Ecological Studies Analysis and Synthesis edited by J. Jacobs Miinchen O. L. Lange Wiirzburg J. S. Olson Oak Ridge W. Wieser Innsbruck Volume 14 with contributions by Elgene Box Institut fiir Physikalische Chemie, KFA, Jiilich; F.R.G., and Curriculum in Ecology, University of North Carolina Chapel Hill, North Carolina 27514 John S. Bunt Australian Institute of Marine Science Townsville, Queensland, 4810 Australia Charles A. S. Hall Marine Biological Laboratory, Woods Hole, Massachusetts, and Section of Ecology and Systematics Cornell University, Ithaca, New York 14850 Helmut Lieth Department of Botany University of North Carolina Chapel Hill, North Carolina 27514 Gene E. Likens Section of Ecology and Systematics Langmuir Laboratory, Cornell University Ithaca, New York 14850 Peter L. Marks Section of Ecology and Systematics Langmuir Laboratory, Cornell University Ithaca, New York 14850 Russell Moll Great Lakes Research Division University of Michigan Ann Arbor, Michigan 48104 Peter G. Murphy Department of Botany and Plant Pathology Michigan State University East Lansing, Michigan 48824 Douglas D. Sharp Department of Botany University of North Carolina Chapel Hill, North Carolina 27514 David Sharpe Department of Geography Southern lllinois University at Carbondale Carbondale, lliinois 62901 Dennis Whigham Department of Biology, Rider College Trenton, New Jersey 08602 Robert H. Whittaker Section of Ecology and Systematics Langmuir Laboratory, Cornell University Ithaca, New York 14850 Primary Productivity of the Biosphere edited by Helmut Lieth and Robert H. Whittaker with 67 figures Springer-Verlag Berlin Heidelberg New York 1975 Library of Congress Cataloging in Publication Data Lieth, Helmut. Primary productivity of the biosphere. (Ecological studies; v. 14) Includes' index. 1. :primary productivity (Biology) I. Whittaker, Robert Harding, 1920- joint author. II. Title. III. Series. QH541.3.L5 574 74-26627 All rights reserved. No part of this book may be translated or reproduced in any form without written permission from Springer-Verlag. © 1975 by Springer-Verlag New York Inc. Softcover reprint of the hardcover 1st edition 1975 Distributed in the British Commonwealth Market by Chapman & Hall Limited, London. ISBN-13:978-3-642-80915-6 e-ISBN-13:978-3-642-80913-2 DOl: 10,1007/978-3-642-80913-2 Preface The period since World War II, and especially the last decade influenced by the International Biological Program, has seen enormous growth in research on the function of ecosystems. The same period has seen an exponential' rise in environmental problems including the capacity of the Earth to support man's population. The concern extends to man's effects on the "biosphere"-the film of living organisms on the Earth's surface that supports man. The common theme of ecologic research and environmental concerns is primary production the binding of sunlight energy into organic matter by plants that supports all life. Many results from the IBP remain to be synthesized, but enough data are available from that program and other research to develop a convincing sum mary of the primary production of the biosphere-the purpose of this book. The book had its origin in the parallel interests of the two editors and Gene E. Likens, which led them to prepare a symposium on the topic at the Second Biological Congress of the American Institute of Biological Sciences in Miami, Florida, October 24, 1971. Revisions of the papers presented at that symposium appear as Chapters 2, 8, 9, 10, and 15 in this book. We have added other chapters that complement this core; these include discussion and evaluation of methods for measuring productivity and regional production, current findings on tropical productivity, and models of primary productivity. The book is directed toward the interests of a range of readers, from'those seeking summaries of research techniques to those concerned with our synthesis of global production. Several institutions and people have helped to complete this work in its present form. The chapters contributed or coauthored by Lieth and Sharpe were supported in part by the Eastern Deciduous Forest Biome US-IBP. The chapters contributed by Whittaker and Hall were supported in part by Brook haven National Laboratory; the contributions by Likens and Whittaker were supported in part by the National Science Foundation. During the final stage of editing this volume, one of the editors (HL) worked as guest researcher at vi Preface the Nuclear Research Center (KFA) in Ji.ilich, West Germany. We gratetully acknowledge the financial and logistic help received at the KFA through Prof. Dr. K. Wagener and his staff at the Institut flir Physikalische Chemie. The index was compiled by Margot Lieth and Cyndi Grossman. We thank them both for their assistance. We gladly give credit to the staff of Springer-Verlag New York for excellent assistance in improving the book. We hope this book will be of value for its characterization of the biosphere as a productive system. We are not confident of man's ability to control the future of the world or even his own existence. Nevertheless, we should be gratified if a focal point of the book-the net primary production of the bio sphere-is one day seen as a figure of real significance to man. If in the future man's population and industry are stabilized, then to biosphere production as a steady-state flow of biological energy in the world will be related two other steady-state flows-of food energy from the biosphere to man and of industrial energy-that will support a human world society living in a durable balance with its environment. Helmut Lieth Robert H. Whittaker Contents Part 1 Introduction Preamble 2 1 Scope and Purpose of This Volume 3 Robert H. Whittaker, Gene E. Likens, and Helmut Lieth 2 Historical Survey of Primary Productivity Ftesearch 7 Helmut Lieth Part 2 Methods of Productivity Measurements 3 Methods of Assessing Aquatic Primary Productivity 19 Charles A. S. Hall and Russell Moll 4 Methods of Assessing Terrestrial Productivity 55 Robert H. Whittaker and Peter L. Marks 5 The Measurement of Caloric Values 119 Helmut Lieth 6 Assessment of Ftegional Productivity in North Carolina 131 Douglas D. Sharp, Helmut Lieth, and Dennis Whigham 7 Methods of Assessing the Primary Production of Regions 147 David M. Sharpe Part 3 Global Productivity Patterns 8 Primary Productivity of Marine Ecosystems 169 John S. Bunt 9 Primary Productivity of Inland Aquatic Ecosystems 185 Gene E. Likens 10 Primary Productivity of the Major Vegetation Units of the World 203 Helmut Lieth 11 Net Primary Productivity in Tropical Terrestrial Ecosystems 217 Peter G. Murphy Part 4 Utilizing the Knowledge of Primary Productivity 12 Modeling the Primary Productivity of the World 237 Helmut Lieth 13 Quantitative Evaluation of Global Primary Productivity Models Generated by Computers 265 Eigene Box 14 Some Prospects beyond Productivity Measurement 285 Helmut Lieth 15 The Biosphere and Man 305 Robert H. Whittaker and Gene E. Likens Index 329 Part 1 Introduction PREAMBLE The last decades of biologic, and especially ecologic, research have made it clear that 1. The notion that man's population and wealth can increase without limit is self-deception and an invitation to self-destruction 2. The unregulated increase of the human population beyond the world's sustainable carrying capacity must be considered a moral crime 3. The relentless increase in the gross natio:J~'J products of the industrial nations, at the expense of the world population, must be considered a social crime 4. The reckless exploitation of our fossil fuel sources for short-term profit and growth, rather than careful planning for a reasonable use for a long-term future, is a crime against our own children Helmut Lieth