTO

CONTENTS

CHAPTER 1 CONCEPTS OF FOREST ECOLOGY 1

Ecology 2Landscape Ecosystems 3Landscape Ecosystem and Community 6Ecosystem Structure 7Ecosystem Function 8Vertical and Horizontal Approaches 8Examples of Landscape Ecosystems 10An Approach to the Study of Forest Ecology 14Applicability in Ecosystem Management 16Suggested Readings 17

PART 1 Ecosystems at Multiple Spatial and Temporal Scales 19

CHAPTER 2 LANDSCAPE ECOSYSTEMS AT MULTIPLESPATIAL SCALES 21

Overview of Spatial and Temporal Scales 21Hierarchical Ecosystem Scales in Space 24Climatic Classification 24Vegetation Types and Biomes 27Physiography 30Distinguishing and Mapping Landscape Ecosystems at Multiple

Spatial Scales 31Regional Landscape Ecosystems 34

Regional Landscape Ecosystems of Michigan 35Local Landscape Ecosystems 38

Local Landscape Ecosystems in Upper Michigan 38Suggested Readings 40

CHAPTER 3 LONG-TERM ECOSYSTEM ANDVEGETATION CHANGE 41

Change Before the Pleistocene Age 42Pleistocene Glaciations 43

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Ecosystem and Vegetational Change Since the Last GlacialMaximum 44

Pollen Analysis 46Eastern North America 46

Overall Migration Sequence and Patterns 48Ecosystem Change in the Southern Appalachians 51

Western North America 51Patterns of Tree Genera and Species Migrations 53Migration Irregularities and Disturbance 55Long-Term Change at a Given Site 56Independent Migration and Similarity of Communities Through

Time 57Adaptations of Conifers and Angiosperms 58Suggested Readings 60

PART 2 The Forest Tree 61

CHAPTER 4 FOREST TREE VARIATION 63

Components of Phenotypic Variation 64Plasticity of the Phenotype 65Sources of Variation 66The Evolutionary Sequence 67Sexual and Asexual Systems 68Genetic Diversity of Woody Species 68Genecology 69 -Patterns of Genecological Differentiation 70

Genecological Categories 71Factors Eliciting Genecological Differentiation 71

Growth Cessation 72Growth Resumption 75

Examples of Genecological Differentiation 76Eastern North American Species 76Scots Pine 78Wide-Ranging Western North American Conifers 78

Ponderosa Pine 79Douglas-fir 81

Local Genecological Differentiation 82 -Factors Affecting Differentiation: Gene Flow and Selection Pressure 83Ecological Considerations at the Species Level 85Niche 87Hybridization 87Polyploidy 90The Fitness-Flexibility Compromise 91Suggested Readings 92

CHAPTER 5 REGENERATION ECOLOGY 94

Regeneration 95Sexual Reproduction 98

Contents IX

Maturation and the Ability to Flower 98Increasing Seed Production 99

Reproductive Cycles 99Pollination 100Periodicity of Seed Crops 102Effects of Reproduction on Vegetative Growth 104Dispersal 105Seed Bank, Dormancy, and Germination 107Establishment Following Sexual Reproduction 108Post-Establishment Development 115Vegetative Reproduction 115Suggested Readings 121

CHAPTER 6 STRUCTURE AND GROWTH 122

Tree Form 122Architectural Models 124

Short and Long Shoots 126Patterns of Intermittent Growth 127Sylleptic and Proleptic Shoots 130

Roots 131Kinds, Forms, and Occurrence 132Fine Root Relations 132Horizontal and Vertical Root Development 135Periodicity of Primary Root Growth 136Root Grafting 137Specialized Roots and Buttresses 137Stems 138Xylem Cells and Growth Rings 140Periodicity and Control of Secondary Growth 141

Control of Early wood and Latewood Formation 143Winter Freezing and Water Transport 144Water Deficits and Tree Growth 145Suggested Readings 148

PART 3 The Physical Environment 149

CHAPTER 7 CLIMATE 153

Climatic Control of Vegetation Distribution 154Radiation 154Air Circulation 156Air Masses 158Water 159Continents and Physiography 163Describing Climate 165Temperature 165Precipitation 167

Fog and Dew 168Snow 170

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Evapotranspiration 170Wind 171Extreme Events 171Storms 172Drought 173Glaze 174Extreme Cold 174Microclimate 175Classifying Climate 175Climatic Change 178Suggested Readings 181

CHAPTER 8 LIGHT 182

Distribution of Light Reaching the Ecosphere 183Interception of Radiation 184Canopy Structure and Leaf Area 186Sun Flecks 189Light Quality Beneath the Forest Canopy 191Light and Growth of Trees 191Light and Tree Morphology and Anatomy 198Leaf Structure and Stratification in Tropical Forests 199Adventitious Buds and Epicormic Sprouting 203Photocontrol of Plant Response 203Light and Ecosystem Change 204Suggested Readings 205

CHAPTER 9 TEMPERATURE 206

Temperatures at the Soil Surface 206Temperatures Within the Forest 208Temperature Variations with Topographic Position 208Temperature and Plant Growth 209Cold Injury to Plants 213Dormancy 214Frost Hardiness and Cold Resistance 215

Thermotrophic Movements in Rhododendrons 218Winter Chilling and Growth Resumption 220Natural Plant Distributions and Cold Hardiness 221Deciduousness and Temperature 222Suggested Readings 223

CHAPTER 10 PHYSIOGRAPHY 224

Concepts and Terms 225Characteristics of Physiography and their Significance 227Physiographic Setting 227

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Specific Landforms 228Elevation 228Form of Landforms 228

Level Terrain 228Sloping Terrain 229Slope Characteristics 229

Position on Slope 229Aspect 230Slope Inclination 230

Parent Material in Relation to Landform 232Position of Landform in the Landscape 232

Multiple Roles of Physiography 233Physiographic Diversity, Landscape Ecosystems, and Vegetation 234Mountainous Physiography 234

Mountainous Terrain of California and the Pacific Northwest 234Physiography and Forests of the Central Appalachians 238

Flatlands 243The Great Plains '243Southwestern Plateaus and Pine Forests 243Pine Savannas of the Western Great Lakes Region 244Till Plains of the Midwest 244Southeastern and Southern Coastal Plain 245Floodplains 245

Physiography and Firebreaks 248Microlandforms and Microtopography 250Tree Uprooting and Pit and Mound Microtopography 250Microtopography and Regeneration in Hardwood Swamps 253Suggested Readings 253

CHAPTER 11 SOIL 255

Parent Material 255Soil Formation 258Soil Profile Development 258Physical Properties of Soil 260Soil Texture 260Soil Structure 262Soil Color 262Soil Water 263

Physical Properties of Water 263Soil Water Potential 264

Chemical Properties of Soil 265Clay Mineralogy 268Cation Exchange and the Supply of Nutrients 270

Soil Acidity 272Soil Organic Matter 274Soil Classification 275Landform, Soil, and Forest Vegetation: Landscape Relationships 276Suggested Readings 278

Contents

CHAPTER 12 FIRE 279

Fire and the Forest Tree 280Causes 280Fire Regime 280

Fire Types, Frequency, and Severity 281Fire Adaptations and Key Characters 284Strategies of Species Persistence 288

Closed-Cone Pines 289Fire and the Forest Site 290Indirect Effects 290Direct Effect 292Detrimental and Beneficial Effects 294Suggested Readings 297

CHAPTER 13 SITE QUALITY AND ECOSYSTEMEVALUATION 298

Direct Measurement of Forest Productivity 300Tree Height as a Measure of Site 300Site-Index Curves 303Comparisons between Species 306Advantages and Limitations 306Vegetation as an Indicator of Site Quality 306Species Groups of Groundflora 308

Indicator Plants of Coastal British Columbia 309Ecological Species Groups 309

Plant Associations and Habitat Types in the Western United States 310Operational Site Classification Based on Vegetation 310

Applications and Limitations of Vegetation 314Environmental Factors as a Measure of Site 314Climatic Factors 315Physiographic Land Classification 315Physiographic and Soil Factors: Soil-Site Studies 316Soil Survey 320Multiple-factor Methods of Site and Ecosystem Classification 320Ecosystem Classification and Mapping in Baden-Wurttemberg 321Applications of Multifactor Methods in the United States and Canada 323

Ecosystem Classification and Mapping in Michigan 323Ecosystem Classification in the Southeastern United States 326Ecological Land Classification in Canada 327

Hills' Physiographic Approach 327Other Canadian Approaches 321

Suggested Readings 329

PART 4 Forest Communities 331

CHAPTER 14 ANIMALS 333

Plant Defense 334

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The Plant Dilemma: To Grow or Defend 334Insects 335

Examples of Injury and Plant Defense 336Between-Plant and Within-Plant Heterogeneity 338Nutrition 338Plant Hybrid Zones as Reservoirs for Insects 339

Mammals 340Roles of Animals in Plant Life History 341Pollination 342Seed Dispersal 342

Fish and Reptiles 343Birds 343Mammals 346

Germination and Establishment 348Decomposition, Mineral Cycling, and Soil Improvement 349Damage and Death 350

Production and Regeneration 352Wildlife Habitat and Fire 352Adaptations to Fire-Dependent Ecosystems 353Kinds and Abundance of Animals 353Factors Affecting Animal Responses to Fire 354Influence of Large Animals on Forest Ecosystems 355Livestock Grazing 356Human Component of Ecosystems 357

Human-Introduced Exotics 358Suggested Readings 360

CHAPTER 15 FOREST COMMUNITIES 361

Community Concept 361Grounding Communities 363

Florida Keys 363Interior Alaska 365Southern Illinois 366

View from the Past: Community Concepts 368Schools and Terminology 368

Concepts of Clements and Gleason 369Phytosociology in Europe 371

Continuum Concept 372Community as a Landscape Property 373Examples of Spatial Variation in Forest Communities 375Discrete Forest Communities 375

Coastal California: Giant and Pigmy Forests 375Forest-Grassland Ecotone 377Alpine Tree-Lines 377

Merging Forest Communities 378Eastern Deciduous Forest-Southern Appalachians 378New England 379

Competition and Niche Differentiation 380Interactions Among Organisms 383

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Mutualisms in Forest Ecosystems 383Symbiotic Mutualisms—Mycorrhizae 384Nonsymbiotic Mutualisms 386

Competition 386Composition, Stand Structure, and Density 387

Vertical Structure 389Stand Density 391

Competition and Overstory Composition 393Competition in the Understory 394

Understory Tolerance 395Characteristics of Understory Tolerant and Intolerant Species 396Tolerance Ratings of Tree Species 397Examples of Understory Tolerance in Forest Ecosystems 398Nature of Understory Tolerance 398

Environmental Factors Relating to Understory Tolerance 398Physiological Processes Relating to Tolerance 403

Suggested Readings 406

PART 5 Forest Ecosystem Dynamics 407

CHAPTER 16 DISTURBANCE 409

Disturbance as an Ecosystem Process 410Source of Disturbance 413Major Disturbances in Forest Ecosystems 414Catastrophic and Local Land Movement 414Fire 414

Role of Fire in Forest Ecosystems 415Pines in New England and the Lake States 417Western Pines and Trembling Aspen 417Southern Pines 421Douglas-fir in the Pacific Northwest 421Giant Sequoia All

Fire History and Behavior 423Northern Lake States 423Boreal Forest and Taiga 426Northern Rocky Mountains 426

Fire Exclusion 427Wind 428

Catastrophic and Local Effects 428Principles of Wind Damage 429Broad-Scale Disturbance by Hurricanes 430

Southern Atlantic Coast 431New England—1938 Hurricane 431

Wave-Regenerated Fir Species 431Floodwater and Ice Storm 432Insects and Disease 433Logging 433Land Clearing 436

Contents XV

Biotic Composition Changes 436Elimination of Species 437Addition of Species 439

Introductions to New Zealand 439Increased Animal Impact 440Climatic Change 441Suggested Readings 442

CHAPTER 17 FOREST SUCCESSION 443

Basic Concepts 444Primary and Secondary Succession 444Biological Legacies 445Successional Pathways, Mechanisms, and Models 445Autogenic and Allogenic Succession 445How Is Succession Determined? 446Evolution of the Concept of Forest Succession 447Formal Ecological Theory 448How Does Succession work? 448Clementsian Succession 450Stages of Succession 451

Primary Succession 451Secondary Succession 454

Successional Causes, Mechanisms, and Models 457Key Characteristics and Regeneration Strategies 457Availability and Arrival Sequence of Species 458Facilitation, Tolerance, and Inhibition 458Diagrammatic Comparison of Models 460

Change in Ecosystems 461End Point of Succession? 462Succession as a Landscape Ecosystem Process 463Biomass and Diversity 466Examples of Forest Succession 468Recently Deglaciated Terrain—A Geoecology Approach 468Succession Following Fire in Ponderosa Pine Forests of Western Montana 471Gap Dynamics 473

Gap Specialists: American Beech and Sugar Maple 477Old-Field Succession in the Eastern United States 479Fire and Oak Dominance—Oaks at Risk 483Suggested Readings 484

CHAPTER 18 CARBON BALANCE OF TREESAND ECOSYSTEMS 486

Carbon Balance of Trees 487Photosynthesis, Dark Respiration, and Leaf C Gain 487Light and Leaf C Gain 489

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Temperature and Leaf C Gain 491Water and Leaf C Gain 492Soil Nitrogen Availability and Leaf C Gain 493Construction and Maintenance Respiration 494Allocation to Structure, Storage, and Defense 496Light and C Allocation 499Soil Nitrogen Availability and C Allocation 499Carbon Balance of Ecosystems 503Biomass and Productivity of Forest Ecosystems 504Measurement of Biomass and Productivity 507Climate and Productivity 510Soil Properties, Forest Biomass, and ANPP 514Biomass Accumulation During Ecosystem Development 515Soil N Availability and Belowground Net Primary Productivity 520Suggested Readings 523

CHAPTER 19 NUTRIENT CYCLING 524

Nutrient Additions to Forest Ecosystems 526Mineral Weathering 526Atmospheric Deposition 527Biological Fixation of Nitrogen 531Nutrient Cycling within Forest Ecosystems 533Nutrient Transport to Roots 534Nutrient Uptake and Assimilation by Roots 535Root Architecture, Mycorrhizae, and Nutrient Acquisition 538

Root Architecture 538Mycorrhizae 539

Plant Litter and the Return of Nutrients to Forest Floor and Soil 540Leaf and Root Litter Production 541Nutrient Retranslocation 543

Nutrients in the Forest Floor 547Organic Matter Decomposition and Nutrient Mineralization 550

Chemical Constituents of Plant Litter 550Dynamics of Decomposition 553Nitrogen Immobilization and Mineralization 557Nitrogen Availability in Forest Ecosystems 559Nitrification 561

Nutrient Loss from Forest Ecosystems 562Nutrient Leaching from Forest Ecosystems 562Denitrification 564The Cycling and Storage of Nutrients in Forest Ecosystems 566Nutrient Storage in Boreal, Temperate, and Tropical Forests 566The Nitrogen and Calcium Cycle of a Temperate Forest Ecosystem 567Ecosystem C Balance and the Retention and Loss of Nutrients 569Forest Harvesting and Nutrient Loss 572Suggested Readings 575

Contents XVil

CHAPTER 20 DIVERSITY 577

Concepts of Biological and Ecosystem Diversity 578Species Diversity and Its Value 578Value of Biodiversity 580Measuring Diversity 582Levels of Diversity 582Measurement 584

Inventory Diversity: Alpha Diversity 584Differentiation or Beta Diversity 588

Diversity of Landscape Ecosystems 588Examples of Diversity 589

Ground-Cover Species Diversity in Northern Lower Michigan 589Ecosystem Groups 589Ecosystem Types 593

Ecosystem Diversity 595Causes of Species Diversity at Multiple Scales 596Diversity at Continental and Subcontinental Scales 597

Paleogeography and Continental Relationships 597Glaciation 600Latitude and Elevation 600

Diversity at Local Scales 602Physiography and Soil 604Community Composition and Structure 605Disturbance and Succession 606

Human Effects and Concerns for Diversity 608Endemics and Rare and Endangered Species 608Keystone Species 608Effects of Forest Management on Diversity 609Conserving Ecosystem and Biological Diversity 610Landscape Ecosystem Approach 611Suggested Readings 612

CHAPTER 21 LANDSCAPE ECOLOGY 613

Concepts of Landscape Ecology 614Old-Growth Forests of the Pacific Northwest 615Coarse Woody Debris and Ecosystem Interactions in Riparian Zones 615Erosion and Geomorphology in Riparian Systems 620Forest-Stream Linkages at Multiple Spatial Scales 621Landscape-Level Disturbance 623Catastrophic Windstorm in New England 623Vegetational Diversity Mediated by Fire in Yellowstone National Park 626Landscape Pattern and Process Using the Landscape Ecosystem

Approach 628Geomorphic Processes and Ecosystem Patterns 629Landscape Ecosystems and Endangered Spaces 630Forest Fragmentation 634

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Ecological Effects 636New Paradigm in Forestry: Sustainable Ecosystems 639Examples of Ecosystem Management 63921st-century Ecosystem Management in Pacific Northwest Forests 640

Variable-Retention Harvest System 642Lifeboating: Refugia and Inocula 642Structural Enrichment and Enhanced Connectivity 642Designing a Variable-Retention Harvest System 644

Simulating Harvests and Forest Fragmentation 645Remote Sensing 649Suggested Readings 652

PART 6 Forests of the World 653

CHAPTER 22 FORESTS OF THE WORLD 655

Evolution of Modern Tree Species 655Present-Day Forests of the World 658Geographical-Climatic Classification 659Physiognomic-Structural Classification 661Floristic Classification 662Multifactor Classification 662A Classification of Forests 663Tropical Forests 663Classification of Tropical Forests 663Swamp Forest 664 ~

Salt-Water Tropical Shorelines 664Fresh-Water Swamps 665

Rain Forest 665Monsoon Forest 667Dry Forest 667Freeze-Hardy Forests 668Boreal Forest Taxa 668

Spruces 669Firs 669Larches 670Pines 670Birches and Aspens 670

Temperate Forest Taxa 671Pines 671

Soft Pines 672Hard Pines 672

Oaks 673Beeches and Maples 675Miscellaneous Hardwoods 676Miscellaneous Conifers 676

Suggested Readings 678

LITERATURE CITED 679SCIENTIFIC NAMES OF TREES AND SHRUBS 753INDEX 763