JEFFREY LEVINTON

A R I N E BIOLOGFUNCTION • B I O D I V E R S I T Y • ECOLOGY

INTERNATIONAL THIRD EDITION

UNIVERSITY PRESS

Ibis text is bosed on a North Amerbeen modified from the original to better serve

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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 KnowThat 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 67VI I

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 WeTellIf We Are at the Edge? 94

Salinity 96

Oxygen 99

GoingDeeper4.1 Quantifying the Relationship Between Body Size and OxygenConsumption 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 Regionsof 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

C O N T E N T S ¡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

P A R T I I I . 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 Changeand 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

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

225

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

P A R T V . ORGANISMS OF THE SEABED

CHAPTER 11. The Diversity of Benthic Marine Invertebrates

Hot Topics in Marine Biology 11.1 From Where Did All This InvertebrateDiversity 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

279

C O N T E N T S

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 Invasiónand Climate Change 383

Spartina Salt Marshes 385

Mangrove Forests 394

HotTopics in Marine Biology 14.2 The Molecular Sleuth, Part I: Invasiónof a Very Aggressive Genotype 395

Estuaries 400

Chapter Summary 407

Further Reading 408

Review Questions 412

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

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 Actívitiesand 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