Ecology 2022 PDF

 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 U...

 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 database or retrieval system, without the prior written consent of McGraw Hill LLC, including, but not limited to, in any network or other electronic storage or transmission, or broadcast for distance learning. Some ancillaries, including electronic and print components, may not be available to customers outside the United States. This book is printed on acid-free paper. 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 All credits appearing on page or at the end of the book are considered to be an extension of the copyright page. The Internet addresses listed in the text were accurate at the time of publication. The inclusion of a website does not indicate an endorsement by the authors or McGraw Hill LLC, and McGraw Hill LLC does not guarantee the accuracy of the information presented at these sites. mheducation.com/highered moL86337_fm_ISE.indd ii 04/29/21 11:37 PM  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 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 Concept 18.3 Review 391 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 Concept 16.4 Review 360 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 Concept 17.1 Review 368 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 Concept 21.1 Review 450 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 Concept 23.2 Review 500 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 Concept 23.3 Review 504 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 Design elements: Anna A. Sher  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 moL22201_ch 05_101-126.indd 101 02/16/21 10:33 AM 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