Forests: Fresh Perspectives from Ecosystem Analysis (Proceedingspf the 40th Annual Biology Colloquium I Edited by Richard H. Waring Oregon State University Press Corvallis, Oregon Library of Congress Cataloging in Publication Data Biology Colloquium, 40th, Oregon State University, 1979. Forests, fresh perspectives from ecosystem analysis. 1. Forest ecologyCongresses. I. \Varing, Richard H.II.Title. QH541 .5.F6B561979 574.5'2642 80-11883 iSBN 0-87071-179-2 © 1980 1w the Oregon State University Press Printed in the United States of America All rights reserved Preface Over the last decade a new perspective sections within this volume. The first on how forest ecosystems operate has section (chapters 1-3) revises classical emerged. Ecosystems appear much more flex- theories on community structure, succession, ible than we once thought. Even the most and ecosystems. The second section (chap- persistent is still evolving in composition. ters 4-8) examines in detail how forest Yet for all their diversity, very similar canopies, soil microbes, and root systems processes are seen as operating in all operate as almost independent subsystems. forests, providing a point for comparative The final section (chapters 9-12) focuses studies. A more balanced time perspective on the impact of the different materials-- brings a greater appreciation of the in- logs and leaves, soil and sediments, water frequent but dominating events that shape and minerals--that move through forests and the course of ecosystem development. into stream ecosystems. Although each Through joint studies of forests and streams section is distinct in scope, they share a we see new roles for living and dead com- common link of matter and energy flow. ponents of both ecosystems. For permitting us the opportunity to For the 40th annual Biology Colloquim share these ideas, we are indebted to the held April 27-28 1979 at Oregon State late Ralph Shay, who originally suggested University, a select group of scientists the topic and helped arrange the colloquim. was invited to share this new perspective To him we dedicate this volume. on forest ecosystems with a wider audience. The colloquium papers are grouped into three Richard H. Waring V - Contributing Authors Daniel B. Botkin R. V. O'Neill Department of Biological Studies Environmental Sciences Division University of California Oak Ridge National Laboratory Santa Barbara, California Oak Ridge, Tennessee George C. Carroll Dennis Parkinson Departnent of Biology Department of Biology University of Oregon University of Calgary Eugene, Oregon Calgary, Alberta Kermit Cromack, Jr. D. E. Reichle Department of Forest Science Environmental Sciences Laboratory Oregon State University Oak Ridge National Laboratory Corvallis, Oregon Oak Ridge, Tennessee Kenneth W. Cummins Dan Santantonio Department of Fisheries and Wildlife School of Forestry Oregon State University Oregon State University Corvallis, Oregon Corvallis, Oregon Jerry F. Franklin Wayne T. Swank U. S. Forest Service Coweeta Hydrologic Laboratory Forest Science Laboratory U. S. Forest Service Corvallis, Oregon Franklin, North Carolina W. F. Harris Fredrick J. Swanson Environmental Sciences Division Pacific Northwest Forest and Range Oak Ridge National Laboratory Experiment Station Oak Ridge, Tennessee Forestry Sciences Laboratory Corvallis, Oregon James A. MacMahon Department of Biology and Ecology Center Frank J. Triska Utah State University U.S. Geological Survey Logan, Utah Menlo Park, California D. McGinty Jack B. Waide Department of Biology Environmental Sciences Division Huntingdon College Oak Ridge National Laboratory Montgomery, Alabama Oak Ridge, Tennessee Richard H. Waring Department of Forest Science Oregon State University Corvallis, Oregon vii Contents Section I: Ecosystem Theory A Grandfather Clock down the Staircase: Stability and Disturbance in Natural Bcosystems ....................................................................... Daniel B. Botkin Dimensions of Ecosystem Theory ........................................................... 11 R. V. O'Neill and D. E. Reichie Ecosystems over Time: Succession and Other Types of Change .............................. 27 James A. MacMahon Section II: Terrestrial Ecosystems Distinctive Features of the Northwestern Coniferous Forest: Development, Structure, and Function .................................................................. 59 Jerry F. Franklin and Richard H. Waring Forest Canopies: Complex and Independent Subsystems ..................................... 87 George C. Carroll Aspects of the Microbial Ecology of Forest Ecosystems .................................... 109 Dennis Parkinson The Dynamic Belowground Ecosystem ........................................................ 119 W. F. Harris, Dan Santantonio, and D. Mccinty Vital Signs of Forest Ecosystems ......................................................... 131 R. H. Waring Section III: Watershed and Stream Ecosystems Interpretation of Nutrient Cycling Research in a Management Context: Evaluating Potential Effects of Alternative Management Strategies on SiteProductivity ........................................................................ 137 Wayne T. Swank and Jack B. Waide Geomorphology and Ecosystems ............................................................. 159 Frederick J. Swanson The Role of Wood Debris in Forests and Streams ........................................... 171 Frank J. Triska and Kermit Cromack, Jr. The Multiple Linkages of Forests to Streams .............................................. 191 Kenneth W. Cummins ix A Grandfather Clock down the Staircase: Stability and Disturbance in Natural Ecosystems Daniel B. Botkin BASIC CONCEPTS book, Man and Nature, that "nature, left un- disturbed so fashions her territory as to In a small village in New Hampshire, an give it almost unchanging permanence of elderly lady lives in an old red brick house form, outline, and proportion, except when crammed with antiques that her parents shattered, by geologic convulsions; and in bought at the turn of the century. At the these comparatively rare cases of derange- top of the staircase stands a wonderful ment, she sets herself at once to repair the grandfather clock, whose pendulum had swung superficial damage, to restore as nearly as rhythmically to and fro for many year practicable, the former aspect of her Several years ago two grandchildren dis- dominion." We tend also to agree with covered the clock and, in the process of in- Marsh that an undisturbed wilderness forest, vestigating it with all the empirical skills the kind that he referred to as occurring in known to small children, managed to edge it "new countries," meaning those not yet sub- over to the staircase. With a push, they ject to civilization's heavy hand, is watched it tumble down the stairs with a characterized by a single, permanent equili- marvelous series of noises.With consider- brium condition the climax forest of able effort, friends and neighbos eventually twentieth century ecology. In Wildernesses, returned the clock to its former position, Marsh wrote, "the natural inclination of the but the disruption had been too great for its ground, the self-formed slopes and levels, internal mechanisms and it has stood silent are graded and lowered or elevated by frost since, outwardly intact, a sad image of and chemical forces and gravitation and the mechanical frailty. flow of water and vegetable deposit and the Nature's biota sometimes seem like that action of the winds until, by a general clock -- able to withstand small disturbances compensation of conflicting forces, a condi- but unable to survive major perturbations. tion of equilibrium has been reached which, In ecological systems, perturbation and without the action of man, would remain, stability seem to oppose one another. In with little fluctuations, for countless both scientific literature and in popular ages" (Marsh, 1864). discussions of the effects of civilization On the other hand, we also seem to on the environment, stability is referred agree with Lucretius, who in the first to as resisting disturbance and perturbation century B.C. wrote in De Rerum Natura that as a disruption of the stability of nature. all things are subject to mutability and We call an undisturbed forest stable and a change; that "time does change the nature of highly perturbed one unstable. the whole wide world; one state follows from Are ecosystems, in these terms, stable another; not one thing is like itself for-- or unstable? Is perturbation common and, ever, all things move, all things are in some sense, natural, necessary, or nature's wanderers, whom she gives no rest, desirable? Or is perturbation always the ebb follows flow." Lucretius used the ero- product of human interference in nature, and sion along riverbanks in forests as an bad something to be avoided in our example of the mutability of nature. Here management of natural ecosystems? the forests, Lucretius said so long ago, In regard to these questions, we seem are "shorn, gnawed by the current" to believe simultaneously in contradictory (Hunphries, 1968). views. On the one hand, we tend to agree We acknowledge, as did Lucretius, that with George Perkins Marsh, who wrote more nature is subject to change, that forests than one hundred years ago in his classic change: trees have mast years when, for H 2 FORESTS: FRESH PERSPECTIVES FROM ECOSYSTEM ANALYSIS reasons we do not now understand, a single (idealized) pendulum, which will oscillate species will produce an abundance