Limnological Analyses. Second Edition

Transcription

1 Limnological Analyses Second Edition

2 Robert G. Wetzel Gene E. li kens limnological Analyses Second Edition Springer Science+Business Media, LLC

3 Robert G. Wetzel, 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 art supplied by R.G. Wetzel. Printed on acid-free paper Library of Congress Cataloging-in-Publication Data Wetzel, Robert A. Limnological analyses/robert G. Wetzel, Gene E. Likens.-2nd ed. p. cm. Includes bibliographical references. Includes index. ISBN ISBN (ebook) DOI / Limnology-Field work. 2. Limnology-Laboratory manuals. I. Likens, Gene E., II. Title. QH96.57.AIW '2632-dc The first edition of this volume was published by W.B. Saunders Co W.B. Saunders Co Springer Science+Business Media New York Originally published by Springer-Verlag New York, Inc. in 1991 Softcover reprint of the hardcover 2nd edition 1991 All rights reserved. This work may not be translated or copied in whole or in part without the written permission of the publisher Springer Science+Business Media, LLC, except for brief excerpts in connection with reviews or scholarly analysis. Use in connection with any form of information and retrieval, electronic adaptation, computer software, or by similar or dissimilar methodology now known or hereafter developed is forbidden. 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 accoi;dingly be used freely by anyone. Typeset by Thomson Press (India) Ltd., New Delhi ISBN

4 We acknowledge with appreciation the early limnological training and inspiration provided by David C. Chandler and Arthur D. Hasler

5 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 a series of interrelated field and laboratory exercises 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 concerned 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 well as some integrated analyses of whole ecosystems. A few exercises address the effects of human activities on aquatic ecosystems. The later exercises usually build on experience gained in the earlier ones. Although most of the exercises can be performed largely in an intensive afternoon, 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 small team projects, with results that may be reported to the entire class in effective learning experiences. Each exercise consists generally of a brief introduction to the topic with a statement of its limnological relevance or importance. We selected methods to illustrate a variety of problems and how they might be solved. Although the methods presented 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 allow some students to pursue a more ambitious course of study. We have been gratified that students often have pursued aggressively many of these suggested extensions of the exercises. Wherever possible, we have attempted to combine and integrate field and

6 viii Preface to the Second Edition 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 students to answer as many of them as possible, for they often generate critical evaluations 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 frustration both in the teaching of courses in limnology and in the preparation of this book. The taxonomic backgrounds and experience of students entering 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 well 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 experiences 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 developmental history. As a result, we are no longer certain of their exact origins. Effective teachers in our background certainly contributed at least indirectly to aspects that appear in some exercises, particularly A.D. Hasler, W.T. Edmondson, G.H. Lauff, D.C. Chandler, and J.R. 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 about and suggested revisions to portions of the first and/or second editions were provided by H.L. Allen, J.R. Barnes, R. Bilby, J. Cole, W.G. Crumpton, H.W. Cunningham, F. denoyelles, J.A. Dickerman, J. Eaton, R. Edwards, P. Godfrey, J.B. Grace, C.A.S. Hall, D.J. Hall, R. Hall, G. Hendrey, M. Mattson, W. McDowell, J. Meyer, S. Nodvin, D.K. Nordstrom, P.H. Rich, B. Riemann, D.W. Schindler, J. Sloane, A. Stewart, D. Strayer, R. Walter, A.K. Ward, G.M. Ward, and W. Youngs, Jr. Special insights were received on specific topics by several individuals, 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 the efforts of Kenneth Wagner and W. Charles Kerfoot. Mark D. Mattson contributed significantly to the revision of the exercise on the inorganic carbon complex. The exercises on manipulation of model ecosystems, diurnal changes in stream ecosystems, and effect of sewage outfall

7 Preface to the Second Edition ix on stream ecosystems contain input from F.1. denoyelles, P. Godfrey, C.A.S. Hall, J. Sloane-Richey, and Raymond Barrett. William D. Taylor helped with facets of phytoplankton biomass analyses, Kathleen Weathers with inorganic nutrients, Michael Pace with zooplankton feeding, Nina Caraco with whole ecosystem analyses, Jonathon Cole with decomposition, and Robert E. Moeller with paleolimnological analyses. Anita J. Johnson prepared many of the final figures. Phyllis Likens provided invaluable assistance with proofreading. We gratefully acknowledge the assistance of these and other persons that have encouraged us with this effort. The final responsibilities for the contents, however, rest with us. Robert G. Wetzel Gene E. Likens

8 Contents Preface to the Second Edition vii EXERCISE 1 Lake Basin Characteristics and Morphometry EXERCISE 2 Light and Temperature 15 EXERCISE 3 Physical Characteristics: Lake Models 31 EXERCISE 4 The Heat Budget of Lakes 43 EXERCISE 5 Morphology and Flow in Streams 55 EXERCISE 6 Dissolved Oxygen 69 EXERCISE 7 Inorganic Nutrients: Nitrogen, Phosphorus, and Other Nutrients 81 EXERCISE 8 The Inorganic Carbon Complex: Alkalinity, Acidity, CO 2, ph, Total Inorganic Carbon, Hardness EXERCISE 9 Organic Matter 129 EXERCISE 10 Composition and Biomass of Phytoplankton 139

9 xii Contents EXERCISE 11 Collection, Enumeration, and Biomass of Zooplankton 167 EXERCISE 12 Benthic Fauna of Lakes 179 EXERCISE 13 Benthic Fauna of Streams 199 EXERCISE 14 Primary Productivity of Phytoplankton 207 EXERCISE 15 Zooplankton Feeding 227 EXERCISE 16 Zooplankton Production 235 EXERCISE 17 Predator-Prey Interactions 241 EXERCISE 18 Enumeration of Fish or Other Aquatic Animals 247 EXERCISE 19 Bacterial Growth and Productivity 255 EXERCISE 20 Decomposition: Relative Bacterial Heterotrophic Activity on Soluble Organic Matter 271 EXERCISE 21 Decomposition: Particulate Organic Matter 281 EXERCISE 22 The Littoral Zone 291 EXERCISE 23 Experimental Manipulation of Model Ecosystems 301 EXERCISE 24 Diurnal Changes in a Stream Ecosystem: An Energy and Nutrient Budget Approach 315

10 Contents xiii EXERCISE 25 Diurnal Changes in Lake Systems 325 EXERCISE 26 Special Lake Types EXERCISE 27 Historical Records of Changes in the Productivity of Lakes 337 EXERCISE 28 Effect of Sewage Outfall on a Stream Ecosystem 345 EXERCISE 29 Estimates of Whole Lake Metabolism: Hypolimnetic Oxygen Deficits and Carbon Dioxide Accumulation 349 ApPENDIX 1 General Chemical Relationships 359 ApPENDIX 2 Basic Definitions Used in Community Analyses 365 ApPENDIX 3 Useful Relationships Relative to the Use of Colorimeters and Spectrophotometers 371 ApPENDIX 4 Characteristics and Taxonomic Sources of Common Freshwater Organisms 373 Index 387