Disturbance and Ecosystems Components of Response

Edited by H. A. Mooney and M. Godron

Contributors D. Auclair F. A. Bazzaz F. S. Chapin III R. T. T. Forman

M. Godron P.-H. Gouyon S. L. Gulmon G. Heim P. Jacquard S. Jain M. Lamotte R. Lee R. Lumaret P. C. Miller

H. A. Mooney M. Rapp WA. Reiners B. Saugier H. H. Shugart G. Valdeyron Ph. Vernet P. M. Vttousek D. A. Weinstein

G. M. Woodwell

With 82 Figures

Springer-Verlag Berlin Heidelberg New York Tokyo 1983

Prof. HAROLD A. MOONEY, Department of Biological Sciences, Stanford University, Stanford, CA 94305-2493, USA

Prof. M. GODRON, Universite des Sciences et Techniques du Languedoc, Laboratoire de Systematique et d'Ecologie Mediterraneennes, Institut de Botanique,

Rue Auguste-Broussonnet, F-34000 Montpellier

ISBN-13: 978-3-642-69139-3 e-ISBN-13: 978-3-642-69137-9 DOI: 10.1007/978-3-642-69137-9

Library of Congress Cataloging in Publication Data. Main entry under title: Disturbance and ecosystems. (Ecological studies; v. 44) I. Ecology. 2. Man - Influence on nature. 3. Pollution - Environmental aspects. 1. Mooney, Harold A.

n. Godron. Michel. III. Series. QH541.D57 1983 574.5 83-4771.

This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcastings, reproduction by photocopying

machine or similar means, and storage in data banks. Under § 54 of the German Copyright Law where copies are made for other than private use, a fee is payable to

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© by Springer-Verlag Berlin . Heidelberg 1983

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Typesetting, printing, and binding: Brühlsehe Universitätsdruckerei, Giessen 2131/3130-543210

Ecological Studies Analysis and Synthesis

Edited by

W. D. Billings, Durham (USA) F. Golley, Athens (USA)

O. L. Lange, Wiirzburg (FRG) 1. S. Olson, Oak Ridge (USA)

H. Remmert, Marburg (FRG)

Volume 44

Preface

The earth's landscapes are being increasingly impacted by the activities of man. Unfortunately, we do not have a full understanding of the consequences of these disturbances on the earth's productive capacity. This problem was addressed by a group of French and U.S. ecologists who are specialists at levels of integration extending from genetics to the biosphere at a meeting at Stanford, California, sponsored by the National Science Foundation and the Centre National de la Recherche Scientifique. With a few important exceptions it was found at this meeting that most man-induced disturbances of ecosystems can be viewed as large­scale patterns of disturbances that have occurred, generally on a small scale, in ecosystems through evolutionary time.

Man has induced dramatic large-scale changes in the environment which must be viewed at the biosphere level. Acid deposition and CO2 increase are two examples of the consequences of man's increased utilization of fossil fuels. It is a matter of considerable concern that we cannot yet fully predict the ecological consequences of these environmental changes. Such problems must be addressed at the international level, yet substantive mechanisms to do this are not available.

Ecologists are now viewing the dynamic processes operating at the level of whole landscapes. The consequences of the patterns of man's use oflandscapes are just beginning to be understood. Networks connecting centers of man's activities often sever the flow of resources of populations between ecosystems. They further serve as pathways for invasions of weedy species. Concepts and tools to deal with landscape phenomena have only been developed recently - they are discussed in this book.

The impact of disturbance by large-scale natural catastrophies, such as fire and hurricanes, is similar to that caused by man's utilization of the landscape's natural productivity. Quantitative information is now available that permits us to predict: 1) the changing structure of forests through time after massive disturbance, 2) the changing status of the nutrient stores in ecosystems, and 3) the changes in water balance of whole ecosystems as they develop following disturbance. Knowledge of the events that immediately follow disturbance, when the link between the regulatory processes of the biota and their resources have been temporarily severed, is crucial to ecosystem management. Losses of system resources such as nitrates can be considerable if management practices discourage natural early successional species. These relationships, as outlined here, point to the value of studies of natural ecosystem processes as guides to resource management.

In a consideration of disturbance it was found that both size and frequency were of considerable importance to subsequent resource patterns and population structures. While concerning ourselves with large-scale patterns and processes, we appreciate that these patterns are the result of events occurring at micro sites. The

VI Preface

size of the disturbance site, the time of its occurrence, the proximity and the kind of potential colonizers are all important in determining subsequent patterns of population and resource development. The few studies available indicate this to be true. We need considerably greater documentation on how "patch size" influences recovery processes before we can make predictions. In particular we need to know the interrelationship between forest patches to determine how, in aggregate, they form a forest landscape.

The past decade has seen an explosive development in the fields of plant physiological ecology, plant demography and plant population genetics. The Stanford conference brought together experts in these subdisciplines to discuss these developments, with particular reference to the features of organisms that are characteristic of disturbance habitats. It became clear that these organisms have a commonality of features that adapt them to these habitats ~ that is, they possess a genetic~demographic~physiological syndrome which can be clearly circumscribed. They generally have distinctive nutrient and water-use and growth patterns. Further, they have characteristic reproductive and genetic patterns of variation. These features are obviously interconnected; however, they have traditionally been studied in isolation, and thus the nature of these interrelationships is poorly understood. It appears, though, that if we can understand the nature of resource exploitation by disturbance species where resources may be briefly abundant, we may begin to develop specific tools for managing habitats of differing frequencies and intensities of disturbance. It is important to gain a sound understanding in this area since ~ due to increased habitat disruptive activities by man ~ the earth's ecosystems are being simplified and in many cases are being increasingly occupied by "weedy" species. In order to control patterns of development subsequent to disruption, we must understand the ecology of the various successful invaders.

We hope the contents of this book provide an initial framework for a comprehensive view of the effects of disturbance on ecosystems and on their components.

Stanford University Centre d'Etudes Phytosociologiques et Ecologiques July 1983

H. A. MOONEY

M. GODRON

Contents

Section 1 Biosphere . . . . . . . . . . . . . . . . . . . . . . .

1.1 The Blue Planet: of Wholes and Parts and Man. G. M. WOODWELL. 2 1.1.1 Introduction 2 1.1.2 The CO2 Problem . 3 1.1.3 Toxification. 5 1.1.4 The Effects: Biotic Impoverishment 6 1.1.5 Conclusions . 8

Resume 8 References 9

Section 2 Landscape 11

2.1 Landscape Modification and Changing Ecological Characteristics. M. GODRON and R. T. T. FORMAN. 12 2.1.1 Ecosystems and Landscapes. 12 2.1.2 Some Ecological Attributes for Comparing Landscapes 13

2.1.2.1 Horizontal Structure 14 2.1.2.2 Stability 14 2.1.2.3 Thermodynamic Characterization 16 2.1.2.4 Chorology 16 2.1.2.5 Minimal Area (or Grain) 16 2.1.2.6 Nutrient Cycling. 17 2.1.2.7 Net Production 17 2.1.2.8 Tactics . 17 2.1.2.9 Phylogeny 18 2.1.2.10 Type of Resistance . 18 2.1.2.11 Conclusion 18

2.1.3 Disturbance Regimes. 19 2.1.4 Effect on Landscape Patch Structure . 20

2.1.4.1 Patch Origins 20 2.1.4.2 Patch Size 22 2.1.4.3 Patch Shape . 22 2.1.4.4 Patch Numbers and Configuration 22 2.1.4.5 Summary for Patch Trends 23

2.1.5 Effect of Linkage Characteristics of the Landscape . 23 2.1.5.1 Line Corridors 23 2.1.5.2 Strip Corridors 23

VIII Contents

2.1.5.3 Stream Corridors 24 2.1.5.4 Networks. 25 2.1.5.5 Habitations . . . 25 2.1.5.6 Matrix. . . . . 25 2.1.5.7 Summary of Trends for Linkage Characteristics 25

2.1.6 Conclusions . 25 Resume . . . . . . . . . . . . . . . . . . . . . 26 References . . . . . . . . . . . . . . . . . . . . 27

2.2 Ecological Modeling of Landscape Dynamics. D. A. WEINSTEIN and H. H. SHUGART . . . . . . . . . . . . . . . 29 2.2.1 Introduction . . . . . . . . . . . . . . . . . . 29 2.2.2 Approaches to Modeling Landscape Dynamics 30 2.2.3 A Differential Equation Model of Landscape Change. 33 2.2.4 Complex Digital Computer Model of Forest Dynamics 36 2.2.5 Evaluating the Impact of Regional Scale Problems on Localized

Stands . . . . . . . . . . . . . . . . . . . . . . . . . 37 2.2.6 Evaluating Impact of Regional Scale Problems Across Landscapes 41 2.2.7 Future Directions in Simulation Analysis of Landscapes 43 2.2.8 Conclusions. 44

Resume . 44 References . 45

Section 3 Ecosystem Functions . 47

3.1 Research on the Characteristics of Energy Flows within Natural and Man-Altered Ecosystems. M. LAMOTTE . . . . . . . . 48 3.1.1 Introduction . . . . . . . . . . . . . . . . 48 3.1.2 Examples of Energy Flows in Selected Ecosystems 49 3.1.3 Primary Production . . . . . . . . . . . . . 53

3.1.3.1 Effect of Rejuvenating Forest Stands. . 53 3.1.3.2 Effect of Cutting on the Production of Grasslands 54 3.1.3.3 Effect of Grazing . . . . . . . . . . . . . . 54 3.1.3.4 Effects of Fire on Grasslands . . . . . . . . . 57 3.1.3.5 Destruction and Transformation of Wooded Areas 58

3.1.4 Diversity of Energy Flows in Animal Populatioqs 59 3.1.4.1 AjI Assimilation Efficiency . 60 3.1.4.2 P jA Tissue Growth Efficiency . . . . . 60 3.1.4.3 PjI Ecological Efficiency . . . . . . . 61 3.1.4.4 Possibility of Changing the Energy-use Efficiency at

the Individual Level . . . . . . . . . . . . . . 62 3.1.4.5 Possibility of Changing the Energy-use Efficiency of

Populations . 65 3.1.5 Conclusions . 65

Resume References .

68 70

Contents IX

3.2 "Natural" Mixed Forests and "Artificial" Monospecific Forests. D. AUCLAIR. . . . . . . . . . . 71 3.2.1 Introduction . . . . . . . 71 3.2.2 Advantages of Mixed Forests 73 3.2.3 The Yield of Mixtures . . . 75 3.2.4 Economic Considerations. . 77 3.2.5 Conclusions - Research on Mixed Forests 78

Resume . . . . . ........ . . References . . . . . . . . . . . . . .

79 82

3.3 Disturbance and Basic Properties of Ecosystem Energetics. W. A. REINERS 83 3.3.1 Introduction . . . . . . . . . . . . . . . . . . . . . . 83 3.3.2 Biomass and Energy Flow in Infrequently Disturbed Ecosystems. 84

3.3.2.1 Net Primary Production and Energy Flow Pathways 84 3.3.2.2 Biomass and Detritus Accumulation . . . 87 3.3.2.3 Net Ecosystem Production . . . . . . . . . .. 88 3.3.2.4 Variation in Infrequent Disturbance Events . . .. 88

3.3.3 Biomass and Energy Flow in MUltiple Disturbance Ecosystems. 89 3.3.3.1 Constant Species Composition and Site Quality 89 3.3.3.2 Changing Ecosystem Structure with Disturbance

Frequency 92 3.3.4 Conclusions: Integration

Resume . References . . . . . .

3.4 Ecosystem Water Balance. R. LEE 3.4.1 Basic Concepts . . . . .

3.4.1.1 Water Budgeting. 3.4.1.2 Energy Budgeting 3.4.1.3 Practical Limitations

3.4.2 Ecosystem Influences. . . . 3.4.2.1 Gross Precipitations 3.4.2.2 Evaporation Losses 3.4.2.3 Discharge Losses

3.4.3 Human Influences . . . . . 3.4.3.1 Major Disturbances 3.4.3.2 Flow Regimes. . . 3.4.3.3 Miscellaneous Influences

3.4.4 Conclusions . Resume References .

3.5 Some Problems of Disturbance on the Nutrient Cycling in Ecosystems.

93 96 96

99 99 99

101 103

105 105 107 108 110 110 113 115 115 116 116

M. RAPP . . . . . 117 3.5.1 Introduction . . . . . . . . . . . . . . . . . . . . . . 117

x Contents

3.5.2 Fire . . . . . . . . . . 3.5.3 Reforestation by Conifers . 3.5.4 Forest Fertilization 3.5.5 Removal of Forest Products 3.5.6 The Mineral Budget and Plant Succession. 3.5.7 Conclusions.

Resume References .

118 118 120 122 123 126 126 127

3.6 Mechanisms of Ion Leaching in Natural and Managed Ecosystems. P. M. VITOUSEK . . . . . . . . . . . 129 3.6.1 Introduction . . . . . . . . . . . . . 129 3.6.2 Leaching of Anions and Cations. . . . . 130

3.6.2.1 Measurement of Leaching Losses 130 3.6.2.2 Leaching Mechanisms - Anion Mobility 131 3.6.2.3 The Major Anions . . . . . . . . . . 132 3.6.2.4 Effects of Management Practices on Nitrate Fluxes 138 3.6.2.5 Leaching Losses in Other Biomes 140

3.6.3 Conclusions. 141 Resume 141 References . 142

Section 4 Species Physiological Characteristics 145

4.1 The Determinants of Plant Productivity - Natural Versus Man-Modified Communities. H . A. MOONEY and S. L. GULMON 146 4.1.1 Introduction . . . . . . . . . . . 146 4.1.2 Comparisons of Productivity . . . . 147 4.1.3 The Components of Plant Productivity 147

4.1.3.1 The Biotic Component . . . 147 4.1 .3.2 Environmental Influences on the Biological Components

of Productivity . . . . . . . . . . . . . . . . . 148 4.1.3.3 Interactions of Productivity Components and Resource

Level in Natural Communities . 150 4.1.4 Succession and Plant Productivity 151 4.1.5 Succession Anomalies . . . . . . . . 152 4.1. 6 Convergence in Prod ucti vi ty . . . . . 154 4.1.7 Agricultural Versus Natural Community Productivity. 155 4.1.8 Conclusions. 155

Resume . 156 References . 157

4.2 Plant Growth and Its Limitations in Crops and Natural Communities. B. SAUGIER . . . . . . . . . . 159 4.2.1 Introduction . . . . . . 159 4.2.2 Plant Growth Parameters . 159

Contents XI

4.2.2.1 Photosynthesis. . . . 159 4.2.2.2 Respiration. . . . . 161 4.2.2.3 Other Growth Processes 162

4.2.3 Comparison of Cultivated and Wild Species. 162 4.2.3.1 The Case of Wheat. . . . . . . . 162 4.2.3.2 Adaptation of Natural Species to a Given Level of

Resources . . . . . . . . . . . . . . . . . . 163 4.2.4 Crops Versus Natural Communities . . . . . . . . .. 167 4.2.5 Towards an Estimate in the Level of Available Resources . 170

4.2.5.1 Light. . 171 4.2.5.2 Water . 171 4.2.5.3 Nutrients 172

4.2.6 Conclusions. 173 Resume . 173 References . 173

4.3 Patterns of Nutrient Absorption and Use by Plants from Natural and Man-Modified Environments. F. S. CHAPIN III . 175 4.3.1 Introduction . . . . . . . . . . . 175 4.3.2 General Patterns and Nutrient Use. . . 176 4.3.3 Successional Changes in Nutrient Use . 177 4.3.4 Nutritional Patterns Related to Disturbance. 179

4.3.4.1 Abandoned Fields . 179 4.3.4.2 Post-Fire Succession . . . . . . . 180 4.3.4.3 Tundra Disturbance . . . . . . . 180 4.3.4.4 Disturbances Causing Reduced Nutrient Availability. 181

4.3.5 Conclusions . Resume . References .

4.4 Comparisons of Water Balance Characteristics of Plant Species in

184 184 185

"Natural" Versus Modified Ecosystems. P. C. MILLER. 188 4.4.1 Introduction . . . . . . . . . . . . . . 188 4.4.2 Theoretical Background . . . . . . . . . . 189

4.4.2.1 Heat and Water Exchange Processes . 189 4.4.2.2 Water Availability and Plant Characteristics. 191

4.4.3 Survey of Plant Characteristics . . . . . . . . . . 196 4.4.3.1 General Relations . . . . . . . . . . . . 196 4.4.3.2 Factors and Processes Affecting Water Loss. 196 4.4.3.3 Factors and Processes Affecting Water Uptake. 197 4.4.3.4 State of Water in the Plant . . . . . . . . . 199 4.4.3.5 Growth and Death in Relation to Plant Water Content. 201

4.4.4 Theoretical Considerations Relating Plant Characteristics and Successional State . . . . . . . . . . . . . . . . . . . . 201

XII Contents

4.4.4.1 Water Availability and Vegetative Recivery in the Semiarid Mediterranean Regions of Southern California 201

4.4.5 Conclusions. 206

Resume . References .

206

207

Section 5 Population Characteristics . 213

5.1 Reproductive Strategies and Disturbance by Man. P. H. GOUYON et al. . 214

5.1.1 Introduction . . . . . . . . . . . . . . . 214 5.l.2 Chemical and Sexual Polymorphism in Thyme. 214

5.l.2.1 Sexual Polymorphism . . . . . . . . 214 5.l.2.2 Chemical Polymorphism . . . . . . 215 5.l.2.3 Environment and Population Genetic Structure 217

5.1.3 Enzymatic Polymorphism in Orchard Grass . 5.1.4 Conclusions .

Resume . References .

5.2 Demographic Strategies and Originating Environment.

220 223

224

224

P. JACQUARD and G. HElM . . . . . . . . . . . 226 5.2.1 Introduction . . . . . . . . . . . . . . . 226 5.2.2 Description of the Originating Environments . 226 5.2.3 Between and Within-Population Variations of Strategies in

Arrhenatherum elatius . . . . . . . . . . . . . . .. 227 5.2.4 Between and Within-Population Variations of Strategies in

D. glomerata 230 5.2.5 Conclusions. 234

References . 238

5.3 Genetic Characteristic of Populations. S. JAIN 240

5.3.1 Introduction . . . . . . . . . . . 240 5.3 .2 Population Studies in Avena spp. . . 241 5.3.3 Rose Clover, a Case History of Recent Colonization 244 5.3.4 Population Dynamics of Species in a Coastal Grassland

Ecosystem . . . . . . . . . . . . . . . . . . . . 246 5.3.5 Variation and Colonization Success of Crop-Weed Hybrids 247 5.3.6 Alternative Strategies of Colonizing Success . 248 5.3.7 Evolutionary Genetics of Adaptive Responses . . 250 5.3.8 Recombination Properties of Genetic Systems . . 251 5.3.9 Interspecific Interactions in Community Dynamics 252 5.3.lO Conclusions. 254

Resume 255

References . 256

5.4 Characteristics of Populations in Relation to Disturbance in Natural and Man-Modified Ecosystems. F. A. BAZZAZ . . . . . . . . . . . . . 259

Contents XIII

5.4.1 Introduction . . . . . . . . . . . . . . . . . . .. 259 5.4.2 The Nature of Disturbance . . . . . . . . . . . . . 260 5.4.3 Disturbance Characteristic with Relevance to Population

Response. . . . . . . . . . . 261 5.4.3.1 Size . . . . . . . . . 261 5.4.3.2 Frequency of Occurrence 262 5.4.3.3 Intensity . . . . . . . 263 5.4.3.4 Time of Disturbance . . 263 5.4.3.5 Level of Environmental Heterogeneity 263 5.4.3.6 Nature of the Biologic Neighborhood. 266

5.4.4 Population Characteristics Responsive to Disturbance . 5.4.4.1 Density, Dispersion and Age Structure 5.4.4.2 Genotypic Variability in Populations 5.4.4.3 Interactions Between Species

5.4.5 Life History Characteristics and Disturbance 5.4.5.1 Life Span . . . . . . . . . . . . 5.4.5.2 Reproductive Strategies. . . . . . 5.4.5.3 Germination, Growth and Response Breadth

5.4.6 Conclusions. Resume . References

Subject Index . . . .

266 266 267 268 268 268 269 270 272 273

273

277

Contributors

AUCLAIR, DANIEL, Station de Recherches sur la Foret et l'Environment, Institut National de la Recherche Agronomique, Olivet, France

BAZZAZ, FAKHRI A., Department of Botany, University of Illinois, Urbana, Illinois, USA

CHAPIN, F. STUART III, Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska, USA

FORMAN, RICHARD T. T., Department of Biological Sciences-Botany, Rutgers University, New Brunswick, New Jersey, USA

GODRON, MICHEL, CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B.P. 5051, Montpellier, France

GOUYON, PIERRE-HENRI, Institute National Agronomique, 16 rue Claude Bernard, Paris, France

GULMON, SHERRY, L., Department of Biological Sciences, Stanford University, Stanford, California, USA

HElM, GEORGES, CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B. P. 5051, Montpellier, France

JACQUARD, PIERRE, CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B. P. 5051, Montpellier, France

JAIN, SUBODH, Department of Agronomy and Range Science, University of California, Davis, California, USA

LAMOTTE, MAXIME, Ecole Normale Superieure, Laboratoire de Zoologie, 46 rue d'Ulm, Paris, France

LEE, RICHARD, Division of Forestry, West Virginia University, Morgantown, West Virginia, USA

LUMARET, R., CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B. P. 5051, Montpellier, France

MILLER, PHILIP c., Systems Ecology Research Group, San Diego State University, San Diego, California, USA

MOONEY, HAROLD A., Department of Biological Sciences, Stanford University, Stanford, California, USA

XVI Contributors

RApp, MAURICE, CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B.P. 5051, Montpellier, France

REINERS, WILLIAM A., Department of Biological Sciences, Dartmouth College, Hanover, New Hampshire, USA

SAUGIER, BERNARD, Laboratoire d'Ecologie Vegetale, Universite de Paris-Sud, Batiment 431, Orsay, France

SHUGART, HERMAN H., Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee, USA

V ALDEYRON, G., CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B.P. 5051, Montpellier, France

VERNET, PH., CNRS, Centre d'Etudes Phytosociologiques et Ecologiques L. Emberger, Route de Mende, B. P. 5051, Montpellier, France

VITOUSEK, PETER M., Department of Botany, University of North Carolina, Chapel Hill, North Carolina, USA

WEINSTEIN, DAVID A., Environmental Sciences Division, Oak Ridge National Laboratory, Oak Ridge, Tennessee, USA

WOODWELL, GEORGE M., The Ecosystems Center, Marine Biological Laboratory, Woods Hole, Massachusetts, USA