JEFFREY LEVINTON ARINE BIOLOG FUNCTION BIODIVERSITY ECOLOGY INTERNATIONAL THIRD EDITION

Transcription

1 JEFFREY LEVINTON ARINE BIOLOG FUNCTION BIODIVERSITY ECOLOGY INTERNATIONAL THIRD EDITION UNIVERSITY PRESS Ibis text is bosed on a North Amer been modified from the original to better serve 1 er locales. It is not for sale n Nortt^Ameríca.

2 Preface PART I. PRINCIPIES OF OCEANOGRAPHY AND MARINE BIOLOGY CHAPTER1. Sounding the Deep 3 Marine Biology as a Discipline 3 Historical Background of Marine Biology 4 Obscrvation, Experimentation, and Hypothcses 9 Habitats and Lite Habits: Some Definitions 13 Going Deeper 1.1 A Glimpse ínto the Variation Problem: How Do We Know That One Estímate Differs from Another? 14 Chapter Summary 16 Further Reading 16 Rcview Questions 16 CHAPTER 2. The Oceanic Environment 17 Trie Ocean and Marginal Seas 17 Topography and Structure of the Ocean Floor 18 The Ocean Above the Seabed 22 Circulation in the Open Sea: Patterns and Causes 26 Seawater Density and Deep Circulation 31 The Grcenhouse Effect and Changing Ocean Climate 32 HotTopícsin Marine Biology 2.1 Warming of Marginal Seas and Estuaries: Has It Occurred and What Does It Mean? 35 The Edge of the Sea 38 Chapter Surnmary 43 Further Reading 43 Revicw Questions 45 CHAPTER 3. Ecológica! and Evolutionary Principies of Marine Biology Ecological Interactions 46 The Ecological Hicrarchv 47 Interactions on the Scale of Individuáis 47 The Population Level 56 Species and Classification 61 Marine Biogeography 67 VI I

3 The Cornmuriity Level: Structure and Interspecies Interactions The Ecosystem Level 77 Chapter Summary 78 Further Reading 79 Review Questions 81 CHAPTER 4. The Chemical and Physical Environment 85 Measures of Physiologícal Performance 85 Temperature 87 HotTopics in Marine Biology 4.1 Global Warrning: How Can WeTell If We Are at the Edge? 94 Salinity 96 Oxygen 99 GoingDeeper4.1 Quantifying the Relationship Between Body Size and Oxygen Consumption Rate 99 Light 103 Cycles: Physiologícal and Behavioral Rcsponses 105 Chapter Summary 106 Further Reading 107 Review Questions 108 CHAPTER 5. Life in a Fluid Médium 109 Introduction 109 Density, Viscosity, and Reynolds Number 109 GoingDeeper5.1 Is Seawater Always Seawater? A Tale from the Regions of Intermedíate Reynolds Number 111 Moving Water 111 Water Moving over Surfaces and Obstructions, Such as Organisms 113 Using Water Motion for Biological Advantage 114 HotTopics in Marine Biology 5.1 Flow Is a Drag, but It Sure Can Smell Good 118 Chapter Summary 121 Further Reading 121 Review Questions 122 CHAPTER 6. Reproduction, Dispersa!, and Migration 123 Ecological and Evolutionary Factors in Sex 123 HotTopics in Marine Biology 6.1 A Lover and a Fighter 125 HotTopics in Marine Biology 6.2 When Shrimp Socialize in the Extreme 134 Reproduction, Demography, and Life Cycles 136 Migration 137 Larval Dispersa!: The Long and the Short Haul 141 Hot Topics ín Marine Biology 6.3 How to Get a Free Ride Out to Sea 148 Planktonic Larvae: Getting Through Major Obstacles to the Final Destination 154

4 CONTENTS X The Macroscale: Major Separations Lead to Biogeographic Structure 155 Planktonic Dispersa!: Why Do Ihey Do It? 158 Chapter Summary 160 Further Reading 160 Review Questions 164 PARTIII. ORGANISMS OFTHE OPEN SEA CHAPTER 7. The Water Column: Plankton 167 Introduction and Definitions 167 Life in the Open Sea 167 Phytoplankton 169 Zooplankton 172 Molecular Techniques to Identify Planktonic Microorganismal Diversity 178 HotTopics in Marine Biology 7.1 Endless Microbial DNA Sequences, Most Beautiful 179 Going Deeper 7.1 DNA: Ihe Basics 183 Chapter Summary 184 Further Reading 184 Review Questions 185 CHAPTER 8. The Water Cotumn: Nekton and Other Marine Vertebrales 187 Cephalopods 187 Fish 189 Mamnials 201 Birds and Reptiles 207 HotTopics in Marine Biology 8.1 Last March of the Penguins? Climate Change and a Bottorn-Up Trophic Cascade 208 Sea Snakes 216 SeaTurtles 216 Chapter Summary 218 Further Reading 219 Review Questions 221 PART IV. PROCESSES IN THE OPEN SEA CHAPTER 9. Critical Factors in Plankton Abundance 225 Patchiness of the Plankton 225 The Seasonal Pattern of Plankton Abundance 227 Water Column Parameters and the Spring Diatom Increase 228 Light 233 Going Deeper 9.1 The Basics of Photosynthesis 234 Nutriente Required by Phytoplankton 235 Rate of Nutrient Uptake 240

5 Phytoplankton Succession and the Paradox of Phytoplankton Coexistence 243 HotTopicsin Marine Biology 9.1 Luminescence, Night Vision, and Death in the Deep 245 The Microbial Loop: Nutrient Cycling by Heterotrophs and Chemoautotrophs 247 Zooplankton Grazing in the Sea 248 Going Deeper 9.2 Quantification of the Effect of Grazing 249 Diurnal Vertical Migration of the Zooplankton 250 Defense Against Predation 252 Chapter Sumrnary 253 Further Reading 254 Review Questions 256 CHAPTER 10. Productivity, Food Webs, and Global Climate Change 258 Productivity and Biomass 258 Food Chains and Food Webs 258 Measuring Primary Productivity 261 Going Deeper 10.1 How to Calcúlate Productivity, Using the Oxygen Technique 262 Going Deeper 10.2 Using the Radiocarbon Technique to Estimate Productivity 263 Geographic Distribution of Primary Productivity 265 Global Climate Change and The Global Carbón Pump 269 HotTopics in Marine Biology 10.1 Carbón Dioxide and Ocean Acidification 271 Going Deeper Box 10.3 Solubility 272 Chapter Summary 274 Further Reading 275 Review Questions 276 PARTV. ORGANISMS OF THE SEABED CHAPTER 11. The Diversity of Benthic Marine Invertebrates 279 Hot Topics in Marine Biology 11.1 From Where Did All This Invertebrate Diversity Come? 280 Kingdom Protista: Single-Celled Organisrns 284 Phylum Porifera: Sponges, Simplest of Animáis 285 Phylurn Cnidaria: Hydrozoans, Jellyfish, Anemones, and Coráis 286 Phylum Platyhelminthes: Flatworms 288 Phylurn Nemertea: Ribbon Worms 288 Phylum Nematoda: The Roundworms 289 Phylum Annelida: Segmented Worms 289 Phylum Sipuncula: Peanut Worms 292 Phylum Ponogonophora: Gutless Wonders 292 Phylum Mollusca: Shelled Invertebrates (Mostly) 293 Phylum Arthropoda: Jointed Appendages 297 The Lophophorate Phyla 299 Phylum Bryozoa: Moss Animáis 300 Phylum Brachiopoda: Lingulas and Lampshells 301

6 CONTENTS Phylum Phoronida: Wormlike Animáis wíth a Lophophore 301 Phylum Echinodermata: Animáis with Fivefold Symmetry 302 Phylum Chordata: The Sea Squirts 305 Chapter Summary 306 Further Reading 307 Review Questions 308 CHAPTER 12. Seaweeds, Sea Grasses, and Benthic Microorganisms 309 Seaweeds 309 Sea Grasses 315 Benthic Microorganisms 316 Fungi 318 Chapter Summary 319 Further Reading 319 Review Questions 320 CHAPTER 13. Benthic Life Habits 321 Life Habit Classification 321 Life in Mud and Sand 322 GoingDeeper 13.1 Measuring Grain Size of Sediments 323 Suspensión Feeding and Life Habits on Hard Surfaces and in Moving Waters 334 Benthic Carnivores 340 Benthic Herbivores 342 HotTopics in Marine Biology 13.1 Diary of a Stinging Snail 343 Chapter Summary 348 Further Reading 349 Review Questions 351 ART VI. COASTAL BENTHIC ENVIRONMENTS CHAPTER 14. The Tidelands: Rocky Shores, Soft-Substratum Shores, Marshes, Mangroves, and Estuaries 355 The Rock)' Shore and Exposed Beaches 355 Soft-Sediment Interactions in Protected Intertidal Áreas 375 Invasions and the Reorganization of Intertidal Communities 381 HotTopics in Marine Biology 14.1 The Powerful Interaction of Invasión and Climate Change 383 Spartina Salt Marshes 385 Mangrove Forests 394 HotTopics in Marine Biology 14.2 The Molecular Sleuth, Part I: Invasión of a Very Aggressive Genotype 395 Estuaries 400 Chapter Summary 407 Further Reading 408 Review Questions 412

7 CHAPTER 15. Sea Grass Beds, Rocky Reefs, Kelp Forests, and Coral Reefs 413 Sea Grass Beds 413 The Rocky Reef-Kelp Forest System 418 Subtidal Rock}- Reefs 419 Kelp Forests 422 HotTopics in Marine Bioíogy 15.1 Reorganization of a Rocky Subtital Ecosystern: A Cod and LobsterTale 423 Coral Reefs 432 HotTopics in Marine Bioíogy 15.2 Global Warming and Acidification: The Endof Coral Reefs? 448* Chapter Summary 454 Further Reading 455 Review Questions 459 RTVII. FROM THE SHELF TO THE DEEP SEA CHAPTER 16. From the Continental Shelf to the Deep Sea 463 Sampling the Subtidal Soft-Bottom Benthos 463 SedimentType and Spatial Distribution 466 The Shelf-Deep-Sea Gradient 471 Deep-Sea Islands of High Diversity 477 Pressure Change 486 Chapter Summary 487 Further Reading 488 Review Questions 491 CHAPTER 17. Biodiversity and Conservation of the Ocean 492 Speciation, Extinction, and Biogeographic Factors 492 Major Gradients of Species Diversity 501 Explanations of Regional Diversity Differences 505 Conserving Marine Biodiversity 510 Marine Invasions 516 HotTopics in Marine Bioíogy 17.1 The Molecular Sleuth Returns: Where Did the Invaders Come From? 519 Chapter Summary 522 Further Reading 522 Review Questions 525 PARTVIli. HUMAN 1MPACTON THE SEA CHAPTER 18. Fisheries and Food from the Sea 529 The Fishery Stock and Its Variability 530 Stocks and Markers 530 Life History and Stock Size 532

8 CONTENTS Stock Health and Production 533 Going Deeper 18.1 A Simple Model to Explain the Máximum Sustainable Yield 536 Fishing Techniques and Their Effects 537 Fisheries Impact and Management 539 Causes and Cures of Stock Reduction 543 Going Deeper 18.2 Age-Based or Life-Histor}' Stage-Based Population Models 547 Overexploitation of Whales: A Case History 549 OtherTypes of Degradation 551 HotTopics in Marine Biology 18.1 The Return of the Molecular Sleuth: Keeping Whalers Honest 552 Disease as a Major Danger to Coastal Fisheries 554 Mariculture 554 Chapter Summary 560 Further Reading 561 Review Questions 563 CHAPTER 19. Environmentai Impacts of Industrial Actívities and Human Populations 564 Human Effects on the Marine Environment 564 Measuring the Impact of Pollutants on Populations and Communities 565 Toxic Substances 570 Nutrient Input and Eutrophication 576 Thermal Pollution and Power Station Fish Mortality 580 Global Environmentai Change and the Ocean 582 HotTopics in Marine Biology 19.1 Is Iron Really a FDC? 585 Chapter Summary 585 Further Reading 586 Review Questions 588 Glossary G-l Marine Biology Journals J-l Index 1-1