Principles of Conservation Biology

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1 3 0^ ^ rrv' \f Principles of Conservation Biology Gary K. Meffe University of Georgia's Savannah River Ecology Laboratory C. Ronald Carroll Institute of Ecology, University of Georgia and Contributors SINAUER ASSOCIATES, INC. Publishers Sunderland, Massachusetts

2 Contents Contributing Chapter Authors Preface xv xiv Part I Introductory Concepts 1. What Is Conservation Biology? 3 Environmental Problems and Human Population Growth 3 A Brief History of Conservation Biology 7 Conservation in the United States 10 Modern Conservation Biology: A Synthesis 13 Guiding Principles for Conservation Biology 15 Some Postulates of Conservation Biology 19 Some Characteristics of Conservation Biology 19 A Crisis Discipline 19 A Multidisciplinary Science 20 An Inexact Science 21 A Value-Laden Science 21 A Science with an Evolutionary Time Scale 22 A Science of Eternal Vigilance 22 A Final Word 22 ESSAY A: The Role of the University in Conservation Biology 5 ESSAY B: Conservation Biology and the Management of Natural Resources 9 ESSAY C: The Role of Science in Defining Conservation Priorities for Nongovernmental Organizations Conservation Values and Ethics 24 The Value of Biodiversity 24 Instrumental Value 24 Intrinsic Value 27 Monetizing the Value of Biodiversity 30 Conservation Ethics 35 Anthropocentrism 35 The Judeo-Christian Stewardship Conservation Ethic 36 Traditional Non-Western Environmental Ethics 37 Biocentrism 42 Ecocentrism 44 ESSAY A: Our Duties to Endangered Species 30 ESSAY B: Monks, Temples, and Trees: The Spirit of Diversity 40 ESSAY C: Discovering Radical Environmentalism in our Own Cultural Backyard The Species in Conservation 50 Views on Species 50 Typological versus Populational Thinking 51 Species Concepts 53 The Species and Conservation 59 How Do Species Concepts Affect Conservation Efforts? 59 Conserve Diversity, Not Latin Binomials 61 Is the Species Category Useful to Conservation? 62 Taxonomic Problems in Conservation 63 Species and the Law: The ESA and CITES 66 How Will Environmental Degradation Affect Speciation? 70, If There Are 30 Million or So Species, Can Conservation Be Based on a Species Approach? 73 ESSAY A: Recognition of Freshwater Mussel Taxa: A Conservation Challenge 57 ESSAY B: Systematics and Conservation 64 ESSAY C: When Endangered Species Hybridize: The U.S. Hybrid Policy 69 ESSAY D: Law and Protection of Biodiversity Global Biodiversity I: Patterns and Processes 78 The Levels of Biodiversity 78 Genetic Diversity 78 Intraspecific Diversity 81 Species Richness 81 Richness of Higher Taxa 85 ; Ecosystem and Biome Diversity 86 Patterns of Species Richness 88 Species Richness over Geologic Time 88

3 Local Species Richness 90 Regional Turnover of Species Richness 92 What Are the Limits to Species Richness? 95 Evolutionary Limits to Species Richness 95 Ecological Limits to Species Richness 97 The Future of Biodiversity Studies 103 ESSAY A: Hierarchical Indicators for Monitoring Changes in Biodiversity 79 ESSAY B: A 50-Year Plan for Biodiversity Surveys 83 ESSAY C: Uncharted Waters: Conserving Marine Biological Diversity 85 ESSAY D: A New Paradigm in Conservation of Biodiversity: Endangered Biological Phenomena Global Biodiversity II: Losses 110 Overview of Mass Extinctions 110 Critical Regional Losses: Hot Spots of Extinction 123 Patterns of Species Vulnerability 124 Rare Species 124 Long-Lived Species 128 Keystone Species 129 Genetic Losses 130 Habitat Fragmentation and Degradation 131 Synergistic Interactions 132 The Problem of Stressed Biotas 134 The Future of Evolution: A Process at Risk 135 Concluding Thoughts 138 Box A: Quantifying Patterns of Deforestation 114 ESSAY A: Extinction of Birds on Barro Colorado Island 118 Box B: Quantifying the Loss of Species Due to Tropical Deforestation 121 ESSAY B: Setting Priorities for Biodiversity Conservation: One Organization's Approach 125 Part II Population-Level Concerns 6. Genetics: Conservation of Diversity within Species 143 Why Genetics? 144 Genetic Variation: What Is It and Why Is It Important? 145 Variation within Individuals 146 Variation within Populations 147 Variation among Populations 148 The Fitness Consequences of Variation 149 Loss of Genetic Variation 153 Management of Genetic Variation in Natural Populations 158 Time Scales of Concern 159 Units of Conservation 159 Hierarchical Gene Diversity Analysis 161 Biogeographic Models of Gene Flow 166 Depths of Evolutionary Separation 169 "Cookbook Prescriptions" in Genetic Conservation 170 Other Uses of Genetic Information in Conservation 172 The Limitations of Genetics in Conservation 173 ESSAY A: Coadaptation, Local Adaptation, and Outbreeding Depression 152 ESSAY B: Genetically Effective Sizes of Grizzly Bear Populations 155 ESSAY C: A Rose Is a Rose Is a Rose 174 ESSAY D: EX Situ Conservation and the Conservation Role of Zoological Parks and Aquariums Demographic Processes: Population Dynamics on Heterogeneous Landscapes 179 What Is Population Demography? 180 Mechanisms of Population Regulation 183 A Hierarchical Approach to Population Regulation 185 Habitat-Specific Demography 186 Sources and Sinks 186 Metapopulation Concepts 189 Population Viability Analysis 190 Empirical Studies of Population Persistence 196 The Landscape Approach 197 Spatially Explicit Models 199 ESSAY A: Contributions of Long-Term Life History Studies to Conservation Biology 181 Box A: The Theory of Sources and Sinks 187 Box B: Thresholds of Population Responses 192 ESSAY B: Population Viability Analysis: Determining Nature's Share 195 Box C: Conservation of Nontarget Species in Managed Landscapes 200 Part III System-Level Concerns 8. Community-Level Conservation: Species Interactions, Disturbance i Regimes, and Invading Species 209 Critical Species Interactions 210 The Role of Keystone Species 210 Mutualisms 212

4 Indirect Effects and Diffuse Interactions 214 Community Changes in Ecological Time: Disturbance Regimes and Invasive Species 216 The Importance of Disturbance Regimes 216 Species Invasions 218 Community Changes in Evolutionary Time 230 ESSAY A: The Evolution and Importance of Species Interactions 213 ESSAY B: Maintaining an Open Mind on Exotic Species Habitat Fragmentation 237 Fragmentation and Heterogeneity 238 The Fragmentation Process 241 Insularization and Area Effects 243 Biological Consequences of Fragmentation 246 Initial Exclusion 247 Barriers and Isolation 247 Crowding Effect 251 Local and Regional Extinctions Edge Effects 254 Changes in Species Composition 256 The Problem of Climate Change 258 Conclusions and Recommendations 260 ESSAY A: Mosaics and Patch Dynamics 242 ESSAY B: Habitat "Shredding" The Design of Conservation Reserves 265 Considerations and Goals of Reserve Design 268 Goals of Reserves 270 Concerns in Reserve Design 272 Six Critical Issues in Reserve Success 273 Reserve Size 273 Heterogeneity and Dynamics 277 Context 281 Connecting Fragmented Habitats 284 Natural and Modified Landscape Elements 290 Buffer Zones 293 The Problem of Certain Change and Uncertain Direction 294 Anthropological and Cultural Implications of Reserve Design 296 Political and Economic Constraints on Reserve Design 300 ESSAY A: University of California Granite Mountains Reserve 267 ESSAY B: A Justification for the Conservation of Large Reserves 276 ESSAY C: Creating Regional Reserve Networks 289 ESSAY D: Designing Landscapes and Regions to Conserve Nature 292 ESSAY E: Pupfish, Politics, and Preserve Management in the Arid Southwest 301 Part IV Practical Applications and Human Concerns 11. Management to Meet Conservation Goals: General Principles 307 Why Is Management Necessary? 307 Five Basic Principles of Good Conservation Management 312 Principle 1: Critical Ecological Processes Must Be Maintained 313 Principle 2: Goals and Objectives Must Comefroman Ecological Understanding of the System 314 Principle 3: External Threats Must Be Minimized and External Benefits Maximized 316 Principle 4: Evolutionary Processes Must Be Conserved 321 Principle 5: Management Must Be Adaptive and Minimally Intrusive 321 Management Principles in Practice: Controlling Invasive Species 323 Different Scales of Management 325 Living with Uncertainty and Risk 326 Four Causes of Uncertainty 330 Cultural Context of Management Decisions 332 ESSAY A: Conservation Biology and Wildlife Management in America 310 ESSAY B: Multiple Objectives in Planning: Economic and Ecological Solutions 315 ESSAY C: Retaining Indigenous Knowledge Systems as a Management Tool 319 Box A: Examples of Species Using Multiple Habitats 326 ESSAY D: Decision Analysis in Conservation Biology Management to Meet Conservation Goals: Applications 336 How Are Management Decisions Made? 336 Managing at the Population Level 341 Managing for Sustained Yields 341 Conflicts in Single-Species Management: Lessons from Game Management 344 Managing Species as Surrogates for the Larger Community 347 Habitat Management 349 Assessing Habitat Quality 351 Managing at the Community and Ecosystem Levels 352 Managing Ecosystem Processes 354 A Caveat: Do We Need to (or Can We) Save All Species? 355 The Landscape Scale \ Critical Management Issues 358. Conflicts Created by Multiple-Use Management 358

5 Mimicking Nature: Disturbance Type, Frequency, Intensity, and Scale 362 Priorities for Management: Sensitive Natural Areas and Vulnerable Processes 364 ESSAY A: The Importance of Value Systems in Management: Considerations in Desert Fish Management 340 ESSAY B: An Emerging Management Tool: Large Mammal Predator Reintroductions 346 ESSAY C: Seasonal, Freshwater Marshes in the Tropics: A Case in which Cattle Grazing is Not Detrimental 352 ESSAY D: Agency Multiple-Use Conflicts 359 ESSAY E: The Use of Fire in Habitat Management Conservation Management Case Studies 368 Case Study 1. Management of an Endangered Species: The Black-Footed Ferret 369 Causes of Decline 369 Recent History 369 Roadblocks to Recovery 372 Conclusions 373 Case Study 2. Sea Turtle Headstarting and Hatchery Programs 374 Case Study 3. Management of the Spotted Owl: The Interaction of Science, Policy, Politics, and Litigation 380 The Northern Spotted Owl 381 The California Spotted Owl 382 Hypotheses Tested and Reserve Design Principles Invoked 382 Why Two Different Conservation Strategies? 384 Issues that Arise after the Conservation Plan Is Put Forward 386 Improving the Role of Science in Conservation Policy 387 Case Study 4. Management of a Multiple-Function Reserve: The La Selva Biological Station 388 Multiple-Function Reserves 388 Identifying the Mission: What Are the Goals? 390 Defining the Mission: Who Has the Power? 390 Mission and Management: How Well Do They Correspond? 391 Reserve Functions and Long-Term Sustainability 392 Conclusion: From General Principles to Real-World Decisions 394 Case Study 5. Management of Conservation Units: The Costa Rican National System of Conservation Areas 395 An Innovative Approach 396 Funding Strategies: A Key to Success 401 Success or Failure? 403 Conclusions 404 ESSAY A: Debt-for-Nature Swaps Ecological Restoration 409 What Is Ecological Restoration? 409 The Role of Ecological Restoration in Conservation 411 Some Central Concerns of Restoration Ecology 412 The Product 412 Feasibility and Authenticity 413 Scale 418 Costs 420 The Restoration Process 420 A Succession Primer 421 Examples of Restoration Projects 423 Restoration Following Mining Activities in Arid Lands 424 Restoration of a Tropical Dry Forest in Guanacaste National Park 426 Restoration of Native Prairies 429 Examples of Aquatic Restoration 432 General Observations on Restoration 434 Box A: Definitions of Common Terms Used in Restoration Ecology 410 ESSAY A: Measuring Biological Integrity 415 ESSAY B: Restoring a Nation's Wetlands: Why, Where, and How? Ecology, Politics, and Economics: Finding the Common Ground for Decision Making in Conservation 439 Economic Philosophy and the Roots of the Biological Crisis 440 Background 440 Adam Smith and the Invisible Hand 441 Thomas Malthus and Population Growth 442 David Ricardo and the Geographic Pattern of Economic Activity 443 John Stuart Mill and the Steady State 444 Karl Marx and the Ownership of Resources 446 A. C. Pigou and Market Failure 447 Common Property Management 450 Harold Hotelling and the Efficient Use of Resources over Time 451 The Rate of Interest 452 Preserving Biodiversity as an Option 453 The Economics of Biodiversity Conservation 454 New Directions in Economic Thought 455 The Rights of Future Generations 455 Distancing and Economic Globalization 458 The Coevolution of Modern Societies 459 Ecological Economics 463 ESSAY A: Steady-State Economics 444 '. ESSAY B: Valuation of Ecological Systems 449 ESSAY C: Valuation of Extractive Medicines in Tropical Forests: Exploring the Linkage to Conservation 457

6 16. The Role of Institutions and Policymaking in Conservation 466 Types of Institutions and Their Roles in Conservation 468 Educational Institutions 468 Governments 471 Nongovernmental Organizations 473 Business 474 Consumers 475 Legal Institutions 475 Religious Institutions 476 The Media 477 Institutional and Policy Challenges for Conservation Biology 478 Defining Appropriate Environmental Research 478 Incorporating Broader Sources of Information 479 Understanding the Policy Process 480 The Nature of Knowledge in Environmental Policymaking 482 The Nature of the University and the Problems of Research 485 The Sufficiency of Knowledge and Some Roles for Environmental Research 487 Conclusions 488 ESSAY A: Liberalizing the Liberal Arts: From Dominion to Design 469 Box A: Improving the Link Between Policy and Science 481 ESSAY B: OTS as an Institution for Conservation Education Sustainable Development Case Studies 491 Case Study 1. Sustainable Tropical Forestry 494 Theoretical Basis for Tropical Forest Management 495 Ydnesha Forestry Cooperative: The Palcazu Project 496 Case Study 2. Costa Rica's INBio: Collaborative Biodiversity Research Agreements with the Pharmaceutical Industry 500 Costa Rica's National Institute of Biodiversity (INBio) 500 Biodiversity Prospecting 501 Pharmaceutical Collaborative Research 502 The INBio/Merck Agreement 503 Future Challenges 503 Case Study 3. Sustainable Development in African Game Parks 504 Valuation of Protected Areas 504 Three Examples of Protected Areas 506 A New Age in African Wildlife Conservation 509 Conclusions 511 Case Study 4. The Role of Ecotourism in Sustainable Development 511 Which Lands Are Suitable for Ecotourism? 513 How Can Ecotourism Be Made Profitable? 513 Can Ecotourism Avoid Damaging Natural Areas? 514 Conclusions 515 Case Study 5. The Sian Ka'an Biosphere Reserve Project 515 What Is a Biosphere Reserve? 515 The Sian Ka'an Biosphere Reserve 516 Sian Ka'an and the Goals of Biosphere Reserves 518 How Is Sian Ka'an Doing? 519 Is Sian Ka'an a Good Example of Sustainable Development? 520 Case Study 6. The Greater Yellowstone Ecosystem 521 Ecological Issues 521 Development Issues 525 Striking the Balance 526 General Conclusions Meeting Conservation Goals in an Uncertain Future 531 The Nature and Centrality of Uncertainty 533 Responding to Short-Term Economic Change 533 Increasing the Perceived Value of Conservation Biology 534 Improving Conservation Management and Development 536 Improving Investment 538 Developing Policy Incentives 539 Responding to Long-Term Planetary Environmental Change 539 Responding to the Demographic Imperative 542 The Tragedy of the Commons 547 Five Major Actions Needed to Conserve Biodiversity 550 ESSAY A: The Postmodern World 532 ESSAY B: The Importance of Communicating with the Public 535 Box A: Scenarios for Global Climate Change 541 ESSAY C: The Bottom Line: Human Population Control 546 ESSAY D: A Latin American Perspective on the Future of Conservation and Development 550 ESSAY E: A New Vision: Humans and the Value of Nature 553 Glossary 559 Bibliography 565 Index 587

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