1 Introduction to Ecology:Historical Foundations and Developing Frontiers 1
Section Ⅰ Natural History and Evolution 9
2 Life on Land 9
3 Life in Water 40
4 Population Genetics and Natural Selection 70
Section Ⅱ Adaptations to the Environment 90
5 Temperature Relations 90
6 Water Relations 114
7 Energy and Nutrient Relations 135
8 Social Relations 156
Section Ⅲ Population Ecology 181
9 Population Distribution and Abundance 181
10 Population Dynamics 196
11 Population Growth 216
12 Life Histories 227
Section Ⅳ Interactions 248
13 Competition 248
14 Exploitative Interactions:Predation,Herbivory,Parasitism,and Disease 268
15 Mutualism 290
Section Ⅴ Communities and Ecosystems 307
16 Species Abundance and Diversity 307
17 Species Interactions and Community Structure 325
18 Primary Production and Energy Flow 344
19 Nutrient Cycling and Retention 361
20 Succession and Stability 381
Section Ⅵ Large-Scale Ecology 403
21 Landscape Ecology 403
22 Geographic Ecology 426
23 Global Ecology 444
Appendix Statistical Tables 464
Chapter 1 Introduction to Ecology: 1
Historical Foundations and Developing Frontiers 1
Overview of Ecology 2
The Ecology of Forest Birds:Old Tools and New 3
Forest Canopy Research:A Physical and Scientific Frontier 5
Climatic and Ecological Change:Past and Future 6
The Scope of Ecology 8
Section Ⅰ NATURAL HISTORY AND EVOLUTION 9
ter 2 Life on Land 9
Terrestrial Biomes 10
Concepts 10
2.1 Large-Scale Patterns of Climatic Variation 11
Temperature,Atmospheric Circulation,and Precipitation 11
Climate Diagrams 11
Concept 2.1 Review 14
2.2 Soil:The Foundation of Terrestrial Biomes 14
Concept 2.2 Review 17
2.3 Natural History and Geography of Biomes 17
Tropical Rain Forest 17
Tropical Dry Forest 18
Tropical Savanna 21
Desert 22
Mediterranean Woodland and Shrubland 25
Temperate Grassland 27
Temperate Forest 29
Boreal Forest 30
Tundra 33
Mountains:Islands in the Sky 35
Concept 2.3 Review 37
Applications:Climatic Variation and the Palmer Drought Severity Index 37
chapter 3 Life in Water 40
Concepts 41
3.1 The Hydrologic Cycle 41
Concept 3.1 Review 41
3.2 The Natural History of Aquatic Environments 41
TheOceans 41
Life in Shallow Marine Waters:Kelp Forests and Coral Gardens 47
Marine Shores:Life Between High and Low Tides 50
Estuaries,Salt Marshes,and Mangrove Forests 54
Rivers and Streams:Life Blood and Pulse of theLand 58
Lakes:Small Seas 63
Concept 3.2 Review 67
Applications:Biological Integrity—Assessing the Health of Aquatic Systems 67
Number of Species and Species Composition 68
Trophic Composition 68
Fish Abundance and Condition 68
ATest 68
Chapter 4 Population Genetics and Natural Selection 70
Concepts 72
4.1 Variation Within Populations 72
Variation in a Widely Distributed Plant 72
Variation in Alpine Fish Populations 73
Concept 4.1 Review 76
4.2 Hardy-Weinberg 76
Calculating Gene Frequencies 76
Concept 4.2 Review 78
4.3 The Process of Natural Selection 78
Stabilizing Selection 78
Directional Selection 80
Disruptive Selection 80
Concept 4.3 Review 80
4.4 Evolution by Natural Selection 80
Evolution by Natural Selection and Genetic variation 80
Adaptive Change in Colonizing Lizards 81
Rapid Adaptation by Soapberry Bugs to New Host Plants 83
Concept 4.4 Review 84
4.5 Change Due to Chance 84
Evidence of Genetic Drift in Chihuahua Spruce 84
Genetic Variation in Island Populations 85
Genetic Diversity and Butterfly Extinctions 86
Concept 4.5 Review 87
Applications:Evolution and Agriculture 87
Evolution of Herbicide Resistance in Weeds 87
SectionⅡ ADAPTATIONS TO THE ENVIRONMENT 90
Chapter 5 Temperature Relations 90
Concepts 91
5.1 Microclimates 91
Altitude 91
Aspect 91
Vegetation 92
Color of the Ground 92
Presence of Boulders and Burrows 93
Aquatic Temperatures 93
Concept 5.1 Review 94
5.2 Temperature and Performance of Organisms 94
The Principle of Allocation 94
Temperature and Animal Performance 95
Extreme Temperatures and Photosvnthesis 96
Temperature and Microbial Activity 97
Concept 5.2 Review 99
5.3 Regulating Body Temperature 99
Balancing Heat Gain Against Heat Loss 99
Temperature Regulation by Plants 100
Temperature Regulation by Ectothermic Animals 102
Temperature Regulation by Endothermic Animals 104
Temperature Regulation by Thermogenic Plants 108
Concept 5.3 Review 108
5.4 Surviving Extreme Temperatures 109
Inactivity 109
Reducing Metabolic Rate 109
Hibernation by a Tropical Species 110
Concept 5.4 Review 111
Applications:Climatic Warming and the Local Extinction of a Land Snail 111
Chapter 6 Water Relations 114
Concepts 115
6.1 Water Availability 116
Water Content of Air 116
Water Movement in Aquatic Environments 117
Water Movement Between Soils and Plants 118
Concept 6.1 Review 120
6.2 Water Regulation on Land 120
Water Acquisition by Animals 120
Water Acquisition by Plants 122
Water Conservation by Plants and Animals 124
Investigating the Evidence 1:Sample Size 125
Dissimilar Organisms with Similar Approaches to Desert Life 127
Two Arthropods with Opposite Approaches to Desert Life 128
Concept 6.2 Review 130
6.3 Water and Salt Balance in Aquatic Environments 130
Marine Fish and Invertebrates 132
Freshwater Fish and Invertebrates 132
Concept 6.3 Review 134
Chapter 7 Energy and Nutrient Relations 135
Concepts 137
7.1 Photosynthetic Autotrophs 137
The Solar-Powered Biosphere 137
Concept 7.1 Review 141
7.2 Chemosynthetic Autotrophs 141
Concept 7.2 Review 141
7.3 Heterotrophs 141
Chemical Composition and Nutrient Requirements 142
Concept 7.3 Review 148
7.4 Energy Limitation 148
Photon Flux and Photosvnthetic Response Curves 148
Food Density and Animal Functional Response 149
Concept 7.4 Review 151
7.5 Optimal Foraging Theory 151
Testing Optimal Foraging Theory 151
Optimal Foraging by Plants 153
Concept 7.5 Review 154
Applications:Bioremediation—Using the Trophic Diversity of Bacteria to Solve Environmental Problems 154
Leaking Underground Storage Tanks 154
Cyanide and Nitrates in Mine Spoils 155
Chapter 8 Social Relations 156
Concepts 158
8.1 Mate Choice 158
Mate Choice and Sexual Selection in Guppies 159
Mate Choice Among Scorpionflies 163
Nonrandom Mating Among Wild Radish 166
Concept 8.1 Review 168
8.2 Sociality 168
Cooperative Breeders 168
Investigating the Evidence 2:Estimating Heritability Using Regression Analysis 172
Concept 8.2 Review 175
8.3 Eusociality 175
Eusocial Species 175
Evolution of Eusociality 177
Concept 8.3 Review 179
Applications:Behavioral Ecology and Conservation 179
Tinbergen's Framework 179
Environmental Enrichment and Development of Behavior 179
Section Ⅲ POPULATION ECOLOGY 181
Chapter 9 Population Distribution and Abundance 181
Concepts 183
9.1 Distribution Limits 183
Distributions of Plants Along a Moisture-Temperature Gradient 183
Distributions of Barnacles Along an Intertidal Exposure Gradient 184
Concept 9.1 Review 186
9.2 Patterns on Small Scales 186
Scale,Distributions,and Mechanisms 186
Distributions of Desert Shrubs 187
Concept 9.2 Review 188
9.3 Patterns on Large Scales 188
Bird Populations Across North America 189
Plant Distributions Along Moisture Gradients 190
Concept 9.3 Review 191
9.4 Organism Size and Population Density 191
Animal Size and Population Density 191
Plant Size and Population Density 192
Concept 9.4 Review 193
Applications:Rarity and Vulnerability to Extinction 193
Seven Forms of Rarity and One of Abundance 193
Chapter 10 Population Dynamics 196
Concepts 197
10.1 Dispersal 197
Dispersal of Expanding Populations 198
Range Changes in Response to Climate Change 199
Dispersal in Response to Changing Food Supply 199
Dispersal in Rivers and Streams 201
Concept 10.1 Review 202
10.2 Metapopulations 202
A Metapopulation of an Alpine Butterfly 202
Dispersal Within a Metapopulation of Lesser Kestrels 203
Concept 10.2 Review 204
10.3 Patterns of Survival 204
Estimating Patterns of Survival 204
High Survi val Among the Young 205
Constant Rates of Survival 207
High Mortality Among the Young 207
Three Types of Survivorship Curves 208
Concept 10.3 Review 208
10.4 Age Distribution 208
Contrasting Tree Populations 208
A Dynamic Population in a Variable Climate 209
Concept 10.4 Review 210
10.5 Rates of Population Change 210
Estimating Rates for an Annual Plant 210
Estimating Rates When Generations Overlap 212
Concept 10.5 Review 213
Applications:Using Population Dynamics to Assess the Impact of Pollutants 214
Chapter 11 Population Growth 216
Concepts 217
11.1 Geometric and Exponential Population Growth 217
Geometric Growth 217
Exponential Growth 218
Exponential Growth in Nature 219
Concept 11.1 Review 220
11.2 Logistic Population Growth 221
Concept 11.2 Review 223
Applications:The Human Population 223
Distribution and Abundance 223
Population Dynamics 224
Population Growth 225
Chapter 12 Life Histories 227
Concepts 228
12.1 Offspring Number Versus Size 228
Egg Size and Number in Fish 229
Seed Size and Number in Plants 230
Seed Size and Seedling Performance 232
Concept 12.1 Review 234
12.2 Adult Survival and Reproductive Allocation 235
Life History Variation Among Species 235
Life History Variation Within Species 236
Concept 12.2 Review 239
12.3 Life History Classification 239
rand K Selection 239
Plant Life Histories 240
Opportunistic,Equilibrium,and Periodic Life Histories 241
Reproductive Effort,Offspring Size,and Benefit-Cost Ratios 242
Concept 12.3 Review 244
Applications:Using Life History Information to Restore Riparian Forests 245
Section Ⅳ INTERACTIONS 248
Chapter 13 Competition 248
Concepts 250
13.1 Intraspecific Competition 250
Intraspecific Competition Among Plants 250
Intraspecific Competition Among Planthoppers 251
Interference Competition Among Terrestrial Isopods 252
Concept 13.1 Review 252
13.2 Competitive Exclusion and Niches 252
The Feeding Niches of Galápagos Finches 253
The Habitat Niche of a Salt Marsh Grass 253
Concept 13.2 Review 255
13.3 Mathematical and Laboratory Models 255
Modeling Interspecific Competition 255
Laboratory Models of Competition 257
Concept 13.3 Review 258
13.4 Competition and Niches 258
Niches and Competition Among Plants 259
Niche Overlap and Competition Between Barnacles 259
Competition and the Habitat of a Salt Marsh Grass 260
Competition and the Niches of Small Rodents 261
Character Displacement 262
Investigating the Evidence 3:Field Experiments 265
Concept 13.4 Review 266
Applications:Competition Between Native and Invasive Species 266
Chapter 14 Exploitative Interactions:Predation，Herbivory,Parasitism,and Disease 268
Concepts 269
14.1 Complex Interactions 269
Parasites and Pathogens That Manipulate Host Behavior 269
The Entangling of Exploitation with Competition 271
Concept 14.1 Review 273
14.2 Exploitation and Abundance 273
A Herbivorous Stream Insect and Its Algal Food 273
A Pathogenic Parasite,a Predator,and Its Prey 274
Concept 14.2 Review 277
14.3 Dynamics 277
Cycles of Abundance in Snowshoe Hares and Their Predators 277
Experimental Test of Food and Predation Impacts 279
Population Cycles in Mathematical and Laboratory Models 279
Concept 14.3 Review 281
14.4 Refuges 281
Refuges and Host Persistence in Laboratory and Mathematical Models 281
Exploited Organisms and Their Wide Variety of“Refuges” 283
Concept 14.4 Review 286
Applications:Using Predators to Control a Parasite 287
Chapter 15 Mutualism 290
Concepts 291
15.1 Plant Mutualisms 291
Plant Performance and Mycorrhizal Fungi 292
Ants and Swollen Thorn Acacias 295
A Temperate Plant Protection Mutualism 299
Concept 15.1 Review 301
15.2 Evolution of Mutualism 301
Facultative Ant-Plant Protection Mutualisms 303
Concept 15.2 Review 303
Applications:Mutualism and Humans 303
Guiding Behavior 304
Section Ⅴ COMMUNITIES AND ECOSYSTEMS 307
Chapter 16 Species Abundance and Diversity 307
Concepts 308
16.1 Species Abundance 309
The Lognormal Distribution 309
Concept 16.1 Review 310
16.2 Species Diversity 310
A Quantitative Index of Species Diversity 310
Rank-Abundance Curves 311
Concept 16.2 Review 312
Investigating the Evidence 4:Estimating the Number of Species in Communities 313
16.3 Environmental Complexity 314
Forest Complexity and Bird Species Diversity 314
Niches,Heterogeneity,and the Diversity of Algae and Plants 315
The Niches of Algae and Terrestrial Plants 315
Complexity in Plant Environments 316
Soil and Topographic Heterogeneity and the Diversity of Tropical Forest Trees 316
Algal and Plant Species Diversity and Increased Nutrient Availability 318
Nitrogen Enrichment and Ectomycorrhizal Fungus Diversity 318
Concept 16.3 Review 319
16.4 Disturbance and Diversity 319
The Nature of Equilibrium 319
The Nature and Sources of Disturbance 319
The Intermediate Disturbance Hypothesis 319
Disturbance and Diversity in the Intertidal Zone 320
Disturbance and Diversity in Temperate Grasslands 321
Concept 16.4 Review 322
Applications:Disturbance by Humans 322
Disturbance by Humans and the Diversity of Chalk Grasslands 323
Chapter 17 Species Interactions and Community Structure 325
Concepts 327
17.1 Community Webs 327
Detailed Food Webs Reveal Great Complexity 327
Strong Interactions and Food Web Structure 327
Concept 17.1 Review 329
17.2 Indirect Interactions 329
Indirect Commensalism 329
Apparent Competition 330
Concept 17.2 Review 330
17.3 Keystone Species 330
Food Web Structure and Species Diversity 330
Experimental Removal of Sea Stars 332
Snail Effects on Algal Diversity 334
Fish as Keystone Species in River Food Webs 336
Concept 17.3 Review 339
17.4 Mutualistic Keystones 339
A Cleaner Fish as a Keystone Species 339
Seed Dispersal Mutualists as Keystone Species 339
Concept 17.4 Review 340
Applications:Human Modification of Food Webs 340
The Empty Forest:Hunters and Tropical Rain Forest Animal Communities 340
Ants and Agriculture:Keystone Predators for Pest Control 341
Chapter 18 Primary Production and Energy Flow 344
Concepts 346
18.1 Patterns of Terrestrial Primary Production 346
Actual Evapotranspiration and Terrestrial Primary Production 346
Soil Fertility and Terrestrial Primary Production 347
Concept 18.1 Review 348
18.2 Patterns of Aquatic Primary Production 348
Patterns and Models 348
Whole Lake Experiments on Primary Production 349
Global Patterns of Marine Primary Production 349
Concept 18.2 Review 351
18.3 Consumer Influences 351
Piscivores,Planktivores,and Lake Primary Production 351
Grazing by Large Mammals and Primary Production on the Serengeti 353
Concept 18.3 Review 353
18.4 Trophic Levels 355
A Trophic Dynamic View of Ecosystems 355
Energy Flow in a Temperate Deciduous Forest 356
Concept 18.4 Review 357
Applications:Using Stable Isotope Analysis to Trace Energy Flow Through Ecosystems 357
Trophic Levels of Tropical River Fish 358
Using Stable Isotopes to Identify Sources of Energy in a SaltMarsh 358
Food Habits of Prehistoric Human Populations 359
Chapter 19 Nutrient Cycling and Retention 361
Concepts 362
19.1 Nutrient Cycles 362
The Phosphorus Cycle 363
The Nitrogen Cycle 364
The Carbon Cycle 365
Concept 19.1 Review 366
19.2 Rates of Decomposition 366
Decomposition in Two Mediterranean Woodland Ecosystems 366
Decomposition in Two Temperate Forest Ecosystems 367
Decomposition in Aquatic Ecosystems 369
Concept 19.2 Review 370
19.3 Organisms and Nutrients 370
Nutrient Cycling in Streams 371
Animals and Nutrient Cycling in Terrestrial Ecosystems 372
Plants and the Nutrient Dynamics of Ecosystems 373
Concept 19.3 Review 375
19.4 Disturbance and Nutrients 375
Disturbance and Nutrient Loss from the Hubbard Brook Experimental Forest 375
Flooding and Nutrient Export by Streams 376
Concept 19.4 Review 378
Applications:Altering Aquatic and Terrestrial Ecosystems 378
Chapter 20 Succession and Stability 381
Concepts 383
20.1 Community Changes During Succession 383
Primary Succession at Glacier Bay 383
Secondary Succession in Temperate Forests 384
Succession in Rocky Intertidal Communities 385
Succession in Stream Communities 385
Concept 20.1 Review 387
20.2 Ecosystem Changes During Succession 387
Ecosystem Changes at Glacier Bay 387
Four Million Years of Ecosystem Change 387
Recovery of Nutrient Retention Following Disturbance 389
Succession and Stream Ecosystem Properties 391
Concept 20.2 Review 392
20.3 Mechanisms of Succession 392
Successional Mechanisms in the Rocky Intertidal Zone 394
Successional Mechanisms in Forests 395
Concept 20.3 Review 396
20.4 Community and Ecosystem Stability 396
Some Definitions 396
Lessons from the Park Grass Experiment 397
Replicate Disturbances and Desert Stream Stability 398
Concept 20.4 Review 400
Applications:Using Repeat Photography to Detect Long-Term Change 400
Section Ⅵ LARGE-SCALE ECOLOGY 403
Chapter 21 LandscaPe Ecology 403
Concepts 405
21.1 Landscape Structure 405
The Structure of Six Landscapes in Ohio 405
The Fractal Geometry of Landscapes 407
Concept 21.1 Review 408
21.2 Landscape Processes 408
Landscape Structure and the Dispersal of Mammals 409
Habitat Patch Size and Isolation and the Density of Butterfly Populations 410
Habitat Corridors and Movement of Organisms 411
Landscape Position and Lake Chemistry 412
Investigating the Evidence 5:Comparison of Two Samples Using a Rank Sum Test 413
Concept 21.2 Review 414
21.3 Origins of Landscape Structure and Change 414
Geological Processes,Climate,and Landscape Structure 414
Organisms and Landscape Structure 417
Fire and the Structure of a Mediterranean Landscape 421
Concept 21.3 Review 421
Applications:Restoring a Riverine Landscape 422
Riverine Restoration:The Kissimmee River 423
Chapter 22 Geographic Ecology 426
Concepts 428
22.1 Area,Isolation,and Species Richness 428
Sampling Area and Number of Species 428
Island Area and Species Richness 428
Island Isolation and Species Richness 430
Concept 22.1 Review 431
22.2 The Equilibrium Model of Island Biogeography 431
Species Turnover on Islands 432
Experimental Island Biogeography 433
Colonization of New Islands by Plants 434
Manipulating Island Area 435
Island Biogeography Update 436
Concept 22.2 Review 436
22.3 Latitudinal Gradients in Species Richness 436
Area and Latitudinal Gradients in Species Richness 438
Continental Area and Species Richness 439
Concept 22.3 Review 440
22.4 Historical and Regional Influences 440
Exceptional Patterns of Diversity 440
Investigating the Evidence 6:Sample Size Revisited 441
Historical and Regional Explanations 441
Concept 22.4 Review 443
Chapter 23 Global Ecology 444
The Atmospheric Envelope and the Greenhouse Earth 445
Concepts 446
23.1 A Global System 446
The Historical Thread 447
El Ni？o andLaNi？a 448
El Ni？o and Marine Populations 449
El Ni？o and the Great Salt Lake 451
E1 Ni？o and Terrestrial Populations in Australia 452
Concept 23.1 Review 453
23.2 Human Activity and the G1obal Nitrogen Cycle 453
Concept 23.2 Review 454
23.3 Changes in Land Cover 454
Tropical Deforestation 454
Concept 23.3 Review 458
23.4 Human Influence on Atmospheric Composition 458
Depletion and Recovery of the Ozone Layer 460
The Future 461
Concept 23.4 Review 462
Applications:Cooperative Research Networks for Global Ecology 462
Appendix Statistical Tables 464
Glossary 468
Credits 478