Marine Ecological Processes

555 Pages·1984·14.044 MB·English

by  Ivan Valiela (auth.)

Springer Advanced Texts in Life Sciences
David E. Reichle, Editor
Photo by Jan Hahn, courtesy of Woods Hole Oceanographic Institution.
Ivan Valiela
Marine Ecological
Processes
With 220 Figures
Springer Science+Business Media, LLC
Ivan Valiela Series Editor:
Boston University Marine Program David E. Reichle
Marine Biological Laboratory Environmental Sciences Division
Woods Hole, Massachusetts 02543 Oak Ridge National Laboratory
U.S.A. and Oak Ridge. Tennessee 37830 U.S.A.
Department of Biology
Boston University
Boston, Massachusetts 02215 U.S.A.
Library of Congress Cataloging in Publication Data
Valiela, Ivan.
Marine ecological processes.
(Springer advanced texts in life sciences)
Bibliography: p.
Includes index.
I. Marine ecology. I. Title. II. Series.
QH541.5.S3V34 1984 574.5'2636 83-20037
Printed on acid-free paper.
© 1984 by Springer Science+Business Media New York
Originally published by Springer-Verlag New York, Inc.in 1984.
Softcover reprint of the hardcover 1st edition 1984
All rights reserved. No part of this book may be translated or reproduced in
any form without written permission from Springer Science+Business Media, LLC.
The use of general descriptive names, trade names, trademarks, etc., in this
publication, even if the former are not especially identified. is not to be taken
as a sign that such names, as understood by the Trade Marks and Merchandise
Marks Act, may accordingly be used freely by anyone.
Typeset by Bi-Comp, Incorporated, York, Pennsylvania.
9 8 7 654 3 2
ISBN 978-1-4757-1835-5 ISBN 978-1-4757-1833-1 (eBook)
DOI 10.1007/978-1-4757-1833-1
Preface
This text is aimed principally at the beginning graduate or advanced
undergraduate student, but was written also to serve as a review and,
more ambitiously, as a synthesis of the field. To achieve these purposes,
several objectives were imposed on the writing. The first was, since ecol
ogists must be the master borrowers of biology, to give the flavor of the
eclectic nature of the field by providing coverage of many of the interdis
ciplinary topics relevant to marine ecology. The second objective was to
portray marine ecology as a discipline in the course of discovery, one in
which there are very few settled issues. In many instances it is only
possible to discuss diverse views and point out the need for further study.
The lack of clear conclusions may be frustrating to the beginning student
but nonetheless reflects the current-and necessarily exciting-state of
the discipline. The third purpose is to guide the reader further into
topics of specialized interest by providing sufficient recent references
especially reviews. The fourth objective is to present marine ecology for
what it is: a branch of ecology. Many concepts, approaches, and methods
of marine ecology are inspired or derived from terrestrial and limnological
antecedents. There are, in addition, instructive comparisons to be made
among results obtained from marine, freshwater, and terrestrial environ
ments, I have therefore incorporated the intellectual antecedents of par
ticular concepts and some non-marine comparisons into the text.
The plan of this book is to present information on specifics about
physiological and populational levels of biological organization in Chap
ters 1~ 7. Notions of how populations relate to each other, and their envi
ronment, are documented (Chapters 8-9) and so community ecology is
introduced. This is followed by Chapters 10-12, where major aspects of
the chemistry of organic matter and nutrients in marine ecosystem are
developed, based on much of the material of previous chapters. Then,
having provided the essentials for understanding the working of various
processes in marine ecosystems, the final chapters (Chapters 12-15) dwell
vi Preface
on how the structure of marine communities and ecosystems may be
maintained over space and time.
Although I am responsible for whatever errors remain, this book has
been greatly improved by many people. I have to thank my colleages in
Woods Hole, especially John Teal and John Hobbie, for many years of
discussion and exchange of ideas. One or more chapters were criticized
by Randy Alberte, Karl Banse, Judy Capuzzo, Hal Caswell, Jon Cole,
Joseph Connell, Tim Cowles, Werner Deuser, Bruce Frost, Joel Gold
man, Charles Greene, Marvin Grosslein, Loren Haury, John Hobbie,
Robert Howarth, Michael Landry, Cindy Lee, Jane Lubchenco, Kenneth
Mann, Roger Mann, Scott Nixon, Mark Ohman, Bruce Peterson, Donald
Rhoads, Amy Schoener, Sybil Seitzinger, Charles Simenstad, Wayne
Sousa, and Richard Wiegert.
The graduate students associated with my laboratory during the writing
of this book have served as a critical sounding board, and have substan
tially contributed in many ways. I therefore have to acknowledge the
contributions of Gary Banta, Donald Bryant, Robert Buchsbaum, Nina
Caraco, Charlotte Cogswell, Joseph Costa, Cabell Davis, William Denni
son, Kenneth Foreman, Rod Fujita, Anne Giblin, Jean Hartman, Brian
Howes, Alan Poole, Armando Tamse, Christine Werme, David White,
and John Wilson. All of them have helped in some fashion with this text,
especially Kenneth Foreman and Anne Giblin, who read and criticized
most of the chapters. Virginia Valiela did much of the work on the index.
Sarah Allen provided technical help throughout the writing of this book,
and Jean Fruci was invaluable in helping put together the final manu
script. Lastly, I want especially to thank Virginia, Luisa, Cybele, and
Julia Valiela for putting up with me while I was writing this book and my
parents for providing a learning environment long ago.
Most of this text was written at the Boston University Marine Pro
gram, Marine Biological Laboratory, in Woods Hole. Arthur Humes and
Richard Whittaker, Directors of BUMP, were always helpful and pro
vided the time and academic environment in which to put this book to
gether. Dorothy Hahn, Mark Murray-Brown, and Dale Leavitt patiently
converted my endless sheets of illegible scribbles into neat piles of read
able word processor output. lowe thanks also to Jane Fessenden and her
staff, especially Lenora Joseph and Judy Ashmore, at the MBL Library
for ever-ready help. The drafting skill of Laurie Raymond is obvious in
the illustrations, and her sharp eye for errors was invaluable.
A necessary and stimulating stint of writing took place during a leave of
absence at the Department of Oceanography, University of Washington.
Karl Banse, Bruce Frost, Mike Landry, Amy Schoener, and the oceanog
raphy graduate students were hospitable and provided stimulation for my
writing.
Contents
Part I Primary Producers in Marine Environments
Chapter 1
Producers and Processes Involved in Primary Production 3
1.1 The Kinds and Amounts of Primary Producers in the Sea 3
1.2 Production: The Formation of Organic Matter 18
1.3 The Measurement of Producer Biomass and Primary
Production 22
1.4 Production Rates by Marine Primary Producers 32
Chapter 2
Factors Affecting Primary Production 38
2.1 Light 38
2.2 The Uptake and Availability of Nutrients 50
2.3 Temperature and Interactions with Other Factors 68
2.4 Distribution of Phytoplankton Production Over the
World Ocean 71
Part II Consumers in Marine Environments
Chapter 3
Dynamics of Populations of Consumers 79
3.1 Elements of the Mathematical Description of Growth
of Populations 79
viii Contents
3.2 Survival Life Tables 81
3.3 Fecundity Life Tables 84
3.4 Some Properties of Life Table Variables 86
3.5 Reproductive Tactics 90
Chapter 4
Competition for Resources Among Consumers 95
4.1 Population Growth in Environments with Finite Resources 95
4.2 The Nature of Competition 96
4.3 Density-Dependent Control of Abundance 104
4.4 Density-Dependent Versus Density-Independent Effects
on Abundance 106
4.5 Resource Partitioning 113
4.6 Niche Breadth and Species Packing 117
Chapter 5
Feeding and Responses to Food Abundance 119
5.1 Introduction 119
5.2 Functional Response to Prey Density 125
5.3 Numerical Responses by the Predator to Density of Prey 140
5.4 Developmental Response to Prey Density 144
Chapter 6
Food Selection by Consumers 147
6.1 Introduction 147
6.2 Behavioral Mechanisms Involved in Finding and
Choosing Food 147
6.3 Factors Affecting Food Selection by Consumers 148
6.4 Examples of Feeding Mechanisms at Work:
Suspension Feeding 159
6.5 Optimization in Food Selection by Consumers 164
6.6 Vulnerability and Accessibility of Food Items 168
6.7 The Importance of Alternate Prey 171
6.8 Predation and Stability of Prey Populations 174
Chapter 7
Processing of Consumed Energy 177
7.1 Flow of Energy Through Consumers 177
7.2 Assimilation 179
7.3 Respiration 186
Contents ix
7.4 Growth 192
7.5 Production 201
7.6 Energy Budgets for Populations 211
7.7 Energy Budgets for Ecosystems 214
Part III Interaction of Producers and Consumers: Competition
and Consumption
Chapter 8
The Consequences of Competition Among Producers
and Consumption by Herbivores 221
8.1 Models and Theory of Consumer-Food Interaction 221
8.2 Herbivores of Macroalgae and Marine Vascular
A~s ~
8.3 Grazers of Microalgae 236
8.4 Basic Impacts of Herbivores on Marine Producers 242
Chapter 9
The Consequences of Competition Among Prey and
Consumption by Predators 244
9.1 The Impact of Predators in Kelp Forests 244
9.2 Interaction of Predation and Competitive Exclusion in
the Rocky Intertidal Zone 248
9.3 Predation and Competition in Soft Sediment Benthic
Communities 250
9.4 Speculations as to Control of Prey Populations by
Predators in the Marine Water Column 254
Part IV Organic Matter, Decomposition, and Nutrient Cycles
Chapter 10
Transformations of Organic Matter: The Carbon Cycle 273
10.1 Inorganic Carbon 273
10.2 The Carbon Cycle in Aerobic Environments 275
10.3 The Carbon Cycle in Anoxic Environments 300
Chapter 11
Nutrient Cycles: Phosphorus, Nitrogen, and Sulfur 312
11.1 Phosphorus 312
11.2 Nitrogen 320
11.3 Sulfur 335