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1 Introduction to Ecology:Historical Foundations and Developing Frontiers1

Section Ⅰ Natural History and Evolution9

2 Life on Land9

3 Life in Water40

4 Population Genetics and Natural Selection70

Section Ⅱ Adaptations to the Environment90

5 Temperature Relations90

6 Water Relations114

7 Energy and Nutrient Relations135

8 Social Relations156

Section Ⅲ Population Ecology181

9 Population Distribution and Abundance181

10 Population Dynamics196

11 Population Growth216

12 Life Histories227

Section Ⅳ Interactions248

13 Competition248

14 Exploitative Interactions:Predation,Herbivory,Parasitism,and Disease268

15 Mutualism290

Section Ⅴ Communities and Ecosystems307

16 Species Abundance and Diversity307

17 Species Interactions and Community Structure325

18 Primary Production and Energy Flow344

19 Nutrient Cycling and Retention361

20 Succession and Stability381

Section Ⅵ Large-Scale Ecology403

21 Landscape Ecology403

22 Geographic Ecology426

23 Global Ecology444

Appendix Statistical Tables464

Chapter 1 Introduction to Ecology:1

Historical Foundations and Developing Frontiers1

Overview of Ecology2

The Ecology of Forest Birds:Old Tools and New3

Forest Canopy Research:A Physical and Scientific Frontier5

Climatic and Ecological Change:Past and Future6

The Scope of Ecology8

Section Ⅰ NATURAL HISTORY AND EVOLUTION9

ter 2 Life on Land9

Terrestrial Biomes10

Concepts10

2.1 Large-Scale Patterns of Climatic Variation11

Temperature,Atmospheric Circulation,and Precipitation11

Climate Diagrams11

Concept 2.1 Review14

2.2 Soil:The Foundation of Terrestrial Biomes14

Concept 2.2 Review17

2.3 Natural History and Geography of Biomes17

Tropical Rain Forest17

Tropical Dry Forest18

Tropical Savanna21

Desert22

Mediterranean Woodland and Shrubland25

Temperate Grassland27

Temperate Forest29

Boreal Forest30

Tundra33

Mountains:Islands in the Sky35

Concept 2.3 Review37

Applications:Climatic Variation and the Palmer Drought Severity Index37

chapter 3 Life in Water40

Concepts41

3.1 The Hydrologic Cycle41

Concept 3.1 Review41

3.2 The Natural History of Aquatic Environments41

TheOceans41

Life in Shallow Marine Waters:Kelp Forests and Coral Gardens47

Marine Shores:Life Between High and Low Tides50

Estuaries,Salt Marshes,and Mangrove Forests54

Rivers and Streams:Life Blood and Pulse of theLand58

Lakes:Small Seas63

Concept 3.2 Review67

Applications:Biological Integrity—Assessing the Health of Aquatic Systems67

Number of Species and Species Composition68

Trophic Composition68

Fish Abundance and Condition68

ATest68

Chapter 4 Population Genetics and Natural Selection70

Concepts72

4.1 Variation Within Populations72

Variation in a Widely Distributed Plant72

Variation in Alpine Fish Populations73

Concept 4.1 Review76

4.2 Hardy-Weinberg76

Calculating Gene Frequencies76

Concept 4.2 Review78

4.3 The Process of Natural Selection78

Stabilizing Selection78

Directional Selection80

Disruptive Selection80

Concept 4.3 Review80

4.4 Evolution by Natural Selection80

Evolution by Natural Selection and Genetic variation80

Adaptive Change in Colonizing Lizards81

Rapid Adaptation by Soapberry Bugs to New Host Plants83

Concept 4.4 Review84

4.5 Change Due to Chance84

Evidence of Genetic Drift in Chihuahua Spruce84

Genetic Variation in Island Populations85

Genetic Diversity and Butterfly Extinctions86

Concept 4.5 Review87

Applications:Evolution and Agriculture87

Evolution of Herbicide Resistance in Weeds87

SectionⅡ ADAPTATIONS TO THE ENVIRONMENT90

Chapter 5 Temperature Relations90

Concepts91

5.1 Microclimates91

Altitude91

Aspect91

Vegetation92

Color of the Ground92

Presence of Boulders and Burrows93

Aquatic Temperatures93

Concept 5.1 Review94

5.2 Temperature and Performance of Organisms94

The Principle of Allocation94

Temperature and Animal Performance95

Extreme Temperatures and Photosvnthesis96

Temperature and Microbial Activity97

Concept 5.2 Review99

5.3 Regulating Body Temperature99

Balancing Heat Gain Against Heat Loss99

Temperature Regulation by Plants100

Temperature Regulation by Ectothermic Animals102

Temperature Regulation by Endothermic Animals104

Temperature Regulation by Thermogenic Plants108

Concept 5.3 Review108

5.4 Surviving Extreme Temperatures109

Inactivity109

Reducing Metabolic Rate109

Hibernation by a Tropical Species110

Concept 5.4 Review111

Applications:Climatic Warming and the Local Extinction of a Land Snail111

Chapter 6 Water Relations114

Concepts115

6.1 Water Availability116

Water Content of Air116

Water Movement in Aquatic Environments117

Water Movement Between Soils and Plants118

Concept 6.1 Review120

6.2 Water Regulation on Land120

Water Acquisition by Animals120

Water Acquisition by Plants122

Water Conservation by Plants and Animals124

Investigating the Evidence 1:Sample Size125

Dissimilar Organisms with Similar Approaches to Desert Life127

Two Arthropods with Opposite Approaches to Desert Life128

Concept 6.2 Review130

6.3 Water and Salt Balance in Aquatic Environments130

Marine Fish and Invertebrates132

Freshwater Fish and Invertebrates132

Concept 6.3 Review134

Chapter 7 Energy and Nutrient Relations135

Concepts137

7.1 Photosynthetic Autotrophs137

The Solar-Powered Biosphere137

Concept 7.1 Review141

7.2 Chemosynthetic Autotrophs141

Concept 7.2 Review141

7.3 Heterotrophs141

Chemical Composition and Nutrient Requirements142

Concept 7.3 Review148

7.4 Energy Limitation148

Photon Flux and Photosvnthetic Response Curves148

Food Density and Animal Functional Response149

Concept 7.4 Review151

7.5 Optimal Foraging Theory151

Testing Optimal Foraging Theory151

Optimal Foraging by Plants153

Concept 7.5 Review154

Applications:Bioremediation—Using the Trophic Diversity of Bacteria to Solve Environmental Problems154

Leaking Underground Storage Tanks154

Cyanide and Nitrates in Mine Spoils155

Chapter 8 Social Relations156

Concepts158

8.1 Mate Choice158

Mate Choice and Sexual Selection in Guppies159

Mate Choice Among Scorpionflies163

Nonrandom Mating Among Wild Radish166

Concept 8.1 Review168

8.2 Sociality168

Cooperative Breeders168

Investigating the Evidence 2:Estimating Heritability Using Regression Analysis172

Concept 8.2 Review175

8.3 Eusociality175

Eusocial Species175

Evolution of Eusociality177

Concept 8.3 Review179

Applications:Behavioral Ecology and Conservation179

Tinbergen's Framework179

Environmental Enrichment and Development of Behavior179

Section Ⅲ POPULATION ECOLOGY181

Chapter 9 Population Distribution and Abundance181

Concepts183

9.1 Distribution Limits183

Distributions of Plants Along a Moisture-Temperature Gradient183

Distributions of Barnacles Along an Intertidal Exposure Gradient184

Concept 9.1 Review186

9.2 Patterns on Small Scales186

Scale,Distributions,and Mechanisms186

Distributions of Desert Shrubs187

Concept 9.2 Review188

9.3 Patterns on Large Scales188

Bird Populations Across North America189

Plant Distributions Along Moisture Gradients190

Concept 9.3 Review191

9.4 Organism Size and Population Density191

Animal Size and Population Density191

Plant Size and Population Density192

Concept 9.4 Review193

Applications:Rarity and Vulnerability to Extinction193

Seven Forms of Rarity and One of Abundance193

Chapter 10 Population Dynamics196

Concepts197

10.1 Dispersal197

Dispersal of Expanding Populations198

Range Changes in Response to Climate Change199

Dispersal in Response to Changing Food Supply199

Dispersal in Rivers and Streams201

Concept 10.1 Review202

10.2 Metapopulations202

A Metapopulation of an Alpine Butterfly202

Dispersal Within a Metapopulation of Lesser Kestrels203

Concept 10.2 Review204

10.3 Patterns of Survival204

Estimating Patterns of Survival204

High Survi val Among the Young205

Constant Rates of Survival207

High Mortality Among the Young207

Three Types of Survivorship Curves208

Concept 10.3 Review208

10.4 Age Distribution208

Contrasting Tree Populations208

A Dynamic Population in a Variable Climate209

Concept 10.4 Review210

10.5 Rates of Population Change210

Estimating Rates for an Annual Plant210

Estimating Rates When Generations Overlap212

Concept 10.5 Review213

Applications:Using Population Dynamics to Assess the Impact of Pollutants214

Chapter 11 Population Growth216

Concepts217

11.1 Geometric and Exponential Population Growth217

Geometric Growth217

Exponential Growth218

Exponential Growth in Nature219

Concept 11.1 Review220

11.2 Logistic Population Growth221

Concept 11.2 Review223

Applications:The Human Population223

Distribution and Abundance223

Population Dynamics224

Population Growth225

Chapter 12 Life Histories227

Concepts228

12.1 Offspring Number Versus Size228

Egg Size and Number in Fish229

Seed Size and Number in Plants230

Seed Size and Seedling Performance232

Concept 12.1 Review234

12.2 Adult Survival and Reproductive Allocation235

Life History Variation Among Species235

Life History Variation Within Species236

Concept 12.2 Review239

12.3 Life History Classification239

rand K Selection239

Plant Life Histories240

Opportunistic,Equilibrium,and Periodic Life Histories241

Reproductive Effort,Offspring Size,and Benefit-Cost Ratios242

Concept 12.3 Review244

Applications:Using Life History Information to Restore Riparian Forests245

Section Ⅳ INTERACTIONS248

Chapter 13 Competition248

Concepts250

13.1 Intraspecific Competition250

Intraspecific Competition Among Plants250

Intraspecific Competition Among Planthoppers251

Interference Competition Among Terrestrial Isopods252

Concept 13.1 Review252

13.2 Competitive Exclusion and Niches252

The Feeding Niches of Galápagos Finches253

The Habitat Niche of a Salt Marsh Grass253

Concept 13.2 Review255

13.3 Mathematical and Laboratory Models255

Modeling Interspecific Competition255

Laboratory Models of Competition257

Concept 13.3 Review258

13.4 Competition and Niches258

Niches and Competition Among Plants259

Niche Overlap and Competition Between Barnacles259

Competition and the Habitat of a Salt Marsh Grass260

Competition and the Niches of Small Rodents261

Character Displacement262

Investigating the Evidence 3:Field Experiments265

Concept 13.4 Review266

Applications:Competition Between Native and Invasive Species266

Chapter 14 Exploitative Interactions:Predation，Herbivory,Parasitism,and Disease268

Concepts269

14.1 Complex Interactions269

Parasites and Pathogens That Manipulate Host Behavior269

The Entangling of Exploitation with Competition271

Concept 14.1 Review273

14.2 Exploitation and Abundance273

A Herbivorous Stream Insect and Its Algal Food273

A Pathogenic Parasite,a Predator,and Its Prey274

Concept 14.2 Review277

14.3 Dynamics277

Cycles of Abundance in Snowshoe Hares and Their Predators277

Experimental Test of Food and Predation Impacts279

Population Cycles in Mathematical and Laboratory Models279

Concept 14.3 Review281

14.4 Refuges281

Refuges and Host Persistence in Laboratory and Mathematical Models281

Exploited Organisms and Their Wide Variety of“Refuges”283

Concept 14.4 Review286

Applications:Using Predators to Control a Parasite287

Chapter 15 Mutualism290

Concepts291

15.1 Plant Mutualisms291

Plant Performance and Mycorrhizal Fungi292

Ants and Swollen Thorn Acacias295

A Temperate Plant Protection Mutualism299

Concept 15.1 Review301

15.2 Evolution of Mutualism301

Facultative Ant-Plant Protection Mutualisms303

Concept 15.2 Review303

Applications:Mutualism and Humans303

Guiding Behavior304

Section Ⅴ COMMUNITIES AND ECOSYSTEMS307

Chapter 16 Species Abundance and Diversity307

Concepts308

16.1 Species Abundance309

The Lognormal Distribution309

Concept 16.1 Review310

16.2 Species Diversity310

A Quantitative Index of Species Diversity310

Rank-Abundance Curves311

Concept 16.2 Review312

Investigating the Evidence 4:Estimating the Number of Species in Communities313

16.3 Environmental Complexity314

Forest Complexity and Bird Species Diversity314

Niches,Heterogeneity,and the Diversity of Algae and Plants315

The Niches of Algae and Terrestrial Plants315

Complexity in Plant Environments316

Soil and Topographic Heterogeneity and the Diversity of Tropical Forest Trees316

Algal and Plant Species Diversity and Increased Nutrient Availability318

Nitrogen Enrichment and Ectomycorrhizal Fungus Diversity318

Concept 16.3 Review319

16.4 Disturbance and Diversity319

The Nature of Equilibrium319

The Nature and Sources of Disturbance319

The Intermediate Disturbance Hypothesis319

Disturbance and Diversity in the Intertidal Zone320

Disturbance and Diversity in Temperate Grasslands321

Concept 16.4 Review322

Applications:Disturbance by Humans322

Disturbance by Humans and the Diversity of Chalk Grasslands323

Chapter 17 Species Interactions and Community Structure325

Concepts327

17.1 Community Webs327

Detailed Food Webs Reveal Great Complexity327

Strong Interactions and Food Web Structure327

Concept 17.1 Review329

17.2 Indirect Interactions329

Indirect Commensalism329

Apparent Competition330

Concept 17.2 Review330

17.3 Keystone Species330

Food Web Structure and Species Diversity330

Experimental Removal of Sea Stars332

Snail Effects on Algal Diversity334

Fish as Keystone Species in River Food Webs336

Concept 17.3 Review339

17.4 Mutualistic Keystones339

A Cleaner Fish as a Keystone Species339

Seed Dispersal Mutualists as Keystone Species339

Concept 17.4 Review340

Applications:Human Modification of Food Webs340

The Empty Forest:Hunters and Tropical Rain Forest Animal Communities340

Ants and Agriculture:Keystone Predators for Pest Control341

Chapter 18 Primary Production and Energy Flow344

Concepts346

18.1 Patterns of Terrestrial Primary Production346

Actual Evapotranspiration and Terrestrial Primary Production346

Soil Fertility and Terrestrial Primary Production347

Concept 18.1 Review348

18.2 Patterns of Aquatic Primary Production348

Patterns and Models348

Whole Lake Experiments on Primary Production349

Global Patterns of Marine Primary Production349

Concept 18.2 Review351

18.3 Consumer Influences351

Piscivores,Planktivores,and Lake Primary Production351

Grazing by Large Mammals and Primary Production on the Serengeti353

Concept 18.3 Review353

18.4 Trophic Levels355

A Trophic Dynamic View of Ecosystems355

Energy Flow in a Temperate Deciduous Forest356

Concept 18.4 Review357

Applications:Using Stable Isotope Analysis to Trace Energy Flow Through Ecosystems357

Trophic Levels of Tropical River Fish358

Using Stable Isotopes to Identify Sources of Energy in a SaltMarsh358

Food Habits of Prehistoric Human Populations359

Chapter 19 Nutrient Cycling and Retention361

Concepts362

19.1 Nutrient Cycles362

The Phosphorus Cycle363

The Nitrogen Cycle364

The Carbon Cycle365

Concept 19.1 Review366

19.2 Rates of Decomposition366

Decomposition in Two Mediterranean Woodland Ecosystems366

Decomposition in Two Temperate Forest Ecosystems367

Decomposition in Aquatic Ecosystems369

Concept 19.2 Review370

19.3 Organisms and Nutrients370

Nutrient Cycling in Streams371

Animals and Nutrient Cycling in Terrestrial Ecosystems372

Plants and the Nutrient Dynamics of Ecosystems373

Concept 19.3 Review375

19.4 Disturbance and Nutrients375

Disturbance and Nutrient Loss from the Hubbard Brook Experimental Forest375

Flooding and Nutrient Export by Streams376

Concept 19.4 Review378

Applications:Altering Aquatic and Terrestrial Ecosystems378

Chapter 20 Succession and Stability381

Concepts383

20.1 Community Changes During Succession383

Primary Succession at Glacier Bay383

Secondary Succession in Temperate Forests384

Succession in Rocky Intertidal Communities385

Succession in Stream Communities385

Concept 20.1 Review387

20.2 Ecosystem Changes During Succession387

Ecosystem Changes at Glacier Bay387

Four Million Years of Ecosystem Change387

Recovery of Nutrient Retention Following Disturbance389

Succession and Stream Ecosystem Properties391

Concept 20.2 Review392

20.3 Mechanisms of Succession392

Successional Mechanisms in the Rocky Intertidal Zone394

Successional Mechanisms in Forests395

Concept 20.3 Review396

20.4 Community and Ecosystem Stability396

Some Definitions396

Lessons from the Park Grass Experiment397

Replicate Disturbances and Desert Stream Stability398

Concept 20.4 Review400

Applications:Using Repeat Photography to Detect Long-Term Change400

Section Ⅵ LARGE-SCALE ECOLOGY403

Chapter 21 LandscaPe Ecology403

Concepts405

21.1 Landscape Structure405

The Structure of Six Landscapes in Ohio405

The Fractal Geometry of Landscapes407

Concept 21.1 Review408

21.2 Landscape Processes408

Landscape Structure and the Dispersal of Mammals409

Habitat Patch Size and Isolation and the Density of Butterfly Populations410

Habitat Corridors and Movement of Organisms411

Landscape Position and Lake Chemistry412

Investigating the Evidence 5:Comparison of Two Samples Using a Rank Sum Test413

Concept 21.2 Review414

21.3 Origins of Landscape Structure and Change414

Geological Processes,Climate,and Landscape Structure414

Organisms and Landscape Structure417

Fire and the Structure of a Mediterranean Landscape421

Concept 21.3 Review421

Applications:Restoring a Riverine Landscape422

Riverine Restoration:The Kissimmee River423

Chapter 22 Geographic Ecology426

Concepts428

22.1 Area,Isolation,and Species Richness428

Sampling Area and Number of Species428

Island Area and Species Richness428

Island Isolation and Species Richness430

Concept 22.1 Review431

22.2 The Equilibrium Model of Island Biogeography431

Species Turnover on Islands432

Experimental Island Biogeography433

Colonization of New Islands by Plants434

Manipulating Island Area435

Island Biogeography Update436

Concept 22.2 Review436

22.3 Latitudinal Gradients in Species Richness436

Area and Latitudinal Gradients in Species Richness438

Continental Area and Species Richness439

Concept 22.3 Review440

22.4 Historical and Regional Influences440

Exceptional Patterns of Diversity440

Investigating the Evidence 6:Sample Size Revisited441

Historical and Regional Explanations441

Concept 22.4 Review443

Chapter 23 Global Ecology444

The Atmospheric Envelope and the Greenhouse Earth445

Concepts446

23.1 A Global System446

The Historical Thread447

El Ni？o andLaNi？a448

El Ni？o and Marine Populations449

El Ni？o and the Great Salt Lake451

E1 Ni？o and Terrestrial Populations in Australia452

Concept 23.1 Review453

23.2 Human Activity and the G1obal Nitrogen Cycle453

Concept 23.2 Review454

23.3 Changes in Land Cover454

Tropical Deforestation454

Concept 23.3 Review458

23.4 Human Influence on Atmospheric Composition458

Depletion and Recovery of the Ozone Layer460

The Future461

Concept 23.4 Review462

Applications:Cooperative Research Networks for Global Ecology462

Appendix Statistical Tables464

Glossary468

Credits478