Produced by The Estuarine Research Federation Sponsored by U.S. Environmental Protection Agency U.S. Fish and Wildlife Service U.S. Geological Survey National Oceanic and Atmospheric Administration Marine Ecosystems Analysis Program National Marine Fisheries Service Proceedings of the Fifth Biennial International Estuarine Research Conference, Jekyll Island, Georgia, October 7-12,1979 ESTUARINE PERSPECTIVES Edited by VICTOR S KENNEDY University of Maryland Center for Environmental and Estuarine Studies Horn Point Environmental Laboratories Cambridge, Maryland ACADEMIC PRESS 1980 A Subsidiary of Harcourt Brace Jouanovich, Publishers lew York London Sydney Toronto San Francisco COPYRIGHT © 1980, BY ACADEMIC PRESS, INC. 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. ACADEMIC PRESS, INC. Ill Fifth Avenue, New York, New York 10003 United Kingdom Edition published by ACADEMIC PRESS, INC. (LONDON) LTD. 24/28 Oval Road, London NW1 7DX Library of Congress Cataloging in Publication Data International Estuarine Research Conference, 5th, Jekyll Island, Ga., 1979. Estuarine perspectives. Papers presented at a conference held by the Estua- rine Research Federation and sponsored by U.S. Environ- mental Protection Agency and others. Includes bibliographies and index. 1. Estuarine ecology—Congresses. 2. Estuaries— Congresses. I. Kennedy, Victor S. II. Estuarine Research Federation. III. United States. Environmen- tal Protection Agency. IV. Title. QH541.5.E8I56 1979 574.526365 80-23291 ISBN 0-12-404060-8 PRINTED IN THE UNITED STATES OF AMERICA 80 81 82 83 9 8 7 6 5 4 3 2 1 LIST OF CONTRIBUTORS Ackleson, S. G., College of Marine Studies, University of Delaware, Lewes, DE 19958 Amezcua Linares, F., Centro de Ciencias del Mar y Limnologia, Univer- sidad Nacional Autonoma de Mexico, Apartado Postal 70-305, Mex- ico 20, D.F. Bartlett, D. S., College of Marine Studies, University of Delaware, Newark, DE 19711 Bâtie, S. S., Department of Agricultural Economics, Virginia Polytechnic In- stitute and State University, Blacksburg, VA 24061 Beal, K. L·, Fisheries Management Operations Branch, National Marine Fisheries Service, State Fish Pier, Gloucester, MA 01930 Black, L. F., Zoology Department, Jackson Estuarine Laboratory, University of New Hampshire, Durham, NH 03824 Boardman, D., Marine Biology, Lamont-Doherty Geological Observatory, Palisades, NY 10964 Bobbie, R. J., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Botkin, D. B., Environmental Studies, University of California, Santa Bar- bara, CA 93106 Boynton, W. R., Chesapeake Biological Laboratory, Center for En- vironmental and Estuarine Studies, University of Maryland, Box 38, Solomons, MD 20688 Bozzo, W. E. Department of Biology, The American University, t Washington, DC 20016 Budgell, W. P., Ocean and Aquatic Sciences, Department of Fisheries and Oceans, Canada Center for Inland Waters, 867 Lakeshore Road, Burlington, Ontario, Canada L7R 4A6 Capone, D. G., Marine Science Research Center, State University of New York, Stony Brook, NY 11794 X! xii List of Contributors Champ, M. A, Department of Biology, The American University, Washington, D.C. 20016 Dame, R., Coastal Carolina College and Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina, Conway, SC 29526 Davis, W. M., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Day, J. W., Jr., Coastal Ecology Laboratory, Center for Wetland Resources, Louisiana State University, Baton Rouge, LA 70803 de la Cruz, A. A., Department of Zoology, Mississippi State University, Mississippi State, MS 39762 DeLaune, R. D., Laboratory for Wetland Soils and Sediments, Center for Wetland Resources, Louisiana State University, Baton Rouge, LA 70803 Dunn, M. L·, Department of Marine Science and Engineering, North Carolina State University, Raleigh, NC 27650 Evans, M. S., Great Lakes Research Division, University of Michigan, Ann Arbor, MI 48109 Fazio, S. D., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Findlay, R. H., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Galloway, G. E., Jr., Department of Earth, Space and Graphic Sciences, U.S. Military Academy, West Point, NY 10996 Gilbert, P. M., The Biological Laboratories, Harvard University, Cambridge, MA 02138 Gould, G. A., Ill, Department of Biology, The American University, Washington, D.C. 20016 Grainger, E. H., Arctic Biological Station, Ste. Anne de Bellevue, Quebec, Canada, H9X 3L6 Grève, W., Biologische Anstalt Helgoland, Meerestation, Haus A, 2192 Helgoland, Federal Republic of Germany Hackney, C. T., Department of Biology, University of Southwestern Loui- siana, Lafayette, LA 70524 Hansen, W. J., Environmental Laboratory, U.S. Army Engineer Waterways Experiment Station, P.O. Box 631, Vicksburg, MS 39180 Hardisky, M. A., College of Marine Studies, 103A Robinson Hall, Universi- ty of Delaware, Newark, DE 19711 Healy, W. B., Minister (Scientific), New Zealand High Commission, New Zealand House, Haymarket, London, England SW14 4TQ Heck, K. J., Jr., The Academy of Natural Sciences of Philadelphia, Benedict Estuarine Research Laboratory, Benedict, MD 20612 List of Contributors xiii Jeffries, H. P., Graduate School of Oceanography, University of Rhode Island, Kingston, RI02881 Katsinis, C, Department of Electrical Engineering, University of Rhode Island, Kingston, RI 02881 Kelley, J. T., Department of Earth Sciences, University of New Orleans, New Orleans, LA 70122 Kemp, W. M., Horn Point Environmental Laboratories, Center for En- vironmental and Estuarine Studies, University of Maryland, Box 775, Cambridge, MD 21613 Kjerfve, B., Belle W. Baruch Institute for Marine Biology and Coastal Research, Department of Geology, and Marine Science Program, University of South Carolina, Columbia, SC 29208 Klemas, V., College of Marine Studies, University of Delaware, Newark, DE 19711 Klose, P. N., Environmental Resources Management, Inc., 999 West Chester Pike, West Chester, PA 19380 Kumpf H. £., National Marine Fisheries Service, Southeast Fisheries Center, Office of Fishery Management, 75 Virginia Beach Drive, Miami, FL 33149 Leuinton, J. S., Department of Ecology and Evolution, State University of New York, Stony Brook, NY 11794 Linthurst, R. A, Department of Botany, North Carolina State University, P.O. Box 5186, Raleigh, NC 27650 Livingston, R. J., Department of Biological Science, Florida State Universi- ty, Tallahassee, FL 32306 Loder, T. C, Department of Earth Sciences and Ocean Process Analysis Laboratory, University of New Hampshire, Durham, NH 03824 Maguire, B., Jr., Department of Biology, University of Texas, Austin, TX 78712 Mahne, T. G, Marine Biology, Lamont-Doherty Geological Observatory, Palisades, NY 10964 Martz, R. F., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Maurer, R., National Marine Fisheries Service, Northeast Fisheries Center, Narragansett, RI 02882 McKellar, H. N., Jr., Belle W. Baruch Institute for Marine Biology and Coastal Research, and Department of Environmental Health Sciences, University of South Carolina, Columbia, SC 29208 Miller, J. M., Department of Zoology, North Carolina State University, Raleigh, NC 27650 Moore, B., Ill Complex Systems Group, O'Kane House, University of New Hampshire, Durham, NH 03824 xw List of Contributors Morowitz, H., Department of Molecular Biophysics and Biochemistry, Yale University, New Haven, CT 06520 Neale, P. J., Marine Biology, Lamont-Doherty Geological Observatory, Palisades, NY 10964 Nelson, D., Coastal Carolina College and Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina, Conway, SC 29526 Nichols, M. M, Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, VA 23062 Nickels, J. S., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Odum, E. P., Institute of Ecology, University of Georgia, Athens, GA 30601 Officer, C. B., Earth Sciences Department, Dartmouth College, Hanover, NH 03755 Orth, R. J., Virginia Institute of Marine Science, and School of Marine Science, College of William and Mary, Gloucester Point, VA 20612 Osborne, C. G., Chesapeake Biological Laboratory, Center for En- vironmental and Estuarine Studies, University of Maryland, Box 38, Solomons, MD 20688 Patrick, W. H., Jr., Laboratory for Wetland Soils and Sediments, Center for Wetland Resources, Louisiana State University, Baton Rouge, LA 70803 Peterson, C. H., Institute of Marine Sciences, University of North Carolina at Chapel Hill, P.O. Drawer 809, Morehead City, NC 28557 Pett, R. J., Department of Zoology, College of Biological Science, Universi- ty of Guelph, Guelph, Ontario, Canada, NIG 2W1 Phillips, J. H., Georgia Department of Natural Resources, Coastal Resources Division, 1200 Glynn Avenue, Brunswick, GA 31520 Philpot, W. D., College of Marine Studies, University of Delaware, Newark, DE 19711 Reed, J. P., The Ecosystems Center, Marine Biological Laboratory, Woods Hole, MA 02543 Reimold, R. J., Georgia Department of Natural Resources, Coastal Resources Division, 1200 Glynn Avenue, Brunswick, GA 31520 Reiners, F., Biologische Anstalt Helgoland, Meerestation, Haus A, 2192 Helgoland, Federal Republic of Germany Reppert, R. T., U.S. Army Engineer Institute for Water Resources, Kingman Building, Fort Belvoir, VA 22060 Richardson, S. E., Environmental Laboratory, U.S. Army Engineer Water- ways Experiment Station, P.O. Box 631, Vicksburg, MS 39180 List of Contributors χυ Roff, J. G, Department of Zoology, College of Biological Science, Universi- ty of Guelph, Guelph, Ontario, Canada, NIG 2W1 Rogers, G. F., Department of Zoology, College of Biological Science, University of Guelph, Guelph, Ontario, Canada, NIG 2W1 Schamberger, M. L., U.S. Fish and Wildlife Service, Office of Biological Services, Western Energy and Land Use Team, 2625 Redwing Road, Fort Collins, CO 80525 Shabman, L. A., Department of Agricultural Economics, Virginia Polytechnic Institute and State University, Blacksburg, VA 24061 Sherman, K, National Marine Fisheries Service, Northeast Fisheries Center, Narragansett, RI02882 Slobodkin, L. B., Department of Ecology and Evolution, State University of New York, Stony Brook, NY 11794 Smith, D. W., School of Life and Health Sciences, University of Delaware, Newark, DE 19711 Smith, G. A., Department of Biological Science, Florida State University, Tallahassee, FL 32306 Smith, N. P., Harbor Branch Foundation, Inc., RR 1, Box 196, Fort Pierce, FL 33450 Stevenson, H., Biology Department, and Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina, Columbia, SC 29208 Stout, J. P., Dauphin Island Sea Lab, University of South Alabama, P.O. Box 386, Dauphin Island, AL 36528 Taylor, B. F., Rosenstiel School of Marine and Atmospheric Science, University of Miami, Miami, FL 33149 Tippie, V. K, Center for Ocean Management Studies, University of Rhode Island, Kingston, RI 02881 Viena, K. C, College of Marine Studies, University of Delaware, Lewes, DE 19958 Virnstein, R. W., Harbor Branch Foundation, Inc., RR 1, Box 196, Fort Pierce, FL 33450 Webb, K. L·., Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, VA 23062 Wetzel, R. L., Virginia Institute of Marine Science, College of William and Mary, Gloucester Point, VA 23062 White, D. C, Department of Biological Science, Florida State University, Tallahassee, FL 32306 Wolaver, T. G., Department of Environmental Sciences, University of Virginia, Charlottesville, VA 22903 χυ'ι List of Contributors Yanez-Arancibia, A., Centro de Ciencias del Mar y Limnologia, Universidad Nacional Autonoma de Mexico, Apartado Postal 70-305, Mexico 20, D.F. Zieman, J. G, Department of Environmental Sciences, University of Virginia, Charlottesville, VA 22903 Zingmark, R., Biology Department, and Belle W. Baruch Institute for Marine Biology and Coastal Research, University of South Carolina, Columbia, SC 29208 FOREWORD Estuaries have a special and historic importance to man. The dense population centers that occur on virtually every major estuary attest to this fact. They are used as fishing grounds, transportation arteries, a source of industrial cooling waters, for waste disposal, and as recreational areas. The continued and increasing development of these areas has presented many problems. Dr. F. John Vernberg, past president of the Estuarine Research Federation, has stated "Estuaries and adjacent environments are ecological systems that are subjected to continual stress by natural and man-induced perturbations " The truth of this statement is obvious. In order to ameliorate or solve these problems, it is necessary to understand the complex systems which interact within an estuary. To that end a group of scientists, students, engineers, resource managers and other specialists met at Jekyll Island, Georgia, in 1964 to hold a symposium on estuaries. The objectives were to provide an opportunity for the exchange of ideas between the various disciplines and individuals interested in estuarine research, to summarize the present knowledge of the char- acteristics of estuaries, and to delineate the direction of current research. Estuaries were defined, described, categorized, systematized, and discussed throughout this historic four-day meeting. About 1000 individuals par- ticipated in this first major estuarine symposium in North America. Since that meeting, the Estuarine Research Federation has arranged four more symposia on estuaries. In 1973 a symposium considering "Re- cent Advances in Estuarine Research" was held in Myrtle Beach, South Carolina. The next symposium examined "Estuarine Processes" and was held in Galveston, Texas in 1975, followed by "Estuarine Interactions" held at Mt. Pocono, Pennsylvania in 1977. In 1979 the Fifth Biennial Interna- tional Estuarine Research Conference was held October 7-12 at Jekyll Island, Georgia. Perhaps our returning to Jekyll Island has completed some sort of cycle and indicates a greater perspective on all our parts about the importance of estuaries. Appropriately, the title of this conference was "Estuarine Perspectives. " xvii