Marine Biology: An Ecological Approach

598 Pages·2004·49.989 MB·English

by  Nybakken, James W., Bertness, Mark D.

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JAMES W. NYBAKKEN
MARK D. BERTNESS
SIXTH EDITION
Digitized by the Internet Archive
in 2021 with funding from
Kahle/Austin Foundation
httos://archive.org/details/marinebiologyeco0000nyba_s3c0

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ne Benjamin
—_ Cummings
ACQUISITIONS EDITOR: Chalon Bridges
EDITOR-IN-CHIEF: Beth Wilbur
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MANAGING EDITOR: Erin Gregg
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TEXT AND COVER DESIGNER: Seventeenth Street Studios
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TEXT PRINTER: R.R. Donnelley, Willard
ISBN 0-8053-4582-5 (Student)
ISBN 0-8053-4586-8 (Instructor)
Library of Congress Cataloging-in-Publication Data
Nybakken, James Willard.
Marine biology: an ecological approach / James W. Nybakken, Mark Bertness.—6th ed.
p. cm.
Includes bibliographical references and index.
ISBN 0-8053-4582-5
1. Marine biology. 2. Marine ecology. I. Bertness, Mark D., 1949- II. Title.
QH91.N9 2004
577.7--dc22
2004052771
On the cover: Photograph of a Blue Chromis (Chromis cyanea) CHAPTER OPENING PHOTO CREDITS: * Chapter 1: © Rick
over Boulder Brain Coral (Colpophyllia natans) Doyle/CORBIS + Chapter 2: © Alfred Pasieka/Photo Researchers, Inc.
in the Caribbean. Copyright © 2003 + Chapter 3: © Amos Nachoum/CORBIS + Chapter 4: © James Nybakken
Norbert Wu/www.norbertwu.com. + Chapter 5: Ralph A. Clevenger/CORBIS + Chapter 6: Art Wolfe/Getty
Images/Stone + Chapter 7: © Andrew Syred/Photo Researchers, Inc.
+ Chapter 8: Phil Schermeister/CORBIS + Chapter 9: © CORBIS
+ Chapter 10: JeffH unter/Getty Images/The Image Bank + Chapter 11:
Paul Hanna/ CORBIS/Reuters
Copyright © 2005 Pearson Education, Inc., publishing as Benjamin Cummings, 1301 Sansome St., San Francisco, CA 94111.
All rights reserved. Manufactured in the United States of America. This publication is protected by Copyright and permission should be obtained
from the publisher prior to any prohibited reproduction, storage in a retrieval system, or transmission in any form or by any means, electronic,
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Many of the designations used by manufacturers and sellers to distinguish their products are claimed as trademarks. Where those designations
appear in this book, and the publisher was aware of a trademark claim, the designations have been printed in initial caps or all caps.
56789 10—DOW—08 07
Preface, ix = COMPARISON OF TERRESTRIAL
AND MARINE ECOSYSTEMS a1
Physical and Chemical Differences, 32
CHAPTER 1
Biodiversity Differences, 34
Introduction to
Life-History Differences, 34
the Marine Environment Structural and Functional Differences, 35
= DIVISION OF THE MARINE
= PROPERTIES OF WATER
ENVIRONMENT 35
Chemical Composition, 1
Major Subdivisions of the World Ocean, 36
Physical and Chemical Properties, 2
Summary of Key Concepts, 38
Seawater, 4
CHAPTER 2
BASIC OCEANOGRAPHY Plankton and Plankton
Geography and Geomorphology
Communities 42
of the Oceans, 7
Plate Tectonics, 9 Definitions, 42
Temperature and Vertical Stratification, 10
Water Masses and Circulation, 11 THE PHYTOPLANKTON 43
Diatoms, 43
Dinoflagellates, 45
SOME ECOLOGICAL PRINCIPLES
The Smaller Phytoplankton, 46
Terms and Definitions, 19
The Viruses, 47
Ecosystem Components, 20
Biogeochemical Cycles, 20
THE ZOOPLANKTON 48
Biotic Structure of Ecosystems, 21
Copepods, 48
Ecological Control and Regulation, 24
Other Zooplankton, 50
LARVAE AND LARVAL ECOLOGY ZS FLOTATION MECHANISMS 56
Larval Types and Strategies, 26 Principles 56
Larval Ecology and Reduction of Overweight, 58
Community Establishment, 29 Changes in Surface of Resistance, 59
Life-History Strategies, 29 Water Movements, 60
iv ¢ CONTENTS
PRIMARY PRODUCTION 61 Surface of Resistance and Body Shape, 110
Primary Productivity, 61 Defense and Camouflage, 114
Measurement of Primary Productivity, 62 Sensory Systems, 116
Standing Crop, 65 Echolocation, 117
Reproduction and Life Cycle, 119
FACTORS AFFECTING PRIMARY Migrations, 125
PRODUCTIVITY 65 Special Adaptations of Marine Birds
Physical and Chemical Factors, 65 and Mammals, 128
Light, 66
= ECOLOGY OF NEKTON 132
Nutrients, 68
Feeding Ecology and Food Webs, 132
Turbulence and Critical Depth, 72
Geographical Variations in Productivity, 72 Ecological Significance of Marine Mammals, 137
Summary of Key Concepts, 141
Temperate Seas, 73
Tropical Seas, 75
Polar Seas, 75 CHAPTER 4
Productivity in Inshore and Coastal Waters, 75
Deep-Sea Biology 144
PRIMARY PRODUCTIVITY
OF THE BIOSPHERE 76
= ZONATION 145
THE OCEAN ECOSYSTEM: THE CLASSIC
= SAMPLING THE DEEP SEA 146
MODEL 7/7
=» ENVIRONMENTAL CHARACTERISTICS 148
Grazing, 77
Copepod Cycles, 80 Light, 148
Copepod Influence on Atlantic and Pacific Plankton Pressure, 150
Community Ecology, 82 Salinity, 151
Vertical Migration, 85 Temperature, 151
Seasonal Succession in Phytoplankton, 87 Oxygen, 152
Food, 153
THE OCEAN ECOSYSTEM: A CHANGING Biomass, 155
MODEL 89
= ADAPTATIONS OF DEEP-SEA
Microorganisms and Productivity, 89
ORGANISMS 155
Nanoplankton, Respiration, and Grazing, 90
Bacteria, Particulates, and Dissolved Bioluminescence in the Deep Sea, 163
Organic Matter, 91
=» COMMUNITY ECOLOGY
The Marine Planktonic Food Web, 94
OF THE BENTHOS 166
Spatial Distribution of Plankton, 94
The Major Plankton Biomes, 96 Faunal Composition, 166
Summary of Key Concepts, 98 Community Structure, 167
Diversity, 169
Seamounts, 172
CHAPTER 3
Life-History Patterns, 174
Oceanic Nekton 103
Hydrothermal-Vent and Cold-Seep
Communities, 175
COMPOSITION OF THE OCEANIC
= MIDWATER COMMUNITY ECOLOGY 187
NEKTON 104
Deep Scattering Layers and Vertical Migration, 187
ENVIRONMENTAL CONDITIONS 105
Species Composition and Distribution, 188
ADAPTATJONS OF OCEANIC NEKTON 106 Food and Feeding, 188
Bric Life-History Patterns of Midwater Organisms, 191
Lo ¢ Summary of Key Concepts, 191
CONTENTS ¢ v
CHAPTER 6
CHAPTER 5
Shallow-Water Subtidal Intertidal Ecology 266
Benthic Associations 196
=» ENVIRONMENTAL CONDITIONS 267
Coverage and Definitions, 196
Tides, 267
= ENVIRONMENTAL CONDITIONS 196 Temperature, 270
Wave Action, 271
= UNVEGETATED SEDIMENTARY
Salinity, 271
ENVIRONMENTS 198
General Patterns, 198
= ADAPTATIONS OF INTERTIDAL
Sampling, 198
ORGANISMS 272
Infaunal Associations, 199
Resistance to Water Loss, 272
Community Pattern and Structure, 201
Maintenance of Heat Balance, 274
Community Structure and Change, 202
Mechanical Stress, 275
Positive Interactions, 209
Respiration, 276
Vertical Distribution and
Feeding, 276
Competition, 209
Salinity Stress, 276
Community Patterns, Stability,
Reproduction, 277
and Variability, 210
= ROCKY SHORES 277
= ROCKY SUBTIDAL COMMUNITIES 214
Atlantic and Pacific Rocky Shores, 277
= KELP BEDS AND KELP FORESTS 221
Zonatio, 278
Structure and Distribution, 223 Causes of Zonation, 280
Ecology and Life Cycle, 225
Physical Factors, 280
Biological Factors, 284
= SEAGRASS COMMUNITIES 253 Patchiness, 297
Composition and Distribution, 233 Succession, 300
Environmental Conditions, 233 Horizontal Distribution Patterns, 302
Productivity, 236 Tide Pools, 302
Structure and Biological Interactions, 236 Tropical Intertidal Shores, 305
Relationships to Other Systems, 239
Ecology, 240 = COBBLE BEACHES 306
=» SANDY SHORES 308
=» SOME SPECIAL COMMUNITIES 24) Environmental Conditions, 309
A Sea-Pen Community, 241 Adaptations of Organisms, 313
A Sea-Pansy Community, 243 Types of Organisms, 315
Feeding Biology, 315
= BIOLOGY OF POLAR SEAS 243 Community Organization on Exposed
Physical Conditions of the Arctic Sand Beaches, 317
and Antarctic Regions, 245 Community Organization on Protected
Biological Comparisons of Arctic Sand Flats, 321
and Antarctic Seas, 248
Ecological Communities =a MUDDY SHORES 325
of Polar Seas, 250 Physical Factors, 325
Adaptations of Organisms, 326
Sea-Ilce Communities, 250
An Antarctic Soft-Sediment Types of Organisms, 327
Community, 257 Feeding Biology and Trophic
An Antarctic Hard-Bottom Community, 258 Structure, 328
Summary of Key Concepts, 260 Zonation and Community Structure, 330
vi ¢ CONTENTS
= INTERTIDAL FISHES 331 = ECOLOGY OF ESTUARIES 374
= BIRDS 332 Productivity, Organic Matter,
and Food Sources, 374
Summary of Key Concepts, 334
Community Structure of Estuaries, 376
Food Webs, 377
CHAPTER 7 Plankton Cycles, 380
Meiofauna 342
= SALT MARSHES 381
Definitions, 342 Definition and Characterization, 382
Environmental Characteristics, 382
= ENVIRONMENTAL CHARACTERISTICS 343 Composition and Distribution, 383
= COMPOSITION OF THE INTERSTITIAL Marsh Development and Zonation, 383
ASSEMBLAGES 345 Causes of Zonation, 388
= SAMPLING AND EXTRACTING Positive Plant-Animal Interactions, 390
Positive Plant Feedback, 392
MEIOFAUNA 349
Productivity, 393
= ADAPTATIONS 351
Consumer Control in Salt-Marsh Systems, 395
Life History, 352
Trophic Interactions and Food Webs, 397
Human Impacts on Salt Marshes, 400
= ECOLOGY 353
Summary of Key Concepts, 401
Meiofauna—Macrofauna Interactions, 356
Species Richness, Distribution, CHAPTER 9
and Dispersal, 357
Summary of Key Concepts, 358 Tropical Communities 407
= CORAL REEFS 407
CHAPTER 8
Reef Distribution and Limiting Factors, 409
Estuaries and Salt Marshes _ 361
Structure of Corals, 410
Types of Reefs, 411
= TYPES OF ESTUARIES 361
Origin of Reefs, 412
= PHYSICAL CHARACTERISTICS Composition of Reefs, 414
Coral Distribution and Reef Zonation, 417
OF ESTUARIES 364
Salinity, 364 Atlantic and Indo-Pacific Reefs, 422
Productivity, 423
Substrate, 366
Biology of Hermatypic Corals, 424
Temperature, 367
Nutrition, 424
Wave Action and Currents, 367
Growth and Calcification, 425
Turbidity, 368
Sexual Maturity, Reproduction,
Oxygen, 368
and Recruitment, 426
Species Interactions and the Ecology
THE BIOTA OF ESTUARIES 368
of Reefs, 428
Faunal Composition, 368
Competition, 429
Estuarine Vegetation, 370
Predation, 430
Estuarine Plankton, 371
Grazing, 432
Positive Interactions, 435
= ADAPTATIONS OF ESTUARINE Regional Differences, 436
ORGANISMS 371 Bacteria in Reef Systems, 436
Role of Algae in Reef Systems, 436
Ecology of Reef Fishes, 437
Coral Reef Cryptofauna and Bioerosion, 447
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