Marine Biology

Acknowledgments xvii Preface xix About the Companion Website xxi Part I The Ocean Domain: Introduction to Planet Ocean 1 1 The Life Aquatic 3 1.1 Introduction 3 1.2 Comparison Between Sea and Land 3 1.3 Fractal Complexity of Marine and Terrestrial Ecosystems 6 2 The Seabed 13 2.1 Ocean Basins 13 2.2 Ocean Bottom: A (Mostly) Static Habitat of Ocean Life 15 2.3 Characteristics of Sediments 20 2.4 Boundary Layers and Their Characteristics 20 2.5 Sediment Movement 23 2.6 Characteristics of Hard Substrata 26 2.7 Characteristics of Soft Sediments 26 3 The Water Column 29 3.1 Properties and Characteristics of Seawater 29 3.2 An Ocean In Motion 44 Part II Life in Seas and Oceans: Fundamentals of Marine Biology 53 4 General Adaptations in Marine Organisms I: From the Ocean Surface to the Seabed 55 4.1 Adaptations to Temperature 55 4.2 Adaptations to Low Oxygen Concentrations 60 4.3 Adaptation to Salinity 64 4.4 Adaptation to Pressure 68 4.5 Adaptations to Light 69 4.6 Adaptations to Nutrients 73 4.7 Electrical Conductivity Adaptations 74 4.8 Ectocrine Adaptations 74 4.9 Adaptations to Produce Sound and Communicate in Water 74 5 Adaptations in Marine Organisms II: Life in a Fluid Habitat 79 5.1 Adaptions to Life in the Water Column 79 6 Adaptations in Marine Organisms III: Benthic Biota between a Rock and a Soft Place 91 6.1 Adaptations to Life on the Seafloor 91 6.2 Support and Protection Structures 91 6.3 Adaptation to Waves and Energy 94 6.4 Feeding and Nutrition 95 6.5 Adaptation to Aerial Exposure 98 6.6 Adaptation to Extreme Temperatures and Potentially Toxic Chemicals 98 Part IIB Life in Seas and Oceans: Fundamentals of Marine Biology 101 7 Marine Biodiversity 103 7.1 Introduction 103 7.2 Origin and Evolution of Marine Life 103 7.3 Mechanisms of Marine Speciation 107 7.4 Quantifying Marine Organism Biodiversity 113 8 Biodiversity Patterns 125 8.1 Broad-Scale Biodiversity Patterns 125 8.2 Processes Controlling the Distribution of Marine Biodiversity 125 8.3 Marine Biogeography 128 8.4 Theories on Evolution and Maintenance of Biodiversity: The Deep-Sea as Examplar 136 9 Biodiversity of the Benthos 141 9.1 Introduction: Benthos and Plankton 141 9.2 Benthic Biota 141 9.3 Classification of Benthos Based on Size 144 Part IIC Life in Seas and Oceans: Fundamentals of Marine Biology 157 10 Ecology of Benthos 159 10.1 Ecology of the Benthos: From Microbes to Megafauna 159 10.2 Trophic Groups: Classification of Benthos Based on Diet 164 10.3 Comparison Between Hard and Soft Bottom Benthos 165 10.4 Ecology of Benthos Inhabiting Soft Bottoms 167 10.5 Changes in Benthos in Space and Time 170 10.6 Organization of Benthic Assemblages 171 10.7 Zonation of Benthic Organisms 172 11 Biodiversity of the Plankton 179 11.1 Introduction to the Plankton 179 11.2 Planktonic Organisms 180 11.3 Planktonic Classification Based on Water Column Distributions 181 11.4 Plankton Classification Based on Life Cycles 182 11.5 Plankton Size Classes 184 11.6 Abundance Comparisons Among Different Planktonic Components 197 12 Ecology of the Plankton 201 12.1 Plankton Distribution 201 12.2 Ecology of Plankton 208 12.3 How Many Phytoplankton Species Coexist in a Volume of Water? “Homage to Santa Rosalia” 211 12.4 Zooplankton Nutritional Mode 211 Part IID Life in Seas and Oceans: Fundamentals of Marine Biology 215 13 Biodiversity of the Nekton 217 13.1 Species Contributing to the Nekton 217 13.2 Main Organisms and Characteristics of Nekton 218 13.3 Fishes 218 13.4 Marine Mammals 224 13.5 Cephalopods 227 13.6 Reptiles – The “Land-based” Marine Species 228 13.7 Seabirds 228 13.7 Patterns of Biodiversity in Nekton 229 14 Ecology of the Nekton 233 14.1 Introduction 233 14.2 Fishes and Formation of Fish Shoals 233 14.3 Ecology of Chondrichthyes 235 14.4 Sharks at Risk of Extinction from Indiscriminate Hunting 236 14.5 Ecology of Cephalopods 236 14.6 Marine Reptile Ecology 239 14.7 Ecology of Seabirds 241 14.8 Ecology of Marine Mammals 242 14.9 Great Migrations of Nekton 247 14.10 Role of Top Predators in Food Webs 255 15 Life Cycles and Larval Ecology 261 15.1 Life Cycles and Reproduction 261 15.2 Larval Ecology 264 15.3 Life History Strategies 267 15.4 Supply Side Ecology 270 15.5 Forms of Resistance and Benthic-pelagic Coupling 270 Part III Comparative Marine Ecology: Habitat Types, Their Biodiversity, and Their Functioning 275 16 Ecosystem Functioning I: Primary and Secondary Production 277 16.1 Introduction 277 16.2 Primary Production 277 16.3 Chemosynthetic Primary Production in the Ocean 278 16.4 Photosynthetic Primary Production 280 16.5 Secondary Production 287 16.6 Respiration 290 17 Ecosystem Functioning II: Organic Matter Recycling 293 17.1 Introduction – Extra-Specific Processes 293 17.2 Organic Matter and Detritus in the Ocean 293 17.3 Dissolved Organic Matter in the Ocean (DOM) 299 17.4 Pelagic-Benthic Coupling 300 17.5 Consequences of Organic Matter Export to the Seabed 307 18 Interspecific Interactions and Trophic Cascades 313 18.1 Biodiversity and Ecosystem Functioning 313 18.2 Facilitation and Cooperation – Positive Interactions 315 18.3 Symbiosis 319 18.4 Complex Biotic Interactions: Trophic Networks and Cascades 323 18.5 Keystone Species 333 18.6 Trophic Cascades 333 Part IIIB Comparative Marine Ecology: Habitat Types, Their Biodiversity, and Their Functioning 341 19 Interspecific Interactions II: Negative Interactions 343 19.1 Predation 343 19.2 Methods to Escape Predation 348 19.3 Competition 349 19.4 Parasitism 351 19.5 Diseases of Marine Organisms 354 20 Intertidal Ecosystems and Lagoons 359 20.1 Rocky Intertidal Habitats 359 20.2 Transitional Environments Between Land and Ocean 372 20.3 Mangroves 380 20.4 Salt Marshes 385 20.5 Summary 388 21 Subtidal Hard Substrata Ecosystems 391 21.1 Introduction 391 21.2 Subtidal Distributions 391 21.3 Kelp Forests 394 21.4 Coral Reefs 399 21.5 Coralligenous Habitats 418 21.6 Rhodolith Beds (Maërl) 422 21.7 Underwater Caves 423 21.8 Summary 429 Part IIIC Comparative Marine Ecology: Habitat Types, Their Biodiversity, and Their Functioning 431 22 Estuarine, Seagrass, and Sedimentary Habitats 433 22.1 Estuaries 433 22.2 Seagrass Beds 438 22.3 Sedimentary Habitats 446 23 Polar Ecosystems 455 23.1 Biogeography and Characteristics 457 23.2 Biodiversity 460 23.3 Biodiversity Within Sea Ice 460 23.4 Pelagic Biodiversity 462 23.5 Fishes 464 23.6 Marine Mammals 465 23.7 Benthic Biodiversity 465 23.8 Food Webs and Functioning 466 23.9 Antarctica 467 23.10 Summary 483 24 Neritic Aquatic Ecosystems 485 24.1 Introduction 485 24.2 Zonation, Extent, and Distribution 485 24.3 Biogeography and Characteristics 485 24.4 Biological Characteristics 487 24.5 Ecosystem Functioning in the Neritic Zone 490 24.6 Fisheries Production 492 24.7 Factors Influencing Functioning of Neritic Systems 493 24.8 Summary 497 Part IIID Comparative Marine Ecology: Habitat Types, Their Biodiversity, and Their Functioning 499 25 Deep-Sea Ecosystems along Continental Margins 501 25.1 Introduction to the Deep Sea 501 25.2 Deep-Sea Biodiversity 508 25.3 Deep-Sea Habitats 511 25.4 Submarine Canyons 512 25.5 Deep-Water Corals 515 25.6 Cold Seep (Hydrocarbon-Based) Ecosystems 517 25.7 Cold Seep Biodiversity and Symbiotic Organisms 518 25.8 Hypoxic and Anoxic Systems (Dead Zones) 522 25.9 Oxygen Minimum Zones, OMZs 522 25.10 Summary 526 26 Deep Ocean Basins 531 26.1 Introduction 531 26.2 Abyssal Plains 531 26.3 Abyssal Biodiversity and Adaptations 531 26.4 Abyssal Gigantism and Dwarfism 533 26.5 Functioning of Abyssal Systems 535 26.6 Deep-Sea Hydrothermal Vents 541 26.7 Whale Carcasses 551 26.8 Affinities Between Vent and Seep Communities 554 26.9 Anoxic Basins 554 26.10 Ocean Trenches 556 26.11 Summary 562 27 Oceanic Ecosystems 567 27.1 Introduction 567 27.2 Factors Influencing the Life and Distribution of Pelagic Organisms 567 27.3 Classification of Pelagic Regions 571 27.4 Functional Classification of Pelagic Systems 573 27.5 Vertical Zonation in Pelagic Ecosystems 575 27.6 Biodiversity of Pelagic Systems 577 Part IV Human Impacts and Solutions for Planet Ocean: Applied Marine Biology 581 28 Human Impacts on Marine Ecosystems 583 28.1 Historical Data 583 28.2 Biodiversity Loss 587 28.3 The Main Threats to Marine Life and Ecosystems 590 28.4 Synergistic Impacts on Marine Ecosystems 597 29 Marine Biodiversity Conservation 603 29.1 Introduction 603 29.2 Conservation Objectives 603 29.3 The Third Dimension of Marine Conservation 606 29.4 Conservation Strategies 606 29.5 Marine Protected Areas 608 29.6 Cumulative Impacts and Biodiversity Conservation 613 29.7 Conservation Frameworks 614 29.8 Legal Instruments 615 29.9 Science Challenges and Solutions – Moving Science to Policy? 616 29.10 How Science Can Contribute 616 30 Restoring Marine Habitats 619 30.1 A Decade For Ecosystem Restoration 619 30.2 Defining Ecological Restoration 620 30.3 A Global Plan for Marine Ecosystem Restoration 623 30.4 Restoring Fragile Marine Habitats 623 30.5 Restoration of Coral Reefs 625 30.6 Restoration of Seagrass Meadows 626 30.7 Restoration of Macroalgal Forests 627 30.8 Restoring Ecosystem Engineers: The Case of Coralligenous Outcrops 629 30.9 Restoration of Deep-Sea Habitats 630 30.10 Perspectives of Marine Ecosystem Restoration 631 31 How Far We Have Come: Past, Present, and Future Research on the Marine Biology of Planet Ocean 633 31.1 Introduction 633 31.2 The Birth of Marine Biology 634 31.3 The History of Ocean Exploration 636 31.4 Present and Future of Marine Biology 637 31.5 Application of Marine Technologies 644 31.6 Marine Biology Research in the Next Decade 645 Glossary 649 Index 675