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 Final PDF to printer

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
of this publication may be reproduced or distributed in any form or by any means, or stored in a
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1 2 3 4 5 6 7 8 9 LWI 26 25 24 23 22 21

ISBN 978-1-265-28633-0
MHID 1-265-28633-7

Cover Image: Anna A. Sher

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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

Design elements: Anna A. Sher

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
Concept 2.1 Review 16
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
Concept 4.1 Review 84
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
Concept 6.2 Review 141
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
Concept 7.1 Review 153
Concept 5.1 Review 105
7.2 Chemosynthetic Autotrophs 154
5.2 Evolutionary Trade-Offs 105
Concept 7.2 Review 156
The Principle of Allocation 105
7.3 Heterotrophs 156
Concept 5.2 Review 106
Chemical Composition and Nutrient Requirements 156
5.3 Temperature and Performance of Organisms 106
Concept 7.3 Review 162
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
Concept 7.4 Review 165
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
Concept 5.5 Review 123
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
Concept 10.3 Review 228
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
Concept 9.1 Review 202
Exponential Growth in Nature 241
9.2 Patterns on Small Scales 202
Concept 11.1 Review 242
Scale, Distributions, and Mechanisms 202
11.2 Logistic Population Growth 242
Distributions of Tropical Bee Colonies 202
Concept 11.2 Review 245
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
Concept 11.3 Review 248
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
Concept 9.4 Review 210
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

Concept 14.3 Review 314
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
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