Limnological Analyses

430 Pages·2000·8.86 MB·English

by  Robert G. Wetzel

Limnological Analyses
Third Edition
Springer Science+Business Media, LLC
Robert G. Wetze)
Gene E. Likens
Limnological Analyses
Third Edition
With 89 Illustrations
" Springer
Robert G. Wetzei, Bishop Professor of Biology, Department of Biology, University of Alabama,
College of Arts and Sciences, Tuscaloosa, AL 35487, USA
Gene E. Likens, Director, Institute of Ecosystem Studies, The New York Botanical Garden,
Cary Arboretum, Millbrook, NY 12545, USA
Cover illustration: Cover art supplied by Gene E. Likens.
Library of Congress Cataloging-in-Publication Data
Wetzei, Robert G.
Limnological analyses/Robert G. Wetzei, Gene E. Likens.-3rd ed.
p. cm.
Includes bibliographical references.
1. Limnology-Field work. 2. Limnology-Laboratory manuals.
I. Likens, Gene E. 11. Title.
QH96.57.A1W48 2000
577.6--dc21 99-042459
Printed on acid-free paper.
The first edition of this volume was published by W.B. Saunders Co. © 1979 W.B. Saunders Co.
© 2000, 1991 Springer Science+Business Media New York
Originally published by Springer-Verlag New York, Inc. in 2000.
Softcover reprint of the hardcover 3nd edition 2000
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9 8 7 6 5 432 1
ISBN 978-1-4419-3186-3 ISBN 978-1-4757-3250-4 (eBook)
DOI 10.1007/978-1-4757-3250-4
We acknowledge with appreciation the early
limnological training and inspiration provided by
David C. Chandler and Arthur D. Basler
Preface to the Third Edition
The continuing positive responses by students and instructors to the first and second
editions of Limnological Analyses over the past two decades are most gratifying.
The exercises in this book were set forth initially because of our frustrations in
teaching such a complex, multifaceted discipline as limnology. As with any ecolo
gical subject, understanding requires experiencing the techniques used for analysis
of properties of the ecosystems being studied and the responses of organisms within
them. Our intention is to provide aseries of coupled field and laboratory exercises
on the basic subjects of limnology. The exercises examine, as before, a broad range
of topics within both standing and running fresh waters. Methods evolve over time,
however, and here we present a revised, updated, and expanded version of the
second edition.
The previous style of presentation of each exercise worked effectively in our
teaching and was very weil received by others. Indeed, arecent book on methods
in analyses of streams imitated our style closely. Each exercise begins with abrief
introduction to the topic and its relevance within the science or to human activities.
Throughout, we attempt to present the details of the methods in relation to impor
tant questions and problem solving. Methods presented are generally of contem
porary research quality. Although the methods are presented for instruction and
problem solving, and not as a rigorous manual on limnological methods, many
sections could be and have been used for research purposes. As previously, some
methods presented are not the best available for the subject but were selected to
introduce the subjects to students within the constraints of time and facilities of the
classroom.
The importance of problem solving is reflected in the structure of the book and
its exercises. Initial exercises address physical components of lakes and streams and
major chemical constituents within them. These topics are then followed by quan
titative methods for analyzing biota and metabolic processes. The later exercises
often build on experience gained in the earlier on es. The previous provision of
several options for doing the exercise continues in expanded form he re in an
attempt to accommodate various logistic, time, and equipment constraints. Simul
taneously, these options offer suggestions for more ambitious students to extend
their inquiries beyond what was undertaken in regular class periods.
The questions set forth at the end of each exercise are important to the learning
experience. We have modified many of these, again based on continuing experience
in the classroom. We urge their use to enhance and hone rationale underlying the
methods, limitations of techniques, and interpretation of the information obtained.
viii Preface to the Third Edition
We do not have a solution to the perplexing problems of needing to identify
taxonomically the dominant aquatic flora and fauna. Students in initial courses in
limnology usually come with diverse backgrounds and experience in taxonomy. We
include only a few rudimentary pictorial keys to get the students with little experi
ence started. Thereafter it is essential for instructors to assist with general and
regional guides to identification.
In addition to the many people cited in the Preface to the Second Edition who
have offered guidance and constructive criticism of our previous editions, we would
particularly like to acknowledge the inputs of the following persons to the present
revision: Jean-Christian Auclair,Arthur C. Benke,Arthur S. Brooks, Paul R. Burten,
Jonathan J. Cole, Gary N. Ervin, Steven N. Francoeur, Stephen K. Hamilton, Richard
Hauer, Stuart H. Hurlbert, Mark D. Johnson, Klaus Jürgens, Winfried Lampert,
Mans LindeIl, Joseph C. Makarewicz, lan D. McKelvie, Michael L. Pace, Vincent
Resh, Richard D. Robarts, David L. Strayer, Keller Suberkropp, Karen Wiltshire,
and J. Wulfhorst. R. Wetzel acknowledges the stimulating atmosphere of the Max
Planck-Institut für Limnologie, Plön, Germany, that served as arefuge where much
of his writing was accomplished. Inputs from student users have been most helpful.
We appreciate greatly the assistance of all those mentioned and other persons who
have encouraged us to prevail in this endeavor. The final responsibilities for the
contents, however, rest with uso
Tuscaloosa, Alabama Robert G. Wetzel
Millbrook, New York Gene E. Likens
Preface to the Second Edition
The response by both instructors and students to the first edition of Limnological
Analyses, which appeared slightly over a decade ago, has been most gratifying. The
intent then was to provide aseries of interrelated field and laboratoty~xercises on
basic subjects of limnology. Those objectives have not changed. Here we present a
revised, updated, and expanded version.
The 29 exercises examine a broad range of topics concemed with both standing
and running fresh waters. The complex mechanisms controlling the metabolism and
dynamics of biotic populations and communities in aquatic ecosystems require an
understanding of physical and chemical variables. About one-third of the exercises
address the major physical components of lakes and streams, important mineral
nutrients, and organic matter. The remainder of the exercises provide rationale and
methods for quantitative analyses of the biota, as weH as some integrated analyses
of whole ecosystems. A few exercises address the effects of human activities on
aquatic ecosystems.
The later exercises usuaHy build on experience gained in the earlier ones.
Although most of the exercises can be performed largely in an intensive aftemoon,
sometimes additional time beyond the scheduled classroom hours may be needed.
Certain subjects, such as decomposition, do not lend themselves to simple, short
term analyses. We urge that exercises requiring more time not be avoided, because
the subjects are of major importance, and we have found them to be particularly
informative in our limnological classes. Often these longer exercises provide an
opportunity for more independent individual or smaH-team projects, with results
that may be reported to the entire class in effective leaming experiences.
Each exercise consists generaHy of abrief introduction to the topic with a state
ment of its limnological relevance or importance. We selected methods to illustrate
a variety of problems and how they might be solved. Although the methods pre
sen ted are nearly always of contemporary research quality, our intentions were not
to provide a rigorous manual of limnological methods, even though many sections
could be and have been used for research purposes. Certain techniques presented
in the book are not the best available, but they provide an introduction to the subject
within the constraints of time, facilities, and experience of the classroom. Where
better methods are available, we have attempted to provide references to these
techniques.
As previously, each exercise provides three or more options in an attempt to
accommodate various time, logistic, and equipment constraints or to aHow some stu-
x Preface to the Second Edition
dents to pursue a more ambitious course of study. We have been gratified that stu
dents often have pursued aggressively many of these suggested extensions of the
exercises. Wherever possible, we have attempted to combine and integrate field and
laboratory analyses. However, some options can be performed completely in the
classroom laboratory. Although we consider these options to be the least desirable,
some topics are more appropriately conducted in the laboratory at this level of
instruction. For example, the important and complex subject of hydrodynamics can
be introduced in a most illuminating and enjoyable manner by means of lake model
experiments.
We have expanded the questions at the end of each exercise. We encourage stu
dents to answer as many of them as possible, for they often genera te critical evalu
ations of the problems being addressed and the efficacy of the techniques being
used. The questions and references provided for each exercise can lead students to
explore further the myriad complexities of freshwater ecosystems.
The taxonomy of dominant aquatic flora and fauna is consistently a source of frus
tration both in the teaching of courses in limnology and in the preparation of this
book. The taxonomic backgrounds and experience of students ente ring an initial
course in limnology are usually diverse. Even superficial keys to common organisms
from bacteria to fish would exceed the length of the present text. In the first edition,
we included a few basic keys to orient students with little taxonomical experience
to major separations. These keys were not received weil by systematists and were
not used extensively by students. Therefore, we include only a few rudimentary
"starter" taxonomic keys in this edition, and we recommend that the instructors use
general and regional guides to the taxonomy of the flora and fauna of particular
regions. We cite example references of useful general taxonomic works.
We recognize that it would be difficult to accomplish all of the exercises in this
book in an intensive course in limnology during a single semester. Hopefully our
efforts, however, will assist in reducing the labors in such courses and enhance
insights into the operational integrity of aquatic ecosystems. We found the experi
ences of others important to the improvements to this book. We welcome further
suggestions on how the exercises might be improved or expanded.
Our ideas for effective instruction of certain topics had a long and diffuse devel
opmental history. As a result, we are no longer certain of their exact origins. Effec
tive teachers in our background certainly contributed at least indirectly to aspects
that appear in some exercises, particularly AD. Hasler, W.T. Edmondson, G.H.
Lauff, o.c. Chandler, and IR. Vallentyne. Gordon L. Godshalk read critically the
entire work and offered many helpful suggestions to both the first edition and parts
of the second edition. Constructive criticism and good counsel have been received
from numerous persons, including students as they grappled with the techniques in
classes. Discussions ab out and suggested revision to portions of the first and/or
second editions were provided by H.L. Allen, IR. Barnes, R. Bilby, 1 Cole, WG.
Crumpton, H.W Cunningham, F. deNoyelles, IA Dickerman, 1 Eaton, R. Edwards,
P. Godfrey, IB. Grace, C.AS. Hall, 0.1 Hall, R. Hall, G. Hendrey, M. Mattson, W
McDowell, 1 Meyer, S. Nodvin, D.K. Nordstrom, P.H. Rich, B. Riemann, o.W
Schindler, 1 Sioane, AStewart, 0. Strayer, R. Walter, AK. Ward, G.M. Ward, and
W. Youngs, Jr. Special insights were received on specific topics by several individu
als, particularly toward improvements in the second edition. Arthur C. Benke
offered great refinements and improvements to the treatments of benthos. Michael
F. Coveney, Clifford Ochs, and Michael Pace counseled us on our new treatment of
bacterial productivity. The new exercise on predator-prey analyses was guided by