Alga,1E cology Freshwater Benthic Ecosystems AQUATIC ECOLOGY Series Series Editor James H. Thorp Department of Biology University of Louisville Louisville, Kentucky Other titles in the series: Groundwater Ecology Janine Gibert, Dan L. Danielopol, Jack A. Stanford Fresh water Ben thic Ecosystems Edited by R. Jan Stevenson Department of Biology University of Louisville Louisville, Kentucky Max L. Both well Environmental Sciences National Hydrological Research Institute Saskatoon, Saskatchewan, Canada Rex L. Lowe Department of Biological Sciences Bowling Green State University Bowling Green, Ohio ACADEMIC PRESS San Diego New York Boston London Sydney Tokyo Toronto Front cover photograph: The filamentous diatom Aulacoseira entangled among the stalked diatoms Cymbella and Gomphonema. (See Chapter 1, Figure 1 for more details.) This book is printed on acid-flee paper. (~ Copyright 1996, Elsevier (USA) All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any means, electronic or mechanical, including photocopy, recording, or any information storage and retrieval system, without permission in writing from the publisher. Requests for permission to make copies of any part of the work should be mailed to: Permissions Department, Harcourt, Inc., 6277 Sea Harbor Drive, Orlando, Florida 32887-6777. Academic Press An imprint of Elsevier 525 B Street, Suite 1900, San Diego, California 92101-4495, USA http://www.academicpress.com Academic Press 84 Theobald's Road, London WC1X 8RR, UK http ://www. academicpress, com Library of Congress Cataloging-in-Publication Data Algal ecology: freshwater benthic ecosystems/edited by R. Jan Stevenson, Max I. Bothwell, Rex L. Lowe. p. cm.-(Aquatic ecology series) Includes bibliographical references and index. ISBN 0-12-668450-2 (alk. paper) 1. Freshwater algae--Ecology. 2. Freshwater ecology. I. Stevenson, R. Jan. II. Bothwell, M.L. (Max L.) III. Lowe, Rex IV. Series. QK570.25.A43 1996 589.3'5---dc20 95-51737 CIP PRINTED IN THE UNITED STATES OF AMERICA 03 04 05 06 9 8 7 6 5 4 Jan and Rex graciously asked me to dedicate this book to my wife Carol R. Bothwell (deceased November 5, 1994) in remembrance of the selfless dedication she gave to me over the years so that my career might prosper. In this, she was not unlike many other wives, including Mariellyn Stevenson and Sheryn Lowe, who spend their lives nurturing their husbands and children so that they might grow and fulfill their own dreams. Max L. Bothwell This Page Intentionally Left Blank Contents Contributors xxi Preface xxiii Acknowledgments XXV SECTION ONE PATTERNS OF BENTHIC ALGAE IN AQUATIC ECOSYSTEMS 1 An Introduction to Algal Ecology in Freshwater Benthic Habitats R. Jan Stevenson vii viii Contents I. The Diversity of Benthic Algae and Their Habitats in Fresh Waters 3 A. The Taxonomic and Morphological Diversity of Benthic Algae 3 B. The Habitats of Freshwater Benthic Algae 8 II. The Niche of Freshwater Benthic Algae 10 A. The Role of Benthic Algae in Ecosystems 10 B. How Are Algae in Benthos and Plankton Different? 11 III. Methods for Characterizing Freshwater Benthic Algae 12 A. Population and Community Structure 12 B. Population and Community Function 18 W. Conceptual Frameworks for Benthic Algal Community Ecology 23 References 26 2 Patterns in Benthic Algae of Streams Barry J. E Biggs I. Introduction 31 A. Conceptual Overview: Disturbance-Resource Supply-Grazer Control of Stream Benthic Algae 32 B. Typical Biomass Values for Streams 34 II. Temporal Patterns 36 A. Short Term 36 B. LongTerm 38 III. Spatial Patterns 42 A. Microscale: Substratum Patterns 42 B. Mesoscale- Within Catchment Patterns 43 C. Broadscale: Intercatchment Patterns 44 IV. Benthic Algal Proliferations 46 V. Concluding Remarks 51 References 51 3 Periphyton Patterns in Lakes Rex L. Lowe I. Introduction 57 II. The Lentic Periphyton Community 58 A. Habitat 58 B. TheBiota 59 Contents ix III. Important Influential Factors 63 A. Overview 63 13. Resources 64 C. Disturbance 68 IV. Conclusions and Missing Pieces 71 References 72 4 Pattern in Wetlands L. Gordon Goldsborough and Gordon G. C. Robinson I. Introduction 78 II. Algal Assemblages in Wetlands 78 III. The Role of Algae in Wetlands 81 IV. Species Composition of Wetland Algae 87 V. Algal Production in Wetlands 89 VI. Factors Affecting Algal Production 90 A. Hydrodynamics 90 B. Nutrients 91 C. Light 94 D. Temperature 96 E. Macrophytes 97 E Herbivory and Other Faunal Influences 99 G. Anthropogenic Factors 101 VII. Conceptual Model of Wetland Algae 103 A. Dry State 103 B. Open State 105 C. Sheltered State 107 D. Lake State 108 VIII. Conclusion 109 References 109 SECTION TWO FACTORS AFFECTING BENTHIC ALGAE 5 Effects of Light Walter R. Hill I. Introduction 121 II. Photosynthetic Processes 122