Table Of ContentEcological Studies
Analysis and Synthesis
Edited by
W.D. Billings, Durham (USA) F. Galley, Athens (USA)
O.L. Lange, Wiirzburg (FRG) J.S. Olson, Oak Ridge (USA)
H. Remmert, Marburg (FRG)
Volume 96
Ecological Studies
Volume 78 Volume 88
Agroecology: Researching the Pinnipeds and El Nino: Responses to
Ecological Basis for Sustainable Environmental Stress (1991)
Agriculture (1990) Edited by F. Trillmich and K.A. Ono
Edited by S.R. Gliessman
Volume 89
Volume 79 Plantago: A Multidisciplinary Study
Remote Sensing of Biosphere (1992)
Functioning (1990) Edited by P.I.C. Kuiper and M. Bos
Edited by R.I. Hobbs and H.A. Mooney
Volume 90
Volume 80 Biogeochemistry of a Subalpine
Plant Biology of the Basin and Range Ecosystem: Loch Vale Watershed (1992)
(1990) Edited by Iill Baron
Edited by C.B. Osmond, L.F. Pitelka,
and G.M. Hidy Volume 91
Atmospheric Deposition and
Volume 81 Forest Nutrient Cycling (1992)
Nitrogen in Terrestrial Ecosystems Edited by D.W. Iohnson and
(1990) S.E. Lindberg
By C.O. Tamm
Volume 92
Volume 82 Landscape Boundaries: Consequences
Quantitative Methods in Landscape for Biotic Diversity and Ecological
Ecology (1990) Flows (1992)
Edited by M.G. Turner and R.H. Edited by A.I. Hansen and
Gardner F. di Castri
Volume 83 Volume 93
The Rivers of Florida (1990) Fire in South African Mountain Fynbos:
Edited by R.I. Livingston Ecosystem, Community, and Species
Response at Swartboskloof (1992)
Volume 84 Edited by B.W. van Wilgen, D.M.
Fire in the Tropical Biota: Ecosystem Richardson, F.I. Kruger, and H.I. van
Processes and Global Challenges (1990) Hensbergen
Edited by J.G. Goldammer
Volume 94
Volume 85 The Ecology of Aquatic Hyphomycetes
The Mosaic·Cycle Concept of (1992)
Ecosystems (1991) Edited by F. Barlocher
Edited by H. Remmert
Volume 95
Volume 86 Palms in Forest Ecosystems of Amazonia
Ecological Heterogeneity (1991) (1992)
Edited by 1. Kolasa and By F. Kahn and 1.·1. DeGranville
S.T.A. Pickett
Volume 96
Volume 87 Ecology and Decline of Red Spruce in
Horses and Grasses: The Nutritional the Eastern United States (1992)
Ecology of Equids, and Their Impact on Edited by C. Eagar and M.B. Adams
the Camargue (1991)
By P. Duncan
Christopher Eagar
Mary Beth Adams
Editors
Ecology and Decline
of Red Spruce in the
Eastern United States
Contributors
M.B. Adams C. Cogbill E.R. Cook D.H. DeHayes
C. Eagar 1.J. Fernandez K.F. Jensen A.H. Johnson
D.W. Johnson R.J. Kohut S.B. McLaughlin
M.M. Miller-Weeks V.A. Mohnen N.S. Nicholas
D.R. Peart G.A. Schier T.G. Siccama P.S. White
S.M. Zedaker
With 90 illustrations, 2 in color
Springer-Verlag
New York Berlin Heidelberg London Paris
Tokyo Hong Kong Barcelona Budapest
Christopher Eagar Mary Beth Adams
USDA Forest Service USDA Forest Service
Northeastern Forest Experiment Northeastern Forest Experiment
Station Station
P.O. Box 640 Timber and Watershed Laboratory
Durham, NH 03824 P.O. Box 404-Nursery Bottom
USA Parsons, WV 26287
USA
Library of Congress Cataloging-in-Publication Data
Ecology and decline of red spruce in the eastern United States
edited by Christopher Eagar and Mary Beth Adams.
p. cm. - (Ecological studies; v. 96)
Includes bibliographical references and index.
ISBN-13: 978-1-4612-7714-9 e-ISBN-13: 978-1-4612-2906-3
001: 10.1 007/978-1-4612-2906-3
1. Red spruce decline-East (U.S.) 2. Red spruce-East (U.S.)
Effect of air pollution on. 3. Red spruce-East (U.S.)-Ecology.
4. Forest declines-East (U.S.) 5. Forest ecology-East (U.S.)
I. Eagar, Christopher. II. Adams, Mary Beth, 1958-
III. Series.
SB608.R33E36 1992
585' .2-dc20 91-46553
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Preface
In the early 1980s there were several published reports of recent,
unexplained increases in mortality of red spruce in the Adirondack
Mountains and the northern Appalachian Mountains of the eastern
United States. These reports coincided with documentation of reductions
in radial growth of several species of pine in the southeastern United
States, and with the severe, rapid, and widespread decline of Norway
spruce, silver fir, and some hardwoods in central Europe. In all of these
instances, atmospheric deposition was hypothesized as the cause of the
decline. (Throughout this volume, we use the term "decline" to refer to a
loosely synchronized regional-scale deterioration of tree health which is
brought about by a combination of stress factors. These may be biotic or
abiotic in nature, and the combinations may differ from site to site.)
Heated public debate about the causes and possible cures for these forest
declines ensued. Through the course of this debate, it became clear that
information about forest health and air pollution effects on forests was
inadequate to meet policymakers' needs. Ecology and Decline of Red
Spruce in the Eastern United States addresses that gap for eastern spruce
fir forests and represents the culmination of a great deal of research
conducted in recent years. The focus is on red spruce because the decline
of red spruce was both dramatic and inexplicable and because of the great
amount of information gathered on red spruce.
Section I of this book examines the characteristics of spruce-fir forests
VI Preface
in the eastern United States, including descriptions of the history and
ecology of these forests, the soils commonly found under these forests,
and the atmospheric conditions these forests experience. An extensive
discussion of recent changes in forest health and growth trends concludes
this section.
Section II presents experimental results from many air pollution effects
studies, and evaluates mechanisms of air pollution effects on red spruce.
Four specific mechanisms are evaluated: soil-mediated effects, foliar
leaching of nutrients and direct foliar injury, altered cold tolerance, and
effects on carbon allocation and associated physiological processes.
Section III develops a synthesis of the state of knowledge regarding
red spruce decline at high elevations in the eastern United States. The
discussion was developed by the authors of the first two sections, under
the leadership of Drs. Arthur Johnson and Samuel McLaughlin, and
represents a consensus of the listed contributors.
Much of the research discussed in this volume was funded by the
Spruce-Fir Research Cooperative of the National Acid Precipitation
Assessment Program's Forest Response Program. The Spruce-Fir Research
Cooperative was formed in March 1985 and was jointly planned, man
aged, and funded by the USDA Forest Service and the United States
Environmental Protection Agency. Its mission was to explain possible air
pollution-related changes in the structure, function and composition of
spruce-fir forests. In particular, the Cooperative addressed two questions:
(1) What are the effects of air pollutants and acidic deposition on eastern
spruce-fir forests? (2) Why are red spruce declining at high elevations?
During its six years of existence, the Spruce-Fir Research Cooperative
funded $16 million of research, involving more than 43 projects and 120
investigators from many different institutions. Affiliated with the Spruce
Fir Research Cooperative were the Mountain Cloud Chemistry Program
and the National Vegetation Survey, which provided valuable informa
tion on atmospheric conditions and forest condition, respectively. Other
organizations that sponsored research on red spruce summarized in this
compendium include the National Council of the Paper Industry for
Air and Stream Improvement, the Electric Power Research Institute
predominantly through its Integrated Forest Study (IFS)-the Tennessee
Valley Authority, and the Mellon Foundation.
Through the efforts of many investigators in these programs and
others, a clearer picture has been drawn of the ecology and decline of
red spruce. The evolution of the information presented in this volume
demonstrates the value and success of diligent scientific endeavor for
addressing critical policy issues.
Christopher Eagar
Mary Beth Adams
Acknowledgments
Production of a book of this scope requires the cooperation and assist
ance of many individuals. The hard work of the individual chapter
authors, and the diligence in particular of Dr. Arthur Johnson and Dr.
Sandy McLaughlin in working toward consensus were essential to the
success of this book.
The following individuals served as peer reviewers for one of the
first nine chapters: David H. Alban, Robert A. Amundson, Ralph
Baumgardner, Russell D. Briggs, Neil Cape, Phillip Dougherty,
George M. Hidy, Jay S. Jacobson, J. Dev Joslin, Richard Klein, James
McClenahan, Imants Millers, Kevin Percy, David Peterson, Daniel D.
Richter, John Skelly, Kim C. Steiner, Richard Tinus, and John Witter.
Their reviews greatly contributed to the high quality of these chapters.
Special thanks go to David Grigal and Ellis Cowling for reviewing the
first draft of the book in its entirety. The dedicated clerical assistance
of Dolores Beckford, Cass Cloak, and Marie Lanza is also greatly appre
ciated. Layne Godwin assisted with word processing during final revisions.
Support for preparation of this book, and for much of the research
discussed in this book, was provided by the Spruce-Fir Research Co
operative, of the joint U.S. Environmental Protection Agency-USDA
Forest Service Forest Response Program of the National Acid Precipita
tion Assessment Program. The support of the Northeastern Forest
Experiment Station of the USDA Forest Service was particularly helpful
Vlll Acknowledgments
during the preparation of this book and is acknowledged. This book has
not been subject to Environmental Protection Agency or Forest Service
policy review and should not be construed to represent the policies of
either agency.
Contents
Preface
V
Acknowledgments
VII
Contributors
XI
Section I. Characterization of the Condition of Eastern U.S.
Spruce-Fir Forests
1. Spruce-Fir Forests of Eastern North America 3
Peter S. White and Charles V. Cogbill
2. Characterization of Eastern U.S. Spruce-Fir Soils 40
Ivan J. Fernandez
3. Atmospheric Deposition and Pollutant Exposure of Eastern
U.S. Forests 64
Volker A. Mohnen
4. Condition and Recent Trends in High-Elevation Red Spruce
Populations 125
David R. Peart, N.S. Nicholas, Shephard M. Zedaker,
Margaret M. Miller-Weeks, and Thomas G. Siccama
5. The Dendroecology of Red Spruce Decline 192
Edward R. Cook and Shephard M. Zedaker
x Contents
Section II. Air Pollution and Atmospheric Deposition Effects on
Red Spruce: Evaluation of Mechanisms
6. Soil Mediated Effects of Atmospheric Deposition on Eastern
U.S. Spruce-Fir Forests 235
Dale W. Johnson and Ivan J. Fernandez
7. Atmospheric Deposition Effects on Foliar Injury and Foliar
Leaching in Red Spruce 271
George A. Schier and Keith F. Jensen
8. Winter Injury and Developmental Cold Tolerance of Red
Spruce 295
Donald H. DeHayes
9. The Effects of Atmospheric Deposition and Ozone on Carbon
Allocation and Associated Physiological Processes in Red
Spruce 338
Samuel B. McLaughlin and Robert J. Kohut
Section III. Why are Red Spruce Declining at High Elevations?
A Synthesis of Epidemiological and Mechanistic Studies
10. Synthesis and Conclusions from Epidemiological and
Mechanistic Studies of Red Spruce Decline 385
A.H. Johnson, S.B. McLaughlin, M.B. Adams, E.R. Cook,
D.H. DeHayes, C. Eagar, 1.1. Fernandez, D.W. Johnson,
R.J. Kohut, V.A. Mohnen, N.S. Nicholas, D.R. Peart, G.A.
Schier, and P.S. White
Index 413
