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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