Marine Biology

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 A wild sea
In the distance over Sado
The Milky Way
Matsuo Basho

First the tide rushes in

Plants a kiss on the shore
Then rolls out to sea
And the sea is very still once more
Carl Sigman and Robert Maxwell

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 FOURTH EDITION

Marine Biology
Function, Biodiversity, Ecology

JEFFREY S. LEVINTON
Stony Brook University

New York Oxford

OXFORD UNIVERSITY PRESS
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Printing Number: 9 8 7 6 5 4 3 2 1
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 For Joan, Nathan, Andy and all others in the choir

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 BRIEF
CO N T E N TS

PREFACE XIV

PRINCIPLES OF OCEANOGRAPHY AND MARINE ECOLOGY

CHAPTER 1 Sounding the Deep 1
CHAPTER 2 The Oceanic Environment 13
CHAPTER 3 Ecological and Evolutionary Principles of Marine Biology 40

MARINE ORGANISMS: FUNCTION AND ENVIRONMENT

CHAPTER 4 The Chemical and Physical Environment 69
CHAPTER 5 Life in a Fluid Medium 91
CHAPTER 6 Reproduction, Dispersal, and Migration 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 Productivity, Food Webs, and Global Climate Change 225

ORGANISMS OF THE SEABED

CHAPTER 11 Benthic Microorganisms, Seaweeds, and Sea Grasses 242
CHAPTER 12 The Diversity of Benthic Marine Invertebrates 255
CHAPTER 13 Benthic Life Habits 283

COASTAL BENTHIC ENVIRONMENTS

CHAPTER 14 The Tidelands: Rocky Shores, Soft-Substratum Shores, Marshes,
Mangroves, Estuaries, and Oyster Reefs 309
CHAPTER 15 The Shallow Coastal Subtidal: Sea Grass Beds, Rocky Reefs,
Kelp Forests, and Coral Reefs 361

FROM THE SHELF TO THE DEEP SEA

CHAPTER 16 Benthos from the Continental Shelf to the Deep Sea 404
CHAPTER 17 Biodiversity and Conservation of the Ocean 434

HUMAN IMPACT ON THE SEA

CHAPTER 18 Fisheries and Food from the Sea 461
CHAPTER 19 Environmental Impacts of Industrial Activities and Human
Populations 492

GLOSSARY 517
MARINE BIOLOGY JOURNALS 535
INDEX 536

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 CO N T E N TS

PREFACE XIV

PRINCIPLES OF OCEANOGRAPHY AND MARINE ECOLOGY

CHAPTER 1 Sounding the Deep 1
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 Questions 12

CHAPTER 2 The Oceanic Environment 13

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
Hypothesized Effects of Climate Change on Sea Level, Circulation, and Ocean
Chemistry 32
Going Deeper 2.1 Solubility 33
The Edge of the Sea 33
Chapter Summary 38
Review Questions 39

CHAPTER 3 Ecological and Evolutionary Principles of Marine Biology 40

Ecological Interactions 40
The Ecological Hierarchy 40
Interactions on the Scale of Individuals 41
The Population Level 48
The Community Level: Structure and Interspecies Interactions 52
The Ecosystem Level 60
Species, Genetic Variation, Evolution, and Biogeography 62
Chapter Summary 67
Review Questions 68

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 viii CONTENTS

MARINE ORGANISMS: FUNCTION AND ENVIRONMENT

CHAPTER 4 The Chemical and Physical Environment 69
Measures of Physiological Performance 69
Temperature 71
Hot Topics In Marine Biology 4.1 Global Warming: How Can We Tell
If We Are at the Edge? 78
Salinity 81
Oxygen 83
Going Deeper 4.1 Quantifying the Relationship Between Body Size
and Oxygen Consumption Rate 83
Light 87
Chapter Summary 89
Review Questions 90

CHAPTER 5 Life in a Fluid Medium 91

Introduction 91
Density, Viscosity, and Reynolds Number 91
Going Deeper 5.1 Is Seawater Always Seawater? A Tale from the Regions
of Intermediate Reynolds Number 93
Moving Water 93
Water Moving over Surfaces and Obstructions, Such as Organisms 95
Using Water Motion for Biological Advantage 96
Chapter Summary 101
Review Questions 101

CHAPTER 6 Reproduction, Dispersal, and Migration 102

Ecological and Evolutionary Factors in Sex 102
Hot Topics In Marine Biology 6.1 A Lover and a Fighter 104
Hot Topics In Marine Biology 6.2 Whos Your Daddy? An Investigation
of 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 the
Serengeti 119
Planktonic Larvae: Getting Through Major Obstacles to the Final
Destination 133
The Macroscale: Major Separations Lead to Biogeographic Structure 134
Planktonic Dispersal: Why Do They Do It? 137
Chapter Summary 139
Review Questions 139

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 CONTENTS ix

ORGANISMS OF THE OPEN SEA

CHAPTER 7 Plankton 141
Introduction and Definitions 141
Marine Viruses 141
Marine Bacteria and Archaea 142
Eukaryotic Phytoplankton 142
Zooplankton 145
Gelatinous Zooplankton 149
Life in the Open Sea 152
Defense Against Predation 153
Diurnal Vertical Migration of the Zooplankton 154
Hot Topics In Marine Biology 7.1 Bioluminescence, Night Vision,
and Death in the Deep 155
Molecular Techniques to Identify Planktonic Microorganismal Diversity 158
Chapter Summary 161
Review Questions 161

CHAPTER 8 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
Hot Topics In Marine Biology 8.2 Last March of the Penguins?
Climate Change and a Bottom-Up Trophic Cascade 187
Marine Reptiles 194
Chapter Summary 196
Review Questions 196

PATTERNS AND PROCESSES IN THE WATER COLUMN

CHAPTER 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 205
Going Deeper 9.1 The Basics of Photosynthesis 207
Nutrients Required by Phytoplankton 208
Rate of Nutrient Uptake 213
Harmful Algal Blooms 215
Phytoplankton Succession and the Paradox of Phytoplankton Coexistence 216
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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 x CONTENTS

The Microbial Loop: Nutrient Cycling by Viruses, Heterotrophs,

and Chemoautotrophs 219
Zooplankton Grazing in the Sea 220
Going Deeper 9.2 Quantification of the Effect of Grazing 221
Chapter Summary 223
Review Questions 223

CHAPTER 10 Productivity, Food Webs, and Global Climate Change 225

Productivity and Biomass 225
Food Webs and Food Chains 225
Measuring Primary Productivity 229
Going Deeper 10.1 How to Calculate Productivity Using the Oxygen
Technique 230
Going Deeper 10.2 Using the Radiocarbon Technique to Estimate
Productivity 231
Geographic Distribution of Primary Productivity 233
Global Climate Change and the Global Carbon Pump 237
Chapter Summary 241
Review Questions 241

ORGANISMS OF THE SEABED

CHAPTER 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
Review Questions 254

CHAPTER 12 The Diversity of Benthic Marine Invertebrates 255

Hot Topics In Marine Biology 12.1 Where Did All This Invertebrate Diversity
Come 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 268
Phylum Ponogonophora: Gutless Wonders 268
Phylum Mollusca: Shelled Invertebrates (Mostly) 269
Phylum Arthropoda: Jointed Appendages 272
The Lophophorate Phyla 275
Phylum Bryozoa: Moss Animals 275

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 CONTENTS xi

Phylum Brachiopoda: Lingulas and Lampshells 276

Phylum Phoronida: Wormlike Animals with a Lophophore 277
Phylum Echinodermata: Animals with Fivefold Symmetry 277
Phylum Chordata: The Sea Squirts 280
Chapter Summary 281
Review Questions 282

CHAPTER 13 Benthic Life Habits 283

Life in Mud and Sand 283
Going Deeper 13.1 Measuring Grain Size of Sediments 284
Life Habits on Hard Surfaces and in Moving Waters 297
Benthic Carnivores 300
Benthic Herbivores 301
Hot Topics In Marine Biology 13.1 Diary of a Stinging Snail 302
Chapter Summary 307
Review Questions 308

COASTAL BENTHIC ENVIRONMENTS

CHAPTER 14 The Tidelands: Rocky Shores, Soft-Substratum Shores, Marshes,
Mangroves, Estuaries, 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 Invasion
and Climate Change 333
Spartina Salt Marshes 335
Mangrove Forests 343
Hot Topics In Marine Biology 14.2 The Molecular Sleuth: Invasion of a Very
Aggressive Genotype 344
Estuaries 349
Oyster Reefs 355
Chapter Summary 359
Review Questions 360

CHAPTER 15 The Shallow Coastal Subtidal: Sea Grass Beds, Rocky Reefs,
Kelp Forests, and Coral Reefs 361
Sea Grass Beds 361
The Rocky ReefKelp Forest System 366
Subtidal Rocky Reefs 366
Hot Topics In Marine Biology 15.1 Reorganization of a Rocky Subtidal
Ecosystem: A Cod and Lobster Tale 367
Kelp Forests 371
Coral Reefs 378
Chapter Summary 401
Review Questions 402

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 xii CONTENTS

FROM THE SHELF TO THE DEEP SEA

CHAPTER 16 Benthos from the Continental Shelf to the Deep Sea 404
The Subtidal Landscape 404
Sampling the Subtidal Soft-Bottom Benthos 404
Sediment Type and Benthic Distribution 406
The Deep-Sea Gradient 410
The ShelfDeep-Sea Gradient 410
Deep-Sea Island Hotspots of High Diversity 417
Pressure Change 428
Polar Environments 428
Arctic Subtidal Benthic Environments 429
The Antarctic Shelf Benthos: A Special Case of Isolation 430
Chapter Summary 432
Review Questions 433

CHAPTER 17 Biodiversity and Conservation of the Ocean 434

Diversity, Speciation, Extinction, and Biogeographic Factors 434
Major Gradients of Species Diversity 442
Explanations of Regional Diversity Differences 446
Conserving Marine Biodiversity 450
Marine Invasions 456
Chapter Summary 459
Review Questions 460

HUMAN IMPACT ON THE SEA

CHAPTER 18 Fisheries and Food from the Sea 461
The Fishery Stock and Its Variability 461
Stocks and Markers 461
Life History and Stock Size 463
Stock Health and Production 464
Going Deeper 18.1 A Simple Model to Explain the Maximum Sustainable Yield 467
Fishing Techniques and Their Effects 468
Fisheries Impact and Management 471
Causes and Cures of Stock Reduction 474
Hot Topics In Marine Biology 18.1 Is Fisheries Policy at Odds with Managing
Marine Ecosystems? 475
Going Deeper 18.2 Age-Based or Life-History Stage-Based Population Models 479
Overexploitation of Whales: A Case History 482
Other Types of Degradation 484
Disease as a Major Danger to Coastal Fisheries 485
Mariculture 485
Drugs from the Sea: Marine Natural Products 489
Chapter Summary 490
Review Questions 491

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 CONTENTS xiii

CHAPTER 19 Environmental Impacts of Industrial Activities and Human

Populations 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 Is the Gulf of Mexico Adapted to Oil? 504
Nutrient Input and Eutrophication 508
Thermal Pollution and Power Station Fish Mortality 512
Global Environmental Change and the Ocean 513
Chapter Summary 515
Review Questions 516

GLOSSARY 517

MARINE BIOLOGY JOURNALS 535

INDEX 536

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P R E FAC E
W
elcome to the fourth edition of Marine Biology:
Function, Biodiversity, Ecology. Th is new edi-
tion reflects my continued excitement at the
teaching of marine biology, a far-reaching science that
manages to excite a broad spectrum of students. We
marine biologists are lucky to have exciting creatures,
great adventures, and continuing discoveries that manage
to enchant students. The challenge in a text is to inject
this excitement into an organized approach to the subject.
Many students find marine biology a little daunting. No
surprise to me, because it is daunting to put it all into a
text. I believe, though, that every student can acquire the
broad spectrum of biological knowledge included in
marine biology while appreciating the organismal diver-
sity of the marine realm. To make that possible, students
must learn concepts along with the facts and begin to think
and reason like scientists. Just as important, they must feel
the pulse of current happenings.
The goal of this text is to appeal to a wide range of
students, while also preparing future specialists with the
knowledge and tools needed to conduct their research.
Marine biology is such a diverse subject that principles
must guide our understanding. I fi rmly believe that, with-
out principles and a sense of excitement, students will see
only an accumulation of facts. They will fail to see how
the same basic processes operate in vastly different marine
communities.
That is why the text addresses three major process-
driven themes: functional biology, ecological processes, and
biodiversity. It is why the text uses full color throughout
in drawings that depict marine biological processes and a
large number of photographs to connect students with
marine environments and organisms. It is also why I have
paid much attention to recent research developments and
have included a series of essays, called Hot Topics in
Marine Biology. These features will put students in con-
tact with the current world of exciting research. In both
the text and the Hot Topics, I try to connect the students
to some of the most important recent research with an
extensive 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 own
classes I point out that a good teacher shows students the
way to good science because library databases are not
user friendly where finding excellent science research is
concerned.
xiv
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The Philosophy of This Text
This text is designed for a one-semester course at the soph-
omore to senior level. Some students will have already
taken a college-level biology course with coverage of or-
ganismal diversity, and they will benefit greatly. A small
number 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 text
for many years, and many students had no background in
organismal biology or ecology. If the book is supplemented
with journal articles, it can also be used in a more advanced
undergraduate course in marine ecology. The new edition
contains many updated references to the primary literature,
which are expanded and now online with a simple link
from the text chapters. This is a valuable resource to get
students started on term papers and essays. The Marine
Biology Web Page (www.oup.com/us/levinton), which I
founded a number of years ago, links students to many
more views of marine biology and to a greater diversity of
organisms. It is a continuing objective to merge principles
with an appreciation of organismal diversity in the ocean.
I have taught Marine Biology for over 30 years and have
always been amazed at the diversity of students who take
the course. Biology majors, marine science majors, geology
majors, and even some humanities majors sit side by side.
At my university, marine science has become a separate
undergraduate discipline in recent years, and these stu-
dents have truly learned the interdisciplinary nature of
marine biology. All learn a great deal, and all seem to
come away with a love for the ocean. You dont have to
convince them to be there: They want to learn about
marine biology.
I do my best to keep that interest alive, and I find that
field trips and the use of color photographs and other
illustrations throughout the text help a great deal. So do
the online resources accompanying this text. Marine
Biology Explorations includes hundreds of photographs
from habitats discussed in the text and still more. As men-
tioned, an expanded and updated reference list is also
online. In class, I keep a large map of the oceans on the
wall for the geographic context of our discussions. I have
taken many of my students with me to marine labs, and
they have launched careers in science or used their back-
grounds to enter other areas. I hope the text will help a
wider audience to get excited about marine life. I hope,
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 P R E FAC E xv

too, that they will understand how the ocean works and hypotheses, as well as making tests practical enough that
why our marine realm is so threatened. they can be put to direct use. From the very beginning,
I introduce the student not only to the scientific method,
A Principles-Driven Approach but also to how it translates into an intellectual tool with
real-world applications.
Marine biology applies the principles of ecology and evolu-
Chapters 1 through 6 introduce basic principles of how
tion, using the crucial tools of cell biology, biomechanics,
the ocean works in a physical, chemical, and ecological
and molecular biology to a wide range of marine biological
context and how marine organisms function with these
studies. These fields of study and their interactions govern
constraints. The second chapter gives the student a com-
the texts three overarching themes: functional biology,
prehensive introduction to oceanography and the impor-
biodiversity, and ecological processes.
tant properties of sea water that might affect marine
Function refers to the way organisms solve problems
organisms. I work in climate change at this stage because
and how physical and chemical factors constrain and select
it will be part and parcel of many discussions throughout
the solutions. What shape should a maneuvering fish have,
the text. This edition greatly expands the effects of tem-
relative to a continuously swimming fish? How does a
perature and especially ocean acidification, and the stu-
small peptide manage to be such an effective poison when
dent sees how an understanding of the ocean and sea water
injected by a snail into a prey? How does this specific
properties will directly affect the fate of marine organisms
biochemical adaptation feed into an understanding of bio-
in the future. I then introduce ecological principles so that
diversity? I believe this textbook is unique in combining
students can work their way through concepts using
effectively functional biology with ecological thinking.
marine examples. This allows all students to be brought up
Biodiversity is an essential part of marine biology, and
to a level of ecological thinking and an understanding of
I introduce the topic both through introductions to the
oceanographic processes. They will see this big picture as
principles used to study and explain biodiversity and to
they read the rest of the text. A crucial chapter then intro-
the factors that strongly affect marine biodiversity. A sep-
duces students to how the physics of fluids shapes the con-
arate chapter discusses diversity and the processes that
straints and adaptations of marine organisms. As far as I
regulate it, both ecological and evolutionary. It is crucial
know, this crucial subject is missing in all other marine
that the student see the historical roots of many current
biology texts, and allows a connection to a complete un-
distributions, which are affected by processes ranging
derstanding of how the marine organisms function in the
from plate tectonics to climate change. This edition adds a
rather complex fluid environment. These chapters con-
great deal of coverage of recent advances, including
clude with a comprehensive introduction to reproductive
molecular tools used to identify microbial organisms and
strategies, larval dispersal, and migration, which sets up
activity in the plankton and molecular methods used to
the big picture of the geographical distribution of marine
study dispersal, speciation, and the rise of marine adapta-
species, down to the microscale of how mobile marine
tions. A chapter devoted to biodiversity makes the con-
larvae succeed in finding a place to live in a turbulent and
nections between evolutionary process, biodiversity, and
stressful world.
biogeography. These areas are related directly to issues of
Chapters 7 through 10 cover the organisms and processes
marine conservation.
important in the open sea. This two-step approach is es-
Last, ecology examines the interactions of organisms
sential so that students will understand the overall econ-
with their environment. Ecology tries to understand the
omy of the marine realm. Chapter 10 uses a global-scale
distribution and abundance of organisms. It involves a
approach to show how biological studies of the ocean lead
series of processes, which I introduce in the context of a
to an understanding of the worlds potential for fisheries
hierarchyfrom individual populations to ecosystems. It
and the global biological impact on the ocean of climate
also involves a discussion of important ecological processes
change. Processes in the water column are also crucial in
along with accounts of major marine habitats and com-
Chapters 11 through 15 for understanding the benthos,
munities. This edition pays special attention to modern
which depends both directly and indirectly on the water
concepts of populations and species interactions, including
world above. In Chapters 11 and 12 I cover benthic crea-
metapopulations, regional genetic differentiation, large-
tures and then go on to discuss the principles necessary to
scale control of dispersal, biological invasions, and alterna-
understand the biology of marine bottom organisms
tive stable states of communities. My background in
(Chapter 13) and the major nearshore marine bottom hab-
geology, ecology, and evolutionary biology allows me to
itats (Chapters 1416). By necessity, I have been selective.
frequently combine ecological and evolutionary thinking
I emphasize those habitats that are not only important and
in discussing marine biology problems.
interesting, but also where important principles can be il-
lustrated to their best advantage. Community-level inter-
Organization actions are emphasized, as is global climate change as it
We begin with a brief historical background. Marine biol- relates to major changes in habitats such as coral reefs.
ogy has a history that is worth understanding, but it is also I discuss a range of geographic locations so that the
crucial to introduce the student to how science works. instructor will find local examples in many instances.
Chapter 1 therefore discusses framing and testing Crucial habitats such as the intertidal, seagrasses, coral

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 xvi P R E FAC E
reefs, mangroves, estuaries, salt marshes, kelp forests, and
others 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 new
section on oyster reefs because of their great worldwide
importance as foci for biodiversity and their role in ecosys-
tem services.
Chapter16 then looks at the important gradient from
the continental shelf to the deep sea, paying special atten-
tion to some of the fascinating discoveries about biological
function and fascinating habitats, from hot vents to deep
water coral mounds to the newly discovered subsurface
bacterial realm over 500 m beneath the sea floor. I have
greatly expanded coverage of Arctic and Antarctic envi-
ronments. The Antarctic is center stage in our focus on the
current and future impacts of climate change, and the text
incorporates a wide range of discussions from the organis-
mal to the ecosystem level. Chapter 17, on gradients in
biodiversity, sums up larger-scale variants in the sea and
includes sections on invasive species, conservation of bio-
diversity, and conservation genetics. More and more, stu-
dents and researchers have focused their attention to the
deteriorating conditions of the ocean, and conservation is
a major field of emphasis.
Finally, Chapters 18 and 19 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 the
ocean. Throughout the text, the effects of climate change
are brought up in many contexts, and how those effects are
related to chemical issues such as acidification and facilita-
tion of biological invasions. I also place strong emphasis on
the reorganization of communities that has been initiated
by the interaction of human activities and strong ecologi-
cal interactions found in natural communities and in food
webs. The impact of overfishing on populations and
trophic cascades is a crucial part of a complete chapter on
fisheries and mariculture. The role of toxic substances,
eutrophication, and hypoxia are discussed clearly and in
depth. I have added to Chapter 18 a section on drug
discovery 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 and
disease, such as cancer.
A Refined Learning Package
This text has a series of pedagogical features designed to
help students absorb a great variety of information by
engaging their imaginations, helping them organize and
prioritize important principles, and keeping them focused
on the big without getting lost in the details.
Hot Topics in Marine Biology essays throughout
the text introduce students to recent advances in the un-
derstanding of marine biology and discuss current issues,
especially marinebiological debates and discoveries.
Instructors can use these essays to kick off discussion, to
expand a students horizons, for course assignments, or as
topics for term papers. Reorganizations of communities
from the Antarctic to the Gulf of Maine are related to
00-Levinton-FM.indd xvi
climate change, overfishing, and other processes that are
now actively being investigated. A new section on the
amazing discoveries of long-distance migration paths
through the Pacific Ocean has been added to Chapter 6.
Ihave also tried to give some additional understanding of
some current problems by adding a Hot Topics feature
that addresses exciting new molecular techniques used to
understand marine mammal population structure (Chap-
ter 8), the vexing mortality of marine mammals on the
Pacific Coast (Chapter 9), the major issue of management
of fisheries (Chapter 18), and what the recent horrific Gulf
Oil 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 of
climate change in the Antarctic Ocean and the impacts on
penguin species in Antarctica.
Key Concept full-sentence summary statements begin
nearly every section of the text to help students identify
central points of discussion and to foreshadow whats to
come. These headings allow students to discern the forest
from the trees and to see the basic progression of material.
Each chapter ends with a bulleted Chapter Summary
and a variety of Thought Questions. Instructors and stu-
dents can use these to follow up on important issues in
marine biology. The combination of these features and the
Key Concept heading sentences successfully guides the
student through a complex subject.
Going Deeper boxes explain equations and related
concepts 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 and
photosynthesis). They also allow instructors who chose to
omit them to press on with no interruptions. An example
is the discussion of Leslie Matrices in Chapter 18, which
give the student an idea of how age-structured population
models help to understand impacts of various factors on
fisheries and management decisions.
Extensive References lists of classic and contemporary
scholarship that instructors may assign as reading and that
can lead students to further assignments are linked online
from the text. These help students see that marine biology
is 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 the
end of the text provides students access to get a quick def-
inition of important concepts, processes, and terms. A list
of journals is a resource for students in writing term
papers and for further research.
What Is New and Noteworthy
in the Fourth Edition?
Expanded illustration program. We continue the fourth
edition using a rich color presentation in order to better
demonstrate marine biological principles and introduce or-
ganismal diversity in a vivid and captivating visual presen-
tation. The new edition includes over 50 new photos and
line drawings (many of the photos generously contributed
26/04/13 10:25 PM

by colleagues), and I believe students will benefit greatly
from now having the color photos integrated directly into
the relevant textual discussion at hand.
More applications. To engage students with the diver-
sity of marine biology today and to highlight the real-world
applications of what they are learning, Ive written many
new in-text examples, including seven new Hot Topics in
Marine Biology. Students will see how molecular tools
can be used to study mating success, how a large-scale pro-
gram of fish tagging has established major migration routes
in the North Pacific Ocean, how a mysterious series of
deaths of marine mammals in California was linked to a
toxic species of diatom, how fishing management practice
can have strong effects on the structure of marine ecosys-
tems, and how microbial degradation may be responsible
for a rapid degradation of some components of hydrocar-
bons following the disastrous Gulf Oil Spill of 2010.
Current and expanded topics maintain the excitement
that underlies my philosophy of teaching and have been
carefully selected to bring the text up-to-date while still
remaining focused on the most important principles
students need to learn.
Ecological interactions. Strong attention is paid to major
ecological interactions that are relevant to ecosystem
structure, such as trophic cascades (Chapters 15 and
18), ecological reorganization in New England and
elsewhere (Chapter15), molecular approaches to
ecology and evolution (Chapters 7, 8, 13, and 14),
natural and human-induced phase shifts (Chapters 14
and 15), biological invasions (Chapters 14, 15, and 17),
and climate change (Chapters 4, 8, and 15).
Evidence and effects of climate change. I have greatly
expanded coverage of climate change, with attention
paid to temperature change in the global ocean and in
coastal areas (Chapter2); the ocean acidification
(Chapters 2, 10 and 15); the role of climate change in
changing species distributions, facilitating biological
invasions, and causing thermal stress (Chapters 2, 4,
14, and 15); and other topics.
Methods of environmental assessment, from remote sensing
to the molecular level. I have also expanded coverage of
the latest methods for remote sensing, estimating world
productivity, and assessing the stress on and change of
ecosystems, including satellite methods and ocean ob-
servatories (Chapters 1 and 10), genetic and molecular
studies of population differentiation (Chapters 3, 6, 17,
and 18), the shifting baseline concept (Chapter18),
diversity gradients and the tropical origins of bio-
diversity (Chapter17), and molecular methods to
trace biological invasions (Chapters 14 and 17).
Human impact on biodiversity. This edition expands
coverage of the decline of coral reefs and adds insights
on other biological impact such as the increase of
sponges at the expense of corals (Chapter15), over-
fishing and the issue of relating management decisions
to management of the basis of ecosystem function
(Chapters 15 and 18), more on declines of sharks and
00-Levinton-FM.indd xvii
P R E FAC E xvii
other apex predators (Chapter18), and the effects of
pollution, especially with coverage of the Deepwater
Horizon well blowout (Chapter19).
Drug discovery. I have added a section on the use of
marine natural products in the development of drugs.
Supplements
Marine Biology, Fourth Edition, is accompanied by a wealth
of electronic resources for both students and instructors,
including a FREE Companion Website, (www.oup.com/
us/levinton) and the Instructors Resource CD-ROM.
Companion Website: Maintained by the author, this
Companion Website (www.oup.com/us/levinton) pro-
vides a multitude of resources for both students and
instructors.
Student Resources
Marine Biology Explorations. Explore the oceans
biodiversity through interactive exercises that will
take you through nine different marine habits; in-
cluding over 450 photos with annotations!
Marine Biology in the News. Frequently updated
current breakthroughs in marine biology research.
Extensive web links to marine biology topics and
research literature. You will also find information
on careers in marine biology and worldwide
marine laboratories.
Instructor Resources (available to adopters of the
text and password-protected)
Electronic Images. All illustrations from the text
available in electronic format for download for
lecture presentations.
PowerPoint Lecture Notes. Over 400 lecture
notes slides organized by chapter.
Test Bank. This comprehensive resource includes
approximately 400 questions written by the author
himself in editable Word files for easy customiza-
tion (available only on Instructors Resource
CD-ROM: contact your Oxford University Press
sales representative for details).
Acknowledgments
Many people have helped metoo many to mention all indi-
vidually. I am especially grateful to the many individuals who
shared their photographs and research experiences with me.
Many sent me preprints, photographs, data, and just plain in-
teresting discussions, which helped me greatly to understand
fields unfamiliar to me. For the current edition, Jason Noe
served as editor and continually moved this project forward.
Keith Faivre worked as an excellent liaison between me and
copy 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, marketing
manager; Frank Mortimer, director of marketing; Michele
Laseau, art director; and Lisa Grzan, managing editor.
I also benefited a tremendous amount from careful review
of the manuscript and additional reviewer recommendations
26/04/13 10:25 PM
 xviii P R E FAC E
based on the third edition. I had a wonderful panel of advi-
sors for the third edition but have continued to be lucky with
the truly excellent panel that helped me with revisions for
the fourth. I could not have done better with such an excel-
lent panel:
Christopher D. Beatty, Santa Clara University
Heather Bennett, Illinois College
Michael Berger, Washington State University
John Berges, University of WisconsinMilwaukee
Patricia M. Biesiot, University of Southern Mississippi
Matthew Bracken, Northeastern University
W. Randy Brooks, Florida Atlantic University
Erin J. Burge, Coastal Carolina University
Deron Burkepile, Florida International University
Marymegan Daly, Ohio State University
Carrie DeJaco, Queens University of Charlotte
Steve Dudgeon, California State University, Northridge
Joy Ferenbaugh, James Madison University
Michael Franklin, California State University, 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 at Jacksonville
Katrin Iken, University of Alaska Fairbanks
Yan Jiao, Virginia Tech University
Catherine Teare Ketter, University of Georgia
George Kraemer, Purchase College, State University
of 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 Gulf
Coast
Linn Montgomery, Northern Arizona University
Anthony Moss, Auburn University
Xiaozhen Mou, Kent State University
Steven Mark Norris, California State University Channel
Islands
Antonios Pappantoniou, Housatonic Community College
00-Levinton-FM.indd xviii
Chris Parsons, George Mason University
Neil Pelkey, Juniata College
Curt Pueschel, Binghampton University
Marjorie Reaka, University of Maryland
Amy Ringwood, UNC Charlotte
Cliff Ross, University of North Florida
Wendy L Ryan, Kutztown University
David Scholnick, Pacific University
Jyotsna Sharma-Srinivasan, University of Texas at San
Antonio
William Stickle, Louisiana State University
Jacqueline Webb, University of Rhode Island
Mary Wicksten, Texas A&M University
Larissa M. Williams, Wheaton College
Tim Wootton, University of Chicago
I also again thank reviewers whose insights contributed to
past editions: Jelle Atema, Boston University; Susan S.
Bell, University of South Florida; Larry E. Brand, Univer-
sity of Miami; Christopher Brown, Florida International
University; James E. Byers, University of New Hampshire;
Edward J. Carpenter, San Francisco State University;
Gerardo Chin-Leo, Evergreen State College; Paul Dayton,
Scripps Institute of Oceanography; Chris DElia, Louisiana
State University; Sean Patrick Grace, Southern Connecti-
cut State University; Larry G. Harris, University of New
Hampshire; William W. Kirby-Smith, Duke University;
Alan J. Kohn, University of Washington; Derek R. Lavoie,
Cuesta College; Larry R. McEdward, University of
Florida; George McManus, University of Connecticut;
Amy Moran, Clemson University; Stephen Norton, East
Carolina 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 University; Keith Walters, Coastal Carolina
University; and Judith S. Weis, Rutgers University.
Some of this edition was prepared at my home institu-
tion, Stony Brook University, but I am very grateful to the
staff of the Friday Harbor Laboratories, University of
Washington, where I spent a summer working on this
text. I am especially grateful to the vision of Arthur
Whiteley for creating, supporting, designing, and nurtur-
ing the Helen Riaboff Whiteley Center, where I worked
and lived. As usual, my wife Joan was supportive and help-
ful with suggestions.
Jeffrey Levinton
Stony Brook University, Stony Brook, New York
26/04/13 10:25 PM