The Ecology of Natural Disturbance and Patch Dynamics

457 Pages·1985·7.225 MB·English

by  Steward T. Pickett and P.S. White (Eds.)

CONTRIBUTORS
Nicholas V. L. Brokaw Mary L. Plumb-Mentjes
Charles D. Canham Kevin Rice
Norman L. Christensen Deborah Rogers
B. S. Collins James R. Runkle
Joseph H. Connell Timothy D. Schowalter
Julie Sloan Denslow S. W. Seagle
K. P. Dunne H. H. Shugart
Kathryn E. Freemark Wayne P. Sousa
Subodh Jain Douglas G. Sprugel
James R. Karr John N. Thompson
Michael J. Keough Thomas T. Veblen
Orie L. Loucks Peter M. Vitousek
P. L. Marks Robert C. Vrijenhoek
S. T. A. Pickett P. S. White
John Wiens
The Ecology
of Natural Disturbance
and Patch Dynamics
Edited by
S. T. A. PICKETT
Department of Biological Sciences
and Bureau of Biological Research
Rutgers University
New Brunswick, New Jersey
P. S. WHITE
Uplands Field Research Laboratory
Great Smoky Mountains National Park
Twin Creeks Area
Gatlinburg, Tennessee
and Graduate Program in Ecology
University of Tennessee
Knoxville, Tennessee
1985
ACADEMIC PRESS, INC.
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ACADEMIC PRESS, INC.
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United Kingdom Edition published by
ACADEMIC PRESS INC. (LONDON) LTD.
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Library of Congress Cataloging in Publication Data
Main entry under title:
The Ecology of natural disturbance and patch dynamics.
Bibliography: p.
Includes index.
1. Natural disasters-Environmental aspects.
2. Biotic communities. I. Pickett, S. T. A.
II. White, P. S. III. Title: Patch dynamics.
QH545.N3E28 1985 574.5'22 84-18599
ISBN 0-12-554520-7 (hardcover) (alk. paper)
ISBN 0-12-554521-5 (paperback) (alk. paper)
PRINTED IN THE UNITED STATES OF AMERICA
86 87 88 9 8 7 6 5 4 3 2
Contributors
Numbers in parentheses indicate the pages on which the authors' contributions begin.
Nicholas V. L. Brokaw1 (53), Department of Biology, Kenyon College, Gambier,
Ohio 43022, and Smithsonian Tropical Research Institute, Balboa, Panama
Charles D. Canham (197), Section of Ecology and Systematics, Cornell Univer­
sity, Ithaca, New York 14853
Norman L. Christensen (85), Department of Botany, Duke University, Durham,
North Carolina 27706
B. S. Collins (217), Department of Biological Sciences and Bureau of Biological
Research, Rutgers University, New Brunswick, New Jersey 08901
Joseph H. Connell (125), Department of Biological Sciences, University of Cal­
ifornia, Santa Barbara, California 93106
Julie Sloan Denslow (307), Departments of Zoology and Botany, University of
Wisconsin, Madison, Wisconsin 53706
K. P. Dunne (217), Department of Biological Sciences and Bureau of Biological
Research, Rutgers University, New Brunswick, New Jersey 08901
Kathryn E. Freemark (153), Department of Biology, Carleton University, Ot­
tawa, Ontario, Canada K1S 5B6
Subodh Jain (287), Department of Agronomy and Range Science, University of
California, Davis, California 95617
James R. Karr2 (153), Department of Ecology, Ethology, and Evolution, Univer­
sity of Illinois, Champaign, Illinois 61820
Michael J. Keough3 (125), Department of Biological Sciences, University of
California, Santa Barbara, California 93106
Orie L. Loucks (71), Butler University,.Indianapolis, Indiana 46208
Present address: Manomet Bird Observatory, Manomet, Massachusetts 02345.
2Present address: Smithsonian Tropical Research Institute, P.O. Box 2072, Balboa, Panama.
3Present address: Department of Biological Science, Florida State University, Tallahassee, Florida
32306.
XJ
xii Contributors
P. L. Marks (197), Section of Ecology and Systematics, Cornell University,
Ithaca, New York 14853
S. T. A. Pickett4 (3, 217, 371), Department of Biological Sciences and Bureau of
Biological Research, Rutgers University, New Brunswick, New Jersey 08901
Mary L. Plumb-Mentjes (71), U.S. Army Corps of Engineers, New Orleans,
Louisiana 70160-0267
Kevin Rice5 (287), Department of Agronomy and Range Science, University of
California, Davis, California 95617
Deborah Rogers (71), Technical Information Project, Pierre, South Dakota 57501
James R. Runkle (17), Department of Biological Sciences, Wright State Univer­
sity, Dayton, Ohio 45435
Timothy D. Schowalter (235), Department of Entomology, Oregon State
University, Corvallis, Oregon 97331
S. W. Seagle (353), Graduate Program in Ecology, University of Tennessee, Knox-
ville, Tennessee 37996
H. H. Shugart6 (353), Environmental Sciences Division, Oak Ridge National
Laboratory, Oak Ridge, Tennessee 37831
Wayne P. Sousa (101), Department of Zoology, University of California, Berke­
ley, California 94720
Douglas G. SprugeF (335), Department of Forestry, Michigan State University,
East Lansing, Michigan 48824
John N. Thompson (253), Departments of Botany and Zoology, Washington
State University, Pullman, Washington 99164
Thomas T. Veblen (35), Department of Geography, University of Colorado,
Boulder, Colorado 80309
Peter M. Vitousek8 (325), Department of Biology, University of North Carolina,
Chapel Hill, North Carolina 27514
Robert C. Vrijenhoek (265), Department of Biological Sciences and Bureau of
Biological Research, Rutgers University, New Brunswick, New Jersey 08901
P. S. White9 (3, 371), Uplands Field Research Laboratory, Great Smoky Moun­
tains National Park, Twin Creeks Area, Gatlinburg, Tennessee 37738, and
Graduate Program in Ecology, University of Tennessee, Knoxville,
Tennessee 37916
John A. Wiens (169), Department of Biology, University of New Mexico, Albu­
querque, New Mexico 87131
4Present address: Institute of Ecosystem Studies, New York Botanical Garden, Mary Flagler
Cary Arboretum, Box AB, Millbrook, New York 12545.
'Present address: Department of Botany, Washington State University, Pullman, Washington
99164-4230.
6Present address: Department of Environmental Sciences, University of Virginia, Charlottesville,
Virginia 22903.
7Present address: College of Forest Resources, AR-10, University of Washington, Seattle,
Washington 98195.
8Present address: Department of Biological Sciences, Stanford University, Stanford, California
94305.
'Present address: Department of Biology, University of North Carolina, Chapel Hill, North
Carolina 27514.
Preface
Ecologists have always been aware of the importance of natural dynamics in
ecosystems, but historically, the focus has been on successional development of
equilibrium communities. While this approach has generated appreciable under­
standing of the composition and functioning of ecosystems, recently many workers
have turned their attention to processes of disturbance themselves and to the evolu­
tionary significance of such events. This shifted emphasis has inspired studies in
diverse systems. We use the phrase "patch dynamics" (Thompson, 1978) to de­
scribe their common focus.
Focus on patch dynamics leads workers to explicit studies of disturbance-related
phenomena—the conditions created by disturbance; the frequency, severity, inten­
sity, and predictability of such events; and the responses of organisms to distur­
bance regimes. The phrase "patch dynamics" embraces disturbances external to
the community as well as internal processes of change. Patch dynamics includes not
only such coarse-scale, infrequent events as hurricanes, but also such fine-scale
events as the shifting mosaic of badger mounds in a prairie. The scope of this book
includes population, community, ecosystem, and landscape levels. The most basic
theme is an evolutionary one: How does the dynamic setting of populations influ­
ence their evolution? What are the implications for communities and ecosystems?
This book seeks to bring together the findings and ideas of workers studying such
varied systems as marine invertebrate communities; grasslands; and boreal, tem­
perate, and tropical forests. Our primary goal is to present a synthesis of diverse
individual contributions. The book is divided into three main sections: (1) examples
of patch dynamics in diverse systems; (2) adaptations of organisms and evolution of
populations in patch dynamic environments; and (3) implications of patch dynamics
for the organization of communities and the functioning of ecosystems. We feel this
approach demonstrates the commonality of disturbance-generated phenomena over
a wide range of scales and levels of organization and thus validates the broad
applicability of the patch dynamic viewpoint. We seek to present clearly a frame­
work that can stimulate the generation of explicit hypotheses and theory and thus
form an alternative to equilibrium concepts of the evolution of populations, com­
position of communities, and functioning of ecosystems. We hope, in addition, that
this volume will help identify areas of future research.
xiii
xiv Preface
This book draws principally on terrestrial and marine systems, in which most
work on patch dynamics has been done. Freshwater environments have received
less emphasis, because studies of patch dynamics have been rare in these systems.
By contrast, there is a rich, and largely recent, literature on a wide variety of
terrestrial systems, examining both biotic and abiotic components and a variety of
trophic levels. This book provides a common focus for this growing body of work,
and aquatic patch dynamics have been included in this synthesis where possible.
Although this book is replete with detailed examples of community dynamics and
organism adaptation, no book of this length could exhaustively treat all systems;
even some terrestrial systems are missing here. For example, descriptions of boreal
forest patch dynamics were omitted in reliance on recent, widely available reviews.
We have sought, rather, to emphasize building a theoretical framework in which to
view disturbance in natural systems. The stress here is on the processes of dynamics
and emerging generalizations from patch dynamic systems, and not a mere catalog
of examples. Treating certain systems in depth, while drawing connections and
parallels with others, will best serve an understanding of processes and successful
generalization.
A state-of-the-art conference on succession held in 1980 and summarized by
West et al. (1981) convinced us that although there is much scattered information
on the subject of disturbance and patch dynamics and many workers are currently
interested in the topic, there is no compilation and synthesis of this information; nor
is there any work which develops a broad framework or theory incorporating distur­
bance and its effect. We feel that this survey, drawing on workers familiar with
many different systems and having interests at various levels of organization, is an
ideal vehicle for meeting these needs.
This book is aimed at ecologists and advanced students working in this rapidly
expanding field. Because we incorporate theoretical, empirical, and applied ap­
proaches to the effects of disturbance on plants, animals, and entire ecosystems, the
book should be useful to workers of diverse backgrounds and interest.
S. T. A. Pickett
P. S. White
Chapter 1
Natural Disturbance
and Patch Dynamics:
An Introduction
P. S. WHITE
Uplands Field Research Laboratory
Great Smoky Mountains National Park
Twin Creeks Area
Gatlinburg, Tennessee
and
Graduate Program in Ecology
University of Tennessee
Knoxville, Tennessee
S. T. A. PICKETT
Department of Biological Sciences and Bureau of Biological Research
Rutgers University
New Brunswick, New Jersey
I. The Dynamics of Biological Systems 3
II. Definitions: Patch Dynamics, Perturbation, and Disturbance 4
A. Patch Dynamics 4
B. Perturbation 5
C. Disturbance 6
D. Endogenous and Exogenous Causes of Disturbance 8
III. Natural Disturbance: The Patch Dynamics Perspective 9
I. THE DYNAMICS OF BIOLOGICAL SYSTEMS
The processes of growth, death, and replacement ensure that biological systems
are dynamic, if only on a local scale. The extreme case in which these dynamics are
entirely a function of individual mortality is illustrated by the monocarpic tropical
THE ECOLOGY Copyright © 1985 by Academic Press, Inc.
OF NATURAL DISTURBANCE All rights of reproduction in any form reserved.
AND PATCH DYNAMICS ISBN 0-12-554520-7
3
4 P. S. White and S. T. A. Pickett
tree Tachigalia versicolor (Foster, 1977) and by the cyclic successions of some
shrub-dominated communities (Watt, 1947; Christensen, Chapter 6, this volume).
In most biological systems, however, other factors besides individual growth and
death contribute to dynamics. Foremost among these factors are natural distur­
bances (White, 1979). The catalog of ecosystems in which natural disturbances play
a fundamental role is a long one, as several reviews attest (Knapp, 1974; Grubb,
1977; Miles, 1979; White, 1979; Pickett, 1980; Oliver, 1981).
Natural disturbances and patch dynamics occur on a wide variety of spatial and
temporal scales (Delcourt et al., 1983). In this volume, dynamics are described that
span a temporal scale of 10° to 103 years and a spatial scale of 10 ~ 4 to 106 m2. The
range of these scales is wide because the biological systems treated in this volume
are varied and because disturbance effects occur on a variety of scales within each
system. Our goal in this volume is the exploration of general themes in this diver­
sity; in this chapter we foster that goal by defining three central concepts: patch
dynamics, perturbation, and disturbance. These definitions lead to a discussion of
endogenous and exogenous factors in community pattern.
II. DEFINITIONS: PATCH DYNAMICS, PERTURBATION, AND
DISTURBANCE
The sources of variation in disturbances include differences in ecosystem scale,
differences in kinds of disturbances, and differences in disturbance regimes. Even
for a single ecosystem and disturbance event, effects vary at different trophic levels
and occur over a wide range of biological levels from suborganismal (e.g., physio­
logical effects; Sousa, Chapter 7, this volume) and organismal (e.g., behavioral
changes; Wiens, Chapter 10, this volume) to ecosystem-wide (e.g., nutrient avail­
ability; Vitousek, Chapter 18, this volume). Most disturbances produce hetero­
geneous and patchy effects; these effects may themselves depend on the state of the
community prior to the disturbance. The consequences of a given disturbance are
strongly dependent on a variety of biotic and physical factors (e.g., regional cli­
matic gradients, topographic gradients, and substrate types). Despite this plurality
of ''disturbances," a definition must be sought in a search for generality.
A. Patch Dynamics
We have adopted the term "patch dynamics" (Thompson, 1978; Pickett and
Thompson, 1978) to label the central organizing theme of this book for the follow­
ing reasons:
1. "Patch" implies a relatively discrete spatial pattern, but does not establish
any constraint on patch size, internal homogeneity, or discreteness.
2. "Patch" implies a relationship of one patch to another in space and to the
surrounding, unaffected or less affected matrix.
3. "Patch dynamics" emphasizes patch change.
1. Introduction 5
We do not suggest that patches are spatially discrete in an absolute sense (note the
descriptions of gap edge in Chapters 2 by Runkle, 4 by Brokaw, and 8 by Connell
and Keough, this volume), nor that patches are internally homogeneous (note the
range of fire effects described by Christensen, Chapter 6, this volume), nor that
patches are necessarily temporally discrete in origin (note that treefalls on gap edges
increase the original gap area; Runkle, Chapter 2, this volume). Particular defini­
tions of "patch" will always be relative to the system at hand. But within a
particular system, we do suggest that community structure and behavior vary locally
and in a relatively patch-wise manner. Organisms, by their very nature, take up
space and use resources; biological systems, on some level, are patchy.
A concept similar to "patch dynamics" is "shifting mosaic" (Bormann and
Likens, 1979; Heinselman, 1981a). This concept connotes, however, a uniformity
of patch distribution in time and space such that an overall landscape equilibrium of
patches applies. Such equilibria are to be expected where feedback occurs between
community characteristics and disturbance events (e.g., when susceptibility to dis­
turbance increases with the time since the last disturbance), where the patch size is
small relative to the homogeneous landscape unit, and where disturbance regimes
are stable. We prefer "patch dynamics" for more general situations in which such
an equilibrium has not been demonstrated and for situations in which patch scale is
large relative to the scale of the relevant landscape (Romme and Knight, 1981).
Further, environmental fluctuation, which may cause a shift in the disturbance
regime, occurs on time scales similar to those of disturbances operating in the same
system (see, e.g., Neilson and Wullstein, 1983). Equilibrium landscapes would
therefore seem to be the exception, rather than the rule (for example, most North
American landscapes have probably been influenced by changing disturbance re­
gimes in the last several thousand to tens of thousands of years [Delcourt et al.,
1983]).
Despite the difficulty in demonstrating the existence of patch equilibria, the
concept of the "shifting mosaic" is important on a theoretical level. For example,
all successional states would be predictable and permanent (if locally fleeting)
features of steady state landscapes under a stable disturbance regime. Size class
distributions of disturbance patches would be invariant. Disturbance regimes could
be calculated from either the temporal or spatial distribution of disturbance patches
because these two aspects of disturbance would be directly linked. The simul­
taneous occurrence of local dynamics and broad-scale equilibrium also underscores
the central importance of scale hierarchies in the interpretation of natural systems
(see, e.g., Allen and Starr, 1982; see also Zedler and Goff, 1973). Dynamics on one
scale determine stasis on another.
B. Perturbation
"Disturbance" and "perturbation" have been used synonymously by some au­
thors and yet possess particular meanings in the work of others. In general, "pertur­
bation" has been used with a whole system orientation in the sense of any change in