new 4th edition of marine biology has

FOURTH EDITION

E*ARINH &Fu nct ion, Biod iversi ty, Ecology

Jeffrey 5. Levinton

lsBN 978-0-1 9-98571 2-8

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I PREFACE XIV

PRINCIPLES OF OCEANOGRAPHY AND MARINE ECOLOGYCHAPTER 1 Sounding the Deep 1CHAPTER 2 The Oceanic Environment 13CHAPTER 3 Ecological and Evolut ionary Pr inciples of Marine Biology 40

MARINE ORGANISMS: FUNCTION AND ENVIRONMENTCHAPTER 4 The Chemical and Physical Environment 69

CHAPTER 5 Li fe in a Fluid Medium 91

CHAPTER 6 Reproduct ion, Dispersal , and Migrat ion 102

ORGANISMS OF THE OPEN SEA

CHAPTER 7 Plankton 141

CHAPTER 8 Marine Vertebrates and Other Nekton 163

PATTERNS AND PROCESSES IN THE WATER COLUMN

CHAPTER 9 Processes in the Open Sea 198

CHAPTER 10 Product iv i ty, Food Webs, and Global Cl imate Change 225

ORGANISMS OF THE SEABED

CHAPTER 11 Benthic Microorganisms, Seaweeds, and Sea Grasses 242

CHAPTER 12 The Diversi ty of Benthic Marine Invertebrates 255

CHAPTER 13 Benthic Li fe Habi ts 283

{CASTAL BENTI-I IC ENVIRONMENTS

{F{AFTtR 14 The Tidelands: Rocky Shores, Soft-Substratum Shores, Marshes,Mangroves, Estuar ies, and Oyster Reefs 309

THAFTHR ' :5 The Shal low Coastal Subt idal : Sea Grass Beds, Rocky Reefs,Kelp Forests, and Coral Reefs 361

FROM THE SHELF TO THE DEEP SEA

CHAPTER 16 Benthos from the Cont inental Shel f to the Deep Sea 4O4

CHAPTER 17 Biodiversi ty and Conservat ion of the Ocean 434

HUMAN IMPACT ON THE SEA

CHAPTER 18 Fisher ies and Food from the Sea 461

CHAPTER 19 Environmental lmpacts of Industr ia l Act iv i t ies and HumanPopulat ions 492

I GLOSSARY G-l

I MARINE BIOLOGY JOURNALS I- I

I INDEX r- l

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I PREFACE XIV

T PRINCIPLES OF OCEANOGRAPHY AND MARINE ECOLOGYCHAPTER r Sounding the Deep 1

CHAPTER 2

CHAPTER 3

Marine Biology as a Discipline 1

Historical Background of Marine Biology 2

Observation and Hypothesis Testing 7

Habitats and Life Habits: Some Definitions 10

Chapter Summary 12

Review Qrestions 72

The Ocea n ic Environ ment 1 3

The Open Oceans and Marginal Seas 13

Topography and Structure of the Ocean Floor 13

The Ocean Above the Seabed 18

Circulation in the Open Sea: Patterns and Causes 22

Seawater Density and Vertical Ocean Circulation 27

The Greenhouse Effect and Changing Ocean Climate 29

HypothesizedEffects of Climate Change on Sea Level, Circulation, and OceanChemistry 32

Going Deeper 2.1 Solubi l i ty 33

The Edge of the Sea 33

Chapter Summary 38

Review Qrestions 39

Ecological and Evolut ionary Pr inciples of Marine Biology 40

Ecologicallnteractions 40

The Ecological Hierarchy 40

Interactions on the Scale of Individuals 4l

The Population Level 48

The Community Level: Structure and Interspecies Interactions 52

The Ecosystem Level 0o

Species, Genetic Variation, Evolution, and Biogeography 62

Chapter Summary 67

Review Qrestions 68

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(O NTE NTS

I MARINE ORGANISMS: FUNci loN AND ENVTRONMENTcHAprER 4 The chemical and physical Environment 69

CHAPTER 5

CHAPTER 6

Measures of Physiological Performance 69

Temperature 7l

Hot Topics In Marine Biology 4.1 Global Climate Change: How Can WeTelllf We Are at the Edge? 78

Salinity 81

Oxygen 83

Going Deeper 4.1 euantifying the Relationship Between Body Sizeand Oxygen Consumption Rate g3

Light s7

Chapter Summary 89

Review Qrestions 90

Life in a Fluid Medium 91

Introduction gI

Density, Viscosity, and Reynolds Number gl

Going Deeper 5.1 ls Seawater Always Seawater? A Tale from the Regionsof Intermediate Reynolds Number 93

Moving Water 93

Water Moving over Surfaces and Obstructions, Such as Organisms gS

Using Water Motion for Biological Advantage 96

Chapter Summary 101

Review Qrestions 101

Reproduct ion, Dispersal , and Migrat ion 1OzEcological and Evolutionary Factors in Sex I02

Hot Topics In Marine Biology 6.1 A Lover and a Fighter 1o4

Hot Topics In Marine Biology 6.2 Who's Your Daddy? An Investigationof Fatherhood in a Snail 112

Reproduction, Demography, and Life Cycles 113

Migration 114

Larval Dispersal: The Long and the Short Haul 118

Hot Topics In Marine Biology 6.3 Great Oceanic Migrations that Dwarf theSerengeti 119

PlanktonicLarvae: Getting Through Major Obstacles to the FinalDestination 133

The Macroscale: Major Separations Lead to Biogeographic Structure ts4

Planktonic Dispersal: Why Do They Do It? r3Z

Chapter Summary l3g

Review Qrestions l3g

CONTENTS

I ORGANISMS OF THE OPEN SEAcHAPTER 7 Plankton 141

CHAPTER B

Introduction and Definitions 141

Marine Viruses l4l

Marine Bacteria and Archaea 143

EukaryoticPhytoplankton 143

Zooplankton 145

GelatinousZooplankton 149

Life in the Open Sea r52

Defense Against Predation 153

Diurnal Vertical Migration of the Zooplankton 154

HotTopics In Marine Biology 7.1 Bioluminescence, NightVis ion,and Death in the Deep 155

Molecular Techniques to Identify Planktonic Microorganismal Diversity

Chapter Summary 161


Marine Vertebrates and Other Nekton 163

Cephalopods 163

Fish 165

Mammals 177

Hot Topics In Marine Biology 8.1 Whales and Wonder Dogs 179

Marine Birds 185

HotTopics ln Marine Biology 8.2 Last March of the Penguins?Climate Change and a Bottom-UpTrophic Cascade 187

Marine Reptiles 194

Chapter Summary 196


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T PATTERNS AND PROCESSES IN THE WATER COLUMNcHAPTER 9 Processes in the Open Sea 198

Critical Factors in Plankton Abundance 198

Patchiness of the Plankton 198

The Seasonal Pattern of Plankton Abundance 200

Water Column Parameters and the Spring Diatom Increase 200

Light 2os

Going Deeper 9.1 The Basics of Photosynthesis 2o7

Nutrients Required by Phytoplankton 208

Rate of Nutrient Uptake 213

Harmful Algal Blooms 215

Phytoplankton Succession and the Paradox of Phytoplankton Coexistence

Hot Topics In Marine Biology 9.1 Angry Birds and Lost Seals:Solution of the Effects of a Mysterious Poison 217

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CONTENTS

CHAPTER 1O

The Microbial Loop: Nutrient Cycling by Viruses, Heterotrophs,andChemoautotrophs 219

Zooplankton Grazing in the Sea 220

Going Deeper 9.2 Quantif ication of the Effect of Grazing 221

Chapter Summary 223


Product iv i ty, Food Webs, and Global Cl imate Change 225

Productivity and Biomass 225

Food Webs and Food Chains 225

MeasuringPrimaryProductivity 229

Going Deeper 10.1 How to Calculate Productivity Using the OxygenTechnique 23O

Going Deeper 10.2 Using the Radiocarbon Technique to Est imateProductivity 231

Geographic Distribution of Primary Productivity 233

Global Climate Change and the Global Carbon Pump 237

Chapter Summary 241


I ORGANISMS OF THE SEABEDcHAPTER 11 Benthic Microorganisms, Seaweeds, and Sea Grasses 242

The Big Picture: Domains of Life 242

Bacteria 243

Eukarya 244

Sea Grasses 252

Chapter Summary 253


cHAPTER t2 The Diversi ty of Benthic Marine lnvertebrates 255

Hot Topics In Marine Biology 12.1 Where Did Al lThis Invertebrate Diversi tyCome From? 256

Kingdom Protista: Single-Celled Organisms 260

Phylum Porifera: Sponges, Simplest of Animals 261

Phylum Cnidaria: Hydrozoans, Jellyfish, Anemones, and Corals 262

Phylum Platyhelminthes: Flatworms 264

Phylum Nemertea: Ribbon Worms 264

Phylum Nematoda: Roundworms 265

Phylum Annelida: Segmented Worms 265

Phylum Sipuncula: Peanut Worms 267

Phylum Pogonophora: Gutless Wonders 268

Phylum Mollusca: Shelled Invertebrates (Mostly) 269

PhylumArthropoda:JointedAppendages 272

The Lophophorate Phyla 275

Phylum Bryozoa; Moss Animals 275

CONTENTS

Phylum Brachiopoda: Lingulas and Lampshells 276

Phylum Phoronida: Wormlike Animals with a Lophophore

Phylum Echinodermata: Animals with Fivefold Symmetry

Phylum Chordata: The Sea Squirts 2gl

Chapter Summary 281


Benthic Li fe Habi ts 283

Introduction 283

Life in Mud and Sand 283

Going Deeper 13.1 Measuring Grain Size of Sediments 294

Deposit Feeding in Soft Sediments 288

Suspension Feeding 293

Life Habits on Hard Surfaces and in Moving Waters 2gZ

Benthic Carnivores 300

Benthic Herbivores 301

Hot Topics In Marine Biology 1 3.1 Diary of a Stinging Snail 302

Chapter Summary 307


I COASTAL BENTHIC ENVIRONMENTSCHAPTER 14 The Tidelands: Rocky Shores, Soft-Substratum Shores, Marsh€s,

Mangrovei , Estuar ies, and Oyster Reefs 309

Rocky Shores and Exposed Beaches 309

Soft-Sediment Interactions in Protected Intertidal Areas 327

Invasions and the Reorganization of Intertidal Communities 330

Hot Topics In Marine Biology 14.1 The Powerful Interaction of Invasionand Climate Change 333

Spartina Salt Marshes 335

Mangrove Forests 343

Hot Topics In Marine Biology 14.2 The Molecular Sleuth: Invasion of a VeryAggressiveGenotype 344

Estuaries 349

Oyster Reefs 355

Chapter Summary 359


The Shal low Coastal Subt idal : Sea Grass Beds, Rocky Reefs,Kelp Forests, and Coral Reefs 361

Sea Grass Beds 361

The Rocky ReefKelp Forest System i66

Subtidal Rocky Reefs 366

Hot Topics ln Marine Biology 15.1 Reorganization of a Rocky SubtidalEcosystem:A Cod and LobsterTale 367

Kelp Forests 371

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CHAPTER 13

CHAPTER 1 5

CONTENTS

Coral Reefs 378

Chapter Summary 401


I FROM THE SHELF TO THE DEEP sEACHAPTER 16 Benthos from the Cont inental Shel f to the Deep Sea 4O4

The Subtidal Landscape 404

Sampling the Subtidal Soft-Bottom Benthos 404

Sediment Typ. and Benthic Distribution 406

The Deep-Sea Gradient 410

The SheltrDeep-Sea Gradient 410

Deep-Sea Island Hotspots of High Diversity 417

Pressure Change 428

Polar Environments 428

Arctic Subtidal Benthic Environments 428

The Antarctic Shelf Benthos: A Special Case of Isolation 430

Chapter Summary 432


Biodiversi ty and Conservat ion of the Ocean 434

Diversity Speciation, Extinction, and Biogeographic Factors 434

Major Gradients of Species Diversity 442

Explanations of Regional Diversity Differences 446

ConservingMarineBiodiversity 450

Marine Invasions 456

Chapter Summary 459


I HUMAN IMPACT ON THE SEAcHAPTER t8 Fisher ies and Food from the Sea 461

The Fishery Stock and Its Variability 46r

Stocks and Markers 467

Life History and Stock Size 463

Stock Health and Production 464

Going Deeper 18.1 A Simple Model to Explain the Maximum SustainableYield

Fishing Techniques and Their Effects 468

Fisheries Impact and Management 471

Causes and Cures of Stock Reduction 474

Hot Topics ln Marine Biology 18.1 ls Fisheries Policy at Odds with ManagingMarine Ecosystems? 475

Going Deeper 18.2 Age-Based or Life-History Stage-Based Population Models

Overexploitation ofWhales: A Case History 482

Other Typ.r of Degradation 484

Disease as a Major Danger to Coastal Fisheries 485

CHAPTER 1 7

CONTENTS

CHAPTER 19

Mariculture 485

Drugs from the Sea: Marine Natural Products 489

Chapter Summary 490


Environmental lmpacts of lndustr ia l Act iv i t ies and HumanPopu lation s 492

Human Effects on the Marine Environment 492

Measuring the Impact of Pollutants on Populations and Communities 493

Toxic Substances 497

Hot Topics In Marine Biology 19.1 ls the Gulf of Mexico Adapted to Oil?

Nutrient Input and Eutrophication 508

Thermal Pollution and Power Station Fish Mortality 512

Global Environmental Change and the Ocean 513

Chapter Summary 515


GLOSSARY G-1

MARINE BIOLOGY JOURNALS J- l

INDEX I-1

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elcome to the fourth edition of Marine Biology:Function, Biodi,uersity, Ecology. This new edi-tion reflects my continued excitement at the

teaching of marine biology, a far-reaching science thatmanages to excite a broad spectrum of students. Wemarine biologists are lucky to have exciting creatures,great adventures, and continuing discoveries that manageto enchant students. The challenge in a text is to injectthis excitement into an organized approach to the subject.Many students find marine biology a little daunting. Nosurprise to me, because it is daunting to put it all into atext. I believe, though, that every student can acquire thebroad spectrum of biological knowledge included inmarine biology while appreciating the organismal diver-sity of the marine realm. To make that possible, studentsmust learn concepts along with the facts and begin to thinkand reason like scientisfs. Just as important, they must feelthe pulse of current happenings.

The goal of this text is to appeal to a wide range ofstudents, while also preparing future specialists with theknowledge and tools needed to conduct their research.Marine biology is such a diverse subject that principlesmust guide our understanding. I firmly believe that, with-out principles and a sense of excitement, students will seeonly an accumulation of facts. Th.y will fail to see howthe same basic processes operate in vastly different marinecommunities.

That is why the text addresses three major process-driven themes: functional biology, ecological processes, andbiodiversity. lt is why the text uses full color throughoutin drawings that depict marine biological processes and alarge number of photographs to connect students withmarine environments and organisms. It is also why I havepaid much attention to recent research developments andhave included a series of essays, called "Hot Topics inMarine Biology." These features will put students in con-tact with the current world of exciting research. In boththe text and the Hot Topics, I try to connect the studentsto some of the most important recent research with anextensive literature section that is now online so that stu-dents can get to the best work for term papers and proj-ects without too much distraction in the text. In my ownclasses I point out that a good teacher shows students theway to good science because library databases are notuser friendly where finding excellent science research isconcerned.

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The Philosophy of This TextThis text is designed for a one-semester course at the soph-omore to senior level. Some students will have alreadytaken a college-level biolog"y course with coverage of or-ganismal diversiry and they will benefit greatly. A smallnumber may even have taken a course in marine inverte-brates or vertebrates and an introductory course in ocean-ography.That said,I have successfully taught from this textfor many years, and many students had no background inorganismal biology or ecology.If the book is supplementedwith journal articles, it can also be used in a more advancedundergraduate course in marine ecology. The new editioncontains many updated references to the primary literature,which are expanded and now online with a simple linkfrom the text chapters. This is a valuable resource to getstudents started on term papers and essays. The MarineBiology Web Page (www.oup.com/us/levinton), which Ifounded a number of years ago, links students to manymore views of marine biology and to a greater diversity oforganisms. It is a continuing objective to merge principleswith an appreciation of organismal diversity in the ocean.

I have taught Marine Biology for over 30 years and havealways been amazed at the diversity of students who takethe course. Biology majors, marine science majors, geologymajors, and even some humanities majors sit side by side.At my university, marine science has become a separateundergraduate discipline in recent years, and these stu-dents have truly learned the interdisciplinary nature ofmarine biology. A11 learn a great deal, and all seem tocome away with a love for the ocean. You don't have toconvince them to be there: Th.y a:ant to learn aboutmarine biology.

I do my best to keep that interest alive, and I find thatfield trips and the use of color photographs and otherillustrations throughout the text help a great deal. So dothe online resources accompanying this text. MarineBiology Exploratiozs includes hundreds of photographsfrom habitats discussed in the text and still more. As men-tioned, an expanded and updated reference list is alsoonline. In class, I keep a large map of the oceans on thewall for the geographic context of our discussions. I havetaken many of my students with me to marine labs, andthey have launched careers in science or used their back-grounds to enter other areas. I hope the text will help awider audience to get excited about marine life. I hope,

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too, that they will understand how the ocean works andwhy our marine realm is so threatened.

A Principles-Driven ApproachMarine biology applies the principles of ecology and evolu-tion, using the crucial tools of cell biology, biomechanics,and molecular biology to a wide range of marine biologicalstudies. These fields of study and their interactions governthe textt three overarching themes: functional biology,biodiversiry and ecological processes.

Function refers to the w^y organisms solve problemsand how physical and chemical factors constrain and selectthe solutions. What shape should a maneuvering fish have,relative to a continuously swimming fish? How does asmall peptide manage to be such an effective poison wheninjected by a snail into a prey? How does this specificbiochemical adaptation feed into an understanding of bio-diversity? I believe this textbook is unique in combiningeffectively functional biology with ecological thinking.

Biodiversity is an essential part of marine biology, andI introduce the topic both through introductions to theprinciples used to study and explain biodiversity and tothe factors that strongly affect marine biodiversity. A sep-arate chapter discusses diversity and the processes thatregulate it, both ecological and evolutionary. It is crucialthat the student see the historical roots of many currentdistributions, which are affected by processes rangingfrom plate tectonics to climate change. This edition adds agreat deal of coverage of recent advances, includingmolecular tools used to identify microbial organisms andactivity in the plankton and molecular methods used tostudy dispersal, speciation, and the rise of marine adapta-tions. A chapter devoted to biodiversity makes the con-nections between evolutionary process, biodiversity, andbiogeography. These areas are related directly to issues ofmarine conservation.

Last, ecology examines the interactions of organismswith their environment. Ecology tries to understand thedistribution and abundance of organisms. It involves aseries of processes, which I introduce in the context of ahierarchy-from individual populations to ecosysterns. Italso involves a discussion of important ecological processesalong with accounts of major marine habitats and com-munities. This edition pays special attention to modernconcepts of populations and species interactions, includingconnectivity, metapopulations, regional genetic differen-tiation, large-scale control of dispersal, biological inva-sions, and alternative stable states of communities. Mybackground in geology, ecology, and evolutionary biologyallows me to frequently combine ecological and evolution-ary thinking in discussing marine biology problems.

OrganizationWe begin with a brief historical background. Marine biol-ogy has a history that is worth understanding, but it is alsocrucial to introduce the student to how science works.Chapter 1 therefore discusses framing and testing

PREFACE

hypotheses, as well as making tests practical enough thatthey can be put to direct use. From the very beginning,I introduce the student not only to the scientific method,but also to how it translates into an intellectual tool withreal-world applications.

Chapters 1 through 6 introduce basic principles of howthe ocean works in a physical, chemical, and ecologicalcontext and how marine organisms function with theseconstraints. The second chapter gives the student a com-prehensive introduction to oceanography and the impor-tant properties of sea water that might affect marineorganisms. I work in climate change at this stage becauseit will be part and parcel of many discussions throughoutthe text. This edition greatly expands the effects of tem-perature and especially ocean acidification, and the stu-dent sees how an understanding of the ocean and sea waterproperties will directly affect the fate of marine organismsin the future. I then introduce ecological principles so thatstudents can work their way through concepts usingmarine examples. This allows all students to be brought upto a level of ecological thinking and an understanding ofoceanographic processes. Theywill see this "big picture" asthey read the rest of the text. A crucial chapter then intro-duces students to how the physics of fluids shapes the con-straints and adaptations of marine organisms. As far as Iknow, this crucial subject is missing in all other marinebiology texts, and allows a connection to a complete un-derstanding of how the marine organisms function in therather complex fluid environment. These chapteis con-clude with a comprehensive introduction to reproductivestrategies, larval dispersal, and migration, which sets upthe big picture of the geographical distribution of marinespecies, down to the microscale of how mobile marinelarvae succeed in finding a place to live in a turbulent andstressful world.

Chapters 7 through L0 cover the organisms and processesimportant in the open sea. This two-step approach is es-sential so that students will understand the overall econ-omy of the marine realm. Chapter L0 uses a global-scaleapproach to show how biological studies of the ocean leadto an understanding of the world's potential for fisheriesand the global biological impact on the ocean of climatechange. Processes in the water column are also crucial inChapters 11 through 15 for understanding the benthos,which depends both directly and indirectly on the waterworld above. In Chapters 11 and \2I cover benthic crea-tures and then go on to discuss the principles necessary tounderstand the biology of marine bottom organisms(Chapter 13) and the major nearshore marine bottom hab-itats (Chapters 14-1.6). By necessity, I have been selective.I emphasize those habitats that are not only important andinteresting, but also where important principles can be il-lustrated to their best advantage. Communitplevel inter-actions are emphasized, as is global climate change as itrelates to major changes in habitats such as coral reefs.I discuss a range of geographic locations so that theinstructor will find local examples in many instances.Crucial habitats such as the intertidal, seagrasses, coral

PREFACE

reefs, mangroves, estuaries, salt marshes, kelp forests, andothers are discussed both from the points of habitat dis-tinctions and ecological processes and the impacts of bio-logical invasions and climate change. I have added a newsection on oyster reefs because of their great worldwideimportance as foci for biodiversity and their role in ecosys-tem services.

Chapter 16 then looks at the important gradient fromthe continental shelf to the deep sea, paying special atten-tion to some of the fascinating discoveries about biologicalfunction and fascinating habitats, from hot vents to deepwater coral mounds to the newly discovered subsurfacebacterial realm over 500 m beneath the sea floor. I havegreatly expanded coverage of Arctic and Antarctic envi-ronments. The Antarctic is center stage in our focus on thecurrent and future impacts of climate change, and the textincorporates a wide range of discussions from the organis-mal to the ecosystem level. Chapter \7, on gradients inbiodiversity, sums up larger-scale variants in the sea andincludes sections on invasive species, conservation of bio-diversity, and conservation genetics. More and more, stu-dents and researchers have focused their attention to thedeteriorating conditions of the ocean, and conservation isa major field of emphasis.

Finally, Chapters L8 and L9 tackle other human inter-actions with the sea, as both a source of food and, unfor-tunately, a waste receptacle. I cover human effects on theocean. Throughout the text, the effects of climate changeare brought up in many contexts, and how those effects arerelated to chemical issues such as acidification and facilita-tion ofbiological invasions. I also place strong emphasis onthe reorganization of communities that has been initiatedby the interaction of human activities and strong ecologi-cal interactions found in natural communities and in foodwebs. The impact of overfishing on populations andtrophic cascades is a crucial part of a complete chapter onfisheries and mariculture. The role of toxic substances,eutrophication, and hypoxia are discussed clearly and indepth. I have added to Chapter L8 a section on drugdiscovery in the ocean, because of the great student inter-est in this subject and the connections between biodiver-sity and the new sources of compounds to combat pain anddisease, such as cancer.

A Refined Learning PackageThis text has a series of pedagogical features designed tohelp students absorb a great variety of information byengaging their imaginations, helping them organize andprioritize important principles, and keeping them focusedon the big without getting lost in the details.

"Hot Topics in Marine Biology" essays throughoutthe text introduce students to recent advances in the un-derstanding oi marine biology and discuss current issues,especially marine-biological debates and discoveries.Instructors can use these essays to kick off discussion, toexpand a student's horizons, for course assignments, or astopics for term papers. Reorganizations of communitiesfrom the Antarctic to the Gulf of Maine are related to

climate change, overfishing, and other processes that arenow actively being investigated. A new section on theamazing discoveries of long-distance migration pathsthrough the Pacific Ocean has been added to Chapter 6.I have also tried to give some additional understanding ofsome current problems by adding a Hot Topics featurethat addresses exciting new molecular techniques used tounderstand marine mammal population structure (Chap-ter 8), the vexing mortality of marine mammals on thePacific Coast (Chapter 9), the major issue of managementof fisheries (Chapter 18), and what the recent horrific GulfOil Spill might tell us about natural processes and oil deg-radation (Chapter 19). Some Hot Topics have been re-tained because they are still "hot," such as the discussion ofclimate change in the Antarctic Ocean and the impacts onpenguin species in Antarctica.

Key Concept full-sentence summary statements beginnearly every section of the text to help students identifycentral points of discussion and to foreshadow what's tocome. These headings allow students to discern the forestfrom the trees and to see the basic progression of material.

Each chapter ends with a bulleted Chapter Summaryand a variety of Thought Qrestions. Instructors and stu-dents can use these to follow up on important issues inmarine biology. The combination of these features and the"K.y Concept" heading sentences successfully guides thestudent through a complex subject.

"Going Deeper" boxes explain equations and relatedconcepts in marine biology. Especially in early chapters,they will help students learn often-difficult material or re-fresh their memory of elementary courses (e.g., DNA andphotosynthesis). Th.y also allow instructors who chose toomit them to press on with no interruptions. An exampleis the discussion of Leslie Matrices in Chapter 18, whichgive the student an idea of how age-structured populationmodels help to understand impacts of various factors onfisheries and management decisions.

Extensive References lists of classic and contemporaryscholarship that instructors may assign as reading and thatcan lead students to further assignments are linked onlinefrom the text. These help students see that marine biologyis a living field of research, not just a static textbook of"known" facts, without interrupting the flow of the text.

A comprehensive Glossary of marine biology at theend of the text provides students access to get a quick def:inition of important concepts, processes, and terms. A listof journals is a resource for students in writing termpapers and for further research.

What ls New and Noteworthyin the Fourth Edit ion?Expanded illustration program. We continue the fourthedition using a rich color presentation in order to betterdemonstrate marine biological principles and introduce or-ganismal diversity in a vivid and captivating visual presen-tation. The new edition includes over 50 new photos andline drawings (many of the photos generously contributed

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by colleagues), and I believe students will benefit greatlyfrom now having the color photos integrated directly intothe relevant textual discussion at hand.

More applications. To engage students with the diver-sity of marine biology today and to highlight the real-worldapplications of what they are learning, I've written manynew in-text examples, including seven new Hot Topics inMarine Biology. Students will see how molecular toolscan be used to study mating success, how a large-scale pro-gram offish tagging has established major migration routesin the North Pacific Ocean, how a mysterious series ofdeaths of marine mammals in California was linked to atoxic species of diatom, how fishing management practicecan have strong effects on the structure of marine ecosys-tems, and how microbial degradation may be responsiblefor a rapid degradation of some components of hydrocar-bons following the disastrous Gulf Oil Spill of 2010.

Current and expanded topics maintain the excitementthat underlies my philosophy of teaching and have beencarefully selected to bring the text up-to-date while stillremaining focused on the most important principlesstudents need to learn.

' Ecological interactions. Strong attention is paid to majorecological interactions that are relevant to ecosystemstructure, such as trophic cascades (Chapters 15 anc1B), ecological reorganization in New England andelsewhere (Chapter 15), molecular approaches toecology and evolution (Chapters 7, B, 13, and 1.4),natural and human-induced phase shifts (Chapters 14and 15), biological invasions (Chapters 14,75,and 1.7),and climate change (Chapters 4, B, and 15).

' Evidence and ffittt of climate change.I have greatlyexpanded coverage of climate change, with attentionpaid to temperature change in the global ocean and incoastal areas (Chapter 2); the ocean acidification(Chapters 2,1.0 and 1,5); the role of climate change inchanging species distributions, facilitating biologicalinvasions, and causing thermal stress (Chapters 2,4,1.4,and 15); and other topics.

' Methods of environmental Assessment,from remote sensingto the molecular level. I have also expanded coverage ofthe latest methods for remote sensing, estimating worldproductiviry and assessing the stress on and change ofecosystems, including satellite methods and ocean ob-servatories (Chapters 1 and 10), genetic and molecularstudies of population differentiation (Chapters 3,6,1.7,and L8), the shifting baseline concept (Chapter 18),diversity gradients and the tropical origins of bio-diversity (Chapter 17),and molecular methods totrace biological invasions (Chapters 14 and 1.7).

' Human impact on biodiaersity.-Ihis edition expandscoverage of the decline of coral reefs and adds insightson other biological impact such as the increase ofsponges at the expense of corals (Chapter 15), over-fishing and the issue of relating management decisionsto management of the basis of ecosystem function(Chapters 15 and 18), more on declines of sharks and

PREFACE

other apex predators (Chapter 18), and the effects ofpollution, especially with coverage of the DeepwaterHorizon well blowout (Chapter 19).

' Drug discovery.I have added a section on the use ofmarine natural products in the development of drugs.

SupplementsMarine Biology, Fourth Edition, is accompanied by awealthof electronic resources for both students and instructors,including a FRtrtr Companion Website, (www.oup.com/us/levinton) and the Instructor's Resource CD-ROM.

Companion Website: Maintained by the author, thisCompanion Website (www.oup.com/us/levinton) pro-vides a r.nultitude of resources for both students andinstructors.

' Student Resources

' Marine BiologyExplorations. Explore the ocearisbiodiversity through interactive exercises that willtake you through nine different marine habits; in-cluding over 450 photos with annotations!

' Marine Biologyin theNews. Frequently updatedcurrent breakthroughs in marine biology research.

. Extensive web links to marine biology topics andresearch literature. You will also find informationon careers in marine biology and worldwidemarine laboratories.

' Instructor Resources (available to adopters of thetext and password-protected)

' Electronic Images. A11 illustrations from the textavailable in electronic format for download forlecture presentations.

' PowerPoint Lecture Notes. Over 400 lecturenotes slides organized by chapter.

' Test Bank. This comprehensive resource includesapproximately 400 questions written by the authorhimself in editable Word files for easy customiza-tion (available only on Instructor's ResourceCD-ROM: contact your Oxford University Presssales representative for details).

AcknowledgmentsMany people have helped me-too many to mention all indi-vidually.I am especially gratefirl to the many individuals whoshared their photographs and research experiences with me.Many sent me preprints, photographs, data, and just plain in-teresting discussions, which helped me gready to understandfields unfamiliar to me. For the current edition, Jason Noeserved as editor and continually moved this project forward.Keith Faiwe worked as an excellent liaison between me andcopy editors and artists to bring the project to completion.I would also like to thank Andrew Heaton, editorial assistant;Patrick Lynch, editorial director; Jason Kramer, marketingmanager; Frank Mortimer, director of marketing; MicheleLaseau, art director; and LisaGrzan, managing editor.

I also benefited a tremendous amount from careful reviewofthe manuscript and additional reviewer recommendations

PREFACE

based on the third edition. I had a wonderful panel of advi-sors for the third edition but have continued to be luckywiththe truly excellent panel that helped me with revisions forthe fourth. I could not have done better with such an excel-lent panel:

Christopher D. Beatry Santa Clara University

Heather Bennett, Illinois College

Michael Berger, Washington State University

John Berges, University of Wisconsin-Milwaukee

Patricia M. Biesiot, University of Southern Mississippi

Matthew Bracken, Northeastern University

W. Randy Brooks, Florida Atlantic University

ErinJ. Burge, Coastal Carolina University

Deron Burkepile, Florida International University

Marymegan Daly, Ohio State University

Carrie DeJaco, Qreens University of Charlotte

Steve Dudgeon, California State Universiry Northridge

Joy Ferenbaugh, James Madison University

Michael Franklin, California State Universiry Northridge

Gary R. Gaston, University of Mississippi

Reuben Goforth, Purdue University

Shannon Gowans, Eckerd College

Fiona M. Harper, Rollins College

Ione Hunt Von Herbing, University of North Texas

Catherine Hurlbut, Florida State College atJacksonville

Katrin Iken, University of Alaska Fairbanks

Yan Jiao, Virginia Tech University

Catherine Teare Ketter, University of Georgia

George Kraemer, Purchase College, State Universityof New York

Tara Maginnis, University of Portland

Katrina Mangin, University of Arizona

Karen Martin, Pepperdine University

Rob Martin, Florida State College

Marion McClary Fairleigh Dickinson University

Scott Milroy, University of Southern Mississippi GulfCoast

Linn Montgomery, Northern Arizona University

Anthony Moss, Auburn University

Xaozhen Mou, Kent State University

Steven Mark Norris, California State University ChannelIslands

Antonios Pappantoniou, Housatonic Community College

Chris Parsons, George Mason University

Neil Pelkey, Juniata College

Curt Pueschel, Binghampton University

Marjorie Reaka, University of Maryland

A-y Ringwood, UNC Charlotte

CliffRoss, University of North Florida

Wendy L Ryan, Kutztown University

David Scholnick, Pacific University

Jyotsna Sharma-Srinivasan, University of Texas atSan Antonio

William Stickle, Louisiana State University

Jacqueline Webb, University of Rhode Island

M"ry Wicksten, Texas A&n4 University

Larissa M. Williams, Wheaton College

Tim Wootton, IJniversity of Chicago

I also again thank reviewers whose insights contributed topast editions: Jelle Atema, Boston University; Susan S.Bell, University of South Florida; Larry E. Brand, Univer-sity of Miami; Christopher Brown, Florida InternationalUniversity;James E. Byers, University of New Hampshire;Edward J. Carpenter, San Francisco State University;Gerardo Chin-Leo, Evergreen State College; Paul Dayton,Scripps Institute ofOceanography; Chris D'Elia, LouisianaState University; Sean Patrick Grace, Southern Connecti-cut State University; Larry G. Harris, LJniversity of NewHampshire; William W. Kirby-Smith, Duke University;AlanJ. Kohn, University ofWashington; Derek R. Lavoie,Cuesta College; Larry R. McEdward, University ofFlorida; George McManus, university of Connecticut;A-y Moran, Clemson University; Stephen Norton, EastCarolina University; Jan A. Pechenik, Tufts University;Kathleen A. Reinsel, Wittenberg University; David Scheel,Alaska Pacific University; Eric P. Scully, Towson Univer-sity; Jayson Smith, California State University Fullerton;Alan E. Stiven, University of North Carolina; Philip Sze,Georgetown LJniversity; Keith Walters, Coastal CarolinaUniversity; andJudith S. Weis, Rutgers University.

Some of this edition was prepared at my home institu-tion, Stony Brook Universitp but I am very grateful to thestaff of the Friday Harbor Laboratories, University ofWashington, where I spent a summer working on thistext. I am especially grateful to the vision of ArthurWhiteley for creating, supporting, designing, and nurtur-ing the Helen RiaboffWhiteley Center, where I workedand lived. As usual, mywifeJoan was supportive and help-ful with suggestions.

Jeffrey LevintonStony Brook University, Stony Brook, New York

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