Forest Ecosystems 3rd Edition Richard H. Waring Download PDF
Forest Ecosystems 3rd Edition Richard H. Waring Download PDF
Waring
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Forest Ecosystems 3rd edition Richard H. Waring Digital
Instant Download
Author(s): Richard H. Waring, Steven W. Running
ISBN(s): 9780123706058, 012370605X
Edition: 3
File Details: PDF, 30.32 MB
Year: 2007
Language: english
Forest Ecosystems
THIRD EDITION
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Forest
Ecosystems
Analysis at Multiple Scales
THIRD EDITION
Richard H. Waring
DEPARTMENT OF FOREST SCIENCE
OREGON STATE UNIVERSITY
CORVALLIS, OREGON
Steven W. Running
DEPARTMENT OF ECOSYSTEM AND CONSERVATION SCIENCES
NUMERICAL TERRADYNAMIC SIMULATION GROUP
UNIVERSITY OF MONTANA
MISSOULA, MONTANA
Cover Image–Gross primary production for June 2–10, 2003, draped over a digital elevation map of the complex
50,000 km2 forested landscape of western Montana. The calculations for this image combined 250 m resolution
spectral data from MODIS of the fraction of absorbed PAR with 1 km daily surface meteorology (Running et al.,
2004).
No part of this publication may be reproduced or transmitted in any form or by any means, electronic or
mechanical, including photocopy, recording, or any information storage and retrieval system, without permission
in writing from the publisher.
Permissions may be sought directly from Elsevier’s Science & Technology Rights Department in Oxford, UK:
phone: (+44) 1865 843830, fax: (+44) 1865 853333, E-mail: permissions@elsevier.co.uk. You may also complete
your request on-line via the Elsevier homepage (http://elsevier.com), by selecting “Customer Support” and then
“Obtaining Permissions.”
Waring, Richard H.
Forest ecosystems analysis at multiple scales / Richard H. Waring, Steven W. Running. – 3rd ed.
p. cm.
Includes bibliographical references and index.
ISBN-13: 978-0-12-370605-8 (pbk. : alk. paper) 1. Forest ecology. 2. Forest management. I. Running,
S. W. II. Title.
QH541.5.F6W34 2007
577.3–dc22 2007017124
ISBN: 978-0-12-370605-8
3. Carbon Cycle
I. Introduction 59
II. Photosynthesis 62
III. Autotrophic Respiration 67
IV. Heterotrophic Respiration 71
vii
viii Contents
4. Mineral Cycles
I. Introduction 99
II. Plant Processes Affecting Nutrient Cycling 100
III. Sources of Nutrients 111
IV. Soil and Litter Processes 119
V. Mass Balance and Models of Mineral Cycles 138
VI. Summary 144
Epilogue 345
Bibliography 347
Index 409
Color plates appear between pages 220–221; 260–261; 268–269; and at the back of the book.
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PREFACE TO THE THIRD EDITION
With notification that the second edition of Forest Ecosystems was out of print, we con-
sidered whether a complete revision of the text was warranted. We recognize that the
modeling of forest ecosystem responses has increased significantly in the last decade to
include biodiversity and climatic limitations, but the underlying principles presented in
the second edition appear to remain sound. At the same time, the network of sites that
continuously monitor seasonal and interannual variation in CO2 and water vapor exchange
has grown. By combining data from many sites, new generalities have emerged that should
be shared. Also, we wished to illustrate that forest disturbance and recovery can be more
accurately documented than was previously possible. Finally, with the publication of the
2007 reports by the Intergovernmental Panel on Climate Change, we recognize the press-
ing need to provide background material to policy makers charged with designing policies
that reduce carbon emissions and perpetuate healthy forests in an unstable climate with
new mixtures of species.
In our decision to publish a third edition of the textbook, we have corrected errors in
the previous edition, updated color plates, and added Chapter 10 that focuses on new
information. Specifically, we document in the added chapter how climatic change has
already affected forests, offer insights gained from an expanded network of eddy-flux sites,
and provide evidence of improvements in remote sensing technology. To keep up with the
expanded role that remote sensing and modeling will play in predicting and monitoring
the effects of future policies, we provide a web site reference to replace the compact disc
available with the previous edition.
R. H. W.
S. W. R.
xi
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PREFACE TO THE SECOND EDITION
The first edition of Forest Ecosystems: Concepts and Management, with my colleague
William Schlesinger of Duke University, was published in 1985. At that time, most of the
information on forest ecosystems consisted of mass balance analyses conducted on stands
or small watersheds for periods up to one year. Few simulation models were available,
and those that could be tested were largely restricted to predictions of streamflow. Today,
new methods and new models provide a much wider basis for extrapolation, in space as
well as time. In 1991 and again in 1997, William Schlesinger demonstrated his unique
abilities to synthesize and expand our understanding of terrestrial and aquatic ecosystems
by publishing Biogeochemistry: An Analysis of Global Change.
The opportunity to expand the scope of analysis of forest ecosystems was clear. Such
an expansion, however, required new techniques and experience beyond those possessed
by either author of the first edition. With my colleague’s support, I sought a new coauthor
with experience that extended to the global scale. The person with the most noteworthy
experience in scaling the analysis of forest ecosystems was Steven Running at the Uni-
versity of Montana. To my pleasure, he agreed to join me in the endeavor of writing a
major revision of the first edition that emphasized quantitative modeling and extrapolations
across large spatial and time scales.
A broadened perspective of management is essential today. The pressing issues include
regional and global analyses of biodiversity, changes in climatic cycles, implications of
wide-scale pollution, and the possibility of fire, floods, insect out-breaks, and other major
disturbances that extend beyond the limits of political boundaries. Organizing the princi-
ples and providing examples for expanding the horizon of ecosystem analyses were the
challenges in writing the new edition. From our own experience in teaching graduate and
undergraduate courses, we recognize the difficulty in presenting material that crosses many
fields, but the success of expanded integration and problem analysis lies in acquiring new
methods and concepts. To that end we have made an attempt to define terms and to explain
concepts in a variety of ways by providing equations, graphs, and tabular examples.
Many critical facets of ecosystem behavior, as well as future changes in the environ-
ment, remain unknown. Perhaps the best we can do is to distinguish those processes that
have a firm basis for analysis from those that require more research. The search for prin-
ciples that scale, matched with appropriate methods, will be required, regardless of the
quest. We hope that students, faculty, research scientists, and managers of natural resources
will gain confidence in their abilities to predict and to monitor the implications of various
changes. We encourage concern about the long-term implications of policies, in the hopes
that the alternatives considered will sustain ecosystem processes to which many organisms
contribute, and on which all life depends.
R. H. Waring
xiii
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PREFACE TO THE FIRST EDITION
Our challenge and reasons for writing this book are to share an emerging insight that there
are key linkages between the processes that operate in forests. We emphasize forests in
this book because we know them best and because their long life permits us to evaluate
the effects of periodic disturbance more readily. Our examples, most often, are drawn from
simple cases in which principles are more easily seen and explained. We believe, however,
the principles apply widely, as we show in predicting transpiration and other hydrologic
properties for a variety of forests in differing climatic settings.
In many cases, scientists cannot accurately predict the effects of acid rain, fire suppres-
sion, short-rotation timber harvest, or increasing carbon dioxide levels in the atmosphere
on forests or other ecosystems. We believe a diagnostic approach linking a variety of
processes is warranted and that with carefully designed experiments the mysteries will
unravel.
We have striven to provide a cosmopolitan flavor to the book. Because most experi-
mental work has been focused on rather simple systems, our examples are drawn mainly
from temperate and boreal forests. However, the same processes operate in more complex
forests, as references denote. The book is written for upper-level students with some
background in general ecology, inorganic chemistry, physics, and plant physiology. We
hope that specialists will see new implications to their work and be encouraged to develop
integrative experiments. Managers of forest resources (wood products, wildlife, and water)
will find explanations for some of their observations and predictions of the effects of
various management policies.
We owe a debt to earlier studies of ecosystems, particularly those sponsored by the
National Science Foundation in the 1970s as part of the International Biological Program,
which established a group of five major ecosystem programs in the United States, in addi-
tion to earlier work at Hubbard Brook in New Hampshire. For almost a decade, balance
sheets were constructed describing how carbon, water, and minerals are stored or trans-
ported in a variety of forest, grassland, desert, tundra, and aquatic systems.
Much of the basic information has been published in books and other periodicals. A
summary of the international program with listings of data from all woodland sites
appeared in 1981, edited by D. E. Reichle.1 Regional efforts at synthesis have also been
made for the other biomes. These references, as well as the open literature, provide a
description of how forest systems operate.
Few of the research programs, however, involved experiments that evaluated linkages
between major processes. The influence of fire, erosion, wind storms, and epidemic out-
breaks of insects or disease organisms could not be rigorously evaluated until a benchmark
1
Reichle, D. E. (1981). “Dynamic Properties of Forest Ecosystmes.” Int. Biol. Programme No. 23, Cam-
bridge Univ. Press, London and New York.
xv
xvi Preface to the First Edition
for rates of normal processes had been established. The foundation was laid for critical
experiments that could test hypotheses involving how and why ecosystems respond to
periodic disturbances of various kinds. We propose that integrated experiments based on
ecosystem-level insight can provide answers to managers. Whether this is the case, as we
emphasize in interpreting the probability of disturbance in forests, awaits future tests.
R. H. W.
W. H. S.
ACKNOWLEDGMENTS
R. H. W.
xvii
xviii Acknowledgments
seemed to have little in common. In addition, I thank Dan Fagre, Glacier National Park,
for funding all work related to the GNP. My Numerical Terradynamic Simulation Group
has had a stream of excellent students who have continued to advance the development
of principles that apply to regional analyses, including Joe Coughlan, George Riggs, Rob
Kremer, Ronni Korol, E. Ray Hunt, Kevin Ryan, Daolan Zheng, Lars Pierce, Joe White,
Peter Thornton, John Kimball, Kathy Hibbard, Galina Churkina, and Mike White. A
number of them built new simulations and images specifically for Chapters 7–9 of this
textbook. Joe Glassy and Saxon Holbrook provided an advanced computing environment
for the laboratory.
Finally, Connie, Trina, and Emily have endured many nights with a husband and father
on the road, rather than at home helping them.
S. W. R.
COMPANION WEB SITE INFORMATION
The companion Web site for this book can be found at:
www.books.elsevier.com/companions/9780123706058
This site contains a link to the authors’ site which contains modeling software, tutorials,
and video clips.
Random documents with unrelated
content Scribd suggests to you:
substance is a true bituminous coal, containing more bitumen than is
found in any other variety. Polished sections of the compact masses
exhibit the peculiar structure of coniferous trees, and prove that the
coal was derived from a species allied to the American Fir.”64 A
similar phenomenon was observed by Doctor Dieffenbach in the
Chathain Islands. In the same bed of peat he was able distinctly to
trace a gradual transition from pure vegetable matter to a mineral
substantially identical with common coal.65
But though Peat may thus, as it should seem, pass directly into pure
Coal, there are many cases in which it first assumes a more
imperfect form, known under the name of Lignite. This substance is
described as of a brownish color, “soft and mellow in consistence
when freshly quarried, but becoming brittle by exposure, the
fracture following the direction of the fibre of the wood.”66 It clearly
occupies an intermediate position between Peat and Coal. Like the
former, it still exhibits the stems and woody fibre of the plants from
which it is derived, very little altered in their structure; while on the
other hand it is already beginning to acquire some of the consistency
and density of Coal; to which also it approaches much more closely
in its chemical composition. It should be remembered, moreover,
that Lignite does not designate a substance of a fixed, invariable
character. On the contrary, under the one general name are
comprised a definite number of varieties, leading from one extreme
to the other by a series of almost insensible gradations; the extreme
variety on one side being scarcely distinguishable from Peat, while
the extreme variety on the other is practically identical with ordinary
Coal. It can hardly be doubted, therefore, that Coal must have the
same origin as Lignite, while it is at least equally certain that Lignite
has been derived from Peat; and we have already seen what
overwhelming evidence may be adduced to show that the origin of
Peat is to be sought for in the sunken swamps and forests of a long
past age.
Not less than thirty such trees, some of them four or five feet in
diameter, and all incrusted with Coal, were laid bare a short time
since, in a Colliery near Newcastle, within an area of fifty yards
square. “In 1830,” writes Sir Charles Lyell, “a slanting trunk was
exposed in Craigleith quarry, near Edinburgh, the total length of
which exceeded sixty feet. Its diameter at the top was about seven
inches, and near the base, it measured five feet in its greater, and
two feet in its lesser, width. The bark was converted into a thin
coating of the purest and finest Coal.” Again, “in South Staffordshire,
a seam of Coal was laid bare in the year 1844, in what is called an
open work at Parkfield Colliery, near Wolverhampton. In the space of
about a quarter of an acre, the stumps of no less than seventy-three
trees, with their roots attached, appeared, some of them more than
eight feet in circumference. The trunks, broken off close to the root,
were lying prostrate in every direction, often crossing each other.
One of them measured fifteen, another thirty feet in length, and
others less. They were invariably flattened to the thickness of one or
two inches, and converted into Coal. Their roots formed part of a
stratum of Coal ten inches thick, which rested on a layer of clay two
inches thick, below which was a second forest resting on a two-foot
seam of Coal. Five feet below this again was a third forest, with
large stumps of Lepidodendra, Calamites, and other trees.”67
Our design from the beginning was to consider the points of contact
between Geology and Revelation; to examine the relations that exist
between these two departments of knowledge,—one resting upon
reason and observation, the other given to us from Heaven; and to
inquire how far it may be possible to adopt the conclusions of the
former, while we adhere, at the same time, with unswerving fidelity,
to the unchangeable truths of the latter. With this end in view, we
proceeded at once to sketch out the more prominent features of
Geological theory; not the particular theory of one writer, or of one
school, but that more general theory which is adopted by all writers,
and prevails in every school. This theory, we were all well aware, is
in many points widely at variance with the common notions of
sensible and even well-informed men who have not devoted much
attention to the study of Physical Science. And it occurred to us that,
possibly, many of our readers might be disposed to cut the
controversy short by rejecting, in a summary way, the whole system
of Geology, and treating it as an empty shadow or an idle dream.
This, we were convinced, would be a mistaken and mischievous
course. Geology is not a house of cards that it may be blown down
by a breath. It is a hypothesis, a theory, if you will; but no one can
in fairness deny that behind this theory there are facts,—
unexpected, startling, significant facts; that these facts, when
considered in their relation to one another, when illustrated by the
present phenomena of Nature, and skilfully grouped together, as
they have been by able men, disclose certain general truths, and
suggest certain arguments, which do seem to point in the direction
of those conclusions at which Geologists have arrived.
When the word Fossil was first introduced into the English language,
it was employed to designate, as the etymology suggests, whatever
is dug out of the earth.69 But it is now generally used in a much
more restricted sense, being applied only to the remains of plants
and animals embedded in the Crust of the Earth and there preserved
by natural causes. When we speak of remains, we must be
understood to include even those seemingly transient impressions,
such as foot-prints in the sand, which having been made permanent
by accidental circumstances, and thus engraved, as it were, on the
archives of Nature, now bear witness to the former existence of
organic life.
Now in every part of the world where the Stratified Rocks have been
laid open to view, remains of this kind are found scattered on all
sides in the most profuse abundance. In Europe, in America, in
Australia, in the frozen wastes of Siberia, in the countless islands
scattered over the waters of the Pacific, there is scarcely a single
formation, from the lowest in the series to the highest, that, when it
is fairly explored, does not yield up vast stores of shells, together
with bones and teeth, nay, sometimes whole skeletons of animals;
also fragments of wood, impressions of leaves, and other organic
substances.
We will suppose, then, that the visitor has gratified his sense of
wonder in gazing at the larger and more striking forms, few in
number, that rise up prominently before him, and seem to stare at
him in return from their hollow sockets: he must next turn his
attention to the cases that stand against the walls, and to the
cabinets that stretch along the galleries in distant perspective. Let
him survey that multitude of bones of every shape and size, and
those countless legions of shells, and then try to realize to his mind
what a profusion and variety of animal life are here represented. And
yet he must remember that this is but a single collection. There are
thousands of others, public and private, scattered over England,
France, Germany, Italy, and beyond the Atlantic, on the continent of
America, and even in Australia; all of which have been furnished
from a few isolated spots,—scarcely more than specks on the
surface of the Globe,—where the interior of the Earth’s Crust has
chanced to be laid open to the explorations of the Geologist.
Lastly, before he leaves this splendid gallery, let him take a passing
glance at the Organic Remains of the vegetable world. There is no
mistaking the forms here presented to his view. He will recognize at
once the massive and lofty trunks of forest trees with their spreading
branches; the tender foliage of the lesser plants; and, in particular,
the graceful fern, which cannot fail to attract his eye by its unrivalled
luxuriance. But if the forms are familiar, how strange is the
substance, of this ancient vegetation! The forest tree has been
turned into sandstone; many of the plants are of the hardest flint;
and the rich green of the fern has given place to the jet black color
of coal. Let him take a magnifying glass and scrutinize the internal
structure of these mineralized remains; for the more closely they are
examined the more wonderful do they appear. He can observe
without difficulty their minute cells and fibres, the exact counterpart
of those which may be seen in the plants that are now growing upon
the earth; he may detect the little seed-vessels on the under surface
of the coaly fern; nay, if he gets a polished transverse section of the
sandstone tree, he may count the rings that mark its annual growth,
and tell the age it attained in its primeval forest.
Fig. 13.—Fossil Wood, from the Carboniferous Limestone of Mayo,
showing the rings of Annual Growth.
CHAPTER XI.
FOSSIL REMAINS—THE EXPLORATION.
Near the town of Aix, the ancient capital of Provence, in the south of
France, is a group of strata, consisting chiefly of Conglomerate, Marl,
Gypsum, and Limestone, which has earned for itself no small fame in
the annals of Geology. Besides many curious relics of an extinct
vegetation, these strata yield also an abundance of Fossil Insects,
which emerge from the rocky bed in which they have slept for ages,
with a surprising freshness and a life-like reality. But the quarries of
Aix, like those of Monte Bolca, are chiefly famous for their Fossil
Fish. And in this case, too, as in the former, it would seem as if vast
multitudes had suddenly perished together from some mysterious
cause, and were then as suddenly entombed. They exhibit no mark
of mechanical violence: and yet they are found, not unfrequently,
crowded together as closely as they can fit, in every variety of
position, on the same slab of limestone. A good example of such a
block is represented in our woodcut.
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