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ECOLOGY
Published by McGraw Hill LLC, 1325 Avenue of the Americas, New York, NY 10019. Copyright
© 2022 by McGraw Hill LLC. All rights reserved. Printed in the United States of America. No part
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About the Authors
Anna A. Sher is a full professor in the Department of Biological Sciences at the Uni-
versity of Denver, where she has been faculty since 2003. Until 2010 she held this position jointly
with the Denver Botanic Gardens as the Director of Research and Conservation. As a student, she
was a double major in Biology and Art at Earlham College, where she has also taught ecology, and
was the co-leader of the Earlham Study Abroad Kenya Program. She received her PhD from the
University of New Mexico, where she also taught botany as a visiting lecturer. As a postdoctoral
researcher, Dr. Sher was awarded a Fulbright postdoctoral research fellowship to conduct research
on plant interactions in Israel at Ben Gurion University’s Mitrani Department of Desert Ecology,
and she also studied the ecology of an invasive grass at the University of California, Davis. She has
also been a visiting professor at the University of Otago, Dunedin, New Zealand.
Dr. Sher’s primary research focus has been on the ecological dynamics associated with
the removal of invasive riparian plants. She is known as a leading expert in the ecology of
Tamarix, a dominant exotic tree, and she was the lead editor of the first book exclusively on
the topic. Her research interests and publications have spanned several areas within ecol-
ogy, including not only restoration ecology, competition, and invasive species ecology, but
also interactions between plants and soil chemistry, mycorrhizae, insect diversity and trophic
cascades, ethnobotany, phenology, climate change, and rare species conservation. She is also
lead author of the textbook series An Introduction to Conservation Biology (Oxford University Courtesy of Anna Sher
Press). Dr. Sher has a particular interest in quantitative ecological methods, with her lab
specializing in multivariate methods and spatial models at both individual organism and regional scales. She is currently principal
investigator of a National Science Foundation award to investigate the human dimension of the restoration of damaged ecosys-
tems, and she has been a TEDx speaker on the way ecosystems can teach us how to solve human problems.
Above all, Dr. Sher loves to teach and mentor students doing research at both undergraduate and graduate levels.
Manuel C. Molles Jr. is an emeritus Professor of Biology at the University of New Mexico, where he has been a
member of the faculty and curator in the Museum of Southwestern Biology since 1975. He received his BS from Humboldt State
University and his PhD from the Department of Ecology and Evolutionary Biology at the University of
Arizona. Seeking to broaden his geographic perspective, he has taught and conducted ecological research
in Latin America, the Caribbean, and Europe. He was awarded a Fulbright Research Fellowship to con-
duct research on river ecology in Portugal and has held visiting professor appointments in the Department
of Zoology at the University of Coimbra, Portugal, in the Laboratory of Hydrology at the Polytechnic
University of Madrid, Spain, and at the University of Montana’s Flathead Lake Biological Station.
Originally trained as a marine ecologist and fisheries biologist, the author worked mainly on river and
riparian ecology at the University of New Mexico. His research has covered a wide range of ecological levels,
including behavioral ecology, population biology, community ecology, ecosystem ecology, biogeography of
stream insects, and the influence of a large-scale climate system (El Niño) on the dynamics of southwestern
river and riparian ecosystems. His current research interests focus on the influence of climate change and cli-
matic variability on the dynamics of populations and communities along steep gradients of temperature and
moisture in the mountains of the Southwest. Throughout his career, Dr. Molles has attempted to combine
research, teaching, and service, involving undergraduate as well as graduate students in his ongoing projects.
At the University of New Mexico, he taught a broad range of lower division, upper division, and graduate
courses, including Principles of Biology, Evolution and Ecology, Stream Ecology, Limnology and Oceanog-
raphy, Marine Biology, and Community and Ecosystem Ecology. He has taught courses in Global Change
and River Ecology at the University of Coimbra, Portugal, and General Ecology and Groundwater and
Riparian Ecology at the Flathead Lake Biological Station. Dr. Manuel Molles was named Teacher of the
Year by the University of New Mexico for 1995–1996 and Potter Chair in Plant Ecology in 2000. In 2014, he
received the Eugene P. Odum Award from the Ecological Society of America based on his “ability to relate
basic ecological principles to human affairs through teaching, outreach and mentoring activities.”
Courtesy of Manuel Molles
Design elements: Anna A. Sher
iii
Dedication
To the Sher Lab and the whole next generation
of ecologists, who inspire me to do this work.
Also, I dedicate this edition to my co-author
and mentor, Manuel.
—Anna A. Sher
Brief Contents
1 Introduction to Ecology: Historical Foundations and Developing Frontiers 1
Section Natural History and Evolution 11
I 2 Life on Land 11
3 Life in Water 44
4 Population Genetics and Natural Selection 78
Section Adaptations to the Environment 101
II 5
6
7
Temperature Relations 101
Water Relations 127
Energy and Nutrient Relations 149
8 Social Relations 172
Section Population Ecology 196
III 9
10
11
Population Distribution and Abundance
Population Dynamics 215
Population Growth 237
196
12 Life Histories 254
Section Interactions 277
IV 13 Species Interactions and Competition 277

14 Exploitative Interactions: Predation, Herbivory, Parasitism, and Disease 299
15 Mutualism 325
Section Communities and Ecosystems 345
V 16
17
18
Species Abundance and Diversity 345
Species Interactions and Community Structure
Primary and Secondary Production 383
365
19 Nutrient Cycling and Retention 403

20 Succession and Stability 423
Section Large-Scale Ecology 445
VI 21 Landscape Ecology 445

22 Geographic Ecology 468
23 Global Ecology 490
Appendix A Investigating the Evidence 514

Appendix B Statistical Tables 541
Appendix C Abbreviations Used in This Text 545
Appendix D Global Biomes 547

v
Contents
Preface xiii
Chapter 3 Life in Water 44
Chapter 1 Introduction to Ecology: Concepts 44
Historical Foundations and Aquatic Biomes and How They Differ 45

Developing Frontiers 1 3.1 Water Cycling 47
The Hydrologic Cycle 47
Concepts 1
The Effects of Wind and Temperature 47
1.1 Overview of Ecology 2 Concept 3.1 Review 48
Concept 1.1 Review 3 3.2 The Natural History of Aquatic Environments 49
1.2 Sampling Ecological Research 3 The Oceans 49
Climatic and Ecological Change: Past and Future 7 Life in Shallow Marine Waters: Kelp Forests and Coral
Concept 1.2 Review 9 Gardens 54
Applications: Ecology Can Inform Environmental Law Marine Shores: Life Between High and Low Tides 57
and Policy 9 Transitional Environments: Estuaries, Salt Marshes,
Mangrove Forests, and Freshwater Wetlands 59
Rivers and Streams: Life Blood and Pulse of the Land 64
Section I Lakes: Small Seas 69

Concept 3.2 Review 73
NATURAL HISTORY AND EVOLUTION Applications: Biological Integrity—Assessing the Health of
Aquatic Systems 73
Chapter 2 Life on Land 11 Number of Species and Species Composition 74
Trophic Composition 74
Concepts 11
Fish Abundance and Condition 74
Terrestrial Biomes and the Importance A Test 74
of Plants 12
2.1 Large-Scale Patterns of Climatic Variation 14
Temperature, Atmospheric Circulation, and Chapter 4 Population Genetics and Natural
Precipitation 14 Selection 78
Climate Diagrams 16
Concepts 78
4.1 Variation Within Populations 81
2.2 Other Factors That Shape Terrestrial
Biomes 17 Variation in a Widely Distributed Plant 81
Concept 2.2 Review 19 Variation in Alpine Fish Populations 83
2.3 Natural History and Geography of Biomes 19
4.2 Hardy-Weinberg Principle 84
Tropical Rain Forest 19
Tropical Dry Forest 21 Calculating Gene Frequencies 84
Tropical Savanna 22 Concept 4.2 Review 86
Desert 25 4.3 The Process of Natural Selection 87
Woodland and Shrubland 27 Stabilizing Selection 87
Temperate Grassland 29 Directional Selection 87
Temperate Forest 31 Disruptive Selection 87
Boreal Forest 32 Concept 4.3 Review 88
Tundra 35 4.4 Evolution by Natural Selection 88
Mountains: A Diversity of Biomes 37 Heritability: Essential for Evolution 89
Concept 2.3 Review 40 Directional Selection: Adaptation by Soapberry Bugs to
Applications: Finer Scale Climatic Variation over Time New Host Plants 90
and Space 40 Concept 4.4 Review 92
vi
Contents vii
4.5 Change due to Chance 92 Water Acquisition by Plants 135

Evidence of Genetic Drift in Island Crickets 93 Water Conservation by Plants and Animals 136
Genetic Diversity and Butterfly Extinctions 95 Dissimilar Organisms with Similar Approaches to Desert
Concept 4.5 Review 96 Life 139
Applications: Evolution and Agriculture 96 Two Arthropods with Opposite Approaches to Desert
Life 140
Evolution of Herbicide Resistance in Weeds 97
6.3 Water and Salt Balance in Aquatic Environments 143
Section II Marine Fish and Invertebrates 143

Freshwater Fish and Invertebrates 144
ADAPTATIONS TO THE ENVIRONMENT Concept 6.3 Review 144
Applications: Using Stable Isotopes to Study Water Uptake by
Chapter 5 Temperature Relations 101 Plants 145
Stable Isotope Analysis 146
Concepts 101
Using Stable Isotopes to Identify Plant Water Sources 146
5.1 Microclimates 102
Altitude 103
Aspect 103
Chapter 7 Energy and Nutrient Relations 149
Vegetation 103 Concepts 149
Color of the Ground 103 7.1 Photosynthetic Autotrophs 150
Presence of Boulders and Burrows 104
The Solar-Powered Biosphere 150
Aquatic Temperatures 104
7.2 Chemosynthetic Autotrophs 154
5.2 Evolutionary Trade-Offs 105
The Principle of Allocation 105
7.3 Heterotrophs 156
Chemical Composition and Nutrient Requirements 156
5.3 Temperature and Performance of Organisms 106
Extreme Temperatures and Photosynthesis 108
7.4 Energy Limitation 163
Temperature and Microbial Activity 109
Concept 5.3 Review 110 Photon Flux and Photosynthetic Response Curves 163
Food Density and Animal Functional Response 164
5.4 Regulating Body Temperature 110
Balancing Heat Gain Against Heat Loss 110
7.5 Optimal Foraging Theory 165
Temperature Regulation by Plants 111
Temperature Regulation by Ectothermic Animals 113 Testing Optimal Foraging Theory 166
Temperature Regulation by Endothermic Animals 115 Optimal Foraging by Plants 167
Temperature Regulation by Thermogenic Plants 119 Concept 7.5 Review 168
Concept 5.4 Review 120 Applications: Bioremediation—Using the Trophic
Diversity of Bacteria to Solve Environmental
5.5 Surviving Extreme Temperatures 120
Problems 168
Inactivity 120
Leaking Underground Storage Tanks 169
Reducing Metabolic Rate 121
Cyanide and Nitrates in Mine Spoils 169
Hibernation by a Tropical Species 121
Applications: Local Extinction of a Land Snail in an Urban Chapter 8 Social Relations 172
Heat Island 123 Concepts 172
8.1 Mate Choice versus Predation 174
Chapter 6 Water Relations 127 Mate Choice and Sexual Selection in Guppies 175
Concepts 127 Concept 8.1 Review 178
6.1 Water Availability 128 8.2 Mate Choice and Resource Provisioning 178
Water Content of Air 129 Concept 8.2 Review 181
Water Movement in Aquatic Environments 130 8.3 Nonrandom Mating in a Plant Population 181
Water Movement Between Soils and Plants 131 Concept 8.3 Review 183
Concept 6.1 Review 132 8.4 Sociality 183
6.2 Water Regulation on Land 133 Cooperative Breeders 184
Water Acquisition by Animals 134 Concept 8.4 Review 189
viii Contents
8.5 Eusociality 189 10.3 Patterns of Survival 224

Eusocial Species 189 Estimating Patterns of Survival 224
Evolution of Eusociality 191 High Survival Among the Young 224
Concept 8.5 Review 193 Constant Rates of Survival 226
Applications: Behavioral Ecology and Conservation 193 High Mortality Among the Young 227
Three Types of Survivorship Curves 227
Tinbergen’s Framework 193
Environmental Enrichment and Development of
Behavior 193 10.4 Age Distribution 228
Contrasting Tree Populations 228
A Dynamic Population in a Variable Climate 229
Section III Concept 10.4 Review 230
10.5 Rates of Population Change 230
POPULATION ECOLOGY Estimating Rates for an Annual Plant 230
Estimating Rates When Generations Overlap 231
Chapter 9Population Distribution and Concept 10.5 Review 233
Abundance 196 Applications: Changes in Species Distributions in Response to
Climate Warming 233
Concepts 196
9.1 Distribution Limits 198
Kangaroo Distributions and Climate 198 Chapter 11 Population Growth 237
Distributions of Plants Along a Moisture-Temperature Concepts 237
Gradient 199
11.1 Geometric and Exponential Population Growth 238
Distributions of Barnacles Along an Intertidal Exposure
Geometric Growth 238
Gradient 200
Exponential Growth 240
Exponential Growth in Nature 241
9.2 Patterns on Small Scales 202
Scale, Distributions, and Mechanisms 202
11.2 Logistic Population Growth 242
Distributions of Tropical Bee Colonies 202
Distributions of Desert Shrubs 204
Concept 9.2 Review 205 11.3 Limits to Population Growth 245
9.3 Patterns on Large Scales 205 Environment and Birth and Death Among Darwin’s
Finches 245
Bird Populations Across North America 206
Plant Distributions Along Moisture Gradients 207
Concept 9.3 Review 208 Applications: The Human Population 248
9.4 Organism Size and Population Density 208 Distribution and Abundance 248
Population Dynamics 249
Animal Size and Population Density 208
Population Growth 250
Plant Size and Population Density 209
Applications: Rarity and Vulnerability to Extinction 210 Chapter 12 Life Histories 254
Seven Forms of Rarity and One of Abundance 210 Concepts 254
12.1 Offspring Number versus Size 255
Chapter 10 Population Dynamics 215 Egg Size and Number in Fish 256
Seed Size and Number in Plants 258
Concepts 215 Seed Size and Seedling Performance 259
10.1 Dispersal 217 Concept 12.1 Review 261
Dispersal of Expanding Populations 217 12.2 Adult Survival and Reproductive Allocation 262
Range Changes in Response to Climate Change 218 Life History Variation Among Species 262
Dispersal in Response to Changing Food Supply 219 Life History Variation within Species 264
Dispersal in Rivers and Streams 220 Concept 12.2 Review 266
Concept 10.1 Review 221 12.3 Life History Classification 266
10.2 Metapopulations 221 r and K Selection 266
A Metapopulation of an Alpine Butterfly 222 Plant Life Histories 267
Dispersal Within a Metapopulation of Lesser Kestrels 223 Opportunistic, Equilibrium, and Periodic Life
Concept 10.2 Review 224 Histories 268
Contents ix
Lifetime Reproductive Effort and Relative Offspring Size: Experimental Test of Food and Predation Impacts 306
Two Central Variables? 270 Population Cycles in Mathematical and Laboratory
Concept 12.3 Review 272 Models 307
Applications: Climate Change and Timing of Reproduction and Concept 14.2 Review 310
Migration 272 14.3 Refuges 310
Altered Plant Phenology 272 Refuges and Host Persistence in Laboratory and
Animal Phenology 273 Mathematical Models 310
Exploited Organisms and Their Wide Variety of
Section IV “Refuges” 312

INTERACTIONS 14.4 Ratio-Dependent Models of Functional Response 314
Alternative Model for Trophic Ecology 314
Chapter 13
Species Interactions and Evidence for Ratio-Dependent Predation 315
Competition 277 Concept 14.4 Review 317
14.5 Complex Interactions 317
Concepts 277
Parasites and Pathogens That Manipulate Host Behavior 317
Competitive Interactions Are Diverse 279 The Entangling of Exploitation with Competition 320
13.1 Intraspecific Competition 280 Concept 14.5 Review 321
Intraspecific Competition Among Plants 280 Applications: The Value of Pest Control by Bats: A Case
Intraspecific Competition Among Planthoppers 281 Study 321
Interference Competition Among Terrestrial
Isopods 282
Concept 13.1 Review 282 Chapter 15 Mutualism 325
13.2 Competitive Exclusion and Niches 282 Concepts 325
The Feeding Niches of Darwin’s Finches 283 15.1 Plant Mutualisms 326
Competition for Caterpillars 284 Plant Performance and Mycorrhizal Fungi 327
Concept 13.2 Review 285 Ants and Swollen Thorn Acacias 330
13.3 Mathematical and Laboratory Models 285 A Temperate Plant Protection Mutualism 334
Modeling Interspecific Competition 285 Concept 15.1 Review 335
Laboratory Models of Competition 288 15.2 Coral Mutualisms 335
Concept 13.3 Review 289 Zooxanthellae and Corals 336
13.4 Competition and Niches 289 A Coral Protection Mutualism 336
Niches and Competition Among Plants 289 Concept 15.2 Review 338
Niche Overlap and Competition Between Barnacles 290 15.3 Evolution of Mutualism 338
Competition and the Niches of Small Rodents 291 Facultative Ant-Plant Protection Mutualisms 340
Character Displacement 293 Concept 15.3 Review 341
Evidence for Competition in Nature 295 Applications: Mutualism and Humans 341
Concept 13.4 Review 295 Guiding Behavior 341
Applications: Competition Between Native and Invasive
Species 296
Section V
Chapter 14 Exploitative Interactions: COMMUNITIES AND ECOSYSTEMS 345
Predation, Herbivory,
Parasitism, and Disease 299 Chapter 16 Species Abundance
and Diversity 345
Concepts 299
Concepts 345
14.1 Exploitation and Abundance 300
A Herbivorous Stream Insect and Its Algal Food 300 16.1 Species Abundance 347
Bats, Birds, and Herbivory in a Tropical Forest 301 The Lognormal Distribution 347
A Pathogenic Parasite, a Predator, and Its Prey 303 Concept 16.1 Review 348
Concept 14.1 Review 304 16.2 Species Diversity 348
14.2 Dynamics 304 A Quantitative Index of Species Diversity 348
Cycles of Abundance in Snowshoe Hares and Their Rank-Abundance Curves 350
Predators 304 Concept 16.2 Review 351
x Contents
16.3 Environmental Complexity 351 18.2 Patterns of Aquatic Primary Production 387
Forest Complexity and Bird Species Diversity 351 Patterns and Models 387
Niches, Heterogeneity, and the Diversity of Algae and Whole-Lake Experiments on Primary Production 388
Plants 352 Global Patterns of Marine Primary Production 388
The Niches of Algae and Terrestrial Plants 353 Concept 18.2 Review 389
Complexity in Plant Environments 354 18.3 Primary Producer Diversity 390
Soil and Topographic Heterogeneity 354 Terrestrial Plant Diversity and Primary Production 390
Nutrient Enrichment Can Reduce Environmental Algal Diversity and Aquatic Primary Production 391
Complexity 355
Nitrogen Enrichment and Ectomycorrhizal Fungus
Diversity 356 18.4 Consumer Influences 392
Concept 16.3 Review 357 Piscivores, Planktivores, and Lake
Primary Production 392
16.4 Disturbance and Diversity 357
Grazing by Large Mammals and Primary Production on
The Nature and Sources of Disturbance 357 the Serengeti 394
The Intermediate Disturbance Hypothesis 357 Concept 18.4 Review 396
Disturbance and Diversity in the Intertidal Zone 358
18.5 Secondary Production 396
Disturbance and Diversity in Temperate Grasslands 358
A Trophic Dynamic View of Ecosystems 397
Top-down Versus Bottom-up Controls on Secondary
Applications: Disturbance by Humans 360 Production 397
Urban Diversity 361 Linking Primary Production and Secondary
Production 398
Chapter 17 Species Interactions and Concept 18.5 Review 399
Community Structure 365 Applications: Using Stable Isotope Analysis to Study Feeding
Habits 399
Concepts 365
Using Stable Isotopes to Identify Sources of Energy in a
17.1 Community Webs Salt Marsh 400
Strong Interactions and Food Web Structure 367
17.2 Indirect Interactions 368
Chapter 19 Nutrient Cycling and
Retention 403
Indirect Commensalism 368
Apparent Competition 369 Concepts 403
Concept 17.2 Review 370 19.1 Nutrient Cycles 404
17.3 Keystone Species 371 The Phosphorus Cycle 405
Food Web Structure and Species Diversity 371 The Nitrogen Cycle 406
Experimental Removal of Sea Stars 373 The Carbon Cycle 407
Snail Effects on Algal Diversity 374 Concept 19.1 Review 408
Fish as Keystone Species in River Food Webs 376 19.2 Rates of Decomposition 408
Concept 17.3 Review 377 Decomposition in Two Mediterranean Woodland
17.4 Mutualistic Keystones 378 Ecosystems 408
A Cleaner Fish as a Keystone Species 378 Decomposition in Two Temperate Forest
Seed Dispersal Mutualists as Keystone Species 379 Ecosystems 409
Concept 17.4 Review 379 Decomposition in Aquatic Ecosystems 411
Applications: Human Modification of Food Webs 380 Concept 19.2 Review 412
Parasitoid Wasps: Apparent Competition and Biological 19.3 Organisms and Nutrients 412
Control 380 Nutrient Cycling in Streams and Lakes 412
Animals and Nutrient Cycling in Terrestrial
Chapter 18 Primary and Secondary Ecosystems 415
Plants and the Nutrient Dynamics of Ecosystems 416
Production 383 Concept 19.3 Review 417
Concepts 383 19.4 Disturbance and Nutrients 417
18.1 Patterns of Terrestrial Primary Production 385 Disturbance and Nutrient Loss from Forests 417
Actual Evapotranspiration and Terrestrial Primary Flooding and Nutrient Export by Streams 418
Production 385 Concept 19.4 Review 419
Soil Fertility and Terrestrial Primary Production 386 Applications: Altering Aquatic and Terrestrial
Concept 18.1 Review 387 Ecosystems 419
Contents xi
Chapter 20 Succession and Stability 423 Chapter 22 Geographic Ecology 468

Concepts 423 Concepts 468
20.1 Community Changes During Succession 425 22.1 Area, Isolation, and Species Richness 470
Primary Succession at Glacier Bay 425 Island Area and Species Richness 470
Secondary Succession in Temperate Forests 427 Island Isolation and Species Richness 472
Succession in Rocky Intertidal Communities 427 Concept 22.1 Review 473
Succession in Stream Communities 428 22.2 The Equilibrium Model of Island Biogeography 473
Concept 20.1 Review 429 Species Turnover on Islands 474
20.2 Ecosystem Changes During Succession 429 Experimental Island Biogeography 475
Four Million Years of Ecosystem Change 429 Colonization of New Islands by Plants 476
Succession and Stream Ecosystem Properties 431 Manipulating Island Area 477
Concept 20.2 Review 432 Island Biogeography Update 478
20.3 Mechanisms of Succession 432 Concept 22.2 Review 478
Facilitation 433 22.3 Latitudinal Gradients in Species Richness 478
Tolerance 433 Latitudinal Gradient Hypotheses 478
Inhibition 433 Area and Latitudinal Gradients in Species Richness 480
Successional Mechanisms in the Rocky Intertidal Zone 434 Continental Area and Species Richness 481
Mechanisms in Old Field Succession 435 Concept 22.3 Review 482
Concept 20.3 Review 436 22.4 Historical and Regional Influences 482
20.4 Community and Ecosystem Stability 436 Exceptional Patterns of Diversity 482
Lessons from the Park Grass Experiment 437 Historical and Regional Explanations 483
Replicate Disturbances and Desert Stream Stability 438 Concept 22.4 Review 484
Concept 20.4 Review 440 Applications: Global Positioning Systems, Remote Sensing,
Applications: Ecological Succession Informing Ecological and Geographic Information Systems 485
Restoration 440 Global Positioning Systems 485
Applying Succession Concepts to Restoration 440 Remote Sensing 485
Geographic Information Systems 487
SECTION VI Chapter 23 Global Ecology 490

LARGE-SCALE ECOLOGY Concepts 490
The Atmospheric Envelope and the Greenhouse
Chapter 21 Landscape Ecology 445 Earth 491
Concepts 445 23.1 A Global System 493
The Historical Thread 493
21.1 Landscape Structure 447
El Niño and La Niña 494
The Structure of Six Landscapes in Ohio 447
El Niño Southern Oscillation and Marine
The Fractal Geometry of Landscapes 449
Populations 495
El Niño and the Great Salt Lake 497
21.2 Landscape Processes 450 El Niño and Terrestrial Populations in Australia 498
Landscape Structure and the Dispersal of Mammals 451 Concept 23.1 Review 499
Habitat Patch Size and Isolation and the Density of 23.2 Human Activity and the Global Nitrogen Cycle 499
Butterfly Populations 452
Habitat Corridors and Movement of Organisms 453
Landscape Position and Lake Chemistry 454 23.3 Changes in Land Cover 500
Concept 21.2 Review 455 Deforestation 500
21.3 Origins of Landscape Structure and Change 455
Geological Processes, Climate, and Landscape 23.4 Human Influence on Atmospheric Composition 504
Structure 456 Depletion and Recovery of the Ozone Layer 507
Organisms and Landscape Structure 458 Concept 23.4 Review 508
Fire and the Structure of a Mediterranean Landscape 462 Applications: Impacts of Global Climate Change 508
Concept 21.3 Review 463 Shifts in Biodiversity and Widespread Extinction of
Applications: Landscape Approaches to Mitigating Urban Heat Species 509
Islands 463 Human Impacts of Climate Change 509
xii Contents
Appendix A Investigating the Evidence

16: Estimating the Number of Species in Communities 531
1: The Scientific Method—Questions and Hypotheses 514 17: Using Confidence Intervals to Compare
2: Determining the Sample Mean 515 Populations 532
3: Determining the Sample Median 516 18: Comparing Two Populations with the t-Test 533
4: Variation in Data 517 19: Assumptions for Statistical Tests 534
5: Laboratory Experiments 518 20: Variation Around the Median 535
6: Sample Size 519 21: Comparison of Two Samples Using a Rank Sum
7: Scatter Plots and the Relationship Between Test 537
Variables 520 22: Sample Size Revisited 538
8: Estimating Heritability Using Regression 23: Discovering What’s Been Discovered 539
Analysis 521
Appendix B Statistical Tables 541
9: Clumped, Random, and Regular Distributions 522
10: Hypotheses and Statistical Significance 523 Appendix C Abbreviations Used in This Text 545
11: Frequency of Alternative Phenotypes in a Appendix D Global Biomes 547
Population 524
12: A Statistical Test for Distribution Pattern 526 Glossary 548
13: Field Experiments 527 References 558
14: Standard Error of the Mean 528
15: Confidence Intervals 530 Index 571

Preface
This book was written for students taking their first under- example of an ecological process. All attempt to engage stu-
graduate course in ecology. We have assumed that students dents and draw them into the discussion that follows.
in this one-semester course have some knowledge of basic
Concepts: The goal of this book is to build a foundation of
chemistry and mathematics and have had a course in general
ecological knowledge around key concepts, which are listed
biology, which included introductions to evolution, physiol-
at the beginning of each chapter to alert the student to
ogy, and biological diversity.
the major topics to follow and to provide a place where the
student can find a list of the important points covered in
Organization of the Book each chapter. The sections in which concepts are discussed
An evolutionary perspective forms the foundation of the focus on published studies and, wherever possible, the sci-
entire textbook, as it is needed to support understanding of entists who did the research are introduced. This case-study
major concepts. The textbook begins with a brief introduction approach supports the concepts with evidence, and intro-
to the nature and history of the discipline of ecology, followed duces students to the methods and people that have created
by section I, which includes two chapters on earth’s biomes— the discipline of ecology. Each concept discussion ends
life on land and life in water—followed by a chapter on popu- with a series of concept review questions to help students
lation genetics and natural selection. Sections II through VI test their knowledge and to reinforce key points made in
build a hierarchical perspective through the traditional sub- the discussion.
disciplines of ecology: section II concerns adaptations to
the environment; section III focuses on population ecology;
section IV presents the ecology of interactions; section V
summarizes community and ecosystem ecology; and finally,
First Pages
section VI discusses large-scale ecology, including chapters
on landscape, geographic, and global ecology. These topics
were first introduced in section I within its discussion of
SEC TIO N
II Adaptations
to the Environ
Chapter
ment
the biomes. In summary, the book begins with an overview

of the biosphere, considers portions of the whole in the
5
middle chapters, and ends with another perspective of the
entire planet in the concluding chapter. The features of this
textbook were carefully planned to enhance the students’
comprehension of the broad discipline of ecology.
Temperature
Features Designed with the Student in Relations
Mind ©Digital Vision
/Getty Images
RF
Japanese maca
ques
body heat in the , Macaca fuscata, huddle toget
All chapters are based on a distinctive learning system, fea- mids
temperature, using t of driving snow. The capa
tions, enables these
behavioral, anat
her, conserving
city to regulate
omical, and phys
their
body
iological adapta-
Japan, site of the monkeys to live through the 5.5 Many orga
turing the following key components: 1998 Winter Olym
pics.
cold winters in
Nagano, nisms surv
temperatures
stage. 120
ive extreme
by entering a
resting
Concept 5.5 Rev
CHAPTER CO iew 123
Student Learning Outcomes: Educators are being asked NCEPTS Applications:
Local Extinct
an Urban Hea ion of a Land
Snail in
increasingly to develop concrete student learning outcomes 5.1 Macroclima
landscape to
te inte racts with the
produce microcl local
Summary 124
Key Terms
t Island 123
variation in tem ima

perature. 102 tic 125
for courses across the curriculum. In response to this need Concept 5.1 Rev
iew 105
Review Questio
ns 125
5.2 Adapting
and to help focus student progress through the content, all to one
conditions gen set of environmental
a population’
erally reduce
s
sections of each chapter in the ninth edition begin with a list
s fitness in oth
environments. er
105 LEARNING OU
Concept 5.2 Rev TCOMES
iew 106 After studying this
of detailed student learning outcomes. 5.3 Most species
narrow range
perform best
in a fairly
5.1
section you shou
Distinguish betw
een temperature
ld be able to do
the
following:
of temperatu 5.2 Explain the ecol and heat.
Concept 5.3 Rev res. 107 ogical significa
iew 110 tal temperature nce of environm
en-
Introduction: The introduction to each chapter presents
s.
5.4 Many orga
T
nisms
ways to compen have evolved he thermometer
the student with the flavor of the subject and important in environmen
regulating bod
sate for variatio
tal temperatu
y temperature.
re by
ns appear in the scie
suring and repo
was one of the
ntific tool kit, and
first instruments
we have been mea
to
what do thermom rting temperatures ever sinc -
background information. Some introductions include Concept 5.4 Rev 110 e. However,
iew 120 eters actually
measure of the quantify? Tem
average kinetic perature is a
the molecules, energy, or ener
in a mass of a gy of motion, of
historical events related to the subject; others present an substance. For
example, water
101
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xiii
xiv Preface First Pages
Illustrations: A great deal of effort has been put into In the mid-190
0’s, darker pep
Chapter 7
Energy and Nu
trient Relatio
ns
the development of illustrations, both photographs moths were less
soot-covered tree
s and so had a
pered
visible to predator
s on
fitness ... whereas by the
161
advantage over
and line art. The goal has been to create more-effec- lighter moths... 1990’s, air qual
improved, and ity
lighter moths beca
better camoufl me the
aged morph...
tive pedagogical tools through skillful design and

use of color, and to rearrange the traditional presen-
tation of information in figures and captions. Much
explanatory material is located within the illustra-
tions, providing students with key information where
they need it most. The approach also provides an
ongoing tutorial on graph interpretation, a skill with
which many introductory students need practice.
1959
7%
Detailed Explanations of Mathematics: The math- 1996
ematical aspects of ecology commonly challenge Dark morph abun
dance 8%
93% Light morph abun
many students taking their first ecology course. dance
This text carefully explains all mathematical expres- 92%
sions that arise to help students overcome these Figure 7.16

... such that the
the morphs reve
relative abundan
ce of
Birds and othe rsed; light beca
challenges. In some cases, mathematical expressions extent to which
wake of environ
they are camouf
men
r predators exer
laged. When tree
t strong selectiv
s wer
e pressure on pop
ulations of pep
common than
dark.
me more
of the lighter mor tal regulations that reduced air e covered with soot due to indu pered moths in
are dissected in illustrations designed to comple- examples of natu
Bill Coster IN/A
ph. Frequency
data shown here
ral selection, alth
oug h rece
pollution, the bark
were collected
37 year
of trees became
s apa
strialization, dark
light-colored agai
Europe and else
er morphs were
more
where, based on
n, resulting in dram abundant, whereas in the
the
lamy Stock Phot nt research sugg rt in West Kirb
ment their presentation in the associated narrative. o/H Lansdown/Al ests that other y, Wirral. This atic shifts in the
amy Stock Phot evolutionary forc remains one of frequency
o/Frank Hecker/A es the mos
First Pages lamy Stock Phot
o
also likely played a
role (data from
t commonly-used
Grant et al. 199
8).
Because predat
ors must catch
often select pre and subdue the
y by size, a beh ir
selective predat avior that ecolog prey, they diff icult to find
ion. Because of ists call size- or catch. As we
significantly cor this behavior, pre size-selective pre shall see later
related with pre y size is often dation may also in chapter 7,
A visualization of a population bottle neck, using data from
solitar y predat dator size, esp Size of the predat have an energe
Chapter 10 Population Dynamics ors. One such ecially among or can affect pre tic basis.
mountain lion solitar y predat Shannon Murph y selection in oth
225 , Puma concolo or, the puma, y, Danny Lewis, er ways.
published research.
Yukon to the r, ranges from or impacts of size and Gina Wim
tip of South Am the Canadian by asking if the p studied the
changes substa erica (fig. 7.18 ralis) in salt ma diets of wolf spid
To allow comparisons to ntially along this ). Puma size rshes changed as ers (Pardosa litto
other tin Iriarte and latitudinal gra Older spiders are they aged (Murp -
studies, number of Dall sheep Subtracting number of his colleagues dient. Augus- as much as 30 tim hy et al. 2020).
deaths from number alive increase in size (1990) found spiders; howeve es larger than new
surviving and dying withi , the average size that as r, in theory sma ly hatched
n each the beginning of each year
at (fig. 7.19). Ma of their prey also pumas same diet becaus ll and large spid
e they digest the ers could have the
year of life is converted to mmals make up increases
gives the number alive at and large mamm over 90% of the have to be larger ir prey externally
number per 1,000 births. the als, especially puma’s diet, than their prey. and so don’t
beginning of the next year. nor thern par t deer, are its ma (see chapter 6) Using nitrogen
of its range in in prey in the of field-collected isotope analysis
are larger. In the North Americ mine that diet did spiders, they wer
tro a, wh ere cha nge: young spid e able to
Number of mainly on mediu pics where pumas are smalle pumas tovores, which occ ers primarily hun deter-
m and small pre r, they feed ur on the soil sur ted detri-
should differe y, especially rod the larger, older face under the tha
Age (years) survivors Number of deaths nt-sized pumas ents. Why wolf spiders wer tch, whereas
One reason is feed on differe open. However, e hunting herbivo
at beginning during year that large pre nt-sized prey? a few small spid res out in the
and may even y ma to large spiders, ers had isotope
of year injure the predat y be diff icult to subdue confirming that
there was nothin
signatures similar
or, while small that kept them g physiological
0–1 1,000 prey may be from eating the
199 her colleagues sam e herbivore diet
1–2 1,000–199 concluded that
the mechanism
. Murphy and Chapter 18 Primary and Secon
801 12 for the differen
2–3 789 801–12 13 ce
3–4 By reducing planktivorous
776 789–13 12 fish
4–5 populations, piscivores indir
ectly and introduced them to
764 30 moL22201_ch0
7_149-171.indd increase populations of large th
5–6 734
161
unmanipulated as a con
6–7 etc. 46 zooplankton and indirectly
reduce
trol
688 48 biomass of phytoplankton. The responses of the
7–8 640 69 Lake food web 02/22/21 02:57
PM tal manipulations suppor
8–9 571 t
132 (see fig. 18.14). Reducing
9–10 439 187 the
10–11 to reduced rates of primary
252 156 p
11–12 96 tivorous minnows, the pred
12–13
90 ace
6 Piscivores more numerous. Chaobor
13–14
3 us
3 3 bivorous zooplankton, and
14–15 0 th
blage shifted in dominan
Planktivorous fish ce fr
Planktivorous presence of abundant, larg
li, a mountain sheep of far
Plotting age on the x-axis e
northern and number of survivors invertebrates plankton biomass and rate
e of the classic studies of
survivor- on the y-axis creates a
of
Adding planktivorous
survivorship curve. m
production
ecological response.
Incre
Dall sheep surviving their Large herbivorous Small herbivorous population led to incr
at that age. For example, of life have a high probability
first year
zooplankton eased
the of zooplankton However, though the rese
e that died before the age surviving to about age 9. arch
ary
of planktivorous fish in this

n that generally dies duri expe
Top-down influences on prim
ng 1,000
ssumption is not likely to unintended way. Despite
be the
l that emerges probably gives a few bass remained. So,
Sheep 10 years by
the population, particula old and older are minnows, they basically
rie’s.
rly fed th
easier prey for food supply combined
Number of survivors
e survival patterns for

100 wolves and die at Large phytoplankton with
Dall a high rate. Small phytoplankton induced a strong numeric
skulls. The upper portion Survivorship curves are al re
of (see chapter 10). The man
that Murie constructed. The plotted using a log ipu
eep, the second column lists scale on the y-axis.10 tive rate of the remaining
large
class, and the third colu 10 an abundance of young
mn largemo
class. Notice that although on zooplankton.
e numbers in the table The lake ecosystem resp
are on
ividuals. This adjustment of planktivorous fish (you
is
er populations. 1 ng la
14 also shows how to tran
at the outset of the experim
s-
0 2 4 6 8 10
Nutrients ent.
bers of survivors. Plotting 12 14 Figure 18.12 The trop decreased sharply, the aver
Age (years) hic cascade hypothesis, a
“cascading” indirect inter result of age si
s against age produces the Figure 10.14 Dall sheep actions. Lines indicate both decreased, and phytopl
: from life table to survivorsh indirect (dashed) relations direct (solid) and ankto
wer portion of the figur
e.
(data from Murie 1944). ip curve hips. production increased.
s of life and death within
opulation of Dall sheep,
overall pattern of survival
y rates are higher: during and mor tality among Dall
Helps
between 9 and students
13 years.
much likeworkthat for awith
varietyand of otheinterpret
r large vertebrate quantitative
sheep is informa- Provides a visual representation of a hypothesis involving ael
The trophic cascade mod
predicts that manipulating
ing red deer, Cervus elaph s, includ-
tion, involving
he aged are higher in this
Odocoileuconverting
s hem ionu numerical
us, Colu mbia
s columbianus, East Afri
n information
black-tailed deer, into a graph. set of complex ecological interactions. pisci vore biomass will lead to
iddle years is lower. The can buffalo, Syn- changes in biomass and
cerus caffer, and humans.
The key characteristics production of planktivores,
of survival
herbivores, and phytopla
nkton.
Planktivores
Preface xv
Applications: Many students are concerned with the practical statistics and study design. They are intended to pres-
side of ecology and want to know more about how the tools of ent a broad outline of the process of science, while also
science can be applied to the environmental problems we face providing step-by-step explanations. The series of fea-
in the contemporary world. Including a discussion of appli- tures begins with an overview of the scientific method,
cations at the end of each chapter can motivate students to which establishes a conceptual context for more specific
learn more of the underlying principles of ecology. In addition, material in the next 21 features. The last reading wraps
it seems that environmental problems are now so numerous up the series with a discussion of electronic literature
and so pressing that they have erased a once easy distinction searches. Each Investigating the Evidence ends with
between general and applied ecology. one or more questions, under the heading “Critiquing
the Evidence.” This feature is intended to stimulate crit-
End-of-Chapter Material:
ical thinking about the content. Appendix B, “Statis-
• Summary The chapter summary reviews the main points tical Tables,” is available to the student as a reference
of the content. The concepts around which each chapter in support of the Investigating the Evidence features.
is organized are boldfaced and redefined in the summary Appendix C, “Abbreviations” is a handy guide to the sci-
to reemphasize the main points of the chapter. entific and other abbreviations used throughout the text,
• Key Terms including units of measurement. Appendix D is a global
• Review Questions The review questions are designed to map of the biomes.
help students think more deeply about each concept and • Glossary List of all key terms and their definitions.
to reflect on alternative views. They also provide a place • References References are an important part of any
to fill in any remaining gaps in the information presented scientific work. However, many undergraduates are dis-
and take students beyond the foundation established in tracted by a large number of references within the text.
the main body of the chapter. One of the goals of a general ecology course should be to
Note: Suggested Readings are located online. introduce these students to the primary literature without
burying them in citations. The number of citations has
End-of-Book Material:
been reduced to those necessary to support detailed dis-
• Appendixes Appendix A, “Investigating the Evidence,” cussions of particular research projects.
offers “mini-lessons” on the scientific method, emphasizing • Index
First Pages First Pages
210
Section III
Population Eco
Appendix A logy
Investigating
the Evidence
531 As in animals,
Investigating plant populatio
n
Applications
density decrease
the Evidence s with increasing
16 plant size acro
ss a wide range
of
plant growth form
Estimating the s.
Number of Sp Duckweed, Lem
ecies in Comm na,
the smallest flow one of Rarity and Vu
unities plants, lives at
ering The coastal redw
ood, Sequoia lnerability to
LEARNING OU
very high sempervirens, one
of LEARNING OU Extinction
population dens
ities. trees, lives at one the largest TCOMES
TCOMES population dens
of the lowest
After studying this
After studying this ities. section you shou
section you shou 12 colleagues (Law ld be able
ld be able
to do the following ton et al. 1998) to do the following
16.1 Explain : biologicaldiver attempted to char 9.15 Summar :
thediff iculties sity alonga distu acterize 106 Annual herbs ize and explain
thetotalnumbe involvedintryin forestofCamer rbancegradient commonness and Rabinowitz's clas
rofspeciesina gtoestimate oon,Africa,usi inthetropical rarity. sification of
16.2 Discuss com munity. birdsandbutterf ngindicatortax 9.16 Explain
waystoreduce lies,Lawtonand a.Inadditionto the relationship
m 2)
ingacompariso the effortnecessary beetles,beetlesl hiscolleaguess 104 rarity and the vuln between the cate
noftherelative formak- ivinginthefore ampledflying Clonal perennial erability of spec gories of
stcanopy,canop herbs
Plant density (per
communities. speciesrichness ants, termites, yants,leaf-litter 9.17 Describ ies to extinction.

of and soil nematod e the objectiv
from 1992 to 199
4 and spent seve
es. They sampled
these taxa 102 relate the informa es of the IUCN Red List, and
Howmanyspeci catalogingthea ral more years tion included in
esarethere?Thi pproximately2,0 sorting and categories of rari this report to the
talquestionstha sis one ofthem required approxi 00s peciescollected Trees ty.
tanecologistca ost fundamen- mately 10,000 .Thiswork
increasingthreats naskabout aco theconclusion scientist hours. 1
tobiologicaldiv mmunity.With attheendofthis Unfortunately, Viewed on a
of the most imp ersity,speciesric groupservesasa massivestudywa long-term, geo
ortant commun hnessisalsoone reliableindicato sthatnoone come and go and logical timescal
Forinstance,esti ity attributes we taxonomic grou rofspeciesrichn 10 –2 extinction seem e, populations
matesofspecies might measure. ps. Lawton and essforother tuation mark at s to be the inev
ing areas suitable richnessarecriti theirsurveyinc his colleagues the itable punc-
populations seem end of a species’ history. How
for conservation calfordetermin ludedfromone-t estimated that
environmentalcha , for - number of spec enthtoone-hund
ngeonacommun diagnosing the impact of ies in their stud
y site. Citing thei
redththetotal 10 –4 10 –2 ers. What mak
to be more vuln
erable to extincti
ever, some
habitat for rare ity,andforidentif ence, they con 1 102 es some populat on
number of spec
or threatened
species. However yingcritical
diversityofjust
clud ed that characte
rizing the full
r own experi- 104 106 108 others persist thro ions likely to disa than oth-
ies in a commun , determining the 1hectareoftro biological Plant mass (g) ugh geologic ages ppear, while
Soundestimates ity is not a simp 100,000to1mil picalforestwould Figure 9.21 ter are patterns ? At the heart
ofspeciesrichn le undertaking. lionscientistho requirefrom Plant size and of distribution of the mat-
ess formost taxa time constraints urs.Asaconseq Whi te 1985). popu lation density (dat are often vulnerab and abundance
fullydesigned,st requireacare- , ecologists will uenceofthese a from . Rare species
andardizedsamp le to extinction,
someofthebasi lingprogram.He studies of dive likely continue are seldom so. whereas abunda
cfactorsthatan rewewillreview rsity on smaller to focus their In order to und
designing such ecologistneeds
toconsiderwhen witharestricted groups of taxa
. However, even extinction, we erstand and, perh nt species
a sampling prog taxonomicfocus, need to underst aps, prevent
speciesrichnessw ram to gather info samplingacross itisimportantto figure 9.21 indi especially in this and the various
ithinandamongc rmation about studycommunit standardize cate a predicta time of rapid clim forms of rarity,
ommunities. ies. size and populat ble relationship ate change.
Sampling Effo Standardized ion density. between plant
rt Sampling Seven Forms
Thenumber ofs Standardizing sam The value of of Rarity and
pling effort and such an empiric One of Abun
peciesrecorded for plants or anim al
increaseswithh in
ighersamplingeff asampleofacommunity
essary to provide
a valid basis for
technique is gene
rally nec- als, is that it pro relationship, whether Deborah Rabinow
itz (1981) devised dance
ort.Wereviewe across commun comparing spec which we can vides a standard ness and rarity, base a classification
plifiedexample dahighlysim- ities. For exam ies richness compare measure against d on combination of common-
ofthis inInves ple, Frode Øde of expected pop d densities and geo
weconsideredho tiga tingtheEviden Nor wegian Institute gaard of the ulation densitie gives an idea grap hic range of a s of three factors:
ofspeciesrichn
wnumbers ofq
uadratsinf luen
ce6,where
standardize sam
for Nature Res
earch took grea pose you go out
into the field
s in nature. For
example, sup- habitat toleranc spec ies (ext ensive versus restr (1) the
essinthebenthi ced estimates pling as he com t care to density of som and e (broad versus icted ), (2)
Mountainstream
.In
ccommunityof
asmallRocky
plant-feedingbee
tleslivinginatr
pared species rich
ness among e species of anim measure the population tion size (large narr
versus small). Hab ow), and (3) local popula-
was required. How thatexample,arelativelysmalls cal rainforestin
Panama(Ødegaa
opicaldryforest
andinatropi-
if the densitie
s you encounter al. How would
you know rang e of conditions in whi itat tolerance is rela
ever, often far amplesize rd2006).Ødega about average were unusually ch a ted to the
example, Petri more effort is bothforestsfrom for an animal of high, low, or some plant spec spec ies can live. For
Martikainen and required. For acanopycrane ard sampled the par ies can tolerate instance,
Jari Kouki (200 areasofforest(~ thatprovidedac Without an emp
thattoverifythe
presenceorabs 3) estimated 0.8ha).Hestan cesstosimilar irical relationship ticular size and taxon? pH, and organic
matter content
a broad range
of soil texture,
ciesintheborea enceofthreatene spentsamplingea dardizedtheam figures 9.20 and such as that are confined to , whereas othe
lforestsof Finl dbeetlespe- chtreeorvine,an ountoftimehe 9.21 or a list of shown in a single soil type r plant species
400beetlespeci andrequireda techniquesinbot dheusedthesa impossible to mak species densitie broa d habitat toleranc . As we
es.They alsosug sample ofover hforests.Ødegaa mesampling e such an assessm s, it would be shall see, tigers
100,000individ gestedthat asa approximately rdalsosampledt might attempt ent. One questio range in Asia live e; however, with have
ualbeetlesmayb mpleofover the samenumbe
rofplantspeci
hebeetleson to answer with n that we s the snow leop in the tiger’s hist
est areas in Finl erequiredtoas thedryforestan es—50speciesin species is rare. a population stud row range of con ard, which is con orical
and for their suit sessjust10for- d52intherain As we shall see y is whether a fined to a nar-
tionareasforthr ability to serve capture ofvery forest.Hisefforts section, rarity in ditions in the high
eatenedbeetles
pecies.Toreduc
as conserva-
forests:35,479i
similarnumbers
ofindividual beet
resultedinthe is a more complex the following Applications Plateau. Small
geographic rang mou ntai ns of the Tibetan
effortrequiredt ethesampling ndryforestversu lesinthetwo seem at face valu consideration low population e, narrow habitat
oestimatespecie s30,352inrainf e. than it might density are attr tolerance, and
gistsandconserv srichness,comm his collections orest.However, ibutes of rarity.
ationistsoften unityecolo- in rain forest incl As shown in figu
thatarereliable focusongroups thanindryforest: uded 37% mor re 9.22, there
indicatorsofsp oforganisms 1,603speciesin e beetle species Concept 9.4 Re nations of thes are eigh
eciesrichness. forest.BecauseØ rainforestversus view e factors, seven t possible combi-
Indicator Tax degaard tookcar 1,165indry attribute of rari of which include
a can conclude that etostandardizes ty. The most abu at
Becauseof the wasprobablyhi
the species richness ampling,we 1. What are som threatened by ndant species and least one
greatcostandt gherathisrainfo of plant-feeding e advantages of extinction hav those least
toriesofspecies imeof making effo rest beetles herbivorous mam Damuth’s stric broad habitat e extensive geo
diversity,ecolog thoroughinven- rtsw ereuneven,weco stud ysite.Ifhissam mals in his ana t focus on tolerances, and graphic ranges,
etyoftaxaasind istshavepropose uldnotreachsu pling between body size lysis som ewh ere within thei larg e loca l pop
icatorsofoveral dawidevari- chaconclusion. of the relationship r range. Some ulations at leas
taxa have gene lbiologicaldiver CRITIQUING THE
EVIDENCE 16 2. How might and pop ulat ion density (see dan delions, Norway of these species, t
rally been well sity.Indicator energy and nutr fig. 9.19)? rats, and house such as
oforganismssuc -known and con 1. Acompletel the lower pop ient relations with humans and sparrows, are asso
hasbirdsandbu spicuous groups istofspecieshas ulation densitie explain are considered ciated
thatindicatortax tterf lies.Howev areaofsubstanti notbeendeterm comparable-size s of birds com of small mammal pests. However
ane er,itappears alhabitatanyw inedforany pared to s, birds, and inve , many species
JohnLawton ofI edtobechosenwithcaution.Fo 2. Why do
most surveys hereonearth.W d mammals (see
fig. 9.20)? humans, such rtebrates not asso
mperial College rexample, of species dive hynot? as the deer mou ciated with
in the United
Kingdomand restrictedgroup rsity focus on the marine zoo se, Peromyscus
sofrelativelyw plan maniculatus, or
asplants,mamm
als,andbutterf
ell-knownorgani
smssuch this most common kton, Calanus finmarchicus,
lies? category. also fall into
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Confirming Pages
xvi Preface
New to the Ninth Edition request. Concepts hydrophilic and hydrophobic introduced. Water
isotope section re-written to clarify per reviewer request, including
Nearly every chapter has significant changes in this edition. a new figure to explain. Applications section was re-written with
To update content and respond to reviewers’ comments, we updated example from the meta-analysis by Evaristo and McDon-
have incorporated the research and ideas of over 140 new cita- nell (2017). Questions and one figure were updated.
tions, the majority of which (73%) were authored by under- Chapter 7 Information about chemosynthesis was
represented scientists. A particular effort was made to cite expanded and updated with example from Naples, Italy. Pep-
cutting-edge ecological research by women of color. With pered moth example re-written and figure replaced with one
each edition, we continue toward the goal of making this text that shows actual photographs and data. Old predation exam-
reflect the true diversity of researchers in the field. ples were replaced with those using wolf spiders and coral reef
There are over 100 updated examples in this edition, with fishes research from teams led by women, including new fig-
42 new figures, plus improvements or updates to 20 existing ures. Questions were updated.
figures. Dozens of new questions have been written to corre- Chapter 8 Opening photo replaced with a more appropri-
spond to the new material and, in response to reviewers, many ate one, six references updated. Section on nonrandom mat-
other questions have been re-written to focus more on con- ing in plants significantly updated and clarified. Paragraph
cepts rather than specific examples. Several new terms have on phylogenies based on genetic analysis added. Updated
also been added in the text and glossary to increase student number of cooperative breeding species. Updated section on
understanding and to reflect the evolving nature of the field. lion cooperation with research by Natalia Borrego. Questions
We have also continued to expand connections with evolution were updated.
and global change in this edition. Chapter 9 New example of gorillas replaces an old exam-
ple, and new paragraph added based on the 2020 Living Planet
Significant Chapter-by-Chapter Changes Report. One new image. More information about the Breeding
Chapter 1 In response to reviewer’s comments, we have cre- Bird Survey with updated references. Term endemic added, with
ated a new section that describes the different tools used by paragraph replaced with new example of bird from Hawaii. Figure
ecologists, introducing five new terms including ex situ and on rarity and vulnerability to extinction significantly improved in
in situ. There are a total of nine new figures. We have revised response to reviewer request. Questions were updated.
figure 1 and added microbial ecology as an important frontier. Chapter 10 Seven citations updated. Research on “killer”
We have added new examples from recent literature, including bees updated with genetics research led by a man of color,
about evolution in alpine chipmunks. Questions were updated. including updated figure. Added information and example of
Chapter 2 Three new figures were added, including from pumas in Patagonia to migration section. Questions updated.
research on habitat conversion in India. Data on tropical forest Chapter 11 Introduction re-written with example from
loss was updated. New examples from publications by women the COVID-19 pandemic, including new figures. Three fig-
of color on soils and on logging of boreal forests were added. ures updated, including one for current numbers of whooping
Wording in several places was clarified in response to reviewers’ cranes and another with human population growth. Questions
comments. An explanation of the distinction between weather were updated.
and climate change was added. Improvements were made to 10 Chapter 12 Paragraph replaced with section on life his-
figures, including updating the drought data in figure 2.41 to tory trade-offs, based on ideas by Anurag Agrawal. Updated
2020 and relating it to fires. Questions were updated. number of species of fish with 2020 data from IUCN. One
Chapter 3 Six citations were updated. Sections added on figure improved. Questions were updated.
United Nations Decade of Ocean Science, microplastics from Chapter 13 Self-thinning section updated with research
research by Chatterjee and Sharma (2019), and updated several from people of color, and new figure added to better explain
examples. One figure was updated with current global ice levels. zero growth isocline, per reviewer request. Existing Lotka-Vol-
Chapter 4 The “applications” section was re-written with terra figure simplified. Competition meta-analysis research
an updated example of herbicide resistance by Sushila Chaud- by Jessica Gurevitch and colleagues added. Extra example of
hari and her colleagues, including a new figure. Questions competition deleted, per reviewer request. Questions updated.
were updated, and an existing figure improved. Chapter 14 Section on research by Utida shortened and
Chapter 5 A total of 13 new citations, including examples simplified per reviewer request, including an improvement to
with current citations were provided of how global warming an existing figure. Questions were re-written to focus on con-
is affecting ecosystems. New example and figure created to cepts rather than specific research. Two citations updated.
describe relationship between water temperature and canopy Chapter 16 The concept of a species rarefaction curve is
cover. Research on endothermic fish updated, with a new introduced. A new example of sampling benthic macroinverte-
figure created and concept of RM endothermy added. Old brates replaces an old example, work done by a man of color
example replaced with new section on comparisons between that also introduces the concept of DNA barcoding, including
endothermic and ectothermic fish with research by a man of new figures. Questions were updated.
color, including another new figure. Questions were updated. Chapter 17 Four examples were updated, all from
Chapter 6 Section on water-harvesting re-written with updated research led by underrepresented scientists. This includes a
information and a new figure adapted from the review by Guera new “Applications” example on hyperparasitoids with a new
and Bhushan (2020). Added concept of cohesion, per reviewer figure. Questions were updated.
moL22201_fm_i-xx.indd xvi 12/08/21 09:37 AM

Preface xvii
Chapter 18 Section on primary productivity of oceans was completed this edition. I am also deeply grateful for pedagogi-
re-written with updated environmental factors and relating this cal expertise of Julie Morris, who also went above and beyond.
to global change. Map on marine primary productivity has Thanks also go to the other members of the Sher Lab who
been updated. Research on top-down vs. bottom-up updated pitched in with research and/or offered feedback on new fig-
with a new section and figure from meta-analysis research ures for the ninth edition, including Ali Alghamdi, Violet But-
conducted by Mayra Vidal, a woman of color, and Shannon ler, Rhys Daniels, Alex Goetz, Annie Henry, Alex Kim, Lily
Murphy. Concept of tri-trophic interactions added. Eleven Malone, Mandy Malone, and Allen Williamson. During the
citations were updated, most of which from papers with under- development of this textbook, many colleagues freely shared
represented lead authors. Paragraph on role of microorgan- their expertise, reviewed sections, or offered the encour-
isms added, per reviewer request. Questions were updated. agement a project like this needs to keep it going: Anurag
Chapter 20 All sections on succession at Glacier Bay sec- Agrawal, Brian Buma, Candice Galen, Diane Marshall, Scott
tion completely re-written to reflect more current research led Nichols, Mayra Vidal, and Dhaval Vyas. I am grateful to Pat-
by Brian Buma that changes interpretation of those research, rick M. Burchfield and Hector Chenge Alvarez for keeping me
including new figures. This case study becomes a more inter- up to date in data and photos of turtles. Special thanks to Jake
esting story about how understanding can evolve with new Grossman for sharing his list of Ecologists of Color and Indig-
information. Questions were updated. enous Ecologists.
Chapter 21 Reference to the 2020 California wildfires We would like to especially thank Shannon Murphy for
was added, including a short paragraph about research from her extensive suggestions for the ninth edition, as well as for
UC Berkeley. Questions were updated. providing us with exciting new case studies to illustrate evolu-
Chapter 23 A total of five new figures added, including one tionary ecology concepts. In addition, we are indebted to the
that refers to the Australian wildfires of 2020. Figure on atmos- many students and instructors who have helped by contacting
pheric CO2 updated with current values. Section on nitrogen us with questions and suggestions for improvements.
pollution re-written with more explanation and more current We also wish to acknowledge the skillful guidance and
research. The forest section was re-written with forest biodiver- work throughout the publishing process given by many profes-
sity data from the FAO 2020 report on the State of the World’s sionals associated with McGraw-Hill Education and Straive
Forests and other current research. Corrections made to use of during this project, including Beth Baugh, Melissa Homer,
Spanish words, per reviewer request. Deforestation in Brazil was Jodi Rhomberg, and Mithun Kothandath.
updated. There were a total of 13 new citations, 9 from under- We gratefully acknowledge the many reviewers who, over
represented scientists. Questions were updated. the course of the many revisions, have given of their time and
expertise to help this textbook evolve to its present ninth edi-
tion. Note that some feedback that did not make it into this
Online Materials
edition will be incorporated into the next one. These review-
Available online are suggested readings and answers to concept ers continue our education, for which we are grateful, and we
review, chapter review, and critiquing the evidence questions. honestly could not have continued the improvement of this
textbook without them.
Finally, I would like to thank my co-author Manuel
Related Title of Interest from McGraw-
Molles for entrusting me with this wonderful series, as well as
Hill Education my wife Fran and our son Jeremy for their support throughout
Ecology Laboratory Manual, by Vodopich the production of the ninth edition.
(ISBN: 978-0-07-338318-7;
Reviewers for the Ninth Edition
MHID: 0-07-338318-X)
Darrell Vodopich, coauthor of Biology Laboratory Manual, Thomas C. Adam University of California, Santa Barbara
has written a new lab manual for ecology. This lab manual Henry D. Adams Oklahoma State University
offers straightforward procedures that are doable in a broad Matthew R. Helmus Temple University
Jodee Hunt Grand Valley State University
range of classroom, lab, and field situations. The procedures
Jamie Lamit Syracuse University, and State University of New York
have specific instructions that can be taught by a teaching Joseph R. Milanovich Loyola University Chicago
assistant with minimal experience as well as by a professor. Shannon Murphy University of Denver
Wiline Pangle Central Michigan University
Acknowledgments Jessica Peebles-Spencer Ball State University
Jonathan Shurin University of California San Diego
First and foremost, I must thank my academic partner John F. Weishampel University of Central Florida
Dr. Eduardo González, without whose help I could have never Lan Xu South Dakota State University

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xx Preface
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Chapter
Introduction
to Ecology
1
Historical Foundations
NPS Photo by Jeff Foott
and Developing Frontiers
A yellow-rumped warbler, Dendroica coronata, feeding young. Ecological
studies of warblers have made fundamental contributions LEARNING OUTCOME
to the growth of ecological understanding. After studying this section you should be able to do the following:
1.1 Discuss the concept of environment as it pertains to

the science of ecology.
CHAPTER CONCEPTS
W
hat is ecology? Ecology is the study of relation-
ships among organisms and between organisms
1.1 Ecologists study environmental and the physical environment. These relationships
relationships ranging from those influence many aspects of the natural world, including the dis-
of individual organisms to factors tribution and abundance of organisms, the variety of species
influencing global-scale processes. 2 living together in a place, and the transformation and flow of

Concept 1.1 Review 3 energy in nature.
Humans are rapidly changing earth’s environment, yet we
1.2 Ecologists design their studies based on do not fully understand the consequences of these changes.
their research questions, the temporal For instance, human activity has increased the quantity of
and spatial scale of their studies, and nitrogen cycling through land and water, changed land cover
available research tools. 3 across the globe, and increased the atmospheric concentra-

Concept 1.2 Review 9 tion of CO2. Changes such as these threaten the diversity of
life on earth and will also endanger our life support system.
Applications: Because of the rapid pace of environmental change in the early
Ecology Can Inform Environmental twenty-first century, it is imperative that we better understand
Law and Policy 9 earth’s ecology.
Summary 10 Behind the simple definition of ecology lies a broad
Key Terms 10 scientific discipline. Ecologists may study individual organ-
isms, entire forests or lakes, or even the whole earth. The
Review Questions 10
measurements made by ecologists include counts of individ-
ual organisms, rates of reproduction, and rates of processes
1
2 Chapter 1 Introduction to Ecology
such as photosynthesis and decomposition. Ecologists often Individuals

spend as much time studying nonbiological components
How do zebras regulate their
of the environment, such as temperature and soil chem- internal water balance?
A
istry, as they spend studying organisms. Meanwhile, the
“environment” of organisms in some ecological studies is
other species. While you may think of ecologists as typically
studying in the field, some of the most important conceptual
advances have come from ecologists who build theoretical Population
models or do ecological research in the laboratory. Clearly, What factors control zebra
our simple definition of ecology does not communicate the B populations?
great breadth of the discipline or the diversity of its practi-
tioners. To get a better idea of what ecology is, let’s briefly
review its scope.
Community
How do species interact?

1.1 Overview of Ecology How do these interactions
C affect coexistence?
LEARNING OUTCOMES
After studying this section you should be able to do the following:
1.2 Describe the levels of ecological organization, for

Ecosystem
example, population, studied by ecologists.
1.3 Distinguish between the types of questions How does fire affect nutrient
addressed by ecologists working at different levels of availability in grassland
D ecosystems?
organization.
1.4 Explain how knowledge of one level of ecological
organization can help guide research at another
level of organization. Landscape
How do vegetated corridors

Ecologists study environmental relationships ranging from E
affect the rate of movement
by mammals among isolated
those of individual organisms to factors influencing global-scale forest fragments?
processes. This broad range of subjects can be organized by
arranging them as levels in a hierarchy of ecological organiza-
tion, such as that embedded in the brief table of contents and Biosphere
the sections of this book. Figure 1.1 attempts to display such a
hierarchy graphically. What role does concentration
of atmospheric CO2 play in
Historically, the ecology of individuals, (fig. 1.1A), has F the regulation of global
been the domain of physiological ecology and behavioral temperature?
ecology. Physiological ecologists have emphasized the evolu-
tion (a process by which populations change over time) of
physiological and anatomical mechanisms by which organ- Figure 1.1 Levels of ecological organization and examples of the
isms adapt to challenges posed by physical and chemical kinds of questions asked by ecologists working at each level. These eco-
variation in the environment. Meanwhile, behavioral ecolo- logical levels correspond broadly to the sections of this book.
(A) Glow Images; (B) cinoby/E+/Getty Images; (C) Mogens Trolle/Shutterstock;
gists have focused principally on evolution of behaviors that (D) Photo by Gary Wilson, USDA Natural Resources Conservation Service;
allow animals to survive and reproduce in the face of environ- (E) Comstock/PunchStock; (F) Calysta Images/Getty Images
mental variation.
There is a strong conceptual linkage between ecological
studies of individuals and of populations particularly where are influenced by nonbiological and biological aspects of
they concern evolutionary processes. Population ecology the environment.
is centered on the factors influencing population structure Bringing biological components of the environment into
and process, where a population is a group of interbreed- the picture takes us to the next level of organization, the com-
ing individuals of a single species inhabiting a defined area munity (fig. 1.1C). A community is an association of interact-
(f ig. 1.1B). The processes studied by population ecologists ing species. Ecologists who study interactions between species
include adaptation, extinction, the distribution and abun- have often emphasized the evolutionary effects of the interaction
dance of species, population growth and regulation, and on the species involved. Other approaches explore the effect of
variation in the reproductive ecology of species. Population interactions on population structure or on properties of ecologi-
ecologists are particularly interested in how these processes cal communities.
Chapter 1 Introduction to Ecology 3
The next level of organization is the ecosystem. An eco-

system is a biological community together with its associ-
ated physical and chemical environment. Community and
ecosystem ecology have a great deal in common, since both
are focused on multispecies systems. However, the objects of
their study differ. While community ecologists concentrate
on understanding environmental influences on the kinds and
diversity of organisms inhabiting an area, ecosystem ecologists
focus on ecological processes such as energy flow and decom-
position (f ig. 1.1D).
To simplify their studies, ecologists have long attempted
to identify and study isolated communities and ecosystems.
However, all communities and ecosystems on earth are sub-
ject to exchanges of materials, energy, and organisms with
other communities and ecosystems. The study of these
exchanges, especially among ecosystems, is the intellectual
territory of landscape ecology (f ig. 1.1E). Landscape ecology
in turn leads us to the largest spatial scale and highest level
(a)
of ecological organization—the biosphere, the portions of the
earth that support life, including the land, waters, and atmo-
sphere (f ig. 1.1F).
While this description of ecology provides a brief preview
of the material covered in this book, it is a rough sketch and
highly abstract. To move beyond the abstraction represented
by figure 1.1, we need to connect it to the work of the scien-
tists who have created the discipline of ecology. To do so, let’s
briefly review the research of ecologists working at a broad
range of ecological levels emphasizing links between historical
foundations and some developing frontiers (fig. 1.2).
Concept 1.1 Review (b)
1. How does the level of ecological organization an ecolo- Figure 1.2 Two rapidly developing frontiers in ecology.
(a) Microbial ecology: the study of the interactions among microorgan-
gist studies influence the questions he or she poses? isms and between them and their environment (e.g., Epps and Arnold
2. While an ecologist may focus on a particular level of 2019). The importance of these organisms for regulating systems
ecological organization shown in figure 1.1, might other from populations to ecosystems is becoming increasingly understood.
levels of organization also be relevant? For example, why (b) Urban ecology: the study of urban areas as complex, dynamic
should an ecologist studying factors limiting numbers in ecological systems, influenced by interconnected, biological, phys-
ical, and social components. As ecologists focus their research on the
a population of zebras consider the influences of inter- environment where most members of our species live, they have made
actions with other species or the influences of global unexpected discoveries about the ecology of urban centers such as the city
processes such as climate change? of Baltimore (see chapter 19). (a) STEVE GSCHMEISSNER/Science Photo
Library/Getty Images; (b) Jon Bilous/Shutterstock
1.2 Sampling Ecological Research Ecologists design their studies based on their research questions,
the temporal and spatial scale of their studies, and available
LEARNING OUTCOMES
research tools. Because the discipline is so broad, ecologi-
After studying this section you should be able to do the following: cal research can draw from all the physical and biological sci-
1.5 List three categories of ecological research, and give ences. The following section of this chapter provides a sample
an example of each. of ecological questions and approaches to research.
1.6 Explain how new tools and technology can be used to
Types of Research
advance each category of ecological research.
1.7 Describe different types of ecological models and In the broadest sense, we can consider ecological research in
explain how are they used. three general categories: observation, experimentation, and
modeling. Each of these types of research is necessary for
understanding the organisms and processes at work in our world; forest; by observing the birds in their natural habitat, he
most ecologists use at least two, if not all three approaches to determined that each warbler species had a distinct feeding
answer ecological questions. zone (fig. 1.3). He concluded that this partitioning of the tree
reduced competition among the warblers, stimulating future
Observation generations of studies of competition.
Observation refers to the collection of data in unmanipu- Another classic example of observational research is the
lated settings, such as counting numbers of birds in a patch work by Nalini Nadkarni (1981, 1984a, 1984b), who changed
of forest or describing types of fungal spores seen through our ideas of how tropical and temperate rain forests are struc-
a microscope. Some of this work takes place in the field, or tured and how they function. Nadkarni was one of the first
in situ, meaning in the habitat where the organisms live, while scientists to study the ecology of the unseen world of the for-
other research uses specimens that have been collected in the est canopy (fig. 1.4). Using mountain-climbing equipment,
field but are observed in a laboratory or other setting. Such she took inventories of the distribution of nutrients in rain
specimens may have been sampled for that specific purpose or forests in both Costa Rica and the Pacific Northwest of the
may have been collected long beforehand and stored for future United States. She discovered through this sampling that as
study, such as those found in an herbarium or museum. For much as four times the nutrient content in trees leaves was
example, many researchers have been able to track the impact found in epiphytes—plants such as orchids, ferns, and mosses
of climate change on plant communities using thousands of that grow on the tree trunks and branches.
plant specimens collected over more than a century (Jones MacArthur’s primary research tool was a pair of binoculars
and Daehler 2018, Piao et al. 2019). Observational research and Nadkarni’s was ropes and harness; however, today there are
can be purely descriptive or may test a hypothesis, such as to many more means by which we can collect observational data.
understand relationships between organisms. This may involve For example, new ways to access the forest canopy range from
comparing observations over time or space. hot air balloons and large cranes (see Investigating the Evidence
An important, historical example of ecological observa- 16 in Appendix A) to unmanned aerial vehicles (UAV) such as
tional research is that by Robert MacArthur, who observed drones. These can carry cameras and other equipment to collect
warblers in spruce forests of northeastern North America data (Waite et al. 2018). Thermal sensors on UAV’s have been
(MacArthur 1958). Theory had predicted that two species used to survey animals in the treetops at night (Kays et al. 2019).
with identical ecological requirements would compete with Another example of a new type of data being collected
each other and that, as a consequence, they would not live is stable isotope analysis (see chapter 6). Isotopes of a chemi-
in the same environment indefinitely. MacArthur wanted to cal element, such as isotopes of carbon, have different atomic
understand how several warbler species with apparently simi- masses as a result of having different numbers of neutrons.
lar ecological requirements could live together in the same Water and nutrients from different sources can have different
Cape May warbler Blackburnian warbler Black-throated

green warbler
New needles and New needles and buds New needles and buds
buds at top of tree of upper branches and some older needles
Bay-breasted Yellow-rumped
warbler warbler
Old needles and bare Bare or lichen-covered lower

and lichen-covered trunk and middle branches
middle branches
Figure 1.3 Warbler feeding zones shown in beige. The several warbler species that coexist in the forests of northeastern North America feed in
distinctive zones within forest trees.
(a)
Figure 1.4 Exploring the rain forest canopy. What Nalini Nadkarni
discovered helped solve an ecological puzzle. Courtesy Nalini Nadkarni,
photo by Dennis Paulson
isotopic signatures, thus allowing us to trace them through eco-

systems. In this way, stable isotope analysis provides ecologists
with a new type of “lens” capable of revealing ecological rela-
tionships that would otherwise remain invisible. Melissa Whita-
ker and colleagues used both stable isotope and genetic analysis
to identify trophic relationships for the Anthene usamba, a
butterfly whose larval stage is found on whistling thorn aca-
cia trees (Vachellia drepanolobium) (Whitaker et al. 2019). For
this species, like many small organisms, there were no direct
observations of feeding behavior in the field. However, because (b)
they were found on acacia trees that have a mutualism with
Figure 1.5 (a) Observational research in Kenya, East Africa, by
ants, it was hypothesized that the butterfly larvae might feed Melissa Whittaker and colleagues on the African lycaenid b utterfly,
upon regurgitations from ants or even eat ants themselves. By Anthene usamba. Since the larvae of the butterfly were found in
analyzing the DNA and nitrogen isotope signatures of the gut acacia tree galls with ants (b), it had been hypothesized that it ate ant
contents of butterfly larvae at field sites on Suyian Ranch in excretions. However, isotope research found that this was incorrect;
because acacia leaves have a particular nitrogen isotope signature,
Laikipia County, Kenya (f ig. 1.5a). Whitaker’s group deter-
researchers were able to identify it in the gut of the butterfly’s larvae.
mined that these A. usamba larvae were, in fact, herbivorous, (a) Julianne Pelaez; (b) Dino Martins
feeding almost exclusively off of the acacia tree itself (f ig. 1.5b).
All of these examples are considered observational studies
because there was no manipulation of variables in the field; nature, it is limited because observation cannot be used to
data were collected on organisms as they existed in their natu- definitively exclude a possible phenomenon (Tilman 1989).
ral environment. That is, just because we have never observed something (like
a butterfly eating an ant) is not enough to say it is theoreti-
Experimentation cally impossible. This is why experiments may also be neces-
While observational research is critically important for sary. Experimentation typically refers to research that involves
the field of ecology because it allows us to define patterns in manipulation of variables of interest while holding others
constant in order to test a hypothesis. In her book co-authored

with Elizabeth Lunbeck about scientific observation, Lorraine
Daston described the relationship between observational
research and experiments thus:
“Observation, by the curiosity it inspires and the gaps

that it leaves, leads to experiment; experiment returns
to observation by the same curiosity that seeks to fill
and close the gaps still more; thus one can regard experi-
ment and observation as in some fashion the conse-
quence and complement of one another” (Datson 2011)
Experiments can occur in the field or in a more controlled

setting, such as in a lab, garden, greenhouse, or outdoor enclo-
sures for animals. Ecological experimentation in the field can
be difficult to interpret because of the number of factors that
may influence the data being collected, including factors
that may be unknown to the researcher. In contrast, ex situ (not (a)
in the natural environment) experimentation has the benefit of
being able to exclude all but the factors of interest in the exper- 1.0
iment; however, it has the limitation of not representing real
life. Both ex situ and in situ observation and experimentation
Relative fitness (Seed set)
0.8
may be necessary to fully understand the ecology of an organ-
ism or system.
In a classic example of experimentation, Candace Galen 0.6
tested the importance of bees for the evolution of the alpine .23
x +0
wildflower Polemonium viscosum growing on Pennsylvania 0.08
0.4 y=
Mountain, Colorado (f ig. 1.6a). Observational research had
suggested that bumblebees prefer larger flowers, and also
0.2
that more visitation by bees meant more pollen transferred
and thus more seeds produced. More seeds means greater fit-
ness. But it was also possible that some other factor such as 0
nutrients produced both big flowers and lots of seeds in some –4 –3 –2 –1 0 1 2 3
plants. Were the bees important for the evolution of flower Flower size
size? Galen was one of the first to experimentally test the (b)
hypothesis that larger flowers would have more seeds specifi- Figure 1.6 Experimental research on Polemonium viscosum, an
cally because they were more attractive to bees (Galen 1989). alpine flower in Colorado, USA (a), demonstrated that increasing flower
To do this, she compared seed set in plants that were exposed size was positively associated with fitness, as measured by seed set, in
to pollination by bumblebees (the treatment) versus plants flowers pollinated by bumblebees. (b) We know that bumblebees are
that were hand-pollinated and bagged to exclude bees (the driving this relationship because no such relationship was found for
f lowers that were hand-pollinated (the control in the experiment)
control). Both groups included plants with a range of flower (the graph is adapted from Galen 1989). (a) Candace Galen
sizes. She found that seed number significantly increased with
flower size when pollinated by bees (fig. 1.6b), but that flower
size did not predict seed number in the control group. That
is, her experiment showed that not only did bees prefer larger compare behavior during darkness (the treatment created by
flowers, but also that the bees were influencing the evolution the eclipse) to daylight (the control) within a single hour by
of larger flowers because they caused those plants to have detecting bee movements using their sound. In order to obtain
greater fitness. a very large sample over a wide geographic area, they used
As with observational research, new tools and technology another new resource: data collected by non-scientists. School
have also advanced ecological experimentation. More than 30 children and other “citizen scientists” assisted in collecting
years after her groundbreaking work on bumblebee pollination, recordings of bees’ buzzing at 11 locations in 3 regions using
Galen is still researching the ecology and evolution of these tiny USB microphones dispersed among flowers (f ig. 1.7a).
systems, most recently using a newly developed tool for analyz- Recordings of the bees buzzing were then digitally analyzed
ing audio recordings of bees’ wing movements. Galen and her to document the dramatic decrease in bee movement during
colleagues took advantage of a natural experiment created by the eclipse (f ig. 1.7b). Statistical analysis was used to deter-
a solar eclipse to investigate the impact of light and tempera- mine that this decrease was due primarily to light, rather than
ture on bees’ behavior (Galen et al. 2019). They were able to temperature.
system works. Conceptual models are those which describe sys-

tems in pictures or diagrams, whereas quantitative models are
mathematical and may involve complex equations. Like obser-
vational research, models can be purely descriptive can be
designed to test a hypothesis. A line, which can be represented
by the equation y = mx + b, is an example of a very simple
quantitative model that has many applications within ecology.
In Galen’s early work, a line could be used to represent the
relationship between flower size and fitness (f ig. 1.6b). The
line does not explain all of the variability in fitness, but it
does generally define the rate at which seed set increases with
increasing flower size. In this way, a model can be a tool used
by ecologists to understand the data they have collected. In
other cases, modeling may describe or predict patterns using
data from published work by others. In chapter 6, we will learn
about a modeling approach that is used to summarize findings
from many different studies at once.
Models may also be used to represent a novel hypoth-
esis that can then be tested with or compared against future
observational or experimental data. In chapter 22, we will learn
about McArthur and Wilson’s Island Biogeography model; this
is a conceptual model based on observed data to explain how
size of islands and their proximity to the mainland affect spe-
cies diversity. This model has been applied to many species and
ecosystems, and it has even helped us understand the dynamics
of protected areas as land-islands (Sher and Primack 2018).
Finally, models can be used to simulate natural systems,
allowing us to test scenarios that would be too difficult, expen-
(a)
sive, or logistically impossible to do in real life. For example,
15 Annette Janssen and others created a computer program that
can be used to predict growth of algae in lake ecosystems,
based on different climate models (Janssen et al. 2019). Jans-
sen’s model includes multiple ecological feedbacks that affect
Buzz density (buzzes min–1)
10
algal growth in deep lakes, making it possible to quantify how
the lake will respond to different nutrient levels under climate
warming. The mathematical equations involved in creating
the simulations can be represented by a conceptual model
(fig. 1.8). Although the Janssen et al. model may seem com-
5
plex, many mathematical simulations of this type are even more
so in their attempt to represent the real world. Such simulations
are important for understanding what has both happened in the
past as well as what may happen in the future.
0
None Low High

(b) Relative Light Intensity During Eclipse Climatic and Ecological Change:
Past and Future
Figure 1.7 (a) Emilia Asante, a graduate student at the University
of Missouri, assembles a microphone to record the buzzing of bees. The earth and its life are always changing. However, many
The white, furry “jacket” screens out wind noise. (b) Recordings of the of the most important changes occur over such long periods
bees buzzing documented a dramatic decrease in bee activity as mea- of time or at such large spatial scales that they are difficult
sured by buzz density during the eclipse. (a) Candice King; (b) adapted from to study. Two approaches that provide insights into long-term
Galen et al. 2019
and large-scale processes are studies of pollen preserved in lake
sediments and of evolutionary change.
Modeling Margaret B. Davis (1983, 1989) carefully searched
Modeling is the creation and analysis of representations of through a sample of lake sediments for pollen. The sediments
data or ideas to provide insight or make predictions. In ecol- had come from a lake in the Appalachian Mountains, and the
ogy, models usually represent a hypothesis regarding how a pollen they contained would help her document changes in
AIR Fish-eating Plant-

birds and humans eating birds
Reaeration
Evaporation
WATER Grazing
O2 Production
Marsh
Inflow Oxygen Zooplankton-eating fish
demand Predatory fish
Outflow Algea-eating fish
Zooplankton
Load Plant-eating fish
Water plants
Grazing
Inorganic
matter Mort. Uptake Uptake
Detritus Algae P, N, Si
Predation
Mineralisation
Diffusion
Exchange Uptake
Algae P, N, Si
Mort.
Organic Mineralisation Denitr.
Zoobenthos Mort.
matter
Feeding
Inorg.
matter Sorption
SEDIMENT
Seepage
Figure 1.8 Ecological modeling research includes creating complex mathematical simulations of systems as a way of making predictions. Arrows
indicate movement of energy or matter. This conceptual model or schematic of a lake ecosystem represents only a portion of a larger, a process-based
mathematical model that was created to predict algal blooms under different environmental conditions (diagram based on Janssen et al. 2019).
the community of plants living near the lake during the past Shaw, and Etterson 2005). As climate changes, plant popula-
several thousand years. Davis is a paleoecologist trained to tions simultaneously change their geographic distributions
think at very large spatial scales and over very long periods of and undergo the evolutionary process of adaptation, which
time. She has spent much of her professional career studying increases their ability to live in the new climatic regime. Mean-
changes in the distributions of plants during the Quaternary while, evidence of evolutionary responses to climate change
period, particularly during the most recent 20,000 years. has been found in many animal groups. One such example is
Some of the pollen produced by plants that live near a evidence of rapid evolution in alpine chipmunks (Tamias alpi-
lake falls on the lake surface, sinks, and becomes trapped in nus) for a gene associated with high elevation stress (fig 1.10).
lake sediments. As lake sediments build up over the centuries, This was discovered by researchers at the Museum of Verte-
this pollen is preserved and forms a historical record of the brate Zoology at the University of California using DNA from
kinds of plants that lived nearby. As the lakeside vegetation historic specimens (Bi et al. 2019). Evolution of adaptations
changes, the mix of pollen preserved in the lake’s sediments to elevation have been shown in other rodents using field col-
also changes. In the example shown in f igure 1.9, pollen from lected animals (Velotta et al. 2020).
spruce trees, Picea spp., first appears in lake sediments about In the remainder of this book, we will fill in the details
12,000 years ago; then pollen from beech, Fagus grandifolia, of the sketch of ecology presented in this chapter. This brief
occurs in the sediments beginning about 8,000 years ago. survey has only hinted at the conceptual basis for the research
Chestnut pollen does not appear in the sediments until about described. Throughout this book we emphasize the conceptual
2,000 years ago. The pollen from all three tree species con- foundations of ecology. We also explore some of the applica-
tinues in the sediment record until about 1920, when chest- tions associated with the focal concepts of each chapter. Of
nut blight killed most of the chestnut trees in the vicinity of course, the most important conceptual tool used by ecologists
the lake. Thus, the pollen preserved in the sediments of lakes is the scientific method (see Investigating the Evidence 1 in
can be used to reconstruct the history of vegetation in the Appendix A).
area. Margaret B. Davis, Ruth G. Shaw, and Julie R. Etter- We continue our exploration of ecology in section I with
son reviewed extensive evidence that during climate change, natural history and evolution. Natural history is the founda-
plants evolve, as well as disperse (Davis and Shaw 2001; Davis, tion on which ecologists build modern ecology for which
Lake profile Chestnut decline Present Pollen

due to blight 100
Chestnut
2,000
Years
Beech
8,000
12,000 Spruce
Sediments
Sediment core
Figure 1.9 The vegetation history of landscapes can be reconstructed using the pollen contained within the sediments of nearby lakes. (Chestnut,
Beech and Spruce) Courtesy of the Gretchen and Stanley Jones Palynological Collection and the Botanical Research Institute of Texas
Applications
Ecology Can Inform Environmental
Law and Policy
LEARNING OUTCOMES
After studying this section you should be able to do the following:
1.8 Describe the purposes of the Convention on Inter-

national Trade in Endangered Species of Wild Fauna
and Flora (CITES) and the U.S. Endangered Species
Act (ESA).
1.9 Discuss how subject areas covered in this text are applic-
Figure 1.10 Studies indicate that alpine squirrels (Tamias alpinus) able to identifying and managing endangered species.
are evolving adaptations to higher elevations (Bi et al. 2019).
Danita Delimont/Getty Images
Because ecological science concerns relationships between
organisms and the environment, it is natural to turn to ecology
evolution provides a conceptual framework. A major premise when environmental concerns arise. Consequently, ecology has
of this book is that knowledge of natural history and evolution contributed prominently to the development of environmental
improves our understanding of ecological relationships. law and policy. For example, ecologists have been essential to
evaluating the effects of pollution on the diversity of species in
terrestrial and aquatic communities and on the functioning of
Concept 1.2 Review ecosystems. One area where ecology has played a particularly
significant role is in evaluating the status of individual species
1. What characterizes each of the three types of ecological threatened by human impacts on the environment.
research? How are they different and how might they be Ecological studies of animal and plant populations are
used together? essential to determining when species populations have declined
2. What are some of the new tools and technology being in numbers to the point where they are in danger of extinction
used in ecological research? Why are they valuable? (see chapter 9). Reports of such declines in the 1960s eventually
3. How are ecologists able to look backwards and forwards led to the establishment of international treaties and national
in time? Why is this important? laws to protect endangered species. Two prominent protec-
tions came into force in 1973. The first was the Convention on
International Trade in Endangered Species of Wild Fauna and (chapters 4–8) as well as community, ecosystem, landscape, and
Flora (CITES), an international treaty to protect endangered geographic ecology (chapters 16–22), into greater prominence
species from the threat of wildlife trafficking and trade. The sec- as tools for endangered species management. Because human-
ond was the U.S. Endangered Species Act, or ESA. caused changes to the environment now extend to the entire
The ESA extended protection to all threatened and endan- planet, global ecology (chapter 23) is increasingly relevant to
gered vertebrate animals, invertebrate animals, and plants in the long-term endangered species protection.
United States and to species elsewhere around the globe listed as Ecological studies are also essential to determining whether
endangered under the CITES treaty. The species protected by the protected populations have recovered sufficiently to be removed
ESA have ranged from the large and charismatic, such as griz- from the ESA’s list of endangered species, a process called delisting.
zly bears and whales, to inconspicuous plants and insects. Later There have been a number of high-profile species that have been
amendments to the ESA required that management agencies, delisted in recent years, including the gray whales of the eastern
such as the U.S. Fish and Wildlife Service, identify “critical habi- North Pacific Ocean and bald eagles of the contiguous 48 states.
tat” for threatened and endangered species. This requirement In summary, ecological science has been essential to identifying,
brought studies of the adaptations of species to the environment protecting, and managing species vulnerable to extinction.
Summary
Ecologists study environmental relationships ranging from those microphones that can measure bee behavior by their buzz-
of individual organisms to factors influencing global-scale pro- ing. Modeling as a tool to understand patterns and make
cesses. The research focus and questions posed by ecologists predictions has also become more complex, as computer
differ across the levels of ecological organization studied. simulations have made the processing of large amounts of
Ecologists use three general approaches to research: data easier and faster. Such simulations are particularly useful
observation, experimentation, and modeling. Many scientific in the context of understanding processes that occur at long
questions will require a combination of approaches. Histori- time scales, such as evolution and climate change. Because
cally, research of all three types used very simple approaches ecological science concerns relationships between organisms
and tools, however advances in technology have greatly and the environment, it is often consulted when environmen-
expanded our capacity to learn about our world. This includes tal concerns arise. Ecological science has been particularly
the use of drones to collect data in tree tops, stable isotope important to identifying, protecting, and managing species
analysis to identify the diet of a butterfly, and highly sensitive vulnerable to extinction.
Key Terms
adaptation 8 ecology 1 experiments 5 population 2
biosphere 3 ecosystem 3 in situ 4 quantitative models 7
citizen scientist 6 epiphyte 4 modeling 6 urban ecology 3
community 2 evolution 2 natural experiment 6
conceptual models 7 ex situ 6 observation 4
Review Questions
1. Faced with the complexity of nature, ecologists have divided the in keeping some American redstarts out of the most productive
field of ecology into subdisciplines, each of which focuses on one feeding areas on their wintering grounds?
of the levels of organization pictured in figure 1.1. What is the 4. How has technology advanced our capacity to collect data over
advantage of developing such subdisciplines within ecology? wider geographic and temporal scales? What other advances
2. What are the pitfalls of subdividing nature in the way it is repre- have benefited the scientific field of ecology?
sented in figure 1.1? In what ways does figure 1.1 misrepresent 5. What do the studies of Margaret Davis tell us about the compos-
nature? ition of forests in the Appalachian Mountains during the past
3. What could you do to verify that the distinct feeding zones 12,000 years (see fig. 1.9)? Based on this research, what predictions
used by the warblers studied by MacArthur (see fig. 1.3) are might you make about the future composition of these forests?
the result of ongoing competition between the different species 6. Ecological models range from the very simple to the very com-
of warblers? How might you examine the role of competition plex. Are complex models always better? Why or why not?
Design elements: (Yellowstone thermal pool):©flickrRF/Getty Images

SECTION
I Natural History
and Evolution
Chapter
Life on Land
2
LifeJourneys/iStock/Getty Images
Despite their often barren appearance, deserts are ecosystems with a

diversity of life, uniquely adapted to the temperature and moisture of LEARNING OUTCOMES
those environments. After studying this section you should be able to do the following:
2.1 Explain why plants are the basis of life on Earth

and thus define terrestrial biomes.
CHAPTER CONCEPTS 2.2 List the main environmental features used to
differentiate the various terrestrial biomes.
2.1 Uneven heating of the earth’s surface by
the sun and the tilt of the earth combine
I
to produce predictable latitudinal and f you are standing on a ground of seemingly barren rock
seasonal variation in climate. 14
and sand, what little vegetation being thick-leaved and/

Concept 2.1 Review 16 or prickly, with a few vultures overhead and a lizard eat-
ing a scorpion at your feet, where are you? Although the details
2.2 While terrestrial biome distribution is
differ, this general description fits areas of the southwestern
strongly associated with latitude, biomes
are also influenced by microclimate and United States, western China, Libya, Australia, and elsewhere;
soil type. 17 we refer to this type of ecosystem as a desert. Desert is one
of several types of biomes, major divisions of the terrestrial
environment, distinguished primarily by their predominant
2.3 Environmental conditions shape each plants (fig. 2.1a). In figure 2.1, the boundaries between biomes
biome’s characteristic biology. 19 appear sharp, whereas in nature these transitions generally occur
gradually over long distances along gradients of environmental
variation. But why do we find the same biome in such disparate
Applications: Finer Scale Climatic Variation over locations across the globe? And conversely, why don’t we find
Time and Space 40 desert at the top of a mountain or at the equator? The study
Summary 41 of how organisms in a particular area are influenced by factors
such as climate, soils, predators, competitors, and evolutionary
Key Terms 42
history is called natural history. In this chapter, we explore the
Review Questions 43 natural history of different types of terrestrial biomes, including
the reasons why they are distributed the way that they are.
11
12 Section I Natural History and Evolution
Arctic Circle
Tropic of Cancer
Equator
Tropic of Capricorn
Arctic Tundra
Boreal Forest
Temperate Grassland
Temperate Forest
Woodland and Shrubland
Desert
Tropical Savanna
Tropical Dry Forest
Tropical Rain Forest
Mountains
Lakes and Inland Seas
(a)
500
450
precipitation
Increasing
400
Tropical
350 Average annual precipitation (cm)
rain forest
Decreasing 300
temperature
t
fores 250 . . . but grass will
p erate dominate at the
Tem same temperatures
Tropical 200
dry forest if there is greater
rainfall.
d
slan 150
g ras
Cacti and
Boreal e rate
similar vegetation mp Tropical 100
forest Te savanna
dominate in places
that are very dry but shrubland
Woodland/ 50
with a range of Tundra
temperatures—what Desert
we call “desert” . . . 0
−15 −10 −5 0 5 10 15 20 25 30
Average annual temperature (°C)
(b)
Figure 2.1 (a) Global distribution of biomes. (b) Biomes are generally defined by the average temperature (x-axis) and precipitation (rain, y-axis) of
a given location. Robert Whittaker plotted the vegetation type of various locations against these two variables (inset) to determine the boundaries of the
various biomes, but boundaries between these are not distinct, due to the many other influences of the environment (adapted from W hittaker 1975).
availability (fig. 2.1b). If we plot the combination of these two vari-

Terrestrial Biomes and the ables for locations across the globe by their dominant plant
Importance of Plants types, we can see clear patterns of how different natural histories
The most important factor for understanding the distribution can be defined by temperature and precipitation (fig. 2.1b inset).
of biomes has to do with climate. Whether an ecosystem is But how does a particular environment result in similar
dominated by cacti, grass, deciduous trees, conifers, or other plants? Returning to our desert scene described above, the
types of plants will primarily depend on temperature and water dry environment will mean that regardless of which continent
Chapter 2 Life on Land 13
we are on, plants in a desert will have certain functional traits: we visited above, the lizard is eating a scorpion that may have
particular characteristics that allow them to survive, such as eaten a grasshopper that survived by eating a succulent plant,
waxy coatings on leaves that prevent water loss. These functional such as a cactus. When all of these organisms die, there will be
traits of plants arise through evolution via selective pressure by fungi and insects and bacteria that eat them. Thus, all of these
the environment, a subject explored in more detail in chapter 4. consumer organisms can be considered secondary producers
The section on deserts in this chapter contains a particularly of energy for the organisms that eat them. We will learn more
compelling example of how selective pressure results in similar about food webs and energy flow in ecosystems in chapters 17
functional traits in unrelated plants on different continents. and 18. Selective pressure by the climate will be felt directly by
But what about all of the animals, fungi, and other organisms consumers in an ecosystem, but also indirectly, as each experi-
in an ecosystem; why are biomes defined by plants? One reason ences the selective pressure of its food source. Each consumer
for this is that plants form the foundation of life on this planet. will have traits that make it able to capture, eat, and digest the
On land, plants make the energy of the sun available to all other food available in that environment. Thus, the type and diversity
life-forms, including us, via photosynthesis (explained in more of plants in a given region, determined primarily by climate, will
detail in chapter 7). For this reason, plants and other photosyn- have far-reaching implications for the rest of the ecosystem.
thesizing organisms are called primary producers (fig. 2.2). The We devote chapters 2 and 3 to an overview of general types
biomass produced by primary producers per unit time is primary of ecosystems and where they occur, that is, the natural history
production, which is discussed in detail in chapter 18. Animals of the biosphere. In chapter 2, we examine the natural history of
and fungi cannot get their energy directly from the sun and so life on land. We will explore why natural history is primar-
depend on this conversion by plants. For example, in the desert ily based on latitude—that is, why we don’t find desert at the
Primary
producers
Secondary
producers
(consumers)
Figure 2.2 How energy moves through an ecosystem; size and direction of arrows indicate the movement and loss of energy through trophic groups.
Plants such as cacti capture sun energy via photosynthesis; animals depend on that energy whether by eating the plants or other animals that have
eaten plants, and so on. In this way, all organisms ultimately get their energy from the sun, forming the basis of all terrestrial biomes.
equator—but also how soil type and elevation can have strong Temperature, Atmospheric Circulation,
influences as well, that is, why we also don’t find desert on and Precipitation
mountaintops. The main goal of chapters 2 and 3 is to take a
large-scale perspective of nature before delving, in later chap- The uneven heating of the earth’s surface results from the
ters, into finer details of structure and process. spherical shape of the earth and the angle at which the earth
rotates on its axis as it orbits the sun. Because the earth is a
sphere, the sun’s rays are most concentrated where the sun
2.1 Large-Scale Patterns is directly overhead. However, the latitude at which the sun
of Climatic Variation is directly overhead changes with the seasons. This seasonal
change occurs because the earth’s axis of rotation is not per-
LEARNING OUTCOMES pendicular to its plane of orbit about the sun but is tilted
After studying this section you should be able to do the following: approximately 23.5° away from the perpendicular (fig. 2.3).
Because this tilted angle of rotation is maintained throughout
2.3 Diagram the position of the sun relative to the
earth’s orbit about the sun, the amount of solar energy received
equator and tropics of Capricorn and Cancer, during
by the Northern and Southern Hemispheres changes season-
the equinoxes and solstices.
ally. During the northern summer, the Northern Hemisphere
2.4 Describe how solar driven air circulation produces
is tilted toward the sun and receives more solar energy than the
regional differences in precipitation.
Southern Hemisphere. During the northern summer solstice on
2.5 Interpret a climate diagram.
approximately June 21, the sun is directly overhead at the tropic of
2.6 Explain the influence of the Coriolis effect on wind
Cancer, at 23.5° N latitude. During the northern winter solstice,
direction.
on approximately December 21, the sun is directly overhead at
the tropic of Capricorn, at 23.5° S latitude. During the northern
Uneven heating of the earth’s surface by the sun and the tilt of winter, the Northern Hemisphere is tilted away from the sun and
the earth combine to produce predictable latitudinal and seasonal the Southern Hemisphere receives more solar energy. The sun is
variation in climate. While the global distribution of biomes is directly overhead at the equator during the spring and autumnal
determined primarily by climate, what determines the distribu- equinoxes, on approximately March 21 and September 22 or 23.
tions of climate? Several attributes of climate vary predictably On those dates, the Northern and Southern Hemispheres receive
over the earth. For instance, average temperatures are lower approximately equal amounts of solar radiation.
and more seasonal at middle and high latitudes. Temperature This seasonal shift in the latitude at which the sun is directly
generally shows little seasonality near the equator, while rain- overhead drives the march of the seasons. At high latitudes, in
fall may be markedly seasonal. Deserts, which are concentrated both the Northern and Southern Hemispheres, seasonal shifts
in a narrow band of latitudes around the globe, receive little in input of solar energy produce winters with low average
precipitation, which generally falls unpredictably in time and temperatures and shorter day lengths and summers with high
space. What mechanisms produce these and other patterns of average temperatures and longer day lengths. In many areas
climatic variation? at middle to high latitudes, there are also significant seasonal
Northern Hemisphere has Northern Hemisphere has

spring equinox as equator faces winter as it tilts away from
the sun. Southern Hemisphere the sun. Southern
has autumnal equinox. Hemisphere has summer as
it tilts toward the sun.
0°
Northern Hemisphere
Northern Hemisphere has has autumnal equinox as
summer as it tilts toward the sun. Constant tilt of 23.5° equator faces the sun.
Southern Hemisphere has winter from plane of orbit Southern Hemisphere
as it tilts away from the sun. has spring equinox.
Figure 2.3 The seasons in the Northern and Southern Hemispheres.

changes in precipitation. Meanwhile, between the tropics of the earth rotates from west to east. We can see how air movement
Cancer and Capricorn, seasonal variations in temperature and changes by this simple demonstration: with your finger, trace a
day length are slight, while precipitation may vary a great deal. line from the equator to the north pole on a globe while it is slowly
What produces spatial and temporal variation in precipitation? turning. Do it again for the south pole. Your finger will draw
Heating of the earth’s surface and atmosphere drives diagonal lines in the same direction as winds will be dragged on
circulation of the atmosphere and influences patterns of pre- the surface of the earth. That is, an observer at tropical latitudes
cipitation. As shown in figure 2.4a, the sun heats air at the observes winds that blow from the northeast in the Northern
equator, causing it to expand and rise. This warm, moist air Hemisphere and from the southeast in the Southern Hemi-
cools as it rises. Since cool air holds less water vapor than sphere (fig. 2.5). These are the northeast and southeast trades.
warm air, the water vapor carried by this rising air mass con- Someone studying winds within the temperate belt between 30°
denses and forms clouds, which produce the heavy rainfall and 60° latitude would observe that winds blow mainly from the
associated with tropical environments. west. These are the westerlies of temperate latitudes. At high lati-
Eventually, this equatorial air mass ceases to rise and tudes, our observer would find that the predominant wind direc-
spreads north and south. This high-altitude air is dry, since the tion is from the east. These are the polar easterlies.
moisture it once held fell as tropical rains. As this air mass flows Why don’t winds move directly north to south? The pre-
north and south, it cools, which increases its density. Eventu- vailing winds do not move in a straight north–south direction
ally, it sinks back to the earth’s surface at about 30° latitude and because of the Coriolis effect. In the Northern Hemisphere, the
spreads north and south. This dry air draws moisture from the Coriolis effect causes an apparent deflection of winds to the right
lands over which it flows and creates deserts in the process. of their direction of travel and to the left in the Southern Hemi-
Air moving from 30° latitude toward the equator completes sphere. We say “apparent” deflection because we see this deflec-
an atmospheric circulation cell at low latitudes. As f igure 2.4b tion only if we make our observations from the surface of the
shows, there are three such cells on either side of the equator. earth. To an observer in space, it would appear that winds move
Air moving from 30° latitude toward the poles is part of the in approximately a straight line, while the earth rotates beneath
atmospheric circulation cell at middle latitudes. This warm air them. However, we need to keep in mind that the perspective
flowing from the south rises as it meets cold polar air flow- from the earth’s surface is the ecologically relevant perspective.
ing from the north. As this air mass rises, moisture picked up The biomes that we discuss in chapter 2 are as earthbound as
at lower latitudes condenses to form the clouds that produce our hypothetical observer. Their distributions across the globe
the abundant precipitation of temperate regions. The air rising are substantially influenced by global climate, particularly geo-
over temperate regions spreads northward and southward at a graphic variations in temperature and precipitation.
high altitude, completing the middle- and high-latitude cells of Geographic variation in temperature and precipitation is
general atmospheric circulation. very complex. How can we study and represent geographic varia-
The patterns of atmospheric circulation shown in figure 2.4b tion in these climatic variables without being overwhelmed by a
suggest that air movement is directly north and south. However, mass of numbers? This practical problem is addressed by a visual
this does not reflect what we observe from the earth’s surface as device called a climate diagram.
Sun heats air at equator.
Subtropical and polar

Moisture in ascending air masses meet, creating
air condenses, forming a moist temperate climate.
clouds.
Dry descending air
absorbs moisture,
Some ascending air Some ascending air forming deserts. Temp
e rate f 60° N
flows to the north. flows to the south. orest
Rising air at the Dese
equator is associated rt
with a moist Trop
Warm Warm ical rain f
tropical climate. o 30° N
air rises. Heavy air rises. rest
rains fall.
Dese 0° equator
rt
Deserts Equator Deserts
(30°) (0° latitude) (30°) 30° S
There are three
Dry air flowing over air circulation
60° S
land absorbs moisture. cells on each
side of the equator.
(a) (b)
Figure 2.4 (a) Solar-driven air circulation. (b) Latitude and atmospheric circulation.
Coriolis effect:
Winds in the
Northern Hemisphere
Polar easterlies deflected to the right
60° N of their direction of
travel
Westerlies
30° N
Northeast trade winds
0° (equator)
Southeast trade winds
30° S
Westerlies
Coriolis effect:
60° S Winds in the Southern
Polar easterlies Hemisphere deflected to
the left
Figure 2.5 The Coriolis effect and wind direction.
Climate Diagrams of the year are plotted on the horizontal axis, beginning
with January and ending with December for locations in
Climate diagrams were developed by Heinrich Walter (1985) as
the Northern Hemisphere and beginning with July and
a tool to explore the relationship between the distribution of
ending with June in the Southern Hemisphere. Tempera-
terrestrial vegetation and climate. Climate diagrams summarize
ture is plotted on the left vertical axis and precipitation
a great deal of useful climatic information, including seasonal
on the right vertical axis. Temperature and precipitation
variation in temperature and precipitation, and the length and
are plotted on different scales so that 10°C is equivalent
intensity of wet and dry seasons.
to 20 mm of precipitation. Climate diagrams for wet areas
As shown in figure 2.6, climate diagrams summarize cli-
such as tropical rain forest compress the precipitation scale
matic information using a standardized structure. The months
for precipitation above 100 mm so that 10°C is equivalent
Mean monthly temperature, as Mean monthly precipitation, to 200 mm of precipitation. With this change in scale, rain-
given on the left axis, plotted as given on the right axis, fall data from very wet climates can be fit on a graph of
with red dots. plotted with blue dots. convenient size.
Because the temperature and precipitation scales are
Locality Elevation (m) constructed so that 10°C equals 20 mm of precipitation,
°C mm
Mean annual Mean annual the relative positions of the temperature and precipitation
temperature (°C) precipitation lines reflect water availability. Theoretically, adequate mois-
40 (mm) 80
ture for plant growth exists when the precipitation line lies
30 60
above the temperature line. When the temperature line
lies above the precipitation line, potential evaporation rate
20 40 exceeds precipitation.
As you can see, climate diagrams efficiently summarize
10 20 important environmental variables. In Concept 2.3, we use cli-
mate diagrams to represent the climates associated with major
0 0 terrestrial biomes.
‒10
‒20
J F M A M J J A S O N D Concept 2.1 Review
January Month December
1. How would seasonality in temperature and precipitation
Red shading indicates mean
be affected if earth’s rotation on its axis were perpen-
No shading indicates mean
minimum temperature for the minimum temperature for the dicular to its plane of orbit about the sun?
month is below freezing. month is above freezing. 2. Why does the annual rainy season in regions near 23° N
latitude begin in June?
Figure 2.6 Structure of climate diagrams.
2.2 Other Factors That Shape Microclimates can have dramatic influences on biome
distribution; the western United States is one such example
Terrestrial Biomes (f ig. 2.8). Not only do we observe particular plant commun-
LEARNING OUTCOMES ities on high-altitude mountaintops, but because of the rain
After studying this section you should be able to do the following: shadow effect, we observe a wide range of biomes on the sides
and base of a mountain, even at the same elevation and lati-
2.7 Explain the concept of rain shadow. tude. Desert, for example, is found on the eastern side of the
2.8 Describe the characteristics of each of the typical soil Sierra Nevada mountains.
horizons. Biomes are not only determined by temperature and mois-
2.9 Discuss how climate, organisms, topography, parent ture, however; in addition to and interacting with climate is the
material, and time can influence the structure and effect of soils. As you will see in the biome descriptions later
development of soils and which organisms can sur- in this chapter, soil type can determine if a region is savanna
vive there. or desert, even at the same amount of precipitation. It is partly
for this reason that there are blurry, rather than crisp, bound-
While terrestrial biome distribution is strongly associated with aries between biomes in f igure 2.1b.
latitude, biomes are also influenced by microclimate and soil Soil is a complex mixture of living and nonliving material
type. From the previous section, we know why deserts are upon which most terrestrial life depends. The nonliving com-
not found at the equator: the angle of the sun’s rays on the sur- ponent of soil comes from its underlying geology; weathering
face of the earth results in this being a wet area of the globe. slowly breaks down parent material, often bedrock, into smaller
However, latitude is not the only determinant of where we find and smaller fragments to produce sand, silt, and clay-sized par-
biomes; we also do not find desert on mountaintops, regard- ticles. The size of particles and the minerals associated with
less of latitude. Also consider the fact that at 35°N we can find them can have profound impacts on what types of plants,
temperate forest in North Carolina, grassland in Oklahoma, microorganisms, and even animals can live in the associated
desert in Arizona, and Mediterranean scrubland in California. soils. Here we summarize the general features of soil structure
The distribution of mountains partly explains why biomes and development. The biome discussions that follow include
do not form perfect horizontal stripes on the earth. It is colder at specific information about the soils associated with each.
higher elevations, of course, but whether it will be wetter or drier Nonetheless, it should be kept in mind that variation within a
depends on which side of the mountain you are on. To under- biome is often due to different types of soils occurring there.
stand this, we can apply what we learned in the previous section Though soil structure usually changes gradually with
about rising air losing its moisture in the form of rain or snow. depth, soil scientists generally divide soils into several dis-
Warm, moist air from the ocean that blows toward a mountain crete horizons. In the classification system used here, the
range will have lost much of its moisture when that air reaches soil profile is divided into O, A, B, and C horizons (fig. 2.9).
the leeward side (fig. 2.7). The dry climate that results is called The O, or organic, horizon lies at the top of the profile. The
the rain shadow effect. When the temperature and moisture dif- most superficial layer of the O horizon is made up of freshly
fer from the prevailing climate, we call these local environments fallen organic matter, including whole leaves, twigs, and other
microclimates, to be discussed in greater detail in chapter 5. plant parts, which become more fragmented and decomposed
Moist, warm Cooling air loses Dry, cool air

air rises. moisture as rain. descends.
Temperature
3,000 –4°C
Region of
rain shadow
2,250 2°C
Prevailing winds
Altitude (m)
1,500 8°C
750 14°C
Evaporation
0 20°C
Warm ocean
Figure 2.7 The rain shadow effect accounts for dramatic differences in climate and therefore biome on opposite sites of many mountain ranges.
Conifer forest is favored when there are both moisture
C
and cool temperatures, here due to high elevation.
as c
Ro
a
ains
d
Only the highest peaks will be cold
e
enough to have tundra at this
ck
latitude.
R
Coastal Mount
a
y
n g
Mo
e
Tundra
Temperate conifer forest

Sierr
un t a i Open conifer woodland
Temperate deciduous forest
Mediterranean woodland and shrubland

a Ne
Grassland
Desert
ns
vada
Barren/water
Temperate deciduous forest and

conifer forest occur on the west
slope of the Sierra Nevada
mountains …
... whereas due to the rain
shadow effect, desert and
open conifer woodland
are found on the east slope.
Figure 2.8 In the western United States, mountain ranges can include
s everal different biomes, determined by both temperature and precipitation
(fig. 2.1b), which can be influenced by the mountains themselves (fig. 2.7)
(data from Thompson and Anderson 2000).
Soil horizons
Organic horizon. Upper layer contains
O loose, somewhat fragmented plant litter.
Litter in lower layer is highly fragmented.
Mineral soil mixed with some organic
A matter. Clay, iron, aluminum, silicates,
and soluble organic matter are gradually
leached from A horizon.
Depositional horizon. Materials leached
B from A horizon are deposited in B horizon.
Deposits may form distinct banding patterns.
Weathered parent material. The C horizon

C may include many rock fragments. It
often lies on bedrock.
Figure 2.9 Generalized soil profile, showing O, A, B, and C horizons.

with increasing depth. Fragmentation and decomposition of 2.3 Natural History and
the organic matter in this horizon are mainly due to the activi-
ties of soil organisms, including bacteria, fungi, and animals
Geography of Biomes
ranging from nematodes and mites to burrowing mammals. LEARNING OUTCOMES
This horizon is usually absent in agricultural soils and After studying this section you should be able to do the following:
deserts. At its deepest levels, the O horizon merges gradually
with the A horizon. 2.10 List the major terrestrial biomes.
The A horizon contains a mixture of mineral materials, such 2.11 Describe the climatic differences among the biomes.
as clay, silt, and sand, and incorporated organic material derived 2.12 Contrast the soils typical of the terrestrial biomes.
from the O horizon. Burrowing animals, such as earthworms, 2.13 Describe the types of vegetation, animals, and other
mix organic matter from the O horizon into the A horizon. The organisms characteristic of the terrestrial biomes.
A horizon is generally rich in mineral nutrients. It is gradually 2.14 Explain variation in human presence in the various
leached of clays, iron, aluminum, silicates, and humus, which is terrestrial biomes.
partially decomposed organic matter. These substances slowly
move down through the soil profile until they are deposited in Environmental conditions shape each biome’s characteristic
the B horizon. biology. Early in the twentieth century, many plant ecologists
The B horizon contains the clays, humus, and other studied how climate and soils influence the distribution of veg-
materials that have been transported by water from the A etation. Later ecologists concentrated on other aspects of plant
horizon. The deposition of these materials often gives the B ecology. Today, as we face the prospect of global warming (see
horizon a distinctive color and banding pattern. The B horizon chapter 23), ecologists are once again studying climatic influ-
gradually merges with the C horizon. ences on the distribution of vegetation. International teams of
The C horizon is the deepest layer in our soil pit and the ecologists, geographers, and climatologists are exploring the
only one not typically dominated by plant roots. It consists influences of climate on vegetation with renewed interest and
of the weathered parent material, which has been worked by with much more powerful analytical tools. Ecologist Osvaldo
the actions of frost, water, and the deeper penetrating roots of Sala and others created a predictive model (for more on models,
plants. Because weathering is incomplete and less intense than see Investigating the Evidence 1 in Appendix A) using biological
in the A and B horizons, the C horizon may contain many and environmental data from each biome to determine where
rock fragments. Under the C horizon, we find unweathered biodiversity is at the most risk. While deserts and tundra were
parent material, which is often bedrock. not expected to change much over the next century, Mediterra-
The soil profile gives us a snapshot of soil structure. How- nean and grassland biomes were found to be highly sensitive to
ever, soil structure is in a constant state of flux as a consequence anticipated human-caused changes to the environment, includ-
of several influences. Those influences were summarized by ing but not limited to climate change (Sala et al. 2000).
Hans Jenny (1980) as climate, organisms, topography, parent In this section, we discuss the climate, soils, and organ-
material, and time. Climate affects the rate of weathering of isms of the earth’s major biomes and how they have been influ-
parent materials, the rate of leaching of organic and inorganic enced by humans.
substances, the rate of erosion and transport of mineral
particles, and the rate of decomposition of organic matter.
Living organisms, which as we know are also affected by cli-
Tropical Rain Forest
mate, influence the quantity and quality of organic matter Tropical rain forest is nature’s most extravagant garden (fig. 2.10).
added to soil and the rate of soil mixing by burrowing animals. Beyond its tangled edge, a rain forest opens into a surprisingly
Topography affects the rates and direction of water flow and
patterns of erosion. Meanwhile, parent materials, such as gran-
ite, volcanic rock, and wind- or water-transported sand, set the
stage for all other influences. Last is the matter of time. Soil
age influences soil structure.
As with many aspects of ecology, it is often difficult to
separate organisms from their environment. The biome discus-
sions that follow provide additional information on soils by
including aspects of soil structure and chemistry characteristic
of each biome.
Concept 2.2 Review

1. The organic horizon is generally absent from agricultural
soils because tilling (e.g., plowing), buries organic matter. Figure 2.10 Tropical rain forest in Ecuador. More species live
Why is an organic horizon generally absent from desert soils? within the three-dimensional framework of tropical rain forests than in
any other terrestrial biome. Elena Kalistratova/Vetta/Getty Images
spacious interior, illuminated by dim, greenish light shining Soils

through a ceiling of leaves. The architecture of rain forests, with Heavy rains gradually leach nutrients from rain forest soils and
their vaulted ceilings and spires, has invited comparisons to rapid decomposition in the warm, moist rain forest climate keeps
cathedrals and mansions. However, this cathedral is alive from the quantity of soil organic matter low. Consequently, rain for-
ceiling to floor, perhaps more alive than any other biome on the est soils are often nutrient-poor, acidic, thin, and low in organic
planet. In the rain forest, the sounds of evening and morning, matter. In many rain forests, more nutrients are tied up in living
the brilliant flashes of color, and rich scents carried on moist tissue than in soil. Some rain forests, however, occur where soils
night air speak of abundant life, in seemingly endless variety. are very fertile such as along rivers. Fungi, bacteria, and soil ani-
mals, such as mites and springtails, rapidly scavenge nutrients
Geography
from plant litter (leaves, flowers, etc.) and animal wastes, fur-
Tropical rain forests straddle the equator in three major regions: ther tightening the nutrient economy in tropical ecosystems.
Southeast Asia, West Africa, and South America (fig. 2.11).
Most rain forest occurs within 10° of latitude north or south Biology
of the equator. Outside this equatorial band are the rain forests Trees dominate the rain forest landscape and average about 40 m
of Central America and Mexico, southeastern Brazil, eastern in height but some reach 80 m. These rain forest giants are often
Madagascar, southern India, and northeastern Australia. supported by well-developed buttresses. The diversity of rain for-
est trees is also impressive. One hectare (100 m × 100 m) of
Climate temperate forest may contain a few dozen tree species; 1 ha
The global distribution of rain forests corresponds to areas of tropical rain forest may contain up to 300 tree species.
where conditions are warm and wet year-round (see fig. 2.11). Primary production in tropical rain forests is the highest of
Temperatures in tropical rain forests vary little from month all terrestrial ecosystems, not only from the trees but also because
to month and often change as much in a day as they do over the three-dimensional framework formed by rain forest trees is
the entire year. Average temperatures are about 25°C to 27°C, festooned with other plant growth forms. The trees are trellises
lower than average maximum summer temperatures in many for climbing vines and growing sites for epiphytes, plants that
deserts and temperate regions. Annual rainfall ranges from grow on other plants (fig. 2.12). This vast amount of converted
about 2,000 to 4,000 mm, and some rain forests receive even energy supports a great biological richness of consumers as well.
more precipitation. In a rain forest, a month with less than 100 A single rain forest tree may support several thousand species
mm of rain is considered dry. of insects, many of which have not been described by scientists.
Moist Dry Mean

minimum
temperature >0 C
Tropic of Cancer
Precipitation exceeds 100 mm Equator
during most months.
Tropic of Capricorn
Annual variation in
temperature is slight.
Belem, Brazil 24 m Kisangani, DR Congo 415 m Kuala Lumpur, Malaysia 27 m

C mm C mm C mm
26.7 C 2,438 mm 25.3 C 1,760 mm 27.5 C 2,685 mm
300 300 300
100 100 100
40 80 40 80 40 80
30 60 30 60 30 60
20 40 20 40 20 40
10 20 10 20 10 20
0 0 0 0 0 0
J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D
Month Month Month
Figure 2.11 Tropical rain forest geography and climate.

Orchid Human Influences

People from all over the globe owe more to the tropics than
Shrub is generally realized. Many of the world’s staple foods, includ-
ing maize (called corn in North America and Australia),
rice, bananas, and sugarcane, and approximately 25% of all
prescription drugs were originally derived from tropical plants.
Many more species, directly useful to humans, may await
discovery. Unfortunately, tropical rain forests are fast disappear-
ing. According to data collected by scientists at the University
of Maryland, in 2019 an astonishing 11.9 million hectares of
tropical forest cover was lost; this is equal to a football field-
Water-holding
bromeliad sized area every 8.25 minutes (www.globalforestwatch.org).
Accumulated dead
This loss diminishes our chances of understanding the extent
organic matter Fern and dynamics of biological diversity.
Tropical Dry Forest

During the dry season, the tropical dry forest is all earth tones;
Cross section
of branch
in the rainy season, it’s an emerald tangle (fig. 2.13). Life in
the tropical dry forest responds to the rhythms of the annual
solar cycle, which drives the oscillation between wet and
dry seasons. During the dry season, most trees in the tropi-
cal dry forest are dormant. Then, as the rains approach, trees
flower and insects appear to pollinate them. Eventually, as the
first storms of the wet season arrive, the trees produce their
Roots from tree branches draw Nutrients are contained in living leaves and transform the landscape.
nutrients from epiphyte mat. plants and dead organic matter.
Geography
Figure 2.12 An epiphyte mat in the tropical rain forest canopy.
Epiphyte mats store a substantial fraction of the nutrients in tropical rain Tropical dry forests make up approximately 42% of tropical and
forests and support a high diversity of plant and animal species. sub-tropical forest area (Hasnat and Hossain 2020) (fig. 2.14).
In Africa, tropical dry forests are found both north and south
of the central African rain forests. In the Americas, tropical
The rain forest is not, however, just a warehouse for a dry forests are the natural vegetation of extensive areas south
large number of dissociated species. Rain forest ecology is and north of the Amazon rain forest. Tropical dry forests
marked by intricate, complex relationships between species. also extend up the west coast of Central America and into
In the tropical rain forest, there are plants that cannot live North America along the west coast of Mexico. In Asia, tropi-
without particular species of fungi, which help them absorb cal dry forests are the natural vegetation of most of India and
nutrients (called mycorrhizae), the hummingbirds or insects the Indochina peninsula. Australian tropical dry forests form
that pollinate their flowers, and the animals that disperse a continuous band across the northern and northeastern por-
their seeds. tions of the continent.
Figure 2.13 Tropical dry forest during the wet and dry seasons. Created by Tomas Zrna/Getty Images; Ralph Lee Hopkins/Science Source
Moist Dry Mean

minimum
temperature >0°C
Temperature more variable Tropic of Cancer

than in tropical rain forest. Equator
Tropic of Capricorn
Climate alternates
between very wet and
very dry seasons.
Acapulco, Mexico 3m Bombay, India 4m Darwin, Australia 32 m

°C mm °C mm °C mm
26.9°C 1,473 mm 26.7°C 1,808 mm 29.4°C 1,491 mm
300 300 300
100 100 100
40 80 40 80 40 80
30 60 30 60 30 60
20 40 20 40 20 40
10 20 10 20 10 20
0 0 0 0 0 0
J F M A M J J A S O N D J F M A M J J A S O N D J A S O N D J F M A M J
Month Month July Month June
Climate diagrams for sites in

Southern Hemisphere order
months from July to June.
Figure 2.14 Tropical dry forest geography and climate.
Climate Human Influences

The climate of tropical dry forests is more seasonal than that Heavy human settlement has devastated the tropical dry forest.
of tropical rain forests. The three climate diagrams shown in While the world’s attention has been focused on the plight of
figure 2.14, for example, show a dry season lasting for 6–7 months, rain forests, tropical dry forests have been quietly disappearing,
followed by a season of abundant rainfall, lasting 5–6 months. including in so-called protected areas. The relatively fertile soil
The climate diagrams also indicate more seasonal variation in of tropical dry forests has attracted agricultural development,
temperature compared to tropical rain forest. including cattle ranches, grain farms, and cotton fields. Tropical
dry forests are more easily converted to agriculture compared to
Soils rain forests, since the dry season makes them more accessible and
The soils of many tropical dry forests are of great age, particu- easier to burn. Using remote sensing data that spanned 40 years,
larly those in the parts of Africa, Australia, India, and Brazil Binita Kumari and colleagues found that areas of a reserve in
that were once part of the ancient southern continent of Gond- eastern India that had more human settlements also had higher
wana. The soils of tropical dry forests tend to be less acidic rates of deforestation (Kumari et al. 2020; fig. 2.15). In other
than those of rain forests and are generally richer in nutrients. areas, protections seem to be working; in Ghana, researchers
However, the annual pulses of torrential rain make the soils of found that rates of deforestation had decreased in recent years
tropical dry forest highly vulnerable to erosion. and new trees are colonizing in some areas (Janssen et al. 2018).
The loss of the dry forest is significant because, while rain
Biology forests may support a somewhat greater number of species,
The plants of the tropical dry forest are strongly influenced by many dry forest species are found nowhere else, as many as
physical factors. For example, the height of the dry forest is highly 40% of the tree and shrub species found there are endemic
correlated with average precipitation. Trees are tallest in the wet- (Hasnat and Hossain 2020).
test areas. In the driest habitats, all trees drop their leaves during
the dry season; in wetter areas over 50% may be evergreen. As Tropical Savanna
in the tropical rain forest, many plants produce animal-dispersed Stand in the middle of a savanna, a tropical grassland dotted
seeds. However, wind-dispersed seeds are also common. Many with scattered trees, and your eye will be drawn to the hor-
dry forest birds, mammals, and even insects make seasonal migra- izon for the approach of thunderstorms or wandering herds
tions to wetter habitats along rivers or to the nearest rain forest. of wildlife (fig. 2.16). The tropical savanna is the kingdom of
India Areas of high human density… …were more likely to be deforested

1,000 km
0 20 40 km
Jharkhand State
200 km
Forest
High: 889 Non-Forest in 1975
Low: 0 Additional deforestation by 2015
A. Settlement density B. Area of deforestation
Figure 2.15 A comparison of human settlement density (A) and deforestation (B) at the Palamau Tiger Reserve, the latter using satellite imagery.
Authors found that loss of forest cover over time was greatest in the central and northern regions, where population densities were highest (based on
data from Kumari et al. 2020).
Figure 2.16 Tropical savanna and herbivores in East Africa. This ecosystem is characterized by a dominance of grasses with a
few woody species. Anna Sher
the farsighted, the stealthy, and the swift and is the birthplace accompanied by intense lightning. This lightning often starts
of humankind. It was from here that we eventually moved out fires, particularly at the beginning of the wet season when the
into every biome. Though most humans live away from this savanna is tinder dry. These fires kill young trees while the
first home, the fascination continues. grasses survive and quickly resprout. Consequently, fires help
maintain the tropical savanna as a landscape of grassland and
Geography scattered trees.
The savanna climate is generally drier than that of tropical
Most tropical savannas occur north and south of tropical dry
dry forest. However, San Fernando, Venezuela (see fig. 2.17),
forests within 10–20° of the equator. In Africa south of the
receives as much rainfall as a tropical dry forest. Other savan-
Sahara Desert, tropical savannas extend from the west to the
nas occur in areas that are as dry as deserts. What keeps the
east coasts, cut a north–south swath across the East African
wet savannas near San Fernando from being replaced by forest
highlands, and reappear in south-central Africa (fig. 2.17). In
and how can savannas persist under desertlike conditions? The
South America, tropical savannas occur in south-central Brazil
answer lies deep in the savanna soils.
and cover a great deal of Venezuela and Colombia. Tropical
savannas are also the natural vegetation of much of northern Soils
Australia in the region just south of the tropical dry forest.
Soil layers with low permeability to water play a key role in
Savanna is also the natural vegetation of an area in southern
maintaining many tropical savannas. For instance, because
Asia just east of the Indus River in eastern Pakistan and north-
a dense, impermeable subsoil retains water near the surface,
western India.
savannas occur in areas of southwestern Africa that would
otherwise support only desert. Impermeable soils also help
Climate savannas persist in wet areas, particularly in South America.
As in the tropical dry forest, life on the savanna cycles to the Trees do not move onto savannas where an impermeable sub-
rhythms of alternating dry and wet seasons (see fig. 2.17). soil keeps surface soils waterlogged during the wet season. In
Here, however, seasonal drought combines with another impor- these landscapes, scattered trees occur only where soils are
tant physical factor, fire. The rains come in summer and are well drained.
Moist Dry Mean

minimum
temperature >0°C
Tropic of Cancer
Equator
Tropic of Capricorn
San Fernando, Venezuela 44 m Tahoua, Niger 386 m Longreach, Australia 196 m

°C mm °C mm °C mm
27.8°C 1,534 mm 28.6°C 386 mm 23.3°C 417 mm
300 300 300
However, the wet season is generally
100 shorter and100
drier than in tropical dry 100
forest.
40 80 40 80 40 80
30 60 30 60 30 60
20 40 20 40 20 40
10 20 10 20 10 20
0 0 0 0 0 0
J F M A M J J A S O N D J F M A M J J A S O N D J A S O N D J F M A M J
Month Month Month
There is tropical savanna in some wet
regions where impermeable subsoil Southern Hemisphere site,
creates conditions more favorable to so months are ordered July
grasses than trees. to June.
Figure 2.17 Tropical savanna geography and climate.

Another random document with
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The drive block or the one-on-one block was developed primarily
for a fast hitting attack, and is one that we use on many occasions.
All blocks consist of the following steps: (1) the approach, (2) the
contact, and (3) the follow through.
Approach:
1. Approach with short, controlled step. First step should be with

the outside foot, and it should not go beyond the forward foot. The
knees should be bent at this time.
2. Split the opponent’s course with the first step, and aim the nose
guard at the middle of the opponent’s target.
3. Keep the eyes on the target at all times.
4. Uncoil off the forward foot. Do not duck the head during the
uncoil.
Contact:
1. Strike a jarring blow with the forehead.

2. Bring the head up as contact is made. (The body will follow the
head; this will help arch the back.)
3. The drive should be forward, then upward. The object is to
knock the opponent off-balance, thus depriving him of his traction;
this will make him easy to move out of the play.
4. Bring the feet up fast with short, choppy, digging steps, but still
keeping the knees bent.
Follow Through:
1. Stay on feet with the knees still bent and keep driving.
2. Keep the feet well spread and under the body.
3. Charge through the opponent.
4. Keep the body between the opponent and the path of the ball
carrier.
The most important single step in the drive block is keeping the
feet driving the instant contact is made.
The Reverse Shoulder Block
This block is used primarily when the blocker already has position
on a hard charging lineman.
Approach:
1. Aim the far shoulder for the opponent’s far hip. This will help
allow for the opponent’s charge.
2. Shoot the head and shoulders in front of the defensive man.
3. This movement must be sharp and on a straight line.
4. Take a short positive step with the inside foot.
Contact:
1. Uncoil off the near foot.

2. Strike a good blow on the opponent’s hip, squeezing with the
neck and head in the stomach.
3. Bring inside foot up to help cut off penetration.
Follow Through:
1. Bring the legs up fast and keep them driving.

2. The charge of the defensive man will put the blocker in a perfect
blocking position.
3. Keep the body weight into opponent.
4. If and when the opponent attempts to spin out, go into a crab
position around the outside thigh to prevent pursuit.
Blocking Linebackers
The percentage of time the offense can block the linebackers will
determine to a great extent how successful the offense will be. If the
linebackers are blocked, the defense will usually break down.
Approach:
1. Get off on snap of the ball.

2. Whenever possible release to the inside to block an inside
linebacker; to the outside when blocking an outside linebacker.
3. Stay low, don’t run in a circle, but take the shortest route to the
linebacker.
4. Get to the point of attack in position to block before the
linebacker can get there to defend it.
5. Keep your body between the linebacker and the ball carrier.
Contact:
1. Split the linebacker right down the middle with initial block.
2. Thrust the body forward and upward, hit the linebacker on the
rise and elevate the shoulders on contact.
3. Strike the blow with a low fundamental position, hit with
forehead and forearm.
4. Make contact off top of the feet. Keep the feet underneath the
blocker, thus making it impossible for the blocker to follow-through
properly if he lunges or leaves his feet before or while making
contact.
Follow Through:
1. As soon as contact is made with the linebacker, continue the leg
drive. The initial blow should knock him off balance and the leg drive
should keep him this way.
2. Keep the weight into the linebacker.
3. Gain position follow through. The blocker should continue to
strive for position to eliminate any possibility of the linebacker being
in on the tackle downfield.
Downfield Blocking
The approach of the downfield block will vary with the play called
and position of the defensive secondary men. The approach we will
discuss will be centered around attacking the defensive halfback.
The downfield block is a definite characteristic of all good football
teams, and it is virtually impossible to have many long runs without
good downfield blocking.
Approach:
1. Release rapidly on the line of scrimmage.

2. Sprint shallowly just beyond the line of scrimmage.
3. When approaching the designated person to block, aim for a
spot in front of his original position; this eliminates running behind
the halfback when he comes up to make the tackle.
4. The blocker must have a clear concept of the point of attack and
the prescribed path of the ball carrier.
5. The blocker should make the defensive man commit himself
before he starts to make contact.
Contact:
1. Once the defensive man has committed himself in a specific

direction, the blocker then starts his contact.
2. The blocker will get as close as possible to the defensive man
(about two yards) and spring off his inside foot, thus swinging his
outside elbow and arm toward the head of the opponent. This will
cause a lifting motion and make it possible for the blocker to extend
the entire length of his body in a horizontal position.
3. The blocker must aim his body at the opponent’s throat, trying
to make contact with his hip.
4. The defensive man will not allow him to hit him in the throat, but
by aiming there he may hit him around the knees.
Follow Through:
1. The follow through will consist of keeping the body extended as

long as possible and jumping up and trying to make another block.
The Post-Lead Block
The post-lead block, or two-on-one block, is used to insure the

blocking of the most dangerous defensive man, and at the same
time give the ball carrier the feeling of confidence that he can
concentrate his efforts on whipping the one defensive man to the
inside or outside of the man being double-teamed. With this in mind,
the blockers are more conscious of turning the defensive man away
from the play rather than drive him straight down the field. By using
this same method, it makes it possible to cut off the pursuit.
Approach:
(Post Man)
1. Drive straight at the defensive man with a short controlled step.

First step is with the outside foot.
2. Aim the nose guard at the middle of the target.
3. Keep the eyes on the target at all times.
4. Uncoil off the forward foot.
(Drive Man)
1. Take a position step with the inside foot and then a good step
with the outside foot to get squared away.
2. From this new position aim the nose guard at the middle of the
opponent and use as an apex. From this position the middle of the
opponent would be below the armpit and just above the waist.
3. Keep eyes on target all the way.
Contact:
(Post Man)
1. Strike a good blow with the forehead.

2. Drive straight through the opponent to stop his charge.
3. Hit on the rise and keep the pressure applied, keeping the
knees bent at all times.
(Drive Man)
1. Strike a jarring blow with the forehead below the armpit of the
opponent.
2. After good contact, slide head past body and continue applied
pressure with the inside shoulder.
3. Keep the tail down, head up, and feet under the body.
Follow Through:
(Post Man)
1. After good contact and feet are well up under the body, turn the
tail toward the drive man.
2. Keep pressure applied, feet moving in short, choppy steps, and
drive man down the line.
(Drive Man)
1. Keep pressure with shoulder and neck, and do not let man spin
out to outside.
2. Keep the tail to inside and do not allow defensive man to split.
3. Keep feet moving in short, choppy steps and move man down
the line.
4. Force defensive man into pursuit to help eliminate it.
The Trap Block
Approach:
1. It is important to take the same stance as if going straight

ahead, but just before the ball is snapped, the weight is shifted to the
outside foot.
2. Take a quick, short, 6-inch step with inside foot and point toe
directly toward spot to be trapped.
3. Never raise the body up. Stay down and in a semicoiled
position.
4. Drive off the forward foot, still taking short steps.
5. Anticipate the person to be trapped, to be filling the inside or
floating in the hole.
Contact: (see below)
Follow Through:
The contact and follow through are exactly the same as on our
drive block, which I explained previously.
The Pass Protection (Drop back pass)
This technique is used by the guards and tackles when protecting

the passer on a straight dropback pass. It is good to go from a pre-
shift position when using this type of block.
Approach:
1. Take a short step backwards with the inside foot, then a longer
step backwards with the other foot.
2. While doing this, turn the tail slightly to the inside and remember
the inside gap must be protected.
3. After the second step backwards, the blocker should be in a
good football position (tail down, head up, knees bent, back straight,
and the body in a cocked position ready to strike a blow).
Contact:
1. Look at the man to be blocked, make him come to the blocker.

When close enough, spring and butt him with the forehead, trying to
make contact with him at the numbers.
2. After contact is made and the charge of the offensive man has
been stopped, the blocker will take a step back, regain his football
position and make contact again.
3. At all times the blocker will be forcing the defensive man to the
outside.
Follow Through:
1. When the blocker sees that he can no longer strike a blow with
his head, due to the position of the defensive man, he will fake a
blow with his head, then slide it and his body in front of the defensive
man.
2. He will lock him with his leg and at the same time apply
pressure with his body.
3. At all times the blocker will be in a crab position and working his
feet around to force the defensive man to the outside, thus keeping
him away from the passer.
The Crack Back Block
This block is used primarily by the ends when they are blocking an
inside linebacker. It can also be used by anyone who might be
blocking directly in front of the ball carrier.
Approach:
1. Release from line of scrimmage as quickly as possible and run

straight at linebacker.
2. Aim slightly above the knees of the linebacker.
3. Get within a yard and a half before going into block.
Contact:
1. Throw outside arm with a quick, jerky motion in front of

linebacker. This quick motion will cause the head and shoulder to get
in front of defensive man.
2. Release off of inside foot to start block.
3. Make good solid contact with the outside hip just above the
defensive man’s knees.
Follow Through:
1. After contact is made, go into a four-point position with the arms

extended, and tail high into the air.
2. Work the feet around and force the defensive man to go in front
of blocker’s head.
3. Do not let defensive man get out behind the blocker’s feet
because this will allow him to pursue the ball from a proper angle.
4. Keep tail up and feet working.
The Junction Block
This block is used primarily by the fullback when he is blocking a

true end or a corner man on a wide play.
Approach:
1. Aim for a spot about a yard in front and outside of man to be

blocked.
2. Run low and hard at spot.
Contact:
1. Extend the head and shoulders in front of and past the

defensive man. (The blocker’s head is pointing downfield.)
2. The blocker will use his inside leg and hip to catch the outside
leg and hip of the defensive man.
3. The blocker will then have his head and shoulders past the
defensive man and his inside leg firmly against the outside leg of the
defensive man, with his hands down and his tail up.
Follow Through:
1. Force defensive man to go inside by keeping feet moving and

driving downfield.
2. Do not extend body and fall flat on the ground.
3. Do not let man spin to blocker’s outside.
4. Keep pressure on defensive man as long as possible.
The Roll Block
This block is used when a small back, in particular, is trying to

block a true end or corner man on the ground.
Approach:
1. Aim for a spot about a yard in front and outside of the man to be
blocked.
2. Run low and hard at spot.
Contact:
1. With the head pointed at the outside hip of the defensive man,
the blocker will get about a yard and a half from him. As the blocker
springs, he will drop his inside shoulder and turn his head down and
to the outside.
2. The blocker will hit the outside leg of the defensive man with the
part of his shoulder pad that covers the scapula of his inside
shoulder.
Follow Through:
1. After contact has been made with the back of shoulder, the
blocker will roll to his inside and give a corkscrew effect.
2. Stay as high as possible (around the knees) and make two or
three complete rolls in the corkscrew fashion, trying to knock the legs
out from under the defensive man.
3. At all times the blocker must keep his head pointing downfield
or no less than a 60 degree angle.
NUMBERING THE DEFENSIVE ALIGNMENTS

Figure 102a
In making our blocking rules or assignments for all of the players

versus all different defensive alignments, there are several factors to
take into consideration. The rules must be simple, and secondly,
they must be brief. We have used several different kinds of rules,
such as, “Inside gap, over, linebacker,” and others. This was a good
method, but it amounted to quite a bit of memory work for the players
because the majority of the blocks were all different and the players
were required to learn a number of different sequences. Trying to
adhere to the theory, “the simpler, the better,” we started numbering
the defensive men as illustrated in Figure 102a versus the 5-4
defensive alignment; 102b versus the wide tackle 6; 102c versus the
gap 8 defense; and Figure 102d versus the Eagle defense.
Figure 102b
Figure 102c
Figure 102d
We start counting with the man over the center and number him
zero, and from there go both left and right numbering every man
within two yards of the line of scrimmage, as illustrated in Figures
102a-b-c-d. We also number the men in the secondary. We do this
by merely continuing with our numbering beyond the end lineman in
the direction the ball is going to go, as illustrated in Figure 103a
versus the Oklahoma 5-4 defense, and Figure 103b versus the wide
tackle 6-2 defensive alignment.
Figure 103a
Figure 103b
Application of Our Blocking Rules
By numbering the defensive men it is simple when making out the

blocking rules to assign each man a number, and the offensive man
merely learns which numbered man he will block. There will be
certain plays in which a particular position must have one or two
options. As an example, a rule might read, “#3 unless outside, then,
#2.”
The following are examples of blocking rules by numbers:
Regular Block—straight ahead
On-E #3
On-T #2
On-G #1
C #0, off-side
Off-G #1
Off-T #5
Off-E #6
Trap Block—
On-E #3
On-T L.B.
On-G Slam man #1 or #2, N/T #0
C Off-side
Off-G Trap
Off-T 1st outside guard
Off-E #6
Counter Block—
On-E #6
On-T #2
On-G #1
C #1, N/T #0, N/T off-side
Off-G #1
Off-T #2
Off-E #3
By using this particular method it is also very easy to incorporate

the backs into the blocking scheme. When using the back to the side
the ball is going, in order to have a backfield man block a particular
defensive player, we merely add two zeroes to the number which
has been assigned to the defender, and the back gets this man. As
an illustration, if we want our halfback to block the #3 defensive man,
the halfback’s block is 300; and the #4 man, the block would be
called 400. Several complete illustrations would be as follows:
300 Block—
Off-E #6
Off-T #5
Off-G #1
C #0, N/T off-side
On-G #1
On-T #2
On-E slam #3, then #5
H.B. #3
400 Block—
Off-E #6
Off-T #5
Off-G #1
C #0, N/T off-side
On-G #1
On-T #2
On-E #3
H.B. #4
Under Block—
Off-E #6
Off-T #5
Off-G #1
C #0, N/T off-side
On-G #1
On-T #2
On-E #3
H.B. #4
Over Block—
Off-E #6
Off-T #5
Off-G #1
C #0, N/T off-side
On-G 1st man on line of scrimmage
On-T 2nd man on line of scrimmage
On-E 3rd man on line of scrimmage
H.B. 1st inside linebacker
ATTACKING THE DEFENSIVE ALIGNMENT

Probably like many other football teams, we have too much
offense. However, in order to do an intelligent job of planning our
attack versus the numerous defensive alignments, one must have
sufficient offense since not all plays are good against all defenses.
Regardless of one’s offense, the first approach is to establish a
good sound middle attack that is based on the trap, fullback hand-
off, and a pass off of the same action. Unless a team can force the
defense to respect the inside power and force the defensive
linebackers and guards to stay “at home,” it is almost an impossibility
to perfect the outside attack. After we establish the inside attack,
then we want to run far enough inside the defensive end to cut down
his quick containment. The next step then is to perfect the wide
attack and the corner passes, in order to have a well-rounded
offense. Bootleg passes, reverses, and an occasional trick play are
also needed in order to keep the defense “honest,” and to make the
above-mentioned plays more effective.
GOING WIDE
It is an offensive must for a team to be able to go wide and to get
the long gainer. Occasionally all of us get a good gainer from the
inside attack, but most long gainers are from passes or some form of
wide attack.
Previously we operated on the assumption if we could gain four
yards on each play we would score with a sustained drive. Statistics
will prove a team will generally stop itself by some error, or the
defense will stop the offense, before the attack can make four
consecutive first downs or gain 50 yards, a majority of the time.
Consequently a team must perfect its wide game.
There are several ways of going wide, but regardless of the
manner attempted, the defensive end or corner man must be
eliminated either by blocking him, optioning him, or throwing the
football over his area.
The Run-Pass Option
When trying to get wide against a corner man, the run or pass
option is one of the best methods to employ. Figure 104 illustrates
the on-guard blocking #1, on-tackle #2, and the fullback blocking the
#3 man. The near halfback and the on-end will run their pass routes.
The #4 man is not blocked, and the passer is going to option from
him. As soon as the passer has possession of the ball, he watches
the #4 man. If he comes up to tackle the passer, the latter throws to
the open receiver. If the #4 man drops back to play pass defense,
the passer is instructed to turn upfield and run with the football. If our
passer feels he can gain at least four yards on the play, we want him
to run with it most of the time. The play is much better against a
team that uses a 9-man front than against an 8-man front alignment.
Figure 104
The Quarterback Option
There are several kinds of option plays, but the one designed to
get wide is the two-way option, in which the quarterback either keeps
or pitches to the far halfback, depending upon the reaction of the
defensive man from whom he is optioning.
The blocking rules on the quarterback option play are very simple,
as illustrated in Figure 105. The offensive guard and tackle to the on-
side block their numbers, respectively. The on-end will block to his
inside, N/T (No One There) halfback, and the fullback will block the
#4 man with a junction block. The quarterback must watch his man
all the time in order for the play to turn the corner successfully. He
must option the #3 man and not permit the latter to option the
quarterback.
Figure 105
The Three-Way Option
Figure 106
The other option play, illustrated in Figure 106, is the three-way

option with the quarterback playing the #4 defender. This option is in
the same series with the outside belly play. The quarterback has the
option of (1) giving the ball to the fullback, who will hit off-tackle, (2)
keeping the ball himself, after faking the fullback off-tackle, or (3)

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Ecology: Concepts and Applications

(Ninth Edition) Manuel C. Molles Jr.

Ecology: Concepts and Applications (Ninth Edition)

Ecology: Concepts And Applications 5th Edition Manuel

Miller s Basics of Anesthesia Manuel C. Pardo Jr.

Primary Mathematics 3A Hoerst

Business Ethics: Concepts and Cases -- Books a la Carte

Structural Geology Principles, Concepts, and Problems

Classical Hopf Algebras and Their Applications Algebra

Cryptocurrency: Concepts, Technology, and Applications

This book is printed on acid-free paper.

Cover Image: Anna A. Sher

moL86337_fm_ISE.indd ii 04/29/21 11:37 PM

Section Natural History and Evolution 11

Section Adaptations to the Environment 101

Section Population Ecology 196

12 Life Histories 254

Section Interactions 277

IV 13 Species Interactions and Competition 277

Section Communities and Ecosystems 345

19 Nutrient Cycling and Retention 403

Section Large-Scale Ecology 445

VI 21 Landscape Ecology 445

Appendix A Investigating the Evidence 514

Design elements: Anna A. Sher

Historical Foundations and Aquatic Biomes and How They Differ 45

Section I Lakes: Small Seas 69

4.5 Change due to Chance 92 Water Acquisition by Plants 135

Section II Marine Fish and Invertebrates 143

8.5 Eusociality 189 10.3 Patterns of Survival 224

Section IV “Refuges” 312

Chapter 20 Succession and Stability 423 Chapter 22 Geographic Ecology 468

SECTION VI Chapter 23 Global Ecology 490

Appendix A Investigating the Evidence

Design elements: Anna A. Sher

the biomes. In summary, the book begins with an overview

variation in tem ima

tive pedagogical tools through skillful design and

This text carefully explains all mathematical expres- 92%

sions that arise to help students overcome these Figure 7.16

of planktivorous fish in this

e survival patterns for

First Pages First Pages

communities. speciesrichness ants, termites, yants,leaf-litter 9.17 Describ ies to extinction.

moL22201_fm_i-xx.indd xvi 12/08/21 09:37 AM

Design elements: Anna A. Sher

Study made personal

Laptop: McGraw Hill; Woman/dog: George Doyle/Getty Images

Affordable solutions, Solutions for

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Study anytime, anywhere “I really liked this

Everything you need in one place

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Learning for everyone

New remote proctoring and browser-locking capabilities,

1.1 Discuss the concept of environment as it pertains to

such as photosynthesis and decomposition. Ecologists often Individuals

How do species interact?

1.2 Describe the levels of ecological organization, for

How do vegetated corridors

The next level of organization is the ecosystem. An eco-

Concept 1.1 Review (b)

Cape May warbler Blackburnian warbler Black-throated

moL86337_fm_ISE.indd ii 04/29/21 11:37 PM

communities. speciesrichness ants, termites, yants,leaf-litter 9.17 Describ ies to extinction.

moL22201_fm_i-xx.indd xvi 12/08/21 09:37 AM

availability (fig. 2.1b). If we plot the combination of these two vari-