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PE5eFM New.qxd4/9/1011:28 AMPage iiiESSENTIALS OFCONSERVATIONBIOLOGYFIFTH EDITIONRichard B. PrimackBoston UniversitySinauer Associates, Inc., PublishersSunderland, Massachusetts U.S.A. Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

PE5eFM New.qxd4/9/1011:28 AMPage viBrief ContentsPART I Major Issues That Define the Discipline 11 What Is Conservation Biology? 32 What Is Biological Diversity? 233 Where Is the World’s Biological Diversity Found? 51PART II Valuing Biodiversity 694 Ecological Economics and Direct Use Values 715 Indirect Use Value 916 Ethical Values 115PART III Threats to Biological Diversity 13178910Extinction 133Vulnerability to Extinction 155Habitat Destruction, Fragmentation, Degradation, and Global Climate Change 173Overexploitation, Invasive Species, and Disease 215PART IV Conservation at the Population and Species Levels 24511121314Problems of Small Populations 247Applied Population Biology 273Establishing New Populations 295Ex Situ Conservation Strategies 313PART V Practical Applications 3391516171819Establishing Protected Areas 341Designing Networks of Protected Areas 367Managing Protected Areas 389Conservation Outside Protected Areas 415Restoration Ecology 437PART VI Conservation and Human Societies 45920 Conservation and Sustainable Development at the Local and National Levels 46121 An International Approach to Conservation and Sustainable Development 49322 An Agenda for the Future 523 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

PE5eFM New.qxd4/9/1011:28 AMPage viiContentsPART I Major Issues That Define the Discipline1CHAPTER 1 What Is Conservation Biology? 3The New Science of Conservation Biology 5Conservation Biology Complementsthe Traditional Disciplines 6Conservation Biology Is a Crisis Discipline 7Conservation Biology’s Ethical Principles 7BOX 1.1 Conservation Biology’s InterdisciplinaryApproach: A Case Study with Sea Turtles 8CHAPTER 2A New Science Is Born 19Conservation Biology: A Dynamic andGrowing Field 1923BOX 2.2 Kelp Forests and Sea Otters: Shaping anOcean Ecosystem 3725BOX 2.1 Naming and Classifying Species27The Origin of New Species 29Measuring Species Diversity 30Genetic Diversity 33Ecosystem Diversity 36What Are Communities and Ecosystems?CHAPTER 3European Origins 13American Origins 16What Is Biological Diversity?Species Diversity 24What Is a Species?The Origins of Conservation Biology 1136Ecological Succession 39Species Interactions within Ecosystems 40Principles of Community Organization 40Keystone Species and Guilds 44Keystone Resources 47Ecosystem Dynamics 48Conclusion 49Where Is the World’s Biological Diversity Found?Two of the Most Diverse Ecosystems onEarth 52Tropical Rain ForestsCoral Reefs 5353Patterns of Diversity 54Variation in Climate and Environment 54Variation in Topography, Geological Age, andHabitat Size 55Why Are There So Many Species in theTropics? 5651How Many Species Exist Worldwide? 58New Species Are Being Discovered All the Time58Recently Discovered Communities 60BOX 3.1 Conserving a World Unknown:Hydrothermal Vents and Oil PlumesDiversity Surveys: Collecting and CountingSpecies 62Estimating the Number of Species 63The Need for More Taxonomists 66 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.62

PE5eFM New.qxdviii4/9/1011:28 AMPage viiiContentsPART II Valuing BiodiversityCHAPTER 4Ecological Economics and Direct Use ValuesWhy Economic Valuation Is Needed 72Cost-Benefit AnalysisDirect Use Values 8174Consumptive Use Value 81Productive Use Value 84Multiple Uses of a Single Resource:A Case Study 87Natural Resource Loss and the Wealth ofSocieties 76BOX 4.1 Industry, Ecology, and Ecotourism inYellowstone Park 79Indirect Use Value91Nonconsumptive Use Value 91Ecosystem Productivity and Carbon Sequestration 93Protection of Water and Soil Resources 94BOX 5.1 Prophecy Fulfilled: How EcosystemServices Became Front Page News 96Waste Treatment and Nutrient RetentionClimate Regulation 98Species Relationships 9998Ethical ValuesEnvironmental Monitors 101Recreation and Ecotourism 101Educational and Scientific Value 104The Long-Term View: Option Value 104BOX 5.3 Mighty Multitudes of Microbes:Not to Be Ignored! 106Existence Value 109BOX 5.2 How Much Are Bats Worth? A Case Studyof Texas Bats 100CHAPTER 6Is Economic Valuation Enough? 111115Ethical Values of Biological Diversity 116BOX 6.2 Religion and ConservationBOX 6.1 Sharks: A Feared Animal in Decline118Deep Ecology 126PART III Threats to Biological DiversityExtinction122Enlightened Self-Interest: Biodiversity andHuman Development 124Ethical Arguments for Preserving BiologicalDiversity 117CHAPTER 771Assigning Economic Value to BiologicalDiversity 80Evaluating Development Projects 74CHAPTER 569131133Past Mass Extinctions 134The Current, Human-Caused Mass Extinction 136Background Extinction Rates 141Extinction Rates on Islands 141Extinction Rates in Aquatic Environments 142BOX 7.1 Invasive Species and Extinction in IslandEcosystems 143Estimating Extinction Rates with the IslandBiogeography Model 145Extinction Rates and Habitat Loss 147Assumptions and Generalizations in the IslandBiogeography Model 149Time to Extinction 149Local Extinctions 150 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

PE5eFM New.qxd4/9/1011:28 AMPage ixContentsCHAPTER 8Vulnerability to Extinction155Endemic Species and Extinction 156Conservation Categories 165Species Most Vulnerable to Extinction 158Natural Heritage Data Centers 169BOX 8.1 Why Are Frogs and Toads Croaking?CHAPTER 9ix163Habitat Destruction, Fragmentation, Degradation,and Global Climate Change 173Human Population Growth and Its Impact 174Pesticide PollutionHabitat Destruction 177BOX 9.1 Pesticides and Raptors: Sentinel SpeciesWarn of Danger 198Threatened Rain Forests 180Other Threatened Habitats 184Marine Coastal Areas 185Desertification 187Water Pollution 198Air Pollution 201Global Climate Change 204Changes in Temperate and Tropical ClimatesPlants and Climate Change 209Rising Sea Levels and Warmer Waters 209The Overall Effect of Global Warming 211Habitat Fragmentation 189Edge Effects 193Two Studies of Habitat Fragmentation195Habitat Degradation and Pollution 196CHAPTER 10Overexploitation197Overexploitation, Invasive Species, and Disease215230Invasive Species in Aquatic Habitats 232The Ability of Species to Become Invasive 234Control of Invasive Species 236Exploitation in the Modern World 217International Wildlife Trade 218BOX 10.1 Endangered Whales: Makinga Comeback? 220Disease 237225Invasive Species 226Invasive Species on Islands215BOX 10.2 GMOs and Conservation BiologyCommercial Harvesting 224What Can Be Done to Stop Overexploitation?208Implications of Invasive Species and Diseases forHuman Health 241Conclusion 242228PART IV Conservation at the Populationand Species Levels 245CHAPTER 11Problems of Small PopulationsEssential Concepts for Small Populations 248Minimum Viable Population (MVP) 248Loss of Genetic Variability 250Consequences of Reduced Genetic VariabilityFactors That Determine Effective PopulationSize 257247Other Factors That Affect the Persistence of SmallPopulations 264Demographic Variation 264254Environmental Variation and Catastrophes 266Extinction Vortices 268BOX 11.1 Rhino Species in Asia and Africa: GeneticDiversity and Habitat Loss 262 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

PE5eFM New.qxdx4/9/1011:28 AMPage xContentsCHAPTER 12Applied Population BiologyMethods for Studying Populations 275Gathering Ecological InformationMonitoring Populations 276Population Viability Analysis 285275Metapopulations 287Long-Term Monitoring of Species andEcosystems 290BOX 12.1 Three Primatologists Who BecameActivists 279CHAPTER 13273Establishing New PopulationsThree Approaches to Establishing NewPopulations 296BOX 13.1 Wolves Return to a Cold Welcome295Learned Behavior of Released Animals302Establishing New Plant Populations 305297The Status of New Populations 309Successful Programs with Animals 299CHAPTER 14Ex Situ Conservation StrategiesEx Situ Conservation Facilities 316ZoosBotanical Gardens and ArboretumsSeed Banks 330316BOX 14.1 Love Alone Cannot Save the GiantPanda 317Aquariums313328BOX 14.2 Seed Savers and Crop Varieties333Conclusion 336326PART V Practical ApplicationsCHAPTER 15Establishing Protected AreasEstablishment and Classification of ProtectedAreas 342Existing Protected Areas 343Marine Protected Areas345BOX 15.1 The Phoenix Islands Protected Area: TheWorld’s Largest Marine Park 346The Effectiveness of Protected AreasCHAPTER 16341Creating New Protected Areas 349Prioritization: What Should Be Protected? 351Determining Which Areas Should Be Protected 352Linking New Protected Areas to ReserveNetworks 360Gap Analysis361347Designing Networks of Protected AreasIssues of Reserve Design 368Protected Area Size and Characteristics 369Reserve Design and Species Preservation 373Minimizing Edge and Fragmentation Effects 374Networks of Protected Areas 375Habitat Corridors339367BOX 16.1 Ecologists and Real Estate Experts Mingleat The Nature Conservancy 377Habitat Corridor Case Studies380Landscape Ecology and Park Design 382Conclusion 386375 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

PE5eFM New.qxd4/9/1011:28 AMPage xiContentsCHAPTER 17Managing Protected Areas389Management and People 402Monitoring as a Management Tool 392Identifying and Managing Threats 394Managing Invasive SpeciesxiBOX 17.2 Managing Leopards Togetherwith People 403394Zoning to Separate Conflicting DemandsManaging Habitat 396BOX 17.1 Habitat Management: The Key to Successin the Conservation of EndangeredButterflies 397404Regulating Activities inside Protected Areas 407BOX 17.3 Is Arctic Wildlife Management Compatiblewith Oil Drilling? 408Challenges in Park Management 410Managing Water 399Managing Keystone Resources 401CHAPTER 18Conservation Outside Protected Areas415The Value of Unprotected Habitat 417Ecosystem Management 427Conservation in Urban Areas 420Case Studies 430Managed Coniferous Forests 430African Wildlife Outside Parks 432Community-Based Wildlife Managementin Namibia 432BOX 18.1 In Defense of Wildlife . . . Send in theSoldiers 421Conservation in Agricultural Areas 423Multiple Use Habitat 425CHAPTER 19Restoration EcologyDamage and Restoration 439Restoration in Urban Areas 445BOX 19.1 Can Many Small Projects Clean Up theChesapeake Bay? 441Ecological Restoration Techniques 442Practical ConsiderationsRestoration of Some Major Communities 447Wetlands447BOX 19.2 The Kissimmee River: Restoring aChannelized River to Its Natural State 448443Case Studies 445Wetlands Restoration in Japan 445The Grand Canyon–Colorado River Ecosystem437445Lakes 449Prairies 451Tropical Dry Forest in Costa Rica453The Future of Restoration Ecology 455 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.

PE5eFM New.qxdxii4/9/1011:28 AMPage xiiContentsPART VI Conservation and Human Societies459CHAPTER 20 Conservation and Sustainable Development at the Local andNational LevelsConservation at the Local LevelLand Trusts461463Conservation Beliefs 478Conservation Efforts That Involve TraditionalSocieties 480463BOX 20.1 How Clean Is “Green”Energy?Local Legislation465BOX 20.2 People-Friendly Conservation in theHills of Southwest India: Successesand Failures 485467Conservation at the National Level 469National Legislation 469The U.S. Endangered Species ActEvaluating Conservation Initiatives That InvolveTraditional Societies 489471Traditional Societies, Conservation, andSustainable Use 477CHAPTER 21 An International Approach to Conservation and SustainableDevelopment493International Agreements to Protect Species 495Reforming Development LendingBOX 21.2 How Much Will the Three Gorges DamBOX 21.1 The War for the Elephant: Is the ArmisticeOver?Really Cost?497National Environmental FundsDebt-for-Nature Swaps 517Marine Environments 518International Earth Summits 502Funding for Conservation 506The Role of International DevelopmentBanks 509BOX 22.1 Conservation Education: Shaping theNext Generation into Conservationists 525The Role of Conservation Biologists 531Glossary523Challenges for Conservation Biologists531BOX 22.2 Environmental Activism Confronts theOpposition 532Achieving the Agenda533543545BibliographyIndexIncreased Funding Is Necessary for the Future539Chapter Opener Photograph Credits516How Effective Is Conservation Funding? 518An Agenda for the FutureOngoing Problems and Possible Solutions 524Appendix512Funding Sources and Programs 515International Agreements to ProtectHabitat 499CHAPTER 22511553587 Sinauer Associates, Inc. This material cannot be copied, reproduced, manufacturedor disseminated in any form without express written permission from the publisher.519

The New Science of Conservation Biology 5 Conservation Biology Complements the Traditional Disciplines 6 Conservation Biology Is a Crisis Discipline 7 Conservation Biology’s Ethical Principles 7 BOX 1.1 Conservation Biology’s Interdisciplinary Approach: A Case Study with Sea Turtles 8 The Origins of

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