BIOMES OF THE EARTH

TEMPERATE
FORESTS
Michael Allaby

Illustrations by
Richard Garratt
Temperate Forests

Copyright © 2006 by Michael Allaby

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Allaby, Michael
Temperate forests / Michael Allaby; illustrations by Richard Garratt.
p. cm.—(Biomes of the Earth)
Includes bibliographical references and index.
ISBN 0-8160-5321-9
1. Forest ecology—Juvenile literature. 2. Forests and forestry—Juvenile literature. I. Garratt,
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Illustrations by Richard Garratt
Photo research by Elizabeth H. Oakes

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 From Richard Garratt:
To Chantal, who has lightened my darkness
 CONTENTS
Preface ix
Acknowledgments xiii
Introduction: What is a temperate forest? xv

CHAPTER 1
GEOGRAPHY OF TEMPERATE
FORESTS 1
Types of temperate forest 1
Why it rains more on mountainsides 3
Where do temperate forests grow? 4
How temperate forests developed and how long
they have existed 7
Forest expansion since the last ice age 10
Ancient forests that survive to this day 14
What trees can tell us about climates of the past 17

CHAPTER 2
GEOLOGY OF
TEMPERATE FORESTS 19
Plate tectonics 19
Alfred Lothar Wegener and continental drift 21
Related temperate-forest trees that occur
in widely separated places 23
Soils of temperate forests 25
How soils are classified 28
How water moves through a temperate forest 29

CHAPTER 3
CLIMATE OF TEMPERATE FORESTS 34
Why there are seasons 34
How oceans affect climates 37
General circulation of the atmosphere 38
Air masses and fronts 41
Vilhelm Bjerknes and the Bergen School 43
Jet streams and depressions 45
Blocking 49
Continental and maritime climates 52
How climates are classified 54
Why clouds form 56
Adiabatic cooling and warming 58
Humidity 59
How clouds are classified 60
Rain, snow, and hail 61
Ice storms 63
Dew and frost 65
Evaporation and transpiration 66
Thunderstorms and gales 69
Lapse rates and stability 70
Forest fires 72

CHAPTER 4
HOW TREES WORK 74
How trees evolved 74
How trees find food 77
Photosynthesis and respiration 79
The nitrogen cycle 80
Photosynthesis 82
How trees reproduce 85
Differences between conifers and flowering plants 87
How seeds disperse 89
What is a flower? 90
How wood forms 94

CHAPTER 5
ECOLOGY OF
TEMPERATE FORESTS 98
How ecological ideas developed 98
Food chains, webs, and pyramids 101
How energy flows through a forest 106
Decomposition and the cycling of nutrients 110
Habitats and niches 112
How trees are adapted to climate 115
Succession and climax 118
Fire, and trees that depend on it 120
Yellowstone and coping with fire 122
Structure of a temperate forest 123
Forest communities 127
Animals that attack trees 130
Diseases and parasites of trees 132
Differences between a natural forest and
a plantation 135

CHAPTER 6
BIODIVERSITY AND
TEMPERATE FORESTS 138
What does biodiversity mean? 138
Carolus Linnaeus and the classifying of plants
and animals 140
Why biodiversity matters 142
The Biodiversity Convention 143
Typical trees of temperate forests 144
Typical animals of temperate forests 148

CHAPTER 7
HISTORY OF
TEMPERATE FORESTS 154
Forest clearance in prehistory 154
Using pollen and beetles to study the past 157
Radiocarbon dating 158
Forests in the days of ancient Greece and Rome 161
How North America nearly lost its forests 163
George Perkins Marsh 164
Forests in folklore 167
History of forestry 169
British geography 172

CHAPTER 8
USES OF TEMPERATE FORESTS 175
Renewable and nonrenewable resources 175
Forests for fuel 178
Logging and timber 181
 Charcoal and how it is made 182
Chipboard, fiberboard, and paper 185
Veneers and plywood 188
Small wood and its uses 190

CHAPTER 9
MANAGING
TEMPERATE FORESTS 193
Traditional management 193
Coppicing and pollarding 195
Park woodland 196
Plantation forestry 199
What are lichens? 202
Pest control 203
Modern forestry 207

CHAPTER 10
HEALTH OF
TEMPERATE FORESTS 210
Are the temperate forests disappearing? 210
Acid rain 214
Forest clearance and climate 217
How will forests respond to global warming? 220
The greenhouse effect 222
Forest clearance and soil erosion 225
Sustainable forest management 227

CONCLUSION: WHAT FUTURE

FOR TEMPERATE FORESTS? 230

SI units and conversions 233

Soil classification: Orders of the soil taxonomy 237
Geologic time scale 241
Glossary 243
Bibliography and further reading 259
Index 261
 PREFACE
Earth is a remarkable planet. There is nowhere else in our
solar system where life can survive in such a great diversity of
forms. As far as we can currently tell, our planet is unique.
Isolated in the barren emptiness of space, here on Earth we
are surrounded by a remarkable range of living things, from
the bacteria that inhabit the soil to the great whales that
migrate through the oceans, from the giant redwood trees of
the Pacific forests to the mosses that grow on urban side-
walks. In a desolate universe, Earth teems with life in a bewil-
dering variety of forms.
One of the most exciting things about the Earth is the rich
pattern of plant and animal communities that exists over its
surface. The hot, wet conditions of the equatorial regions
support dense rain forests with tall canopies occupied by a
wealth of animals, some of which may never touch the
ground. The cold, bleak conditions of the polar regions, on
the other hand, sustain a much lower variety of species of
plants and animals, but those that do survive under such
harsh conditions have remarkable adaptations to their test-
ing environment. Between these two extremes lie many
other types of complex communities, each well suited to the
particular conditions of climate prevailing in its region.
Scientists call these communities biomes.
The different biomes of the world have much in common
with one another. Each has a plant component, which is
responsible for trapping the energy of the Sun and making it
available to the other members of the community. Each has
grazing animals, both large and small, that take advantage of
the store of energy found within the bodies of plants. Then
come the predators, ranging from tiny spiders that feed upon
even smaller insects to tigers, eagles, and polar bears that sur-
vive by preying upon large animals. All of these living things

IX
X TEMPERATE FORESTS

form a complicated network of feeding interactions, and, at

the base of the system, microbes in the soil are ready to con-
sume the energy-rich plant litter or dead animal flesh that
remains. The biome, then, is an integrated unit within which
each species plays its particular role.
This set of books aims to outline the main features of each
of the Earth’s major biomes. The biomes covered include the
tundra habitats of polar regions and high mountains, the
taiga (boreal forest) and temperate forests of somewhat
warmer lands, the grasslands of the prairies and the tropical
savanna, the deserts of the world’s most arid locations, and
the tropical forests of the equatorial regions. The wetlands of
the world, together with river and lake habitats, do not lie
neatly in climatic zones over the surface of the Earth but are
scattered over the land. And the oceans are an exception to
every rule. Massive in their extent, they form an intercon-
necting body of water extending down into unexplored
depths, gently moved by global currents.
Humans have had an immense impact on the environ-
ment of the Earth over the past 10,000 years since the last Ice
Age. There is no biome that remains unaffected by the pres-
ence of the human species. Indeed, we have created our own
biome in the form of agricultural and urban lands, where
people dwell in greatest densities. The farms and cities of the
Earth have their own distinctive climates and natural history,
so they can be regarded as a kind of artificial biome that peo-
ple have created, and they are considered as a separate biome
in this set.
Each biome is the subject of a separate volume. Each richly
illustrated book describes the global distribution, the climate,
the rocks and soils, the plants and animals, the history, and
the environmental problems found within each biome.
Together, the set provides students with a sound basis for
understanding the wealth of the Earth’s biodiversity, the fac-
tors that influence it, and the future dangers that face the
planet and our species.
Is there any practical value in studying the biomes of the
Earth? Perhaps the most compelling reason to understand
the way in which biomes function is to enable us to conserve
their rich biological resources. The world’s productivity is the
 PREFACE XI

basis of the human food supply. The world’s biodiversity

holds a wealth of unknown treasures, sources of drugs and
medicines that will help to improve the quality of life. Above
all, the world’s biomes are a constant source of wonder,
excitement, recreation, and inspiration that feed not only
our bodies but also our minds and spirits. These books aim to
provide the information about biomes that readers need in
order to understand their function, draw upon their
resources, and, most of all, enjoy their diversity.
 ACKNOWLEDGMENTS
Richard Garratt drew all of the diagrams and maps that
appear in this book. Richard and I have been working togeth-
er for many years in a collaboration that succeeds because
Richard has a genius for translating the weird electronic
squiggles I send him into clear, simple artwork of the highest
quality. As always, I am grateful to him for all his hard work.
I also wish to thank Elizabeth Oakes for her fine work as a
photo researcher.
I must thank Frank K. Darmstadt, Executive Editor, at
Chelsea House. Frank shaped this series of books and guided
them through all the stages of their development. His
encouragement, patience, and good humor have been
immensely valuable.
I am especially grateful to Dorothy Cummings, project edi-
tor. Her close attention to detail sharpened explanations that
had been vague, corrected my mistakes and inconsistencies,
and identified places where I repeated myself. And occasion-
ally Dorothy was able to perform the most important service
of all: She intervened in time to stop me making a fool of
myself. No author could ask for more. This is a much better
book than it would have been without her hard work and
dedication.
Michael Allaby
Tighnabruaich
Argyll
Scotland
www.michaelallaby.com

XIII
 INTRODUCTION
What is a temperate forest?
Long, long ago a sheet of ice thousands of feet thick covered
North America from southern Alaska to a line running
roughly from Seattle, Washington, to Boston, Massachusetts.
Ice also covered most of northern Europe and the sea was
frozen, so a person could have walked from America to
Greenland and from there to Europe. To the south of the ice
sheets, the land was covered in tough grasses, sedges, and
small shrubs—tundra vegetation like that of northern
Canada today. The weather was cold, dry, and windy, and the
landscape was bleak.
Then, starting about 14,000 years ago in some places and
12,000 years ago in others, the air grew warmer and the ice
began to melt. As it melted, the edges of the ice sheets retreat-
ed northward, leaving bare ground where no plant had
grown for tens of thousands of years. Snow still blanketed the
surface each winter, but in summer the dark-colored rocks
absorbed the warmth of the sunshine and the warmth gradu-
ally penetrated deeper, to where the soil was still frozen. The
frozen soil—called “permafrost”—slowly thawed.
Water from the melting ice filled rivers. In summer, when
the snows of winter melted, water soaked into the ground.
Tiny plant seeds and the spores of ferns, mosses, and fungi
arrived, carried on the wind from lands far to the south. They
had always arrived in this way, but in the past they had per-
ished when they fell onto the barren ice. Now there was just
enough warmth and moisture for some of them to germi-
nate, and enough nourishment for some of the young plants
to survive and grow. Here and there, patches of green
appeared between the boulders and the sheets of bare rock
that had been scraped smooth by the ice. Each summer the
green returned, and each year it covered a bigger area.

XV
XVI TEMPERATE FORESTS

At first it was the tundra that spread northward, following

the edges of the retreating ice. Then, as the ground continued
to grow warmer and the tundra plants helped to build the
soil, other plants began to arrive. There were shrubs and then
trees that stood tall and shaded the ground. Little by little,
many of the tundra plants disappeared, unable to tolerate the
shade or to compete successfully with the newcomers for
moisture and nutrients. The landscape changed, as tundra
gave way to forest. It was a gradual process that took several
thousand years, but eventually it produced vast forests of
birch, larch, spruce, fir, and pine trees.
A forest is an area of land where trees are the most conspic-
uous plants. The word also means a large number of trees
that are growing so closely together that the leaves of one
overlap those of its neighbors, and together the trees shade at
least 60 percent of the surface. Coniferous trees, such as
birch, larch, spruce, fir, and pine, have leaves that are
reduced to needles or tiny scales. These contrast with the
broad leaves of such trees as oak and maple.
The change was not permanent, however, for the climate
continued to grow warmer and the ice maintained its retreat.
Conditions suited other trees, such as oak, ash, and elm, and
these began to predominate in the forest. The original forest
still survives in the north. Today it extends as a belt across
Alaska, Canada, Scandinavia, and Siberia. It is known as the
boreal forest—“boreal” simply means northern—or by its
Russian name, the taiga.
South of the taiga, the natural forest was quite different in
character. Although the climate was now much warmer than
it had been, the ground still froze in winter and when the
ground is frozen, plant roots cannot find the moisture they
need. The newly arrived trees coped with the dry soils of win-
ter by shedding their leaves and effectively shutting down
until spring. A tree or shrub that sheds all of its leaves at the
same time is said to be deciduous.
Deciduous trees have delicate leaves that absorb sunlight
efficiently all summer, but as the days begin to grow shorter,
the trees withdraw some of their chemical contents and cease
supplying others. The resulting chemical changes alter the
color of the leaves. For a few weeks before the winds of the
 INTRODUCTION XVII

FPO
Fig. 1

late fall strip their branches and fill the air with flying, twist- A trail through a
ing, circling leaves, the trees are a riot of red, orange, yellow, temperate forest in late
and brown. summer, just as the
Early spring is another colorful time. As the soil tempera- leaves are starting to
ture starts to rise, there is a brief interlude before the leaf buds change color (Courtesy
open when the Sun shines directly onto the ground. During of Fogstock)
this short interval, smaller plants of the forest floor seize
their opportunity to grow, flower, and produce seed. Then
the trees burst into bright green leaf and in a week or two the
forest floor is shaded from the sunlight.
The spring flowers, bright green young leaves, and spectac-
ular fall colors are typical of the broad-leaved deciduous
forests of North America, Europe, and eastern Asia. Such
forests grow in regions where the weather is neither so hot as
it is closer to the equator nor so cold as it is closer to the
XVIII TEMPERATE FORESTS

North or South Pole. The climate is temperate and these are

temperate forests.
Farther from the equator, plants do not need to survive the
winter cold, for the ground seldom freezes for more than a
few days at a time, but the rainfall is intense in one part of
the year and sparse during a dry season. The trees of these
forests do not shed all of their leaves at the same time of year,
but instead produce leaves that last longer. They are tough
and leathery, with a waxy outer coat that makes them shine.
Trees of this type—holly and live oak are typical examples—
are broad-leaved, but evergreen. Evergreen trees do shed their
leaves, but only when they are worn out, so the tree is never
bare.
The ice sheets have now retreated from North America,
Europe, and Asia, and temperate forest—mainly deciduous
but evergreen in some places—is the natural vegetation over
a large area. Any type of vegetation occupying a substantial
part of the Earth’s surface comprises a biome.
Although the temperate forest constitutes a biome, this
does not mean it is everywhere the same. The tree types vary
greatly. In some places all the trees are broad-leaved and
deciduous. In others the deciduous trees are mixed with
broad-leaved evergreen trees—where oak and holly grow side
by side, for example. There are many forests in which broad-
leaved deciduous trees grow alongside coniferous trees such
as evergreen pines and deciduous larches, and some temper-
ate rain forests are dominated by coniferous species.
A climate that suits a temperate forest is also suitable for
farming. Much of the forest that once covered the eastern
United States and western Europe was cleared long ago to
provide farmland. It is only here and there that patches of
the original forest can still be found. The forest will not dis-
appear, however. Modern farms produce much more food
than the farms of earlier times, so people may feed them-
selves from a smaller area of land. We can allow some of the
less fertile farmland to return to forest. It will not be identical
to the original primeval forest, but if it is planted with the
original species, in time it will become very similar.
 CHAPTER 1

GEOGRAPHY OF
TEMPERATE FORESTS
Types of temperate forest
Broad-leaved deciduous trees dominate the most extensive
type of temperate forest. Often called “summer deciduous
forest,” this is the forest that produces the dazzling colors of
spring and fall. At one time it covered about 3 million square
miles (7.8 million km2) of eastern North America and Europe.
It was the forest our ancestors encountered when first they
migrated into these regions—and the forest they cleared
away, either to hunt game more easily or to make way for
their livestock and farm crops.
A different type of temperate forest grows along the west-
ern coasts of Chile and of South Island, New Zealand. These
are regions with a maritime climate (see “Continental and
maritime climates” on page 52). Winters are wet and mild,
with temperatures seldom remaining below freezing for very
long. Summers are warm and usually drier than the winters,
and fog is very common. Temperatures seldom rise above
80°F (27°C). Although northern Chile is one of the driest
places in the world, the region south of 37°S is one of the
wettest, with an average annual rainfall ranging from about
102 inches (2,591 mm) on the coast to more than 200 inches
(5,080 mm) in the mountains. South Island, New Zealand,
has a generally moist climate, but the western coastal belt
receives as much rain as southern Chile.
The forests here are rain forests that grow in a temperate
climate: temperate rain forests. The most conspicuous trees in
the Chilean forest are southern beeches (Nothofagus species),
Patagonian cypress (Fitzroya cupressoides), and Chile pine
or monkey puzzle (Araucaria araucana). Southern beeches
are also common in the New Zealand forest (see “Related
temperate-forest trees that occur in widely separated places”
on page 23). In the north the beeches are mixed with kauri

1
2 TEMPERATE FORESTS

pines (Agathis australis). There are also small areas of temper-

ate rain forest in southern China and Japan, mostly dominat-
ed by evergreen oaks (Quercus species) and magnolias
(Magnolia species), and in western Scotland, where the native
trees are birch (Betula species), oak, hazel (Corylus avellana),
and holly (Ilex aquifolium).
The most magnificent trees of any temperate rain forest are
found in North America, however. An intermittent coastal
belt that is nowhere more than about 125 miles (200 km)
wide, extending from California to southern Alaska, is home
to Douglas fir, western hemlock, western red cedar, Sitka
spruce, giant sequoias, and coastal redwoods. These are some
of the biggest trees in the world.
The North American forest differs from the temperate rain
forests found in other parts of the world in being dominated
by coniferous trees, rather than broad-leaved species. There
are probably several reasons for this. In the first place, the
summers are fairly dry. On average, less than three inches (76
mm) of rain falls in Vancouver, British Columbia, between
the end of June and beginning of September. Seattle,
Washington, is even drier, with only six inches (152 mm) of
rain from the beginning of May until the end of September.
The lack of rain in the middle of summer allows the ground
to dry and this may impose an intolerable stress on broad-
leaved trees. Evergreen coniferous trees, with needles rather
than leaves, are better able to thrive in such conditions. The
dry period has a second consequence: fire. The redwoods and
other trees of these forests can survive fires.
Western hemlock (Tsuga heterophylla), western red cedar
(Thuja plicata), and Sitka spruce (Picea sitchensis) grow to a
height of 100–200 feet (30–60 m). Douglas fir (Pseudotsuga
menziesii, also called Oregon fir and Oregon pine) and the
giant redwood (Sequoiadendron giganteum, also called the
Sierra redwood, big tree, and wellingtonia) can reach a height
of 320 feet (98 m). These are tall trees, but the coast redwood
(Sequoia sempervirens) can be as much as 390 feet (119 m)
tall—the height of a 28-story apartment building. In many
places the forest canopy—the topmost level of the tree
crowns—is 200–230 feet (60–70 m) above ground level.
The General Sherman tree, a giant redwood growing in the
Sequoia National Park, is believed to be the most massive tree
 GEOGRAPHY OF TEMPERATE FORESTS 3

in the world. It is 274.9 feet (83.8 m) tall and 27 feet (8 m) in

diameter measured eight feet (2 m) above ground level. Its
circumference at ground level is 102.6 feet (31.1 m) and its
crown is an average 106.5 feet (32.5 m) across. Its trunk
weighs an estimated 1,385 tons (1,258 tonnes). The General
Sherman is thought to be 2,000–2,200 years old and it is still
growing vigorously.
It is not true, however, that the General Sherman is the
largest living organism in the world. The biggest single
organism ever discovered is a honey fungus (Armillaria ostoy-
ae) living in the soil and among the tree roots in the Malheur
National Forest, in the Blue Mountains of eastern Oregon. It
covers 2,200 acres (890 hectares) and is estimated to be at
least 2,400 years old.

Why it rains more on mountainsides

When air rises, its temperature falls. Warm air can hold more water vapor than cold air
can, so reducing the temperature of the air raises its relative humidity (RH), which is the
amount of water vapor present in the air measured as a percentage of the amount need-
ed to saturate the air at that temperature.
The lifting condensation level is an altitude at which the RH in rising air reaches 100 per-
cent and the air becomes saturated. When air rises above the lifting condensation level, its
water vapor starts condensing to form clouds. When the clouds are big enough, their
droplets or ice crystals merge until they are heavy enough to fall as rain or snow.
Air is forced to rise as it approaches a mountain. The air is dry until it is lifted above the
lifting condensation level; that is, all of the moisture it carries is present as water vapor (a
gas) rather than liquid droplets, so the RH is less than 100 percent. Cloud starts to form at
the lifting condensation level, and precipitation begins to fall on the mountainside. The air
continues to rise, and more cloud forms at higher elevations. This intensifies the precipita-
tion on the mountainside.
When the rising air reaches the top of the mountain it may continue to rise or it may
subside down the opposite (lee) side of the mountain. If it continues rising, eventually it
will lose enough water for precipitation to end, although cloud may extend some distance
downwind. If the air subsides, it will sink below the lifting condensation level once more.
Precipitation will then cease and the cloud will dissipate as its droplets evaporate.
The overall result is that mountainsides receive more precipitation than the low ground
surrounding the mountain.
4 TEMPERATE FORESTS

Temperate forests change in composition and appearance

with increasing elevation. Temperature decreases with
height, and mountains have a wetter climate than the sur-
rounding low-lands (see the sidebar “Why it rains more
on mountainsides” on page 3). On some mountainsides, al-
though not everywhere, the lowland forest gives way to a for-
est of smaller, often stunted trees that are more widely
spaced. The moist conditions favor mosses, ferns, and lichens
and the trees are usually festooned with lichens, giving the
forest an enchanted, magical appearance. It is called krumm-
holz—a German word meaning “crooked wood.”
Higher still, the trees become more widely scattered up to
an elevation beyond which no tree can tolerate the climate.
The timberline is the upper boundary of the forest. The height
is determined by the temperature, so it varies with latitude—
the lower the latitude, the higher the timberline. The timber-
line in the Sierra Nevada is at about 11,500 feet (3,500 m), in
the Canadian Rockies it is at about 6,500 feet (1,980 m), and
in the central Alps of Europe it is at about 6,800 feet (2,000
m). In contrast, the timberline in the mountains of New
Guinea, which are much closer to the equator, is at about
12,600 feet (3,800 m).

Where do temperate forests grow?

The tropic of Cancer is a line of latitude located at 23.5°N.
The tropic of Capricorn is at 23.5°S. The part of the world
lying between these two lines is known nowadays simply as
the Tropics. Two other lines of latitude, at 66.5°N and 66.5°S,
mark the Arctic and Antarctic Circles. The regions between
these latitudes and the North and South Poles are called the
Arctic and the Antarctic.
These were not always their names, however. The Greek
philosopher Aristotle (384–322 B.C.E) was the first person to
divide the world into climatic regions. He called the Tropics
the torrid zone and the regions to the north and south of the
Arctic and Antarctic Circles the frigid zones. Aristotle called the
part of the world lying between these two zones in each hemi-
sphere the temperate zone. We no longer use the terms torrid
zone and frigid zone, but we have kept the name temperate zone.
 GEOGRAPHY OF TEMPERATE FORESTS 5

tropic of Cancer

equator

tropic of Capricorn

broad-leaved/mixed forest
coniferous (boreal) forest

Obviously, temperate forests are found within the temper- Extent of the temperate
ate zone—they are forests that grow in a temperate climate. forest biome. Temperate
This is broadly true, but as the map shows, temperate forests forest is the natural
occur only in certain parts of this zone. vegetation in the areas
There are few areas of temperate forest in the Southern colored pale green,
Hemisphere, and they are quite small. Temperate forest although large parts of
occurs predominantly in the Northern Hemisphere. This is these areas have been
because there is much less land in the Southern Hemisphere converted to farmland.
than there is in the Northern Hemisphere, and much less of
that land lies within the temperate zone. Cape Horn, the
southernmost tip of South America, at 55.78°S, is as far from
the equator as Copenhagen, Denmark, and Edinburgh,
Scotland (at 55.68°N and 55.92°N, respectively). Large areas
of land lie to the north of these cities. At 45.87°S, the city of
Dunedin, in the south of South Island, New Zealand, is the
same distance from the equator as Portland, Oregon, and
Venice, Italy. Temperate forest occurs only in the southern-
6 TEMPERATE FORESTS

most part of South America and in New Zealand. In the

Northern Hemisphere, in contrast, it covers most of the east-
ern half of the United States, Europe as far eastward as the
Black Sea, eastern Russia and China, and Japan. To the north
of the temperate forest there lies a wide belt of coniferous for-
est. This is the boreal forest, or taiga.
Although temperate forest covers a substantial area, it does
not extend throughout the temperate zone, because temper-
ate forest requires a mild, moist climate. Despite its name,
the climate in many parts of the temperate zone is too
extreme.
Most trees find it difficult to survive where hot, dry weath-
er persists for more than about eight months of the year and
the annual rainfall is less than about 16 inches (400 mm).
Temperate forest grows in those areas where the climate is
influenced by the ocean. Air moving inland from the ocean
is moister than air over the interior of a continent, and it
is cooler in summer and warmer in winter (see “Continental
and maritime climates” on page 52). The central plains of
the United States, the interior of Asia, and Australia are too
dry.
Broad-leaved deciduous trees will not grow where there is
an average of fewer than 120 days each year when the tem-
perature is above 50°F (10°C). Coniferous trees can tolerate
much colder conditions, so this temperature requirement
defines the boundary between the temperate broad-leaved
forest and the boreal forest. There are no large areas of com-
parable coniferous forest in the Southern Hemisphere,
because none of the continents projects far enough to the
south.
Many forests have sharply defined edges, but these are
unnatural. Land that was once forested has been cleared of its
trees and put to other uses—usually agriculture—producing a
clear boundary between the forest and the deforested area.
Natural forests merge gradually with the adjacent land.
Moving away from the forest, the trees become increasingly
scattered until the landscape is treeless. Where summers are
long, hot, and dry, the forest gives way to scrub and finally to
desert. Where summers are wet but the winters are dry, the
forest gives way to grassland.
 GEOGRAPHY OF TEMPERATE FORESTS 7

How temperate forests developed and how

long they have existed
There are many species of broad-leaved deciduous trees to be
found in temperate forests. They vary in size and shape and
in their leaves, but all of them share certain features. The
most important of these is that they produce flowers (see
chapter 4). In common with many of the herbs that grow on
the forest floor, the grasses and wildflowers that grow in the
fields and by roadsides, and all of the crop plants that farmers
grow, these forest trees are flowering plants. Botanists classify
all of them in the subphylum called Angiospermae, thus sep-
arating them from nonflowering plants, such as the conifer-
ous trees, ferns, and mosses.
Flowers first appeared about 130 million years ago, during
the Cretaceous period, when dinosaurs walked the Earth.

petal

anther stigma

stamen

filament style carpel

ovary

sepal

receptacle

Structure of a flower.
The anther and filament,
together comprising the
bract stamen, are the male
parts of the flower; the
female parts are the
stigma, style, and ovary,
together comprising
the carpel.
8 TEMPERATE FORESTS

Climates everywhere were warmer and wetter then than they

are today, and the first flowering plants lived under conditions
similar to those of the present-day Tropics. The very earliest of
them were water lilies. These were soon joined by buttercups
and the first flowering tree: a magnolia. The magnolia family
(Magnoliaceae) also includes a number of other species,
including the tulip tree or tulip poplar (Liriodendron tulipifera).
Members of the Magnoliaceae occur naturally in North and
South America and in Asia (see “Related temperate-forest trees
that occur in widely separated places” on page 23).
Water lilies, buttercups, magnolias, the tulip tree, and all
the other members of this group of plants produce very dis-
tinctive flowers (see the diagram). These are large, have many
petals and sepals—called tepals because they are similar in
shape and size—and also many carpels and stamens with the
stamens arranged in a spiral. The flowers are symmetrical and
bisexual—they contain both male and female organs (that is,
the stamens and carpels). Scientists consider these features to
be primitive, in the sense that in more advanced flowers,
which evolved later, the features are modified and some of
the flower parts are often missing.
Flowers are big and showy in order to attract pollinating
insects. Nonflowering plants must rely on the wind or water
to bring male sperm and female eggs together. Insect pollina-
tion is much more reliable and, not surprisingly, flowering
plants spread rapidly and evolved into the many species that
exist today. Until then forests had consisted of giant tree
ferns, conifers, and other nonflowering plants. By the end of
the Cretaceous period, 65 million years ago, flowering trees
dominated most forests and flowering herbs grew on the for-
est floor beneath them.
Many of the most ancient species of flowering trees still
exist, little changed. Both magnolias and tulip trees grow nat-
urally in the forests of the southeastern United States, but
although these plants are natives of America and Asia, they
are just as familiar to Europeans and Australians. They have
been planted all over the world. Their spectacular flowers
make them popular ornamental plants, grown in gardens
and parks, and tulip trees are also cultivated for their timber,
known as yellow poplar or canary whitewood.
 GEOGRAPHY OF TEMPERATE FORESTS 9

Forests change over long periods. New tree species arrive,

old ones die out, and although the forest continues, its com-
position does not remain the same. If the climate becomes
warmer or cooler, wetter or drier, some trees suffer and others
thrive. Forests respond to changes in their environment.
Over the last few centuries, people have exerted an entire-
ly new force for change: They have introduced new tree
species. Magnolias and tulip trees are now grown all over the
world, but they remain confined to gardens and parks. Other
introduced species have escaped and become naturalized in
their new surroundings. Some of them have then invaded
the natural forest and altered it.
North American old-growth forests have proved resistant
to invasion by introduced trees, but managed forests are
more vulnerable. Perhaps the most invasive of all tree
species, however, is the European sycamore or great maple
(Acer pseudoplatanus)—a type of maple and quite unrelated
to American sycamores, which are plane trees (Platanus
species). The European sycamore was introduced into
England sometime prior to 1500, but it was not planted
widely until the 18th century. It is a handsome tree, highly
valued during the period when large landscaped parks,
designed to display individual trees and other features, were
fashionable among wealthy British landowners. The tree
also produces valuable timber. It escaped from the parks and
plantations and European sycamore now produces seed
more regularly and prolifically than do most native trees,
and its seeds sprout very readily. Sycamore colonizes dis-
turbed ground and invades neglected woodland, where
there are openings in which it can grow. Most British forests
now contain some sycamore and in many it is the most
abundant tree.
There are other ways in which people can affect the com-
position of a forest. For instance, deer will eat the leaves from
young saplings, killing them. Changes in hunting or culling
policy, or the elimination of animals that hunt deer, can lead
to a rapid increase in the deer population, resulting in
changes in any forest to which they can gain access.
Separately, imported timber can bring disease that escapes
into the forests with devastating effect. Chestnut blight and
10 TEMPERATE FORESTS

Dutch elm disease are modern examples (see “Diseases and

parasites of trees” on page 132).

Forest expansion since the last ice age

No plants of any kind, much less forests, grow in the interiors
of Greenland and Antarctica for the obvious reason that in
these places the ground lies beneath ice that in some places is
more than two miles (3.2 km) thick. If the climate were to
grow warmer and these ice sheets were to melt, after a time
plants would begin to appear. The first arrivals would be tiny
and inconspicuous, surviving in crevices and hollows where
they were sheltered from the freezing and drying wind. Then
grasses would appear, and as water from the melting ice
soaked the ground, sedges would establish themselves in the
wetter places. Eventually, low-growing shrubs would arrive,
along with trees that spread horizontally and were no more
than about two feet (60 cm) tall. These would be followed by
taller trees and finally by forests.
Neither Greenland nor Antarctica is likely to lose its ice
sheet any time soon, but there was a time when the ice
extended much farther than it does now. As the map shows,
during the coldest part of the most recent ice age, the
Laurentide ice sheet covered all of Canada and a large part of
the United States.
There have been many ice ages. Scientists call them glacials
and they are separated by periods of warmer conditions called
interglacials. We are living in an interglacial at present. All
glacials and interglacials have names. These are usually
derived from the places where evidence for them has been
found. Consequently, they have different names in different
places.
The differing names also reflect the fact that the glacials
and interglacials began and ended at slightly different times
in different parts of the world. Our present interglacial breaks
the rule by not being named after a place. It is called the
Holocene, a name that means “entirely new” (from the Greek
holos, “whole” or “entire,” and kainos, “new”). The Holocene
also marks the commencement of the Holocene epoch, a
period of geologic time that follows the Pleistocene epoch,
 GEOGRAPHY OF TEMPERATE FORESTS 11

ice sheet

The Laurentide ice sheet.

The map shows the area
of North America
covered by ice at the
time of its greatest
extent, about 20,000
years ago.

during which ice ages and interglacials followed one another

for about 1.64 million years (see the appendix “Geologic time
scale” on page 241). The sidebar “Holocene, Pleistocene, and
late Pliocene glacials and interglacials, on page 13,” gives the
North American, British, and Northwest European names of
the glacials and interglacials, together with their ages and
durations.
During the Pleistocene ice ages, ice sheets spread to cover
much of Europe and Asia as well as North America. Conditions
were cold enough that they would have covered more of Asia,
but the climate there was too dry for sufficient snow to accu-
mulate. As the map shows, a tongue of ice extended deep into
the interior of the continent from the Pacific coast.
At the start of an ice age, the trees die when the tempera-
ture falls below the minimum they can tolerate. Species by
12 TEMPERATE FORESTS

3
2

The world during an

ice age. During the
coldest part of the
most recent ice age, Ice sheets:
ice sheets covered 1 Cordilleran
much of northwestern 2 Laurentide
Europe and North 3 Greenland
America, and the sea 4 Fennoscandian
was covered by ice.

species, the forests disappear. They survive away from the ice,
where the climate is warmer. Thus as ice ages intensify and
then give way to interglacials, the forests slowly move back
and forth in response to the climatic changes. Remnants of
the forest manage to survive in some places (see “Ancient
forests that survive to this day” on page 14). During ice ages,
these refugia contain the plants that supply seeds from which
the forests regenerate when the ice age ends.
By about 14,000 years ago, the temperature was rising
steadily in North America. The Laurentide ice sheet had start-
ed to retreat and by 12,000 years ago a belt of forest, compris-
ing spruce (Picea species) and larch (Larix species), with some
ash (Fraxinus species) and birch (Betula species), ran from the
Atlantic coast along the southern margin of what are now the
Great Lakes. Broad-leaved deciduous woodland was estab-
lished in Florida. As the ice continued to retreat northward,
 GEOGRAPHY OF TEMPERATE FORESTS 13

the forest followed it. The coniferous forest advanced behind

the retreating ice and the broad-leaved forest advanced
behind the coniferous forest. The eastern part of the ice sheet
was slow to move, however, with the result that the conifer-
ous forest grew right up to the edge of the ice. Ice sheets do
not melt at a steady rate. From time to time they halt and
then advance before resuming their retreat. Two Creeks,
Wisconsin, is one of a number of places where a temporary
advance of the ice buried a forest about 11,800 years ago.
In eastern North America the spruce and larch forest gave
way to forest dominated by spruce and pine (Pinus species),

Holocene, Pleistocene, and

late Pliocene glacials and interglacials
Approximate date Northwestern
(1,000 years BP) North America Great Britain Europe

Holocene
10–present Holocene Holocene (Flandrian) Holocene (Flandrian)

Pleistocene
75–10 Wisconsinian Devensian Weichselian
120–75 Sangamonian Ipswichian Eeemian
170–120 Illinoian Wolstonian Saalian
230–170 Yarmouthian Hoxnian Holsteinian
480–230 Kansan Anglian Elsterian
600–480 Aftonian Cromerian Cromerian complex
800–600 Nebraskan Beestonian Bavel complex
740–800 Pastonian
900–800 Pre-Pastonian Menapian
1,000–900 Bramertonian Waalian
1,800–1,000 Baventian Eburonian

Pliocene
1,800 Antian Tiglian
1,900 Thurnian
2,000 Ludhamian
2,300 Pre-Ludhamian Pretiglian
BP means “before present” (present is taken to be 1950). Names in italic refer to interglacials. Other names refer to
glacials (ice ages). Dates become increasingly uncertain for the older glacials and interglacials and the period before
about 2 million years ago. Evidence for these episodes has not been found in North America; in the case of the
Thurnian glacial and Ludhamian interglacial the only evidence is from a borehole at Ludham, in eastern England.
14 TEMPERATE FORESTS

and from about 9,000 years ago by pine and birch forest with
lesser amounts of alder (Alnus species) and spruce. In the
Northwest, birch forest arrived about 11,000 years ago, mixed
with poplar and aspen (both Populus species), willow (Salix
species), juniper (Juniperus species), and spruce. This devel-
oped into alder-birch-spruce forest by about 5,500 years ago.
The Fennoscandian ice sheet began to retreat from Europe
about 13,000 years ago, and sedges and grasses colonized the
newly exposed ground. The first trees to arrive were juniper
(Juniperus communis), arctic willow (Salix herbacea), and dwarf
birch (Betula nana), low-growing plants that could survive
the cold winds. Downy birch (B. pubescens), silver birch (B.
pendula), and aspen (Populus tremula) were growing across
northern Europe about 12,000 years ago. The landscape was
fairly open in the north, like parkland, with patches of wood-
land separated by tundra. Farther south, the trees formed
forests with downy birch the most abundant species.
About 11,000 years ago the advance was halted when the
temperature fell once more and the ice age returned. A cold
period that is shorter and milder than a full ice age is called a
stadial. This one was the Younger Dryas (or Loch Lomond)
stadial, named either for mountain avens (Dryas octopetala),
an arctic and alpine plant typical of cold climates, or for the
lake in western Scotland. Dryas pollen was discovered in
soils, at levels that have been dated, in places that are now
much too warm for the plant to occur naturally. When the
stadial ended, about 10,000 years ago, the forest resumed its
advance. Hazel (Corylus avellana) was widespread in northern
Europe by 9,000 years ago and it was soon followed by birch
and Scots pine (Pinus sylvestris).
The resulting birch-pine-hazel forest is still the natural veg-
etation over much of Europe immediately to the south of the
taiga, although most of it has been cleared to provide land
for farming or commercial forestry. Farther south the forests
were mainly of elm (Ulmus species) and oak (Quercus species).

Ancient forests that survive to this day

Climate changes do not happen smoothly. They proceed
faster in some places than in others and occasionally they
 GEOGRAPHY OF TEMPERATE FORESTS 15

reverse direction for a time. Ice that had been retreating

advances or an advancing ice sheet retreats.
Plants take some time to colonize new areas. After the last
ice age, birch and aspen advanced at about 0.5 mile (1 km) a
year. Beech (Fagus species) and spruce followed them, but at
less than 550 yards (500 m) a year. The time that it takes
plants to respond to climate change can leave individual
species and even some communities isolated when change
happens too quickly for them. If the climate change is un-
even over a large region, isolated species or communities may
survive in places where the change has had least effect.
The four-toed salamander (Hemidactylium scutatum) may
be one such species. This two- to four-inch (5–10-cm) am-
phibian occurs in eastern North America, mainly from Nova
Scotia to New England and west to Minnesota, but also in
several places in the southeastern United States. It lives in
wet places in temperate forests. Scientists believe that as the
ice sheets advanced during the last (Wisconsinian) ice age,
four-toed salamanders along with other animals gradually
moved south to where the climate was warmer. When the ice
age ended, however, grasslands and other treeless areas devel-
oped between the forests in the south and those in the north.
Larvae of fourtoed salamanders live in water, and these dry,
treeless areas formed a barrier the salamanders were unable to
cross. Consequently they remained isolated. The map on
page 16 shows their approximate location.
Species that become isolated in this way are known as
relicts, and the places where they survive are refugia (singular
refugium). There are four areas in the United States, also
shown on the map, where coniferous forests survived the ice
age. They grew in two places that remained free of ice in
Alaska, and above the ice in the Cascade, Rocky, and
Appalachian Mountains. Visit these places today and you
will see forests that have been there since before the last ice
age began 75,000 years ago.
These are large, well-known refugia, but there are many
smaller ones. As the forests advance they encounter places
where trees cannot grow because it is too wet or the soil is
unsuitable. The existing vegetation is not crowded out by the
new arrivals and so it remains as small relict of earlier times.
16 TEMPERATE FORESTS

Alaskan

Cascade Rocky Mountain

Appalachian

relict salamander
populations
forest refugia

North American refugia Rising temperatures may make plants in mountainous

regions move to higher elevations. The new plant communi-
ties colonize the lower parts of the mountains, but the old
communities survive in refugia above them.
Mountains themselves provide many different environ-
ments. There are sheltered hollows that are protected from
the wind but also permanently shaded. While plants charac-
teristic of warmer climates establish themselves on sunnier
parts of the mountain, the change from glacial to interglacial
conditions may allow ice-age plants that once covered a
much larger area to survive in the sheltered hollows. These
hollows are also refugia.
 GEOGRAPHY OF TEMPERATE FORESTS 17

What trees can tell us about

climates of the past
Ivy (Hedera helix) is an Old World plant that has been intro-
duced into North America (poison ivy is a different species).
Cultivated varieties are grown as ornamental plants, but wild
ivy climbs across the faces of buildings, across the ground,
and uses trees for support, eventually covering them com-
pletely. It is a woody climber that resembles the lianas of
tropical forests.
This is no coincidence, for that is what ivy really is. It
belongs to a plant family (Araliaceae) comprising several
hundred species, most of which grow in the Tropics. They are
especially common in Southeast Asia.
Ivy has broad, tough leaves with smooth surfaces, and it is
evergreen. Broad-leaved evergreen plants are typical of the
Tropics, and despite thriving so far from the equator, ivy has
not lost all of its tropical characteristics. It cannot survive a
long, hard winter, and it does not flower or produce seeds if
the average temperature in the coldest month is lower than
35°F (1.6°C). It cannot tolerate drought and will survive, but
not thrive, if the average temperature in summer is below
about 55°F (13°C).
Its precise climatic requirements make ivy very useful to
scientists called paleoclimatologists, who discover what cli-
mates were like in the distant past. Like all flowering plants,
ivy produces pollen, which survives for a very long time in
the soil (see “Using pollen and beetles to study the past” on
page 157). When scientists find ivy pollen in a layer of soil of
a known age, they can be certain that the climate at that time
was mild and wet.
Holly (Ilex aquifolium) is another broad-leaved evergreen
plant belonging to a mainly tropical family, the Aquifolia-
ceae. It also has precise requirements. Healthy holly trees
mean that the climate is wet and the average temperature
does not fall below about 31°F (–0.5°C) in winter and 55°F
(13°C) in summer.
Holly and ivy are plants associated with Christmas be-
cause they are evergreen and symbolize eternal life—holly
means “holy.” Mistletoe (Viscum album) is another Christmas
evergreen. Its presence indicates that the average summer
18 TEMPERATE FORESTS

temperature is higher than about 63°F (17°C), although it can

tolerate winter temperatures as low as 19°F (–7°C).
English yew (Taxus baccata) is a handsome evergreen tree
that lives for a very long time. It grows naturally in many
parts of Europe, North Africa, and the Middle East, but not in
the center of continents or in the far north. It requires a
moist climate and it cannot tolerate hard winters.
These are just a few of many plants that have clear climat-
ic preferences. Evidence that in the past they grew in a par-
ticular area provides scientists with an important clue to the
climate at that time.
 CHAPTER 2

GEOLOGY OF
TEMPERATE FORESTS
Plate tectonics
Antarctica is a barren, icy wilderness, yet there was a time
when forests grew there. There were once tropical swamps in
what is now Pennsylvania, and parts of England were once a
hot desert, like the Sahara.
Plants are able to spread because their seeds are blown by
the wind or carried by animals. They need not travel far. If
seeds give rise to new plants just a short distance from their
parent, the plant’s descendants will travel far over hundreds
of years. Seeds will sprout and grow only where conditions
are suitable, however. Wind-borne tree seeds might land in
Antarctica, but they would perish there, just as seeds from
mangrove, a tropical tree, would perish in Pennsylvania. The
fact that plants once grew in places where they could not sur-
vive today suggests that the climates of those places were
once very different. The Pennsylvanian swamps, for example,
existed during a time that geologists call the Carboniferous
period (see the appendix “Geologic time scale” on page 241).
Climate change cannot explain how tropical swamps once
existed in Alaska, however, or forests in Antarctica. Alaska is
too close to the North Pole and Antarctica to the South Pole.
Sunshine in these high latitudes is not warm enough for trop-
ical plants, and the winters are too long and dark. No amount
of climate change can compensate for their geographic posi-
tion. Some other factor must be at work—and it is.
The continents are moving. At one time Alaska, Antarctica,
and Pennsylvania lay in the Tropics. Tropical plants grew in
lands that lay in the Tropics and that have since moved.
About 400 million years ago, toward the end of the Devonian
period, Antarctica lay in the temperate regions and temperate
forests grew there. The illustration shows where the conti-
nents were 135 million years ago and 65 million years ago,

19
20 TEMPERATE FORESTS

135 Ma

65 Ma

Continental drift. The

maps show the
arrangements of the
continents 135 million
years ago, 65 million
today
years ago, and today.

compared with their positions today (“Ma” in the illustration

means “millions of years ago”). The arrows indicate the direc-
tion they were and are moving.
 GEOLOGY OF TEMPERATE FORESTS 21

Alfred Lothar Wegener and continental drift

Since the first realistic maps of the world were published in the 16th century, many geog-
raphers had puzzled over the fact that the continents on each side of the Atlantic Ocean
looked as though they might fit together. Some thought it mere coincidence, but others
suggested ways a single continent might have split into two parts that then moved apart.
The German meteorologist Alfred Lothar Wegener (1880–1930) went much further.
Wegener compiled a mass of evidence to support what he called “continental displace-
ment.” This phenomenon came to be called continental drift. He studied the scientific lit-
erature for descriptions of rocks that were similar on each side of the Atlantic. He found
plants with limited distribution that are separated by vast oceans and fossil organisms that
are also distributed in this way.
Finally he proposed that about 280 million years ago, during the Upper Paleozoic
subera, all the continents were joined, forming a single “supercontinent,” which he called
Pangaea (from the Greek pan, meaning all, and ge, meaning Earth), surrounded by an
ocean called Panthalassa (thalassa means ocean). He theorized that Pangaea broke apart
and the separate pieces drifted to their present locations; the continents are still drifting.
In 1912 Wegener published a short book outlining his theory, Die Entstehung der
Kontinente und Ozeane (The Origin of the continents and oceans). He was drafted into the
German army in 1914 at the start of World War I but was wounded almost at once. He
developed his ideas further while recovering in a hospital, and in 1915 he published a
much longer edition of his book (it was not translated into English until 1924).
His idea found little support. Geologists at the time believed the mantle—the material
beneath the Earth’s crust—to be solid, and they could not imagine any way that conti-
nents could move. They also found that some of Wegener’s calculations of the rate of con-
tinental displacement were incorrect.
But support for Wegener’s idea began to grow in the 1940s, when for the first time sci-
entists were able to study the rocks on the ocean floor. These studies indicated that the
oceans had grown wider by spreading outward from central ridges, where underwater
volcanoes were erupting, laying down new rock. Wegener’s theory was generally accept-
ed by the late 1960s, but by then Alfred Wegener was dead. He had died in 1930 during
his third expedition to study the climate over the Greenland ice sheet.

At present, the continued widening of the Atlantic Ocean

is carrying North and South America westward at a rate of
about 0.8 inch (20 mm) a year. India is moving northward
into Eurasia, crumpling the rocks of the crust into the
22 TEMPERATE FORESTS

Himalayan Mountains, which are still rising. As Africa and

Europe move northward and eastward, the Red Sea is widen-
ing. Eventually it will grow into a wide ocean. The move-
ment of the continents is called continental drift, and the the-
ory explaining it was first suggested in 1912 by the German
meteorologist Alfred Wegener (see the sidebar on page 21).
Evidence supporting Wegener’s theory began to emerge in
the 1940s from studies of the rocks on the ocean floor.
Scientists found that some oceans are expanding. The Ame-
rican oceanographer Robert Sinclair Dietz (1914–95) called
this process seafloor spreading, and in 1967 Dan McKenzie
The major crustal plates. (born 1942) of Cambridge University brought together all
The plates move very that was known at the time and proposed a new theory: plate
slowly, carrying the tectonics. A scientific theory is an explanation of a natural
continents with them. process, with solid evidence to back it.

Eurasian Plate North American Plate Eurasian Plate

Juan de Fuca Plate

Caribbean Plate
Arabian Plate
Philippine Plate
Cocos Plate Indian Plate

Pacific Plate African Plate

Nazca Plate
Australian Plate South American Plate

Antarctic Plate
Antarctic Plate Scotia Plate
 GEOLOGY OF TEMPERATE FORESTS 23

The theory of plate tectonics holds that the Earth’s solid

crust is composed of a number of blocks, or plates, that move
in relation to each other. A tectonic process is one related to
the deformation of the crust or to structures produced by
such deformation. There are seven major plates: the African,
Eurasian, Pacific, North American, South American, Ant-
arctic, and Australian plates. There are also several lesser
plates: the Cocos, Caribbean, Nazca, Arabian, Indian, Phil-
ippine, and Scotia plates. In addition, there are minor plates
such as the Juan de Fuca plate, microplates, and fragments of
former plates that have broken apart. The map shows the
present location of the major and lesser plates.
The theory of plate tectonics explains many aspects of the
Earth’s geology, such as the reason why earthquakes and vol-
canic eruptions happen only in certain parts of the world. It
also explains why some plants—and some temperate forests
—are found in places thousands of miles apart, and nowhere
else.

Related temperate-forest trees that occur

in widely separated places
Alfred Wegener (see the sidebar “Alfred Lothar Wegener and
continental drift” on page 21) was struck by the fact that cer-
tain plants occur naturally only in certain parts of the world
and that those locations are separated by vast expanses of
ocean. This type of scattered occurrence is called a disjunct
distribution. There are a great many examples of disjunct dis-
tribution among plants and animals that are now extinct and
known only as fossils. There are also many examples among
living plants and animals. For example, marsupial mammals,
the group that includes opossums and kangaroos, occur in
Australia and New Guinea, and also in North and South
America. Southern beeches (35 species of Nothofagus) are now
grown in many parts of the world. They are handsome trees
that produce valuable timber. They occur naturally, however,
in New Guinea, southeastern Australia, Tasmania, and New
Zealand, and also along the southwestern coast of South
America. The monkey puzzle tree or Chile pine (Araucaria
araucana) is a popular ornamental tree. As its name suggests,
24 TEMPERATE FORESTS

(opposite page) Disjunct distribution. Members of certain plant

families occur in widely scattered parts of the world. This is called
disjunct distribution, and it suggests that the regions where the plants
occur were once much closer together. The maps show the distribution
of the Canellaceae (white cinnamon); Caricaceae (pawpaw);
Magnoliaceae (magnolias and tulip tree); and Platanaceae (plane
and buttonwood trees).

it is native to South America, where it forms forests. It also

grows naturally in New Guinea, northeastern Australia, and
on islands in the South Pacific Ocean. These are individual
species, but there are entire families of plants that have a dis-
junct distribution. The illustration shows four of these.
The Canellaceae is a small family of five genera and about
17 species of aromatic trees. Canella bark, obtained from
Canella winterana, known as white cinnamon, is used as a
tonic and flavoring, and in Puerto Rico it is used as a fish poi-
son. The scented wood of Cinnamosma fragrans is used in reli-
gious ceremonies. Warburgia species produce a resin that is
used to fasten handles to tools, leaves that are used in curries,
and bark that is used as a purgative. As the map shows, this
family occurs in Central America and the West Indies, where
there are three genera, and in East Africa and Madagascar,
where there are two genera.
The Caricaceae is a family of four genera and about 30
species of trees, one of which is Carica papaya, the papaya or
pawpaw tree. Members of the family are most abundant in
South America, with some in Central America, but one
genus, Cylicomorpha, grows in tropical Africa.
About 200 species of trees and shrubs belong to the mag-
nolia family, the Magnoliaceae. These include the tulip trees
Liriodendron tulipifera, which is native to eastern North Ame-
rica, and L. chinense, which is native to central China. The
Magnoliaceae occurs in North, Central, and South America
and in Asia, but not in Europe or Africa.
Plane trees and buttonwood trees belong to the family
Platanaceae. It is a small family, with only one genus (Plata-
nus) and about seven species. Most species occur naturally in
the United States, but P. orientalis is a native of the Balkan
 GEOLOGY OF TEMPERATE FORESTS 25

Canellaceae Caricaceae

Magnoliaceae Platanaceae

Peninsula of southeastern Europe and the Himalayas, and P.

kerrii is native to Southeast Asia.
Disjunct distributions come about when a species or group
of species—genus or family—evolves on a particular conti-
nent and the continent then splits into two or more parts,
each carrying part of the population. As Wegener observed,
continental drift is the only plausible explanation for the
occurrence of closely related plants in widely separated parts
of the world.

Soils of temperate forests

In a temperate forest, a layer of dead leaves covers the ground
except on some well-trodden paths. Mixed with the leaves
there are small twigs, remains of other plant material, and
many small animals. Ants, beetles, snails, slugs, spiders, cen-
tipedes, and countless others, some of them too small to see
26 TEMPERATE FORESTS

except with a magnifying glass, live among this surface layer.

Underneath the leaves, the bottom of the layer comprises
plant matter that is damp, dark in color, and broken into
fragments. This is plant material that has been almost com-
pletely decomposed by the activities of bacteria, fungi, and
all the small animals. Beneath that layer is soil.
There are some places where more of the soil is visible. A
deep layer is often exposed on the side of a steep bank and
the rain may have washed away so much of the soil that the
roots of a nearby tree stand out.
A vertical section of soil that is exposed in this way is
called a soil profile. The side of a bank that has been exposed
for a long time will be coated with soil that has washed down
over it, so it will look all the same, but if you could find a
freshly exposed profile you would notice that it comprises
several distinct layers. These soil layers are known as horizons.
Soil scientists, called pedologists, have divided an idealized
soil into five levels, labeled O, A, B, C, and R, with a total of
10 horizons. The illustration shows all of these principal
horizons, with their labels and descriptions. Pedologists
often add much more detail than the illustration shows, to
indicate important aspects of the chemical composition of
the soil in each horizon. Soils vary widely in their physical
and chemical composition, and many types of soil lack one
or more of the full complement of horizons. Some soils pos-
sess no horizons at all.
It is obvious why soils must vary. They are made from min-
eral particles derived from the underlying rock. There are dif-
ferent types of rock and so there are types of mineral particle
that differ in size, shape, and chemical composition. The
leaves lying on a forest floor become incorporated into the
soil, but forests do not grow everywhere. Grasses add a differ-
ent kind of plant matter to the soil and grassland soils differ
from forest soils. The floor of a coniferous forest is carpeted
with needles. These decompose much more slowly than
broad leaves. So the underlying rock and the type of vegeta-
tion determine the type of soil.
Forests require a fairly moist climate, grasslands a drier
one, and rain hardly ever falls in a desert. Average tempera-
tures also influence the type of vegetation growing in a par-
 GEOLOGY OF TEMPERATE FORESTS 27

O1 organic debris

O2 partly decomposed organic debris

mineral material with fine particles

A1 of organic matter

mineral material into which nutrients have

A2 moved from above

A3 transitional layer

B1 transitional layer

B2 layer in which most nutrients accumulate

B3 transitional layer

C parent material

Soil profile. The diagram

shows all the layers,
called horizons, from the
R bedrock surface down to the
underlying bedrock.
28 TEMPERATE FORESTS

ticular area. Through its influence on the vegetation, climate

indirectly influences the development of the soil. Climate
also has a more direct effect because chemical reactions work

How soils are classified

Farmers have always known that soils vary. There are good soils and poor soils, heavy soils
containing a large proportion of clay, sandy soils that dry out rapidly, and light, loamy soils
that retain moisture and nutrients. Loam is a mixture of sand, silt, and clay—mineral parti-
cles of different sizes. In the latter part of the 19th century Russian scientists were the first to
attempt to classify soils. They thought that the differences between soils were due to the
nature of the parent material—the underlying rock—and the climate. They divided soils
into three broad classes. Zonal soils were typical of the climate in which they occur, intra-
zonal soils were less dependent on climate for their characteristics, and azonal soils were
not the result of climate. Azonal soils include windblown soils and those made from silt
deposited by rivers on their floodplains. Individual soil types were placed in one or other of
these broad groups. This system remained in use until the 1950s, and some of the Russian
names for soils are still widely used, such as Chernozem, Rendzina, Solonchak, and Podzol.
American soil scientists were also working on the problem, and by the 1940s their work
was more advanced than that of their Russian colleagues. By 1975 scientists at the United
States Department of Agriculture had devised a classification they called “Soil Taxonomy.”
It divides soils into 10 main groups, called orders. The orders are divided into 47 subor-
ders, and the suborders are divided into groups, subgroups, families, and soil series, with
six “phases” in each series. The classification is based on the physical and chemical prop-
erties of the various levels, or horizons, that make up a vertical cross section, or profile,
through a soil. These were called “diagnostic horizons.”
National classifications are often very effective in describing the soils within their bound-
aries, but there was a need for an international classification. In 1961 representatives from
the Food and Agriculture Organization (FAO) of the United Nations, the United Nations
Educational, Scientific, and Cultural Organization (UNESCO), and the International Society
of Soil Science (ISS) met to discuss preparing one. The project was completed in 1974 and
is known as the FAO-UNESCO Classification. Like the Soil Taxonomy, it was based on diag-
nostic horizons. It divided soils into 26 major groups, subdivided into 106 soil units. The
classification was updated in 1988 and has been amended several times since. It now com-
prises 30 reference soil groups and 170 possible subunits. The FAO has also produced a
World Reference Base (WRB), which allows scientists to interpret the national classification
schemes.
 GEOLOGY OF TEMPERATE FORESTS 29

faster in warm, moist conditions than when it is cold or dry.

Dead organic matter decomposes faster in warm climates
than in cold ones.
Soils also change over time. When the ice age ended, the
retreating ice sheets left behind a surface of bare rock, much
of it shattered, with some finer particles filling crevices and
hollows. Little by little this material became a soil. It was a
young soil, newly formed. Temperate forests grow in mature
soils that are deep enough for the roots of big trees. The
nutrients that sustain the trees dissolve from the minerals
into the water moving through the soil. They are recycled
many times, but the water also carries them away, out of the
forest, and so the soil gradually becomes less fertile. It grows
older. The temperate forests have not yet existed long
enough for their soils to become ancient, but one day they
will be so. Many tropical soils are very deep, but they have
lost most of their plant nutrients. The Tropics were not cov-
ered by ice during recent ice ages and so their soils have con-
tinued to age, and now they are very old.
Depending on their type and age, soils look different. Some
are pale, some dark, and some gray. Some have a fine, almost
powdery texture, some feel gritty, and some are so sticky you
can shape them, like modeling clay. The characteristics of a
soil affect its fertility—its capacity for growing farm crops. This
makes the scientific study of soils, called pedology, important.
Since the 19th century, pedologists have made many
attempts to simplify the study of soils by dividing them into
clearly defined types. There are now several national schemes
of soil classification and an international scheme that is used
by the Food and Agriculture Organization (FAO) of the
United Nations. Temperate forests often grow in soils classi-
fied as Inceptisols, Mollisols, and Spodosols in the U.S. Soil
Taxonomy, or Chernozems, Kastanozems, and Podzols in the
FAO system. The history of soil classification is summarized
in the sidebar.

How water moves through a temperate forest

When it rains, puddles collect on the ground in some places,
but in other places the water soaks into the ground and
30 TEMPERATE FORESTS

disappears at once. After a time, water may start to flow

down gullies on hillsides that until then had been dry. A few
hours later, provided there is no more rain, the ground once
again looks and feels dry.
On city streets, the rainwater flows into storm drains that
carry it away, and the puddles—in depressions so the water
cannot flow to the drains—evaporate. This is not what hap-
pens in the forest. There, the rain is much gentler. The leaves
shelter the forest floor so that the rain drips from them and
falls slowly, and water also runs down the trunks of the trees.
Some of the water evaporates before reaching the ground, so
the rainfall is lighter inside a forest than it is outside. The
raindrops are more likely to fall vertically—straight down—
than they are in the open, beyond the forest edge, because
the trees also slow the wind.
Once rainwater reaches the ground it continues to flow
downward—to soak into the ground. It is able to move
downward because soil is composed of solid particles
derived from the underlying rock. The particles pack togeth-
er closely, but there are small spaces, called pores, between
them. Air fills the soil pores, but it can be displaced by water
flowing through the soil. If water fills most of the pores, the
soil is waterlogged. Walking across it will churn the soil into
mud.
The total amount of pore space determines the amount of
water a given volume of soil can hold. This is known as the
porosity of the soil. Surprisingly perhaps, although soil parti-
cles vary greatly in size, their size does not affect the porosity.
The diagram illustrates this. It shows three compartments of
equal volume, containing one, eight, and many spherical
particles. It looks as though there is more empty space in the
compartment with just one particle—or perhaps in the one
with eight particles. In fact, there is precisely the same
amount of empty space in all three compartments. If the
compartments held different soils, all three would be equally
porous.
Equal porosity does not mean that water moves with equal
ease through all three soils, however. Water passes more easi-
ly through a few large spaces than through many small
spaces. The ease with which water passes is measured as the
 GEOLOGY OF TEMPERATE FORESTS 31

permeability of a soil, and it depends on the size of the pore Porosity and
spaces rather than their total volume. Two soils can be equal- permeability. Each box
ly porous without being equally permeable. The more perme- contains the same
able the soil, the more quickly water passes through it, and amount of air, so all
the sooner it dries after rain. three are equally porous.
Water draining downward through the soil eventually Water would pass more
encounters a layer of impermeable material. This is usually easily through those
the underlying bedrock or a layer of tightly compacted clay. containing fewer
Water cannot pass through it and so it accumulates above the particles, so they are
layer, filling all the pore spaces and saturating the soil. The not equally permeable.
level below which the soil is completely saturated is called
the water table.
Perhaps surprisingly, water in soil travels not only down
but up. Water travels a short distance upward through the
smallest pore spaces by a process called capillarity. Con-
sequently, above the saturated layer there is a capillary fringe
in which some of the pore spaces contain air and others con-
tain water rising by capillarity. The capillary fringe is thicker
in soils made from fine particles than it is in soils made
from coarse particles, because water moves farther by capil-
larity through small spaces than it does through large
spaces.
32 TEMPERATE FORESTS

mixed broad-leaved trees mixed broad-leaved trees and conifers meadow

Effects of drainage Capillarity acts in all directions. Most of the water draining
on vegetation downward through the soil moves by capillarity rather than
by gravity. It sinks by gravity only where the pore spaces are
large—as they are in sand, for instance.
If water continues to drain downward, the water table
will rise. If it rises all the way to the surface, the ground
will be waterlogged and additional water will lie on the sur-
face in pools. It may even cover a large area. The land is then
flooded.
Some plants are more tolerant of wet soils than others, but
very few can survive flooding because their roots need air for
respiration. Consequently, the rate at which drainage re-
moves surplus water from the soil affects the composition of
a forest. As the diagram shows, a variety of broad-leaved trees
grows on well-drained soil, different species mixed with
coniferous trees grow on excessively or poorly drained soil,
and only meadow, comprising sedges and grasses, will grow
in soil that is sometimes flooded.
The underlying impermeable layer is unlikely to be hori-
zontal, and if it is inclined, the water in the saturated layer
will flow down the slope. The moving water is then known as
groundwater, and the permeable soil through which it flows is
an aquifer. In most soils, groundwater moves at speeds rang-
 GEOLOGY OF TEMPERATE FORESTS 33

ing from a few feet a day to a few feet a month. If the water
reaches a place where the overlying soil is very thin and
poorly drained, the water bubbles to the surface as a spring. If
it reaches low-lying ground, it may also rise to the surface
and continue its journey to the sea as a river.
CHAPTER 3

CLIMATE OF
TEMPERATE FORESTS
Autumn, when the Why there are seasons
leaves change color Winters are cold. The ground freezes and in the forest the
and the forest blazes trees are bare. Sometimes the weight of ice and snow snaps a
with patches of red and branch. Winter passes, and in spring the forest comes back to
yellow. Soon the leaves life. Summer is warm, and as it draws to a close the leaves
will fall and the trees change color and then fall.
will be bare. (Courtesy These are the seasons. They are very familiar, but why do
of Fogstock) they happen? Why doesn’t the weather remain the same all

34
 CLIMATE OF TEMPERATE FORESTS 35

March N

June N N

Sun

S S December

S September

year round, as it does near the equator? And why do the sea- How Earth’s tilted axis
sons become more pronounced farther from the equator? produces the seasons.
The answer is that the Earth is not upright. Earth turns on As Earth orbits the Sun,
its own axis, making one complete turn every 24 hours. Earth its tilted axis means that
also orbits the Sun, taking one year to complete each orbit. in June the Northern
Imagine that the path of the Earth’s orbit marks the edge of a Hemisphere, and in
flat disk, with the Sun at its center. This disk is known as the December the Southern
plane of the ecliptic. If the Earth’s own axis were at right angles Hemisphere, is exposed
to the plane of the ecliptic, the Sun would always be directly to more sunlight.
overhead at the equator and it would shine equally onto
both hemispheres. The axis is tilted about 23.5° with respect
to the plane of the ecliptic, however, and the diagram shows
the consequence of that tilt. As the Earth orbits the Sun, first
the Northern Hemisphere and then the Southern Hemi-
sphere is tilted to face the Sun.
The effect is most extreme at the solstices, on June 21–22
and December 22–23. At the June solstice the Sun is directly
overhead at latitude 23.5°N. This latitude marks the tropic of
Cancer. At latitude 66.5°N the Sun remains above the horizon
for 24 hours. This latitude marks the Arctic Circle. At the
December solstice the Sun is directly overhead at the tropic of
Capricorn, at latitude 23.5°S, and remains above the horizon
for 24 hours at the Antarctic Circle, at 66.5°S. The two solstices
36 TEMPERATE FORESTS

are also known as midsummer’s day and midwinter’s day. The

hours of daylight reach a maximum at midsummer’s day and
a minimum at midwinter’s day, and the number of hours of
daylight and darkness varies according to the latitude.
Midway between the solstices, the Sun is directly overhead
at the equator. Consequently, on these days—March 20–21
and September 22–23–the Sun is above the horizon for 12
hours and below it for 12 hours everywhere in the world.
These dates are called the equinoxes.
While the Sun is shining the surface of the Earth absorbs
its radiation and grows warmer. At night, the surface loses the
Angle of sunlight. warmth it absorbed by day and grows cooler. Between the
Latitude affects the spring and autumn equinoxes there is more daylight than
intensity of sunlight. darkness and so the surface has more time to absorb heat
A. At the equator the than it does to lose it. The temperature rises, the plants
Sun is almost directly respond to the warmth, and during the middle part of this
overhead and its light period we enjoy summer. Between the autumn and spring
is concentrated. B. In equinoxes the opposite happens. There are more hours of
high latitudes the Sun is darkness than there are of daylight and so the surface cools
low in the sky and its down and we have winter.
light is spread over a Although everyone experiences seasons, average tempera-
larger surface area, tures in the Tropics are higher than temperatures in higher
making it less intense. latitudes, and the coldest parts of the Earth are around the

A
 CLIMATE OF TEMPERATE FORESTS 37

North and South Poles. The illustration shows why this is so.
Sunshine falls more intensely in places where the Sun is
approximately overhead (A in the picture) than it does where
the Sun is low in the sky (B), because the Sun overhead illu-
minates a smaller area. Air over the Tropics is warmer than
air over any other region of the Earth, and the contrast in
air temperature sets up a worldwide movement of air called
the general circulation of the atmosphere. This circulation, ex-
plained in the sidebar, transports heat from the equator into
high latitudes and produces the world’s climates.

How oceans affect climates

Air moves away from the North Pole as northeasterly winds
and from the South Pole as southeasterly winds. Air moves
toward the equator as the northeasterly trade winds in the
Northern Hemisphere and southeasterly trade winds in the
Southern Hemisphere. In temperate regions the winds are
predominantly from the west.
Even the most constant winds do not blow all the time or
always from the same direction, but these are the prevailing
winds—the directions from which the wind blows more often
than it does from any other directions. They are the winds
produced by the general circulation of the atmosphere (see
the sidebar “General circulation of the atmosphere” on page
38).
A person walking outdoors on a windy day can feel the
wind pushing on him or her and can see the leaves, scraps of
paper, and dust that it blows around. The wind exerts a pres-
sure. The wind over the ocean exerts this pressure on the sur-
face of the water. It pushes the water along the way it blows
leaves and paper on land, but the wind blows more constant-
ly at sea than it does over land, so its pressure is steadier. The
water that it pushes along forms ocean currents, like rivers
flowing through the sea.
On either side of the equator, the North and South
Equatorial Currents flow from east to west. As they approach
land they turn away from the equator and enter middle lati-
tudes, where westerly winds carry them eastward. They con-
tinue to turn, however, until eventually they are flowing
38 TEMPERATE FORESTS

General circulation of the atmosphere

The tropics of Cancer in the north and Capricorn in the south, mark the boundaries of the
belt around the Earth where the Sun is directly overhead on at least one day in the year.
The Arctic and Antarctic Circles mark the boundaries of regions in which the Sun does not
rise above the horizon on at least one day of the year and does not sink below the horizon
on at least one day in the year.
A beam of sunlight illuminates a much smaller area if the Sun is directly overhead than
it does if the Sun is at a low angle in the sky. The amount of energy is the same in both
cases, but the energy is spread over a smaller area directly beneath the Sun than it is when
the Sun is lower. This is why the Tropics are heated more strongly than any other part of
the Earth and the amount of heat falling on the surface decreases with increasing distance
from the equator (increasing latitude).
The Sun shines more intensely at the equator than it does anywhere else, but air move-
ments transport some of the warmth away from the equator. Near the equator, the warm
surface of the Earth heats the air in contact with it. The warm air rises until it is close to the
tropopause, which is the boundary between the lowest layer of the atmosphere (the tro-
posphere) in which air temperature decreases with height, and the layer above (the strat-
osphere), where the temperature remains constant with increasing height. The height of
the tropopause is around 10 miles (16 km), and at this height the air moves away from the
equator, some heading north and some south. As it rises, the air cools, so the high-level air
moving away from the equator is very cold—about –85°F (–65°C).
This equatorial air subsides around latitude 30°N and S, and as it sinks it warms again.
By the time it reaches the surface it is hot and dry, so it warms this region, producing sub-
tropical deserts. At the surface, the air divides. Sometimes called the horse latitudes, this is
a region of light, variable winds or no winds at all. Most of the air flows back toward the
equator and some flows away from the equator. The air from north and south of the equa-
tor meets at the Intertropical Convergence Zone (ITCZ), and this circulation forms a num-
ber of vertical cells called Hadley cells, after George Hadley (1685–1768), the English mete-
orologist who first proposed them in 1735.
Over the poles, the air is very cold. It subsides, and when it reaches the surface it flows
away from the poles. At about latitude 50–60°N and S, air moving away from the poles
meets air moving away from the equator at the polar front. The converging air rises to the
tropopause, in these latitudes about seven miles (11 km) above the surface. Some flows
back to the poles, forming polar cells, and some flows toward the equator, completing
Ferrel cells, discovered in 1856 by the American climatologist William Ferrel (1817–91).
 CLIMATE OF TEMPERATE FORESTS 39

Warm air rises at the equator, sinks to the surface in the subtropics, flows at low level to
around latitude 55°, then rises to continue its journey toward the poles. At the same time,
cold air subsiding at the poles flows back to the equator. The diagram below shows how
this circulation produces three sets of vertical cells in each hemisphere. It is called the
“three-cell model” of the atmospheric circulation.
If it were not for this redistribution of heat, weather at the equator would be very much
hotter than it is, and weather at the poles would be a great deal colder.

polar cell

polar front

Ferrel cell

horse latitudes

Hadley
cell

ITCZ

Hadley
cell

horse latitudes

Ferrel cell

polar front

polar cell

General circulation of the atmosphere. The movement of air transports heat away from the
equator, forming three sets of circulation cells in each hemisphere. In the Hadley cells, warm
air rises over the equator and subsides over the Tropics; some of the air flows away from the
equator and some returns to the equator. Cold air subsides over the poles, flows toward the
equator, and rises at the polar front; this forms the polar cells. The Hadley and polar cells
drive the Ferrel cells, in which air rises at the polar front, flows toward the equator until it
meets air in the Hadley cell, then subsides with the Hadley-cell air, producing the calm
conditions of the horse latitudes.
40 TEMPERATE FORESTS

N. Equatorial C. California C. Equatorial Equatorial Japan C.

Counter C. Counter C.

N. Atlantic C.

Canary C.
Gulf Stream
N. Equatorial C.
Guinea C.
S. Equatorial C.

S. Equatorial C.

warm currents
West Wind Drift Humboldt C. Falkland C. Brazil C. Benguela C. Agulhas C. cool currents

Ocean currents. In back toward the equator. In each of the oceans the wind-
each of the oceans driven currents follow approximately circular paths called
the currents flow gyres. Every current has a name. The map shows the princi-
approximately in pal currents.
circles, called “gyres,” Despite consisting of water moving through water, the
moving clockwise in the ocean currents are perfectly real. In some cases they are even
Northern Hemisphere visible. Kuroshio means “black water” in Japanese, and it is the
and counterclockwise name of a current of dark-colored water that flows northward
in the Southern from the Philippines, past the Japanese coast, and then east-
Hemisphere. ward into the North Pacific Ocean. The current is less than 50
miles (80 km) wide and moves at up to seven MPH (11 km/h).
The Kuroshio Current flows away from the equator, carry-
ing warm water into higher latitudes. It crosses the North Pa-
cific Ocean and turns southward as it approaches the North
American coast, returning to the Tropics as the California
Current carrying cold water.
All of the ocean gyres carry warm water away from the
equator, past the east coasts of the continents. They are
 CLIMATE OF TEMPERATE FORESTS 41

called western boundary currents because they are on the west-

ern sides of the oceans, and they are deep, narrow, and fast-
moving. Eastern boundary currents, on the opposite sides of
the oceans, are wide, shallow, slow-moving, and carry cold
water toward the equator.
Air is warmed or cooled by contact with the surface
beneath it. Consequently, the ocean gyres have a great influ-
ence on the air crossing oceans and on the climates of the
lands bordering the oceans. Ocean gyres help to distribute
the Sun’s warmth more evenly, but the oceans have another
influence on climate that is even more important. This
occurs because water has a much higher specific heat capacity
than dry land.
The specific heat capacity of a substance is the amount of
heat it must absorb for its temperature to rise by one degree.
Dry land is made from rocks of various types, and from soils
made from rock particles. The specific heat capacity of water
is about five times that of rock. This means that water
absorbs five times as much heat as rock to make the same rise
in temperature. Consequently, the oceans warm up much
more slowly than the land in summer, which is why the sea
or a lake feels cold even on a scorching summer’s afternoon.
Air crossing the ocean in summer is cooler than air crossing a
continent. if there were no oceans, summer temperatures
would be much higher than they are.
Water also cools much more slowly than land, for the same
reason. In the middle of winter the sea is often warmer than
the land—and the air over the land. If there were no oceans,
winter temperatures would be much lower than they are.
The oceans therefore affect the climates of the world in
two ways. They store heat, accumulating it slowly through
the summer and releasing it slowly through the winter. In
this way they moderate air temperatures, making summers
cooler and winters warmer. They also transport heat directly.
Ocean gyres carry warm water away from the equator and
cold water back to the equator.

Air masses and fronts

As air moves slowly across a continent it acquires certain
characteristics. It becomes drier, because each time it crosses
42 TEMPERATE FORESTS

high ground some of its moisture condenses to form clouds,

and some of the clouds deliver rain, hail, or snow. There are
few large expanses of water in midcontinent to replenish the
air with water vapor.
The temperature of the ground surface determines the
temperature of the air. Air absorbs very little of the solar radi-
ation passing through it. Sunshine is absorbed by the land
and sea surface. The surface grows warmer and the air above
the surface is warmed by contact with it. Warm air rises and
cooler air takes its place, to be warmed in its turn, so the
warmth of the ground is spread throughout the air above it.
In winter the ground is cold and so the air is also cold.
Because it is cold, the air contracts—the way the air in a party
balloon shrinks and makes the balloon shrivel if it is placed
in a refrigerator for an hour or two. When air contracts, its
molecules move closer together, making the air denser and
therefore heavier. This increases the surface air pressure—the
pressure the weight of the atmosphere exerts on the Earth’s
surface. Air pressure over continents is high in winter.
In summer, the ground becomes very warm. The air over
the continent also becomes warm—although it remains dry.
As it warms, so the air expands. Its molecules move farther
apart, making the air less dense and less heavy. Surface pres-
sure is low over continents in summer.
Eventually the air reaches the coast and moves over the
ocean, and before long its characteristics start to change.
Water evaporates into the dry air, making it moister. Contact
with the sea surface warms very cold air in winter and cools
very hot air in summer. As the temperature of the air rises or
falls, its surface pressure also changes.
During World War I, a group of meteorologists at Bergen,
Norway, led by Vilhelm Bjerknes, studied the physical aspects
of the air. They concluded that the air over a large area was
often at approximately the same temperature, pressure, and
humidity everywhere, but that adjacent to it there would be
air that was warmer or cooler, moister or drier. They gave the
name air masses to these vast volumes of air (see the sidebar).
Adjacent air masses do not mix easily. This is because they
are at different temperatures and therefore densities—cool air
is denser than warm air. Where two air masses meet, either
 CLIMATE OF TEMPERATE FORESTS 43

Vilhelm Bjerknes and the Bergen School

Vilhelm Frimann Koren Bjerknes (1862–1951) was one of the founders of modern mete-
orology. He was born in Oslo, the son of a professor of mathematics, and held posts at
several universities in Norway, Sweden, and Germany. In 1912 he was appointed pro-
fessor of geophysics at the University of Leipzig, Germany, where he founded the
Leipzig Geophysical Institute. While there he developed his ideas on weather forecast-
ing, recruiting a team of assistants and colleagues who came to be known as the
“Leipzig School.”
Bjerknes returned to Norway in 1917 to found the Bergen Geophysical Institute and
become its first director. It was there that he did his most important work.
As in Leipzig, Bjerknes surrounded himself with a team of colleagues, who became the
“Bergen School.” The team included Vilhelm’s son Jakob Aall Bonnevie Bjerknes
(1897–1975), who later became professor of meteorology at the University of California,
Los Angeles, and Tor Harold Percival Bergeron (1891–1977), who later proposed a way
raindrops might form in clouds containing large amounts of ice crystals.
The team established a network of weather stations throughout Norway, from which
observers sent their observations and measurements to Bergen. Members of the Bergen
School plotted these reports on maps, producing pictures of weather conditions over a
large area at particular times.
Studying these pictures, they concluded that the characteristics of the air—its tempera-
ture, pressure, density, and humidity—remained constant over large areas, but that the air
over one area was markedly different from the air over another. They gave the name air
mass to these large bodies of air.
They also noticed that air masses did not readily mix with each other. Where two air
masses met, there was a clearly defined boundary between them. At the time—it was dur-
ing World War I—the newspapers were full of stories about opposing armies and the
fronts separating them. It struck the Bergen meteorologists that air masses were a little like
opposing armies, so they called the boundaries separating them fronts. They then devel-
oped a frontal theory to explain how fronts form, develop, and disappear, and how they
produce the weather that is associated with them. This theory also explained how depres-
sions form in air crossing the North Atlantic Ocean.
Modern meteorology and weather forecasting is largely based on the work of Bjerknes
and his colleagues. Bjerknes was a popular teacher. He attracted talented scientists
and made sure they received full recognition for their achievements. Bjerknes later joined
the staff of the University of Oslo, where he remained until he retired in 1932. He died
in Oslo.
44 TEMPERATE FORESTS

cold, dense air pushes beneath warm, less dense air, or the
warm air rides over the cool air.
There is some mixing at the boundary, or front, between
the air masses. A front appears as a line on a weather map,
Cross section through but in fact it is a region called the frontal zone, 60–120 miles
a frontal system. (100–200 km) wide. A front is named for the air behind it. It
The system extends is a warm front if its passage means that warm air replaces
from the surface to cool air. If cool air replaces warm air it is a cold front. Warm
the tropopause and and cold are relative terms. They mean only that the air is
does not affect air in the warmer or colder than the air adjacent to it.
stratosphere. A mass of Fronts slope upward from the surface and they move across
warm air is rising above the surface. Warm fronts slope at an angle of about 0.5°–1.0°
the surrounding colder and cold fronts at about 2°. The illustration shows the
air, with fronts marking sequence of events when a warm front passes, followed by
the boundaries between warm air, and then a cold front arrives heralding cooler air
warm and cold air. The behind the front. An observer on the surface on the right of
air behind the front the diagram is in a cold air mass. As the warm front approach-
defines the front: es, cloud associated with the front appears high overhead, and
A warm front brings as the front draws closer the cloud base becomes lower and
warm air and a cold the type of cloud changes. When the front passes, the ob-
front brings cold air. server is in a mass of air that is warmer than the air ahead of

stratosphere

tropopause

cold front warm front

cold air warm air cold air

direction of movement
 CLIMATE OF TEMPERATE FORESTS 45

the warm front. In this warm sector, there are often sheets of
dull, gray cloud and light but persistent rain or snow. Then
the cold front arrives, heralding a mass of colder air. When it
has passed the observer sees heaped clouds, sometimes bring-
ing showers. As the cold front recedes the clouds associated
with it appear progressively higher in the sky.
Warm fronts travel at an average speed of about 15 MPH
(24 km/h) and cold fronts at about 22 MPH (35 km/h).
Because cold fronts move faster than warm fronts, the cold
air usually pushes beneath the warm air, progressively lifting
it clear of the surface. Eventually all of the warm air has been
lifted and the two fronts merge—they are said to be occluded
or to form an occlusion. As the warm air rides up the cold
front, its temperature decreases and its water vapor condens-
es to form clouds. These are of the heaped type that can pro-
duce showers or sometimes thunderstorms, and they develop
in air that is rising rapidly. The cold front is steeper than the
warm front and so air is forced to rise faster there than it does
up the shallower slope of the warm front. At the warm front,
condensation in the rising air produces sheets of smooth-
looking cloud, often with drizzle or persistent rain or snow.
Frontal systems are responsible for much of the changeable
weather of temperate climates.

Jet streams and depressions

Air pressure is caused by the weight of the column of air
above a particular place, all the way to the top of the atmos-
phere. It cannot be felt because it is always there, but it is as
real as the pressure an elephant would exert if it sat on the
hood of a car.
Warm air and cold air lie side by side, separated by a
weather front. Warm air is less dense than cold air. This
means that there is less air—there are fewer air molecules—in
a column of warm air than there is in a similar column of
cold air. Pressure is therefore higher at the base of the cold air
column than it is at the base of the warm air column.
Air pressure decreases with height within both air columns
because there is less air above to exert pressure. Since the
densities of the warm and cold air are different, however, pres-
46 TEMPERATE FORESTS

sure decreases more rapidly in the denser cold air than it does
in the less dense warm air. The difference in pressure between
the two columns therefore increases with height, and it is
greatest at the tops of the columns. These are located at the
tropopause. This is the boundary between the lower atmos-
phere—called the troposphere—and the stratosphere above, and
it is the level above which the temperature ceases to decrease
with increasing height. Rising air stops when it reaches the
tropopause, because it is then at the same temperature and
therefore density as the air immediately above it.
Air tries to flow from a region of high pressure to a region
of low pressure at a speed proportional to the difference in
pressure, known as the pressure gradient. This is rather like the
gradient of a hill. Imagine that the dense, high pressure air
rises to form hills and the less dense, low pressure air forms
valleys and hollows. Air then flows from high to low pres-
sure, just as water flows down a hillside. Maps show the ele-
vation of the land by means of contour lines—lines joining
places that are all at the same height above sea level. If the
contour lines are close together it means the ground rises
steeply—a big increase in elevation for a short distance hori-
zontally.
Instead of variations in the height of the surface above sea
level, a weather map shows variations in atmospheric pres-
sure. Weather stations measure the pressure and meteorolo-
gists then adjust these measurements to show the sea-level
pressure. This is necessary because air pressure decreases with
altitude and so the measurements from weather stations are
affected by their elevations. Converting the pressures at all
the stations to sea-level pressure makes it easy to recognize
areas of high and low pressure and the rate at which pressure
changes between them.
Places where the air pressure is the same are joined togeth-
er by lines called isobars—from the Greek isos meaning
“equal” and baros meaning “weight.” If the isobars are close
together it means the pressure changes rapidly over a short
horizontal distance, and if they are widely spaced it means
the pressure remains fairly constant over a large area. This is
very similar to the way contour lines show hills, valleys, and
plains, but isobars illustrate the changes across an invisible
 CLIMATE OF TEMPERATE FORESTS 47

pressure surface rather than the solid ground surface. The clos-
er together the isobars are, the steeper the pressure gradient,
and because air flows down the pressure gradient like water
down a hillside, the stronger the wind.
Because of the rotation of the Earth, however, the air does
not move at right angles to the pressure gradient but parallel
to it. On a weather map, the wind blows parallel to the iso-
bars. In the Northern Hemisphere the air moves counter-
clockwise around a center of low pressure and clockwise
around a center of high pressure. These directions are
reversed in the Southern Hemisphere. When warm and cold
air lie side by side, the pressure gradient becomes steeper
with increasing height, because pressure decreases faster in
the cold air than in the warm air. Maps of the pressure surface
at different altitudes would show the isobars closing up.
Therefore the wind speed increases. It is called a thermal wind
because it is caused by a difference in temperature.
The biggest difference in temperature occurs where cold air
moving away from the poles meets warm air moving toward
the poles. Cold and warm air meet at several latitudes (see the
sidebar “General circulation of the atmosphere” on page 38),
but the sharpest and most continuous boundary occurs at
about 30°–70°N and S, where tropical air meets polar air.
At this boundary, wind speed increases with height and
reaches a maximum between 30,000 feet and 50,000 feet
(9,000–15,000 m). That is where a ribbon of wind, 60–300
miles (100–500 km) wide but only a mile or two thick, blows
at speeds of 120–240 MPH (200–400 km/h) and sometimes
faster in winter, when the temperature contrast is greatest.
The ribbon of wind is called the jet stream. In fact there are
several jet streams at different latitudes, but this is the most
constant one.
Thermal winds blow with the cold air to the left in the
Northern Hemisphere and to the right in the Southern
Hemisphere. In both hemispheres, therefore, the jet streams
blow from west to east (apart from an easterly jet stream that
forms in summer over Asia).
Even the most reliable jet stream does not blow all the time
or invariably from west to east. Jet streams twist and snake
about, and this can have dramatic effects on the weather
48 TEMPERATE FORESTS

below (see “Blocking” on page 49). Waves in the jet stream

produce ridges curving toward the pole and troughs curving
toward the equator. Places on the surface below a ridge are in
tropical air and those below a trough are in polar air. The
illustration shows how ridges and troughs in the jet stream
can affect the weather across the United States. Arrows in the
diagram show the direction of the wind.
Air is drawn into the jet stream on the upstream side of
ridges and flows out from it on the downstream side. Where
air converges to enter the jet stream, air subsides, producing
Ridges and troughs in high pressure at the surface. Divergence draws air upward,
the jet stream across producing low surface pressure. Fronts form between the
North America. areas of high and low pressure and the center of low pres-
The arrows show the sure becomes a depression. As the waves in the jet stream
wind direction and move eastward, like waves in a rope that is secured at one
shading indicates end and shaken up and down at the other, they drag the
the wind speeds in frontal systems and depressions with them, as shown in the
the jet stream. illustration.

100 mph
100 mph

cold air 150 mph

ridge

100 mph

ridge
trough

150 mph
200 mph

warm air warm air

 CLIMATE OF TEMPERATE FORESTS 49

L L

H
L
H

Jet stream and fronts.

Frontal depressions
develop beneath
undulations in the jet
stream, and as the
undulations move
from west to east they
jet stream cold front warm front occluded front drag the frontal
systems with them.

It is the front between tropical and polar air and the behav-
ior of the jet stream this produces that make the weather in
temperate latitudes so changeable and difficult to forecast.

Blocking
About 40,000 feet (12.2 km) above sea level, the jet stream
blows from west to east, but high mountain ranges, such as
the Rocky Mountains, slow the air passing over them and
slightly alter its horizontal course. The resulting deviations
affect the overlying air, all the way up to the jet stream,
producing very long waves—the distance between one wave
crest and the next is between about 2,500 miles (4,000 km)
50 TEMPERATE FORESTS

and 3,700 miles (6,000 km). The crests project to the north
and the troughs to the south.
Atmospheric movement transfers heat away from the
equator (see the sidebar “General circulation of the atmos-
phere” on page 38), but the efficiency of the process varies.
When it is relatively inefficient, warm air accumulates in the

Blocking. These four diagrams show how waves in the jet stream
develop over a period of a few weeks, over the course of the index
cycle. 1. The jet stream flows west to east, with only slight
undulations. 2. The undulations become more pronounced. 3. The
undulations are now very pronounced; in some places the jet stream
blows in a northerly and in other places southerly direction. 4. The
flow has broken up and the jet stream circulates around cells of high
pressure in the north and low pressure in the south; these cells remain
stationary for some time, deflecting weather systems to the north or
south. This is called “blocking.”
 CLIMATE OF TEMPERATE FORESTS 51

south and air in the north becomes colder, increasing the

contrast in temperature on either side of the front separating
tropical and polar air. It is the temperature contrast that
drives the jet stream, so when the contrast increases, the jet
stream accelerates.
At this point the effect of the Earth’s rotation becomes
important. It is called the Coriolis effect, abbreviated as CorF
(because it was once thought to be a force), and in the
Northern Hemisphere it deflects moving air to the right.
CorF is greatest at the poles, and at the equator its value is
zero. Its magnitude is directly proportional to the speed of
the moving air. As the jet stream accelerates, it temporarily
overcomes the influence of the CorF and swings to the left.
This carries it northward, where the CorF increases, swinging
it to the right again. The result is a series of swings that make
the waves bigger and closer together. The diagram shows the
sequence of events.
By stage 3 in the sequence, the waves have become ex-
treme. The troughs are bringing polar air far to the south. In
winter this produces cold waves, with a sudden drop in tem-
perature. Elsewhere, the ridges carry tropical air far to the
north, producing unusually warm weather. The jet stream is
blowing from the north in some places and from the south in
others, and the two streams are fairly close together.
In stage 4, some of the air has taken a shortcut. The jet
stream is once more blowing from west to east, but to its south
there are sections of the jet stream that have been cut off from
the main flow. They now form isolated pools of air, about 900
miles (1,450 km) across, with winds flowing around them.
Those in the north, at about 55°N, consist of tropical air, and
those in the south, at about 33°N, of polar air. Air circulates
clockwise around the pools of tropical air and counterclock-
wise around the pools of polar air. In the Northern Hemi-
sphere, air circulates counterclockwise around centers of low
pressure and clockwise around centers of high pressure. The
pools of tropical air are therefore anticyclones or highs, and the
pools of polar air are depressions or lows. Anticyclones bring
fine, settled weather. Depressions bring wet weather.
The sequence takes three to eight weeks to complete, but at
the end, the isolated anticyclones and depressions remain,
52 TEMPERATE FORESTS

while the jet stream resumes its westerly flow to the north.
The anticyclones and depressions are stationary, but weather
systems continue moving from west to east, usually traveling
at about 20 MPH (30 km/h) in summer and 30 MPH (50
km/h) in winter. When they reach the stationary highs and
lows their way is blocked and so they divert around them.
The process is called blocking, and the pools of air responsible
for it are blocking highs and blocking lows.
Blocking highs and lows can persist for days or even weeks.
They bring prolonged spells of unchanging weather. In the
southern United States, at about the latitude of Dallas, Texas,
this is most likely to be cool, overcast, wet weather. In
Canada, at about the latitude of Edmonton, Alberta, it is usu-
ally fine weather that can contribute to the development of a
drought. At the same time, frontal systems are diverted into
regions to the north and south of the block that might other-
wise have missed them.

Continental and maritime climates

Broad-leaved deciduous trees cannot tolerate prolonged
drought. They need moisture. This requirement for water
means that temperate forests do not grow deep inside conti-
nents, far from the ocean. They prefer a maritime climate, also
called an oceanic climate.
As the name suggests, a maritime climate occurs near
coasts and it is produced by maritime air—air masses that
have formed over the ocean. As well as being moist, maritime
air is relatively mild in winter and cool in summer, so the dif-
ference between summer and winter temperatures is much
smaller than it is in the interior of a continent. At Belfast,
Northern Ireland, for example, the average temperature in
the warmest month is 58°F (14.5°C) and that of the coldest
month 39°F (4°C). The difference between the two, or the
temperature range for Belfast, is 19°F (10.5°C). Contrast this
with Dodge City, Kansas, where the average temperature is
78°F (25.5°C) in the warmest month and 29°F (–1.5°C) in the
coldest month, giving a temperature range of 49°F (27°C).
Belfast has a maritime climate and Dodge City has a conti-
nental climate. The most extreme examples of maritime cli-
 CLIMATE OF TEMPERATE FORESTS 53

mate are found on oceanic islands. The Azores, a group of

islands in the North Atlantic Ocean about 800 miles (1,287
km) to the west of Portugal, are in approximately the same
latitude as Dodge City. The average temperature in the Azores
ranges from 57°F (14°C) to 71°F (21.5°C) in the warmest
month, a difference of 14°F (7.5°C).
Maritime climates are also wet. Moist air from the ocean is
forced to rise as it crosses the coast. Some of its water vapor
condenses to form clouds and it rains (see “Why clouds
form” on page 56). The Azores receive an average 47 inches
(1,186 mm) of rain a year and Belfast 33 inches (846 mm),
but Dodge City receives only 20 inches (518 mm). Not every
oceanic island has a wet climate, however. Seymour Island,
for example, has a desert climate. One of the Galápagos
Islands, about 650 miles (1,045 km) from the coast of
Ecuador, it is low-lying and surrounded by the cold water of
the Peru Current. Seymour Island receives only four inches
(102 mm) of rain a year, and no rain at all from the end of
April until the beginning of January.
These contrasts make it sound as though a line could be
drawn between an area with a continental climate and an
adjacent area with a maritime climate and a person could
cross from one to the other. It would be like crossing the
road—continental on one side and maritime on the other. It
is not like that, however. With increasing distance toward or
away from the coast, the climate changes quite gradually.
The change is more abrupt where a range of mountains runs
parallel to the coast and not very far inland. Approaching air
loses a large proportion of its moisture as it crosses the moun-
tains (see the sidebar “Why it rains more on mountainsides”
on page 3) and the region on the lee side lies in a rain shadow
and has an extremely continental climate.
The Coast Ranges in California demonstrate the effect. San
Diego, California, has a temperature range of 14°F (8°C), and
Phoenix, Arizona, on the inland side of the mountains, has a
range of 39°F (21.5°C). There is less contrast in rainfall.
Phoenix receives an average 7.5 inches (190 mm) of rain a
year and San Diego receives 10 inches (264 mm). San Diego
owes its relatively dry climate to the fact that air approaching
from the Pacific Ocean has to cross the cold California
54 TEMPERATE FORESTS

Current. Contact with the cool water reduces the temperature

of the air, causing some of its water vapor to condense and
thus partly drying the air shortly before it reaches the coast.
Scientists who study climates need a convenient way to
describe the extent to which the climate in a particular place
is continental or maritime. The most widely used method
produces a number called the Conrad index, devised in 1946
by the American scientist V. Conrad. A Conrad index of 0
means a climate is fully maritime and 100 means it is fully
continental. The table lists the Conrad index values for the
places mentioned.
The table shows that the Azores, Belfast, and San Diego
have maritime climates, and that Dodge City and Phoenix
have climates of a more continental type.

Conrad index of continentality

Place Index value

Azores 2.98
Belfast 5.75
Dodge City 48.03
Phoenix 39.13
San Diego 6.04

How climates are classified

Temperate forests prefer a maritime climate, but maritime cli-
mates are not all the same. In some, approximately the same
amount of rain or snow falls in each month. Others have a
dry season in which there is very little rain, and the dry sea-
son may be in the summer or the winter. There are climates
with a hot summer and those with a warm or even cool sum-
mer. Describing a climate merely as maritime or continental
provides some information, but it does not amount to a com-
plete description, because it does not reveal what kind of
maritime or continental climate it is. A system is needed for
classifying climates, like the system for classifying plants and
animals. Such a system will allow people to distinguish
between the different types of maritime and continental cli-
 CLIMATE OF TEMPERATE FORESTS 55

mates, just as the system of biological classification allows

people to distinguish between different kinds of mammals or
birds. Then, when a climate type is named, everyone will
know precisely what is meant.
In fact, there are many classification systems. The ancient
Greeks devised the earliest ones, but those in use today have
grown from systems that were developed in the 19th century.
Botanists and plant geographers made up most of them.
Plant geographers are scientists who define vegetation types
and plot their geographic distributions, and the names they
used to describe climates reflect this. They described “savan-
na climate,” “tropical rainforest climate,” and “tundra cli-
mate,” for example. Some of these names are still used, but
others, such as “penguin climate,” have been dropped.
The only information available to scientists in the early
days referred to average temperature and rainfall and to veg-
etation type. Much more detailed information is available
nowadays, about both the climate itself and its effect on
plants, animals, and people living in it. There are many more
recording stations, satellites monitor the surface and atmos-
phere constantly, and computers make it possible to process
large amounts of data. As a result, climate classifications have
steadily grown more detailed and complicated.
The variety of classification systems is not due to dis-
agreements among scientists or to the search for a perfect
scheme. There can be no single classification that satisfies
the needs of every user. Some schemes help geographers
studying plant distribution. Others relate to crop require-
ments and are designed for agricultural use. There are also
some that simply provide a list of descriptive names and
there are highly technical classifications for the use of cli-
mate scientists.
Many of the most widely used schemes are based on one
devised by the Russian-born German meteorologist and cli-
matologist Wladimir Peter Köppen (1846–1940). The first
version was published in 1900, and in 1936 Köppen issued a
revised version of his classification.
The Köppen classification first divides climates into six
general types, defined principally by temperature and desig-
nated by letters.
56 TEMPERATE FORESTS

A Tropical rainy climates; the average temperature

never falls below 64.4°F (18°C).
B Dry climates.
C Warm temperate rainy climates; temperatures in the
warmest month are higher than 50°F (10°C) and in
the coldest month are between 26.6°F (–3°C) and
64.4°F (18°C).
D Cold boreal forest climates; temperatures in the
warmest month are higher than 50°F (10°C) and in
the coldest month are lower than 26.6°F (–3°C).
E Tundra climate; temperatures in the warmest month
are between 32°F (0°C) and 50°F (10°C).
F Perpetual frost climate; the average temperature
never rises above 32°F (0°C).
These categories relate to plant growth. For example, trees
will not grow where summer temperatures remain below
50°F (10°C), some plants will not grow where winter temper-
atures fall below 64.4°F (18°C), and an average temperature
of 26.6°F (–3°C) means there will be frost and probably snow.
Köppen then divided each of these six categories into a
number of further types based mainly on the amount and
distribution of rain that they receive. This produced a total of
24 climate types. For example, a Cfb climate is a warm, tem-
perate rainy climate that is mild in winter, warm in summer,
and moist throughout the year. A Bwh climate is characteris-
tic of a hot desert.
Finally, 24 additional letters are used to add more detailed
information. Adding f, for example, means there is sufficient
rain for plants to grow healthily in all seasons. Adding s
means there is a dry season in summer, and adding n means
that fog is frequent.
The Köppen classification is the one most widely used by
geographers. When you see a map of the world showing cli-
mate types, it is most likely to be based on the Köppen scheme
and it may use the Köppen letters in addition to colors.

Why clouds form

Water vapor is a gas consisting of separate water molecules,
all moving rapidly in every direction. When two molecules
 CLIMATE OF TEMPERATE FORESTS 57

collide they bounce off each other and fly apart. The more
energy the molecules possess, the faster they travel and the
harder they strike when they collide. If the molecules lose
energy they slow down, and if they slow down sufficiently,
molecules cling to one another when they collide, rather
than bouncing apart. When the molecules begin to join
together in small groups they cease to be a gas. They are then
droplets of liquid water. The process by which water vapor
turns into liquid water is called condensation. When it occurs
in the air, the resulting mass of minute water droplets is visi-
ble as cloud or fog.
For cloud to form, the water vapor in the air must lose so
much energy that its molecules join together. Air loses energy
when it expands and gains energy when it is compressed.
When you pump up a bike tire, the energy you expend in
compressing the air entering the tire is transferred to the air
molecules. They move faster and therefore they strike the sides
of the pump harder. That makes the molecules in the pump
casing vibrate faster, and you feel that increased vibration as a
rise in temperature. In other words, when you compress air its
temperature rises. If you then undo the tire valve and let out
the air, the escaping air will feel cold. As the air expands, its
molecules move farther apart and slow down, so the air tem-
perature falls. When air expands, its temperature falls.
Water vapor loses energy if it expands, and it will expand if
it moves into a region of the atmosphere where the sur-
rounding air pressure is lower. Air pressure decreases with
height, because the weight of overlying air decreases. The
reduction in pressure allows the air to expand, expansion
causes the temperature to fall, and therefore the temperature
of the air decreases with height. Walkers preparing to climb
high into the hills pack extra sweaters because they know it
will be colder up there than it is down in the valley. The tem-
perature falls by an average of 3.5°F every 1,000 feet
(6.5°C/km). Air also expands and cools if it is forced to rise,
for example by crossing a mountain, and sinking air is com-
pressed and its temperature increases. These changes are said
to be adiabatic (see the sidebar).
Air temperature decreases with height, and the amount of
water vapor that air is able to hold varies with the temperature.
58 TEMPERATE FORESTS

Adiabatic cooling and warming

Air is compressed by the weight of air above it. Imagine a balloon partly inflated with air
and made from some weightless substance that totally insulates the air inside. No matter
what the temperature outside the balloon, the temperature of the air inside remains the
same.
Imagine the balloon is released into the atmosphere. The air inside is squeezed between
the weight of air above it, all the way to the top of the atmosphere, and the denser air
below it.
Suppose the air inside the balloon is less dense than the air above it. Denser air will push
beneath it and the balloon will rise. As it rises, the distance to the top of the atmosphere
becomes smaller, so there is less air above to weigh down on the air in the balloon. At the
same time, as the balloon moves through air that is less dense, it experiences less pressure
from below. This causes the air in the balloon to expand.
When air (or any other gas) expands, its molecules move farther apart. The amount of
air remains the same, but it occupies a bigger volume. As they move apart, the molecules
must “push” other molecules out of their way. This uses energy, so as the air expands its
molecules lose energy. Because they have less energy they move more slowly.
When a moving molecule strikes something, some of its energy is transferred to what-
ever it strikes, and part of that energy is converted into heat. This raises the temperature of
the struck object by an amount related to the number of molecules striking it and their
speed.
In expanding air, the molecules are moving farther apart, so a smaller number of them
strike an object each second. They are also traveling more slowly, so they strike with less
force. This means the temperature of the air decreases. As it expands, air cools.
If the air in the balloon is denser than air below, it will sink. As it sinks, the pressure on
the air will increase, its volume will decrease, and its molecules will acquire more energy.
Its temperature will increase.
This warming and cooling has nothing to do with the temperature of the air surround-
ing the balloon. It is called adiabatic warming and cooling, from the Greek word adia-
batos, meaning “impassable,” suggesting that the air is enclosed by an imaginary bound-
ary through which heat is unable to pass.

The warmer the air is, the more water vapor it can hold. At
80°F (27°C), for example, one pound of air at sea-level pres-
sure can hold about 0.3 ounces of water vapor (20 g/kg), but
 CLIMATE OF TEMPERATE FORESTS 59

at 32°F (0°C) it can hold only 0.06 ounces (3.5 g/kg). When
the concentration of water vapor reaches the maximum the
air can contain, the air is saturated. If more water vapor is
then added, or if the temperature falls, some of the water
vapor condenses into liquid droplets. The amount of mois-
ture present in the air is called the humidity. There are several
ways to measure this, explained in the sidebar.
Atmospheric water vapor will condense if the air tempera-
ture decreases. Since air temperature decreases with height,
water vapor condenses when moist air rises.
There are three reasons why air may rise. It rises at weather
fronts, where warm air rises up a cold front (see “Air masses
and fronts” on page 41), and when a moving air mass crosses

Humidity
The amount of water vapor air can hold varies according to the temperature. Warm air
can hold more than cold air. The amount of water vapor present in the air is called the
humidity of the air. This is measured in several ways.
The absolute humidity is the mass of water vapor present in a unit of volume of air,
measured in grams per cubic meter (1 gram per cubic meter = 0.046 ounces per cubic
yard). Changes in the temperature and pressure alter the volume of air, however, and this
changes the amount of water vapor in a unit volume without actually adding or removing
any moisture. The concept of absolute humidity takes no account of this, so it is not very
useful and is seldom used.
Mixing ratio is more useful. This is a measure of the amount of water vapor in a unit
mass of dry air—air with all the water vapor removed. Specific humidity is similar to mixing
ratio but measures the amount of water vapor in a unit mass of air including the moisture.
Both are reported in grams per kilogram. Since the amount of water vapor is always very
small, seldom accounting for more than 4 percent of the mass of the air, specific humidi-
ty and mixing ratio are almost the same thing.
The most familiar term is relative humidity. This is the measurement you read from
hygrometers, either directly or after referring to tables—and it is the one you hear in
weather forecasts. Relative humidity (RH) is the amount of water vapor in the air expressed
as a percentage of the amount needed to saturate the air at that temperature. When the
air is saturated the RH is 100 percent (the “percent” is often omitted).
60 TEMPERATE FORESTS

high ground. Air also rises when, on warm, sunny days, the
ground is heated. Air next to the ground is warmed by con-
tact and rises by convection. Water vapor in air rising up a
front condenses to form frontal cloud. Condensation in air ris-
ing over high ground forms orographic cloud. Moist air rising
by convection forms convective cloud—often seen on fine
summer afternoons as puffy, white, fair-weather cumulus.
There are different ways for clouds to form, and they form
at varying heights because of variations in the amount of
water vapor present in the air. Consequently, clouds vary

How clouds are classified

There is an internationally accepted scheme for classifying clouds on the basis of their
height and appearance. Clouds are grouped into 10 distinctive types called genera (singu-
lar genus). The genera are subdivided into species and species into varieties. There are also
accessory clouds (small clouds seen in association with a bigger cloud of a different type)
and supplementary cloud features (extensions or protrusions from a cloud). Genera and
species names have standard abbreviations. Stratocumulus (Sc), for example, may form in
an almond or lens shape, producing the species lenticularis (len), abbreviated as Sclen.
Cloud genera are described as low-level, medium-level, or high-level according to
the height at which they most commonly form, although clouds can form at higher or low-
er levels. Large storm clouds, which have a low-level base but extend to a great height, are
counted as low-level clouds, mainly for convenience. Most medium-level clouds have names
beginning with the prefix alto- and the names of high-level clouds have the prefix cirr-.

CLOUD GENERA
Low-level clouds. Cloud base from sea level to 1.2 miles (2,000 m).

Stratus (St). An extensive sheet of featureless cloud that will produce drizzle or fine
snow if it is thick enough.
Stratocumulus (Sc). Similar to St, but broken into separate, fluffy-looking masses. If
thick enough, it also produces drizzle or fine snow.
Cumulus (Cu). Separate white, fluffy clouds, usually with flat bases. There may be
many of them, all with bases at about the same height, and they may merge into
a single cloud.
 CLIMATE OF TEMPERATE FORESTS 61

greatly in appearance. A way is needed to describe these dif-

ferent kinds of cloud clearly, so meteorologists use a system
of cloud classification (see the sidebar) based on the height
and appearance of the clouds.

Rain, snow, and hail

A cloud is made from water droplets, ice crystals, or a mixture
of both. Cloud droplets are tiny—typically about 0.0004 inch
(0.01 mm) across. There are usually about 283 droplets in

Cumulonimbus (Cb). Very large cumulus, often towering to a great height. Because
they are so thick, Cb clouds are often dark at the base. If the tops are high
enough they will consist of ice crystals and may be swept into an anvil shape (the
supplementary feature incus).

Medium-level clouds. Cloud base from 1.2–2.5 miles (2,000–4,000 m) in polar

regions, 4–5 miles (6,000–8,000 m) in temperate and tropical regions.

Altocumulus (Ac). Patches or rolls of cloud joined to make a sheet. Ac is sometimes

called “wool-pack cloud.”
Altostratus (As). Pale, watery, featureless cloud, forming a sheet through which the
Sun may be visible as a white smudge.
Nimbostratus (Ns). A large sheet of featureless cloud, often with rain or snow, that is
thick enough to obscure the Sun, Moon, and stars completely. It makes days dull
and nights very dark.

High-level clouds. Cloud base from two to five miles (3,000–8,000 m) in polar
regions, three to 11 miles (5,000–18,000 m) in temperate and tropical regions. All high-
level clouds are made entirely from ice crystals.

Cirrus (Ci). Patches of white, fibrous cloud, sometimes swept into strands with curl-
ing tails (“mares’ tails”).
Cirrocumulus (Cc). Patches of thin cloud, sometimes forming ripples, fibrous in
places, with no shading that would define their shape.
Cirrostratus (Cs). Thin, almost transparent cloud forming an extensive sheet and just
thick enough to produce a halo around the Sun or Moon.
62 TEMPERATE FORESTS

every cubic foot of air (100,000/L) and they fall very slowly.
Droplets are constantly evaporating and water vapor is con-
stantly condensing to produce new ones. A cloud looks very
still and peaceful, but there is a great deal going on inside it.
Cloud droplets are too small to fall as rain, because
droplets falling from the base of the cloud enter much drier
air and evaporate long before they have time to reach the
ground. Even raindrops can evaporate between the cloud and
the ground. You sometimes see showers of them, looking like
a gray veil, called virga or fallstreaks, hanging from the base of
a cloud.
There are two ways for raindrops to form, depending on
whether or not the cloud contains ice crystals. Most of the
clouds in middle latitudes contain ice because they extend
above the altitude at which the temperature reaches freezing.
If ice crystals are present, the cloud will also contain liquid
droplets that are below freezing temperature but remain
unfrozen because there are too few solid particles onto which
ice crystals can grow. Instead of freezing, these supercooled
droplets evaporate and the water vapor is then deposited
onto the ice crystals. The ice crystals grow bigger and the
supercooled droplets become fewer.
As they grow bigger, the ice crystals begin to collide with
each other, forming snowflakes, and when the snowflakes are
big and heavy enough they start to fall. As they fall, the
snowflakes collide with more supercooled liquid droplets
that freeze onto them, making the snowflakes still larger.
Buffeting by air currents breaks some of the snowflakes apart,
releasing tiny ice splinters that are carried aloft by rising air.
These are solid particles onto which more ice crystals can
form.
Other snowflakes continue falling, and growing larger as
they do so. When they enter warmer air in the lower part of
the cloud or between the base of the cloud and the ground,
the snowflakes melt and become raindrops. Most of the rain
that falls in temperate latitudes consists of melted snow, even
in the middle of summer.
If the air temperature between the cloud base and the
ground is lower than about 39°F (4°C), the snowflakes will
not melt. They will fall as snow, and the colder the air, the
smaller the snowflakes will be.
 CLIMATE OF TEMPERATE FORESTS 63

Some clouds contain no ice crystals. They do contain some

much larger droplets, however. These fall faster than the
small droplets and as they fall they gather up and merge with
the smaller droplets they meet along the way. Whether it
forms in this way or as a snowflake that melts, a raindrop
consists of about 1 million cloud droplets.
Raindrops do not always fall from their clouds. If the cloud
is a big cumulonimbus storm cloud, the currents of rising air
inside it may be strong enough to sweep up raindrops from
near the bottom of the cloud and raise them to the top,
where they freeze into ice pellets. The pellets then fall again
and are caught in rising air currents and carried back to the
top of the cloud. As it moves through the cloud, a pellet col-
lides with supercooled water droplets that freeze onto it.
Large droplets spread over the surface of the pellet and freeze
as a layer of clear ice. Small droplets freeze faster, forming
white, opaque ice made from tiny crystals with pockets of air
trapped between them. The pellet is then a hailstone.
Eventually it will either enter a current of air moving down-
ward that will throw it out of the base of the cloud, or it will
grow so big that the rising air currents cannot support its
weight and so it falls from the cloud.
Most hailstones are no bigger than a pea, but a severe
storm can drop hailstones the size of golf balls and occasion-
ally they can be even larger. Hailstones the size of softballs
have been recorded, and on September 3, 1970, a hailstone
5.5 inches (14 cm) across, weighing 1.67 pounds (766 g), fell
at Coffeyville, Kansas.
A hailstone grows by acquiring ice during its travels
between the top and base of the cloud. The more of these cir-
cuits it makes, the bigger the hailstone will be. It falls from
the cloud when it is too heavy to be carried upward, so the
stronger the vertical air currents, the more circuits the hail-
stone can make and the bigger it can become. The size of the
hailstones therefore indicates the size and energy of the
cloud producing them.

Ice storms
Hailstorms can batter farm crops to destruction, but forest
trees can withstand their blows. A tree is more likely to be
64 TEMPERATE FORESTS

damaged by a layer of clear ice that is heavy enough to snap

its branches. Ice is surprisingly heavy. A layer of ice three
inches (7.5 cm) thick on a branch one foot (30 cm) thick and
15 feet (4.6 m) long would weigh about 83 pounds (182 kg).
Clear ice can accumulate very quickly, with startling
results: Roosting birds have been frozen onto tree branches,
ground-nesting birds have had their wings coated with ice so
they could not fly, and prowling cats have been frozen to the
ground. When this happens it is called an ice storm. Ice forms
on roads, airfield runways, railroad tracks, power and tele-
phone lines, and radio and TV masts as well as on trees. An
ice storm can bring transport to a standstill, cause power out-
ages and telephone failures, and do structural damage to
transmitter towers.
Ice storms usually happen ahead of an approaching warm
front (see “Air masses and fronts” on page 41) when the air
ahead of the front is below freezing temperature, the warm
air behind the front is just above freezing, and there is a
strong wind.
At the front, warm air is rising above the adjacent cold air,
but at a very shallow angle of 0.5° to 1°. The shallowness of
the angle means that the front overhangs a large surface area.
When the base of the cloud overhead is about 6,500 feet (2
km) above the ground, the line where the warm front touch-
es the surface may be up to 170 miles (275 km) away.
If the temperature of the air at ground level behind the
front is well above freezing, the cloud along the lower part of
the front will be warm enough to deliver rain rather than
snow, but some of this rain will be barely above freezing tem-
perature. The rain falls through the frontal zone and into the
colder air ahead of the front—and in fact below it. This air is
well below freezing temperature, and because the air is very
cold, so are all the solid objects on the ground, including the
trunks and branches of the trees. Rain that is barely above
freezing temperature cools a little more as it falls through the
cold air. When it strikes a solid surface it immediately starts
to freeze.
Raindrops are too large to freeze instantaneously. The part
of the drop that makes contact with the surface freezes at
once. The remainder of the drop then spreads outward as a
 CLIMATE OF TEMPERATE FORESTS 65

liquid coating that freezes on contact with the surface. Each

raindrop freezes into a thin sheet of ice.
Freezing releases latent heat, however. This warms the sur-
rounding air and will prevent more ice from forming unless
the warm air is quickly removed. The strong wind achieves
this, sweeping away the warmed air and replacing it with
cold air.
As more and more raindrops strike the surface and freeze,
together they rapidly build a thick coating of clear ice.
Because of the wind, the rain falls at an angle and so the ice
coats the sides of objects facing into the wind much more
thickly than it does the opposite, sheltered sides.

Dew and frost

When a cool night with only the lightest of winds follows
a warm day, water vapor often condenses onto surfaces,
especially the leaves of plants growing close to the ground.
This is dew. If the plants are covered with dew early in the
morning it often means the coming day will be fine and
sunny.
During the day, the ground and plant surfaces absorb the
warm sunshine. As their temperature rises, the surfaces also
radiate their own warmth into the sky, but they absorb heat
faster than they lose it, so they grow steadily warmer. Air
next to them is warmed by contact. Water enters the warm
air by evaporation and transpiration (see “Evaporation and
transpiration” on page 66).
After sunset the ground and plants cease to absorb heat,
but they continue to radiate their own heat, so they cool
down. Their temperature falls steadily until about an hour
before dawn, when it reaches its lowest point. A shallow layer
of cool air collects close to the ground. If there is more than
the slightest breeze, this cold air will be swept away and
replaced by warmer air from above, but if the air is fairly still
the temperature in this surface layer remains low.
The relative humidity (RH) increases as the temperature
falls (see the sidebar “Humidity” on page 59). If the RH
exceeds 100 percent, water vapor will condense out of the air,
provided there are surfaces onto which it can condense.
66 TEMPERATE FORESTS

Leaves inside the layer of cool air and at the same tempera-
ture are ideal.
The air must not be completely still. If it is, condensation
will quickly remove enough water vapor from the surface layer
to reduce the RH to below 100 percent. Condensation will
then cease and, although dew will form, the amount will be
too small to be visible. A gentle movement of air will replenish
the surface layer with a steady supply of moist air from above.
In winter it sometimes happens that the temperature drops
to below freezing after dew has formed. When this happens
the dewdrops freeze, coating surfaces with clear ice, often
with round lumps due to the shape of the dewdrops.
Water vapor condenses to form dew at the dew-point tem-
perature. If the air is fairly dry, the dew-point temperature
may be below freezing. Dew will not form in summer under
these conditions, but in winter the temperature in the surface
layer may well fall to below freezing. Water vapor will then
be deposited directly as ice crystals that cover surfaces with
white hoar frost.
Frost protects plants against the cold. This is because ice
conducts heat poorly, so it reduces the amount of warmth
they radiate away. At the same time, the freezing of dewdrops
and deposition of water vapor as ice crystals release latent
heat that warms the underlying surface.
Plants suffer damage when the air is dry and the tempera-
ture falls low enough to freeze the liquid inside their tissues.
This produces black frost. There is no surface coating of ice,
but the frozen leaves die and turn black.
Weather forecasts often warn of air frost or ground frost. Air
frost is the one that can harm plants. It happens when the air
temperature falls below freezing. Ground frost occurs when
the ground temperature falls below freezing, but the air tem-
perature remains above freezing.

Evaporation and transpiration

A water molecule comprises two atoms of hydrogen (H) and
one of oxygen (0), chemical formula H2O. The molecule is
arranged with both hydrogen atoms on the same side of the
oxygen atom. This gives the molecule a negative electromag-
 CLIMATE OF TEMPERATE FORESTS 67

netic charge on the oxygen side and a positive charge on the

hydrogen side. The charges balance, so the molecule is neu-
tral overall, but it carries a charge at each end.
When two water molecules are close together, an attach-
ment called a hydrogen bond forms between a hydrogen atom
of one molecule and the oxygen atom of the other. Molecules
form small groups held together by hydrogen bonds. The
groups are constantly breaking apart and forming again; they
move around freely, sliding past one another, and they will
fill every corner of a container. This is liquid water.
If a molecule absorbs enough energy, it will vibrate so vio-
lently that it breaks away from its group. It can then enter the
air, where it is able to move freely. Free water molecules mov-
ing among the air molecules are water vapor. The transfor-
mation of liquid into vapor is evaporation. It is the opposite of
condensation.
The energy the molecules absorb from their surroundings,
which breaks the hydrogen bonds, does not alter the temper-
ature of the water. It is latent heat, and when water vapor con-
denses precisely the same amount of latent heat is released.
Molecules are constantly escaping from every exposed
water surface. When they leave the surface they enter a thin
boundary layer of air, where they join other water molecules.
Together these molecules exert a pressure that drives mole-
cules back into the liquid. Consequently, water molecules are
entering the liquid as well as leaving it.
If the water surface is at a higher temperature than the air
beyond the boundary layer, water molecules will have
enough energy to escape into the air. More molecules will
leave the surface than enter it, and the water will evaporate.
Water will also evaporate if there is a wind. This is because
the wind removes moist air from above the water surface and
replaces it with dry air. Consequently, wind has a powerful
drying effect—which is why people hang out laundry to dry
in the wind. Where the wind blows mainly from a particular
direction, it can dry the leaf buds and the growing tips of
stems on the exposed side of a tree. This slows or even pre-
vents growth on that side of the tree, while the sheltered side
grows normally. The effect is to shape the tree in a way that
makes it look as though it is bending before the wind.
68 TEMPERATE FORESTS

Trees, shrubs, and the smaller plants that grow on the for-
est floor obtain water from the soil and use it to transport
nutrients and sugars to all parts of the plant (see “How trees
find food” on page 77). Water finally enters cells lying just
below the surface of leaves. These are the cells from which
the plant expels oxygen and absorbs carbon dioxide (see
“Photosynthesis and respiration” on page 79) through small
pores, called stomata. While its stomata are open, however,
water molecules also escape through them. The water is
replaced by water from the ground, and there is a steady flow
of water from the ground, through the plant, and into the air.
This release of water into the air is called transpiration.
Transpiration releases a substantial amount of water into
the air. In summer, an oak tree transpires about 185 gallons
(700 liters) of water a day. It is possible to measure the rate of
transpiration under laboratory conditions, but in the open
air it is extremely difficult to separate the water entering the
air by transpiration from that evaporating from wet surfaces.
Scientists measure the two together, calling them evapotran-
spiration.
Transpiration absorbs latent heat from the leaves, cooling
them. This prevents them from overheating in hot weather.
At the same time, evapotranspiration absorbs so much latent
heat that it has a significant effect on the climate inside a for-
est. Depending on the tree species, the air inside a temperate
forest can be up to 8°F (4°C) cooler than the air outside. The
difference is due partly to the shade cast by the trees but part-
ly to evapotranspiration.
The amount of water entering the air through evapotran-
spiration also makes the air inside a forest moister than air
outside. The relative humidity (see the sidebar “Humidity”
on page 59) varies with the seasons, but on average it is 4–9
percent higher inside a forest than it is in adjacent open
country. Air above the forest is also moister than the sur-
rounding air, so forests encourage cloud formation and rain-
fall.
Forests move water from the ground to the air. Forests on
hillsides help to prevent soil erosion and flooding in the val-
ley, and temperate forests everywhere have an important
influence on the local climate.
 CLIMATE OF TEMPERATE FORESTS 69

Thunderstorms and gales

Evaporation absorbs latent heat, and the condensation of
water vapor into liquid water releases precisely the same
amount of latent heat. Just as evapotranspiration cools the
air inside a forest, condensation inside a cloud warms the air,
sometimes with dramatic results.
Warm air rises and, as it does so, it expands and its temper-
ature falls adiabatically (see the sidebar on page 58). Air tem-
perature decreases with increasing height, but the rate at
which this happens is not necessarily the same in still and ris-
ing air. If the temperature of the rising air falls to that of the
surrounding air, all the air will have the same density and
the air will rise no farther. This air is said to be stable. While
the rising air remains warmer than the surrounding air, it will
continue rising. Air in this condition is unstable. It is also pos-
sible for stable air to become unstable while it is rising. This
air is conditionally unstable (see the sidebar).
Water vapor continues condensing as unstable air rises,
producing “heaped” clouds. If the air is sufficiently moist
and sufficiently unstable it will produce cumulonimbus
storm clouds (see the sidebar “How clouds are classified” on
page 60).
Air is rising strongly inside a big cumulonimbus. Conden-
sation and deposition produce ice crystals, snowflakes, hail-
stones, and raindrops (see “Rain, snow, and hail” on page 61).
Snow, hail, and rain fall from high in the cloud and as they fall
they drag cold air down with them, producing downdrafts.
Eventually, the cold downdrafts chill the air in the updrafts,
preventing air from rising. The cloud then dissipates, some-
times dropping all of its water in a brief cloudburst.
While the cloud remains active, particles carrying a posi-
tive electrical charge accumulate near the top of the cloud
and particles with negative charge accumulate near the base.
Scientists are uncertain of just why this happens, but after a
time the upper part of the cloud carries a positive electrical
charge and the lower part carries a negative charge. The neg-
ative charge at the base of the cloud then induces a positive
charge on the ground surface below. When these charges
have accumulated sufficiently to overcome the electrical
resistance of the air, a spark travels between them.
70 TEMPERATE FORESTS

Lapse rates and stability

Air temperature decreases (or lapses) with increasing height. The rate at which it does so
is called the lapse rate. Although all air contains some water vapor, air that is not saturat-
ed with moisture—all of its moisture is present as vapor rather than liquid droplets or ice
crystals—is said to be dry. When dry air cools adiabatically, it does so at 5.4°F for every
1,000 feet (9.8°C/km) that it rises. This is known as the dry adiabatic lapse rate (DALR).
When the temperature of the rising air has fallen sufficiently, its water vapor will start
to condense into droplets. Condensation commences at the dew-point temperature and
the height at which this temperature is reached is called the lifting condensation level.
Condensation releases latent heat, which warms the air. Latent heat is the energy that
allows water molecules to break free from each other when liquid water vaporizes or ice
melts. It does not change the temperature of the water or ice, which is why it is called
latent, meaning “hidden.” The same amount of latent heat is released, warming the sur-
roundings, when water vapor condenses and when liquid water freezes. Consequently,
the rising air then cools at a slower rate, known as the saturated adiabatic lapse rate
(SALR). The SALR varies, depending on the rate of condensation, but it averages 3°F per
1,000 feet (6°C/km).
The actual rate at which the temperature decreases with height in air that is not
rising is called the environmental lapse rate (ELR). It is calculated by comparing the

The spark is lightning. It may travel between different

parts of a cloud or between one cloud and another. These are
seen as sheet lightning. It may also travel between the cloud
and the ground, as forked lightning.
Lightning releases so much energy that it heats the air next
to the spark by up to 54,000°F (30,000°C) in less than one
second, making the air expand so violently that it explodes
and then contracts again. The explosion is heard as thunder.
Lightning can instantly vaporize water inside the trunk
and branches of a tree. The sudden expansion of the water
makes the tree explode. The explosion may destroy the tree,
but more commonly it strips away some of the bark.
When lightning strikes the forest floor, the electric current
may not be sustained for long enough for the heat to ignite
dry material. This is called cold lightning. Hot lightning sustains
 CLIMATE OF TEMPERATE FORESTS 71

surface temperature, the temperature at the tropopause (it is about –85°F; –65°C at
the equator), and the height of the tropopause (about 10 miles; 16 km over the
equator).
If the ELR is less than both the DALR and SALR, rising air will cool faster than the sur-
rounding air, so it will always be cooler and will tend to subside to a lower height. Such
air is said to be absolutely stable.
If the ELR is greater than the SALR, air that is rising and cooling at the DALR and later
at the SALR will always be warmer than the surrounding air. Consequently, it will contin-
ue to rise. The air is then absolutely unstable.
If the ELR is less than the DALR but greater than the SALR, rising air will cool faster than
the surrounding air while it remains dry but more slowly once it rises above the lifting
condensation level. At first it is stable, but above the lifting condensation level it becomes
unstable. This air said to be conditionally unstable. It is stable unless a condition (rising
above its lifting condensation level) is met, whereupon it becomes unstable.
Stable air brings settled weather. Unstable air produces heaped clouds of the cumu-
lus type. The base of these clouds is at the lifting condensation level, and the cloud
tops are at the altitude where the rising air has lost enough water vapor to make it dry
once more, so it is cooling at the DALR. If the air is sufficiently unstable, however, the
clouds can grow into towering cumulonimbus storm clouds. Equatorial air is usually
unstable.

the current for just a little longer. Dry material catches fire,
smolders below the surface until the storm has passed, then
bursts into flame.
If wind fans the flames, the result may be a forest fire,
although fires are much less common in broad-leaved decid-
uous forests than they are in coniferous forests. Wind can
cause serious damage, even without fires.
Trees absorb the force of the wind and those around the
edge of a forest shelter the interior. The wind speed 100 feet
(30 m) inside a forest is less than 80 percent of that outside.
Wind speed is reduced to 50 percent 200 feet (60 m) from the
edge, and 400 feet (120 m) from the edge it is less than 10
percent of the speed outside.
Often it is the trees on the edge that suffer, but not always.
Trees that stand taller than those around them are at risk, but
72 TEMPERATE FORESTS

there are few of those, because exposed trees suffer badly

from the drying effect of the wind (see “Evaporation and
transpiration” on page 66). It can also happen that the wind
brings down trees deep inside the forest while those at the
edge remain standing.
Wind is the movement of air around areas of low or high
pressure. Its strength is proportional to the rate at which the
pressure changes with distance from the center of the system
(see “Jet streams and depressions” on page 45). Technically, a
gale is defined as any wind with a speed greater than 32 MPH
(51.4 km/h), but a gale strong enough to break or uproot
trees blows at more than 55 MPH (88.4 km/h).

Forest fires
Lightning strikes are the most common cause of natural fires.
Fires are rare in broad-leaved deciduous forests, however.
These forests grow in moist climates, so the fuel they provide
is usually too wet to ignite easily, and the living trees do not
burn readily. Their leaves contain water that makes them dif-
ficult to ignite and their wood burns slowly. This is not to say
that broad-leaved deciduous forests are immune to fire. They
are not. A prolonged spell of hot weather with no rain will
dry out the dead leaves and branches lying on the forest floor
until any small spark may set them alight.
Fires are much more common in coniferous forests.
Coniferous trees contain resins, which are highly flammable,
and their needles contain much less water than broad leaves,
so they catch fire more readily.
Forest fires look terrifying, but they are not necessarily
harmful. Some forest trees depend on fire to allow their seeds
to sprout (see “Fire, and trees that depend on it” on page
120).
The truly frightening fires blaze through the crowns of the
trees, leaping from tree to tree. Most forest fires do not spread
in this way. They start among the dry litter on the forest floor
and spread rapidly along the floor and just below the surface
as a ground fire. A ground fire produces smoke, but no flames,
and most ground fires travel only a short distance.
 CLIMATE OF TEMPERATE FORESTS 73

A ground fire will kill herbs and shrubs, as well as tree

seedlings and young saplings, but the larger trees suffer noth-
ing more serious than some scorching of their bark. The fire
is not very hot. The temperature is usually between about
194°F (90°C) and 248°F (120°C) in the surface litter, but de-
creases rapidly below the surface. Most of the organisms liv-
ing a few inches below the surface remain unharmed as the
fire passes above them.
If the fire should burst into flame, however, it can become
more serious, because there is a chance that it may spread
upward into the crowns. A crown fire blazes fiercely and is
spread from tree to tree by burning cinders that are carried
aloft in rising hot air and then blown forward by the wind
before falling back into the canopy.
Once it starts, a crown fire generates its own wind. Hot air
rises rapidly, producing very strong vertical air currents, and
the removal of air produces low atmospheric pressure near
ground level. Surrounding air rushes into the low-pressure
area, to be heated in its turn.
As it approaches the center of low pressure, the converging
air starts to turn. In the Northern Hemisphere it turns coun-
terclockwise (it turns clockwise in the Southern Hemisphere),
until instead of flowing directly into the low-pressure region,
the air spirals into it. The spiral tightens toward the center.
The air turns in ever-smaller circles and this makes it acceler-
ate. It becomes a twisting wind, like a tornado, and it can
blow with great force. The fire has then generated a firestorm.
The hotter the fire, the fiercer is the storm that it generates.
Like a blacksmith’s bellows, the wind feeds oxygen to the
flames, making them burn faster and hotter—and therefore
making the wind blow even harder. The wind will pick up
any material lying loose on the ground. Most of this will be
flammable plant material and the wind will feed it into the
core of the fire. A firestorm will die down only when it
exhausts its supply of fuel.
Dry weather and lightning combine to start forest fires and
the wind helps them to spread. Weather causes the fire, but if
it blazes out of control a forest fire produces weather of its
own.
CHAPTER 4

HOW TREES WORK

How trees evolved
Life on Earth began in water, and plants are descended from
green single-celled organisms called algae (singular alga) that
Oaks (Quercus sp.) drifted near the surface. Their descendants are still there, and
are magnificent trees, thriving, as the algae that give some seawater its green color.
and never more so Other algae developed further and became much bigger
than when silhouetted plants. These are seaweeds, and some species, known as kelp,
against a night sky lit grow like vast, dense forests on the seabed, their “leaves”
by the rising full Moon. waving gently as the currents move them like trees bending
(Courtesy of Fogstock) in the wind. Several seaweed species grow in this way.

74
 HOW TREES WORK 75

Oarweed, also called tangle (Laminaria digitata), is up to 13

feet (4 m) tall, and furbelows (Saccorhiza polyschides) can
reach 14.75 feet (4.5 m), but the biggest of all are some
Macrocystis and Nereocystis species, known as giant kelps,
which can grow to a height of 100 feet (30 m).
Kelp seaweeds are very treelike. They have stems resem-
bling tree trunks topped by blades that look like leaves. The
stems are not rigid, however. When the plants are exposed at
low tide they lie limply on the shore and their rubbery stems
bend easily. They have no need to be rigid because kelps float
in water, with holdfasts anchoring their flexible stems secure-
ly to rocks on the seafloor.
When plants first moved onto land, about 450 million
years ago, they had to adapt to very different conditions. The
first arrivals were most likely algae, in which the cells are
linked to make long filaments. You can still see algae like
this, called blanket weed (usually Cladophora species),
attached to stones in fairly narrow, slow-moving rivers, its
dark green filaments gently waving in the current like long
hair billowing in the wind. Algae like these grew on the edges
of lakes and marshes.
Algae cannot survive for long out of water, because their
cells dry out. Plants that grow on dry land, exposed to the air,
have a waterproof cuticle that covers them completely, like a
skin, with pores called stomata to allow gases to enter and
leave the plant.
Cooksonia plants were among the first to have a protective
cuticle and were growing around marshes by 428 million
years ago. They were soon joined by species of Rhynia and
Zosterophyllum. These small plants—the biggest was about 20
inches (50 cm) tall—incorporated another important innova-
tion: They were the first vascular plants. That means they had
a system of tubular cells through which water, nutrients, and
the sugars produced by photosynthesis (see “Photosynthesis
and respiration” on pages 79–85) could move through the
plant. Plants that live in water do not need to transport sub-
stances, because everything they need is dissolved in the
water around them and they can absorb nutrients directly
into their cells. Land plants must take carbon dioxide from
the air and water and a range of nutrients from the soil. This
76 TEMPERATE FORESTS

is possible only if different parts of the plant specialize and

substances are transported to where they are needed.
The first vascular plants were at an early stage in specializa-
tion, and they did not have true roots. Instead they absorbed
water and nutrients through hairlike structures called rhizoids
that lay along the ground surface. This was not very efficient,
which is why the plants were small.
Progress was rapid, however, and the ancestor of all gym-
nosperms—the firs, pines, spruces, hemlocks, larches, cedars,
and related plants—was growing by 360 million years ago.
Called Archaeopteris, this first tree was up to 33 feet (10 m)
tall, with a trunk up to five feet (1.5 m) in diameter at the
base and leaves resembling the fronds of a fern. Within a few
million years there were forests of Archaeopteris and other
early trees and tree ferns.
Trees have been spectacularly successful at overcoming
gravity. Buoyancy, which keeps kelp upright, is of no help to
land plants that seek to raise their leaves above their neigh-
bors. Watertight plant cells become rigid, or turgid, when they
are filled with water, and water pressure—the scientific term is
turgor pressure—is sufficient to hold small plants upright. It is
not able to support the weight of a large plant, however. Tall
plants, such as shrubs and trees, have cells that produce a
hard substance called lignin. When the cells die and collapse
the lignin remains. The process is called lignification and its
product is wood (see “How wood forms” on pages 94–97).
Woody plants are able to grow tall—and they did. Those early
forests contained Lepidodendron, with a straight trunk up to
6.5 feet (2 m) in diameter and up to 130 feet (40 m) tall, and
Calamites, a giant ancestor of modern horsetails (Equisetum
species) that grew to about 50 feet (15 m).
Trees are very distinctive. You may not know the names of
many plants, but you can tell whether or not a plant is a tree.
It is tall. When fully grown a tree is at least 33 feet (10 m) tall.
Most trees—though there are many exceptions—have only
one stem, or trunk. Branches emerge from the trunk, and the
lowest branch usually joins the trunk at least four feet (1.2 m)
above the ground. The trunk and branches are hard to the
touch. If the plant is smaller than a tree and has several
woody stems it is a shrub.
 HOW TREES WORK 77

How trees find food

Animals move around in search of food. They find plants
they can eat or, if they are meat eaters, other animals they
can catch. Plants cannot move around—they are literally
rooted to the spot, and it is their roots that feed them as well
as anchoring them securely to the ground. A tree that towers
majestically into the sky is a giant, its branches spreading
wide so its millions of leaves are able to absorb the sunshine,
but only part of the plant is visible. Below ground the tree
has a system of roots that extends far beyond the crown
above ground. The root system accounts for 20–25 percent of
the total mass of the tree.
It is not easy to examine the roots of a full-grown tree.
When a tree is blown down by the wind its roots are
exposed, but individual roots break as the tree falls, and the
part that remains—the root plate—is only a small fraction of
the original root system. It is not practicable to dig out an
entire tree and then carefully wash all the soil from the
roots. Until recently scientists had no choice but to assume
that the roots of a mature tree resembled those of a seed-
ling—which they could see, measure, and scale up to an ap-
propriate size.
Nowadays it is possible to use ground-penetrating radar,
which identifies the roots and allows scientists to map their
location and extent without damaging them. When this was
first done the results came as a surprise: Tree roots are not at
all how they had been pictured.
Rainwater dripping from leaves at the edges of the crown
marks a drip line surrounding the tree. Botanists and foresters
had supposed that a tree spread its roots sideways approxi-
mately as far as this line. The radar studies revealed that in
fact the tree roots extend far beyond the drip line, in some
cases as much as 10 times farther.
Scientists had also believed that most broad-leaved trees,
such as oaks, had thick roots, called taproots, that descended
vertically to a considerable depth. When a seed germinates,
the radicle, which later becomes the root, grows vertically
downward, and tree seedlings of these species do develop
taproots. It was assumed that these continued to grow as the
tree matured. In fact, the taproots do not develop. Almost all
78 TEMPERATE FORESTS

Extent of the root

system of a fully grown
tree. Note that the
roots extend a long
way from the trunk
but are quite shallow.

of the tree’s roots are within the uppermost three feet (1 m)

of the soil, and most of the fine roots through which the tree
absorbs nutrients are within about eight inches (20 cm) of
the ground surface. The illustration shows a complete tree,
including the roots, as these have been revealed by modern
studies.
Tree roots are woody, like the trunk and branches. They
never bear leaves or flowers, but they produce many branch-
es as they push their way through the soil. A root cap, made
from a layer of tough tissue, protects the tip of each root.
Abrasion with the soil strips away the root cap as the tip
advances, but the tissue is constantly renewed and the cells
of the active growing region never lose their protection.
On the surface of the root behind the tip, a layer of cells
produce a fine “fur” of root hairs, each no more than about
0.2 inch (4 mm) long. These are the roots through which the
tree absorbs most of its nutrients. The tree’s root hairs would
stretch for hundreds of miles if all of them were laid end to
end. The root hairs are always located just behind the root
tip. As the root grows longer, old hairs are shed and new
ones appear. Nutrients enter the plant mainly near the root
 HOW TREES WORK 79

tips, although some enter along the entire length of the

roots.
The roots seek moisture in the soil (see “How water moves
through a temperate forest” on pages 29–33), and root hairs
cling tightly to soil particles. Various minerals are dissolved
in the water coating these particles, and as this water passes
through the root hairs and into the main body of the root,
the dissolved minerals go with it. Inside the root a complex
filtering system allows some minerals to pass while excluding
and excreting others. The root is very selective and is able to
extract the important nutrient minerals from the dilute soil
solution and concentrate them before passing them to the
trunk, branches, buds, leaves, and flowers, where the nutri-
ents are needed.
Trees—indeed, most plants—have help in acquiring nutri-
ents. Their roots develop close relationships, called mycor-
rhizae, with soil fungi. The fungi obtain carbohydrates from
the tree and supply minerals to the tree root, so both organ-
isms benefit.
All plants require a range of mineral nutrients, known as
essential elements. Those that are needed in relatively large
amounts are known as macronutrients and include nitrogen,
phosphorus, potassium, sulfur, magnesium, and calcium. A
variety of minerals are needed in much smaller amounts, and
not all of them by every species of plant. These are called
micronutrients and include iron, copper, zinc, molybdenum,
manganese, cobalt, and boron. The essential elements are
constantly recycled, moving between the air, water, and liv-
ing organisms. Plants also need carbon, hydrogen, and oxy-
gen for the production of carbohydrates through the process
of photosynthesis.

Photosynthesis and respiration

Throughout the cold, dark winter the forest lay dormant.
Last year’s leaves lay as a carpet across the ground and the
trees were bare. The branches were bare, but there were buds
on all the twigs, enclosing and protecting next year’s leaves.
Winter passed, and as the days grew longer and the sunshine
warmer those buds swelled. Then, in the course of a few
80 TEMPERATE FORESTS

The nitrogen cycle

Nitrogen is a colorless, odorless gas that composes 78 percent of the air. It is an ingre-
dient of all proteins and consequently an essential macronutrient for all living organ-
isms, but it is useless to most of them in its gaseous form. Before plants can absorb and
utilize it, the nitrogen must be fixed—made to react with another element to form a
compound.
Lightning makes nitrogen (N) react with oxygen (O), producing nitrogen oxides that
dissolve in raindrops (H2O) to form weak nitric acid (HNO3). When nitric acid reaches the
soil it is converted into nitrate (NO3 ), a soluble form in which plant roots are able to
absorb it. Certain soil bacteria are able to utilize nitrogen gas directly. In doing so, they
convert the nitrogen to nitrate; this bacterial process is called nitrification.
Plants use the nitrogen they absorb to manufacture proteins. Animals obtain their pro-
teins by eating plants. Plant and animal wastes and the remains of dead plants and ani-
mals decompose through the action of soil animals, fungi, and bacteria. The decomposers
convert the proteins to ammonia (NH3 ). Ammonia dissolves in water, and some of it is
absorbed by plant roots and used to make more protein. Ammonia evaporates readily (it
boils at –29°F; –34°C) and a small amount returns to the air. The remainder of the ammo-
nia combines with carbon dioxide (CO2) to form ammonium carbonate ((NH4 )CO3 ).
Nitrification by other bacteria converts ammonium carbonate to nitrous acid (NHO2),
with the release of energy: (NH4 )CO3 + 3O2 → 2NHO2 + CO2 + 2H2O + energy. (The
arrow indicates the direction in which the chemical reaction moves; in this case ammoni-
um carbonate reacts with oxygen to produce nitrous acid, carbon dioxide, and water with
the release of energy.)
Nitrous acid reacts with magnesium (Mg) or calcium (Ca) to form magnesium or calci-
um nitrite (Ca(NO2)2 or Mg(NO2)2), which other bacteria nitrify to produce more nitrate:
2Ca(NO2)2 (or 2Mg(NO2)2) + 2O2 → 2Ca(NO3)2 (or 2Mg(NO3)2) + energy.
Finally, another type of bacteria converts the nitrogen compounds to either nitrogen
gas or ammonia; the process is called denitrification. Ammonia that evaporates into the air

weeks, all of them burst open and suddenly the forest was
green. The trees were in leaf—and working.
Green leaves are factories that manufacture sugar, using
carbon dioxide and water as the raw materials and sunlight
as a source of energy. The leaves are green because they con-
tain chlorophyll, a chemical substance that traps light energy.
 HOW TREES WORK 81

is quickly oxidized to nitrate and returns to the soil, where it is absorbed by plants.
Eventually, all the nitrogen that enters the soil returns to the air.
The diagram summarizes the cycle in which atmospheric nitrogen passes through liv-
ing organisms and returns to the air.

nitrogen fixation atmospheric nitrogen fixation

by lightning nitrogen by bacteria

atmospheric
nitrogen denitrification nitrogen
oxides in bacteria

nitrification

nitrates
in soil
uptake by plant roots

nitrification protein protein

in plants feeding in animals

nitrites death death

ammonia in
nitrification dead organic
matter The nitrogen
cycle

Chlorophyll is green. It is one of several photosynthetic pig-

ments, each of which absorbs light at a slightly different
wavelength. Two varieties of chlorophyll a are also known as
P700 and P680. “P” is for “pigment” and the numbers refer to
the wavelengths (in nanometers) of the red light they absorb.
Chlorophyll a is blue-green. Chlorophyll b is very similar, but
82 TEMPERATE FORESTS

it is yellow-green. Xanthophylls and carotenoids are the

most important of the other photosynthetic pigments.
Xanthophylls are yellow and carotenoids are various shades
of orange through red. Each pigment absorbs light at a slight-
ly different wavelength, so the presence of several pigments
widens the range of light the leaf cells can absorb. The two
varieties of chlorophyll a are the most abundant and the
most important.
All of the photosynthetic pigments reside on stacks of
membranes, called thylakoid membranes, inside bodies called
chloroplasts. Chloroplasts once lived independently and they

Photosynthesis
Green plants and some bacteria are able to use energy from sunlight (photo-) to assemble
(synthesize) sugars. The process is called photosynthesis and it depends on a pigment
called chlorophyll. Chlorophyll is green and it is what gives plants their green color.
Photosynthesis proceeds in two stages. The first stage depends on light, and it is called
the light-dependent or light stage. The second stage does not use light energy, so it is called
the light-independent or dark stage (although it also takes place in the light).

Light-dependent stage. When a photon (a unit of light) possessing precisely the

right amount of energy strikes a chlorophyll molecule, the photon disappears and
its energy is absorbed, allowing an electron (a particle carrying negative charge) in
the molecule to break free. This leaves the chlorophyll molecule with a positive
charge. The free electron immediately attaches to a neighboring molecule, there-
by ejecting another electron that moves to a neighboring molecule. In this way
electrons pass along an electron-transport chain of molecules. Each plant cell con-
tains a number of chloroplasts and each chloroplast contains many molecules of
chlorophyll, so while the plant is exposed to light there is a constant stream of pho-
tons being captured and electrons moving along the electron-transport chain.
Some of the transported energy is used to convert adenosine diphosphate
(ADP) to adenosine triphosphate (ATP) by the addition of phosphate, after which
the electron then returns to the chlorophyll. Converting ADP to ATP absorbs ener-
gy; converting ATP to ADP releases the energy. The ADP ↔ ATP reaction (the dou-
ble arrow indicates the reaction can move in either direction) is used by all living
organisms to transport energy and release it where it is needed.
 HOW TREES WORK 83

still have their own DNA allowing them to reproduce, but

very early in the development of life on Earth they moved
inside other cells. Today the chloroplasts live inside the meso-
phyll cells that form the inside of leaves. Each mesophyll cell
contains 30–40 chloroplasts.
Chloroplasts are the “factories” in which the leaves take
carbon dioxide (CO2 ) from the air and water (H2O) carried to
them from the ground below as the raw materials for the pro-
duction of sugars, such as glucose (C6H12O6). The process by
which light (photo) supplies the energy to assemble (synthe-
size) sugars is called photosynthesis (see the sidebar).

Energy that is not used to convert ADP to ATP is used to split a water mole-
cule (H2O) into a hydrogen ion, which bears a positive charge (H+), and a
hydroxyl ion, which has a negative charge (OH–). (An ion is an atom that has
gained or lost one or more electrons, so it bears a positive or negative charge.)
The H+ attaches to a molecule of nicotinamide adenine dinucleotide phosphate
(NADP), converting it to reduced NADP (NADPH). The OH– passes one electron
to the chlorophyll molecule, restoring the neutrality of both chlorophyll and
hydroxyl. Hydroxyls then combine to form water (4OH → 2H2O + O2↑). (The
upward arrow in this chemical formula indicates that the oxygen is released
into the air.) This completes the light-dependent stage.

Light-independent stage. Using ATP from the light-dependent stage as a source

of energy, the first in a series of chemical reactions attaches molecules of car-
bon dioxide (CO2) obtained from the air to molecules of ribulose biphosphate
(RuBP), a substance present in the chloroplast. The enzyme RuBP carboxylase
(the name is usually abbreviated to rubisco) catalyzes the reaction. In a cycle of
reactions the carbon atoms, originally from the carbon dioxide, are combined
with hydrogen obtained from NADPH; the NADP then returns to the light-
dependent stage. The cycle ends with the synthesis of molecules of glucose
and of RuBP. The RuBP is then available to commence the cycle again.
Glucose, a simple sugar, is the most common source of energy for living
things; its energy is released by the process of respiration. Glucose is also used to
synthesize complex sugars: starch and cellulose in plants and glycogen (also
called animal starch) in animals. Plants use cellulose to build cell walls; starch and
glycogen can be converted to glucose, releasing energy.
84 TEMPERATE FORESTS

Photosynthesis is the process that assembles glucose.

Glucose has a number of uses, one of which is as a fuel.
Burning fuel such as coal, wood, oil, or gas creates the con-
ditions in which carbon in the fuel can combine with oxy-
gen in the air. This reaction releases energy—the heat of
the fire. When the carbon in glucose combines with oxygen,
the energy that is released is used to make muscles move, to
drive chemical reactions, and to move molecules into and
out of cells. The process by which it does so is called respira-
tion and it is the opposite of photosynthesis. Respiration
should not be confused with breathing. Breathing is the
physical activity by which terrestrial vertebrates draw in
air into the lungs so that oxygen from the air can enter
the bloodstream. This supplies the oxygen needed for res-
piration, but breathing and respiration are not the same
thing.
Photosynthesis takes place in chloroplasts, located in the
leaves. Respiration takes place in mitochondria, bodies (organ-
elles) that are found in every living cell. Like chloroplasts,
mitochondria were once independent organisms, but they
became incorporated into other cells.
In the overall reaction glucose (C6H12O6) reacts with oxy-
gen (O2 ) to produce carbon dioxide (CO2 ) and water (H2O). It
is easy to see how respiration is the precise opposite of pho-
tosynthesis when the two reactions are compared:
6CO2 + 6H2O + energy → C6H12O6 + 6O2 (photosynthesis)

C6H12O6 + 6O2 → 6CO2 + 6H2O + energy (respiration)

Respiration does not convert glucose to energy directly,

but does so through a process called phosphorylation, in
which phosphate groups are moved from one molecule to
another. When ATP loses a phosphate group, that group is
transferred to another molecule. Phosphorylation may cause
a muscle cell to move, a molecule to enter or leave a cell, or a
chemical reaction to take place. The phosphorylation that
activates one molecule also converts ATP to ADP. Respiration
releases the energy that is needed to phosphorylate ADP,
changing it back into ATP and leaving it ready to do more
useful work.
 HOW TREES WORK 85

How trees reproduce

Trees spread their roots widely, and for some species this pro-
vides an opportunity for new trees to grow. The roots are not
far beneath the ground surface, and although roots do not
produce leaves or flowers, shoots develop at intervals along
the roots of some species. They are called adventitious shoots
because they grow from an unusual position on the plant.
The shoots grow into new trees, resulting in a group of trees
all of the same species. In fact, they are all parts of the same
tree, sharing the same root system. This type of reproduction
is said to be asexual or vegetative. Not all species are able to
reproduce in this way. Elms (Ulmus species), lindens and bass-
woods (Tilia species), and aspens (Populus species) often do
so. Aspens can produce a grove of thousands of trees, all from
a single parent tree.
The ability to reproduce asexually makes a tree virtually
immortal. Individual trees grow old and die, but there are
always more growing from the original root system. Provided
they do not succumb to a fatal disease or are unable to toler-
ate a change in their environment—and provided no one
cuts them down, of course—a tree can live in this way for
thousands of years.
More important, perhaps, is the fact that asexual reproduc-
tion allows trees to thrive in places where the climate is too
cold for them to produce ripe seeds. All of the trees that
reproduce asexually are capable of reproducing sexually,
from seed, but to produce seeds they need more sunshine
and warmth than northern regions can guarantee. They pro-
duce healthy seeds only in warm years, and the amount of
seeds is highly variable. Without an alternative means of
reproduction they would be unable to compete with trees
that are better suited to the climate.
Sexual reproduction involves the combining of chromo-
somes—threadlike structures carrying genes that are present
in the nucleus of all living plant cells—from a male and
female parent. Most cells carry two copies of each chromo-
some. Such cells are said to be diploid. When diploid cells
divide in order to repair damaged tissue or to allow the plant
to grow, they do so by a process called mitosis. Mitosis pro-
duces two identical daughter cells, each of which is diploid.
86 TEMPERATE FORESTS

The cells involved in reproduction—sperm and egg cells

(called ovules)—are known as gametes. Gametes are haploid,
which means they possess only one set of chromosomes.
They are produced from diploid cells by a form of cell divi-
sion called meiosis, in which the cell divides twice to produce
four haploid cells.
Trees, like all other plants, have a life cycle that takes them
through two distinct forms, one diploid and the other hap-
loid. Each form gives rise to, or generates, the other. Thus the
two forms are generations—each is both parent and offspring
of the other—and the plant life cycle consists of an alterna-
Many trees produce tion of generations. The diploid generation is called a sporo-
edible fruits. These are phyte and the haploid generation a gametophyte. One of these
English black walnuts forms is much bigger and more prominent than the other. In
(Juglans nigra). bryophytes—mosses, liverworts, and hornworts—the plant
(Courtesy of Christ one sees is the gametophyte. In all larger plants, including
Jongman, Foto Natura, trees, the plant one sees is the sporophyte. Pollen grains, con-
Minden Pictures) taining sperm, are the male gametophytes and egg cells are
 HOW TREES WORK 87

the female gametophytes. The gametophyte generation is

sheltered in the cones of coniferous (cone-bearing) trees and
in the flowers of flowering trees (see “Differences between
conifers and flowering plants” below).
When a pollen grain reaches the stigma of a flower or an
ovule in the female cone of a conifer, a tube grows from the
pollen grain and reaches the egg cell. Sperm then travel from
the pollen grain along the tube and into the ovule, where the
chromosomes from the two gametes unite. The fusion of
gametes that produces the diploid cell that will become a
new plant is called fertilization.
The resulting diploid cell is called a zygote. It divides by
mitosis. As it does so, different cells begin to specialize and
the zygote develops into an embryo. The embryo contains
simple leaves, called cotyledons or seed leaves; a plumule that
will grow into a shoot; and a radicle that will grow into a root.
The embryo is surrounded by a store of nutrients that will
sustain the young plant until it has grown large enough to
feed itself, and the embryo and its food store are enclosed by
a protective seed coat. The complete structure is then a seed,
from which the next sporophyte generation may grow.

Differences between conifers and

flowering plants
Two very distinct types of trees—conifers and broad-leaved
species—grow in temperate forests. The two look so very dif-
ferent that it is easy to tell them apart. Conifers produce
cones—conifer means “cone-bearing.” No broad-leaved tree
produces cones. Broad-leaved trees bear flowers. No conifer
bears true flowers, although the male cones of some species
look a little like flowers and are sometimes called flowers.
Conifers and broad-leaved trees have very different leaves.
Those of conifers are small and either very narrow and
often sharp like needles, or flattened to form scales. As their
name suggests, broad-leaved trees have broad leaves. Many
conifers, although not all, have an overall conical shape,
with a single straight trunk and straight branches that are
shorter the higher they are. Broad-leaved trees have a more
spreading, rounded shape.
88 TEMPERATE FORESTS

These are obvious differences. There are other, more im-

portant differences in the way the two types of tree grow and
reproduce. All conifers belong to a group of plants called
gymnosperms that appeared on Earth about 400 million years
ago. Broad-leaved trees belong to a group called angiosperms,
and the first of them appeared about 130 million years ago.
Garden flowers, grasses, farm crops, and shrubs as well as
broad-leaved trees are angiosperms. The angiosperms proved
so much more successful than the gymnosperms that there
are now approximately 235,000 species of angiosperms, com-
pared with only about 735 species of gymnosperms. About
550 of these are conifers. The others are cycads, sometimes
called sago palms although they are not true palm trees; the
ginkgo; and the gnetophytes, a group of tropical plants that
includes trees, shrubs, lianas, and smaller plants.
Both gymnosperms and angiosperms are vascular plants.
That is to say, water, dissolved nutrients, and sugars produced
by photosynthesis are transported through the plants along
channels. The channels that carry water and nutrients from the
soil to every part of the plant are known as xylem tissue. Those
that transport sugars from the leaves are called phloem tissue.
Xylem cells are dead. In gymnosperms the xylem tissue
consists of long, tubular cells called tracheids, with tapered
ends perforated by small openings called pits. Tracheids grow
end to end with their tapered ends overlapping. Rings or spi-
rals of a tough substance called lignin strengthen their walls.
As well as conducting water, tracheids help to support the
trunk and branches of the plant.
Angiosperm xylem consists of vessel elements, which are
shorter and wider than tracheids. They conduct water more
efficiently than tracheids but give little mechanical support.
The plant relies on fibers for support. These are long, narrow
cells rich in lignin.
Gymnosperms possess some fiber cells and angiosperms
possess some tracheids, but vessel elements are found only in
angiosperms. Botanists believe that vessel elements and fiber
cells both evolved from tracheids.
Phloem cells are alive, although they possess no nucleus.
In both gymnosperms and angiosperms, phloem consists of
cells called sieve elements. These are cylindrical and have per-
 HOW TREES WORK 89

forated ends called sieve plates. The cells are joined end to end
to form sieve tubes. In angiosperms, each sieve element is
linked to one or more companion cells. Companion cells have
nuclei and are intensely active; they probably supply the
sieve elements with energy and nutrients. Gymnosperm
phloem lacks companion cells. Instead, parenchyma cells sur-
round the sieve elements and may serve the same purpose.
Parenchyma cells are unspecialized. They manufacture and
store a range of substances.
Significant though they are, these are not the most impor-
tant differences between the two groups of plants—the differ-
ence that gives them their names. That difference is the
flower itself. At the base of the flower (see the sidebar on page
90) is the ovary, which contains and protects the ovules.
Angiosperm means “seed vessel,” from angeion (“vessel”) and
sperma (“seed”).
Gymnosperms do not produce flowers. They have male
cones that produce pollen and female cones that hold the
ovules on their scales. The ovules are not contained in an
ovary, and consequently the seeds that develop from the fer-
tilized ovules are not protected by the ovary walls: They are
naked. The Greek word gymnos means “naked,” and so “gym-
nosperm” means “naked seed.”
Lacking flowers to attract insects or other pollinators, gym-
nosperms rely on the wind for pollination, and their life
cycle is more complicated than that of angiosperms. It takes
three years for a gymnosperm to produce seeds from the time
the cones first appear.

How seeds disperse

Once it has produced seeds a tree needs to disperse them. It
could simply release them as soon as they were ready to ger-
minate. This would be wasteful, however, because most of
the seeds would fall to the ground around the base of their
parent, where they would have to compete for light, water,
and nutrients with their overwhelmingly bigger parent. Not
all the seeds land beside their parent, of course. Some drift
with the wind, landing farther away where conditions may
be more favorable.
90 TEMPERATE FORESTS

What is a flower?
A flower is the reproductive structure of a flowering plant, containing and protecting the
gametophyte generation (the reproductive stage in the plant’s life cycle). Flowers vary
greatly in appearance. Rafflesia arnoldii, a plant that parasitizes the roots of tropical trees,
has flowers up to 32 inches (80 cm) across that smell of rotting meat (to attract the flies
that pollinate it). Scarlet pimpernel (Anagallis arvensis) flowers are 0.4–0.6 inches (10–15
mm) across. A sunflower (Helianthus species) is a composite of hundreds of tiny flowers,
called florets. Some flowers are brightly colored and others, such as grass flowers, have no
petals and are inconspicuous.
Despite the great variety of forms, however, all flowers can be related to a generalized
description. A particular flower may not possess all of the parts of the general flower
shown in the drawing, but those it does possess will be included in the description.
Small green leaves at the base of the flower where the flower stem, or peduncle, joins
the main body of the plant are called bracts. Some plants have bracts that are shaped dif-
ferently from the plant’s other leaves.
The upper end of the peduncle is swollen to form the receptacle. The other parts
of the flower are attached to the receptacle. Nectaries on the surface of the receptacle
are glands that secrete nectar, a sweet-tasting liquid; as pollinating insects (and in
some parts of the world birds and bats) probe flowers for nectar, pollen clings to their
bodies.
Sepals are modified leaves attached to the receptacle below the main part of the flower.
The sepals enclose the inner parts of the flower, forming the outer layer of the flower bud.
They protect the flower until it is ready to open. Sepals are usually green, but in some
species, such as Poinsettia and marsh marigold (Caltha palustris), they are colored.
Together, the set of sepals composes the calyx.
Above the receptacle (and attached to it) there are the female and male reproductive
organs. The female organ is called the carpel and the male organs are the stamens.
The carpel consists of the stigma, style, and ovary. The stigma has a sticky surface onto
which pollen grains are deposited. The style links the stigma to the ovary, which contains
the ovules (female gametes). Flowers evolved from modified leaves, and the carpel may
have originated as a rolled-up leaf.

One solution to the difficulty is for fallen seeds to lie dor-

mant in the ground until competing trees die. Most tree
seeds need to germinate within 18 months or less, but some
 HOW TREES WORK 91

The stamen consists of a filament at the top of which there is a club-shaped anther,
where pollen grains are produced. Pollen grains mature into male gametophytes. The
drawing shows only six, but a flower usually has many filaments and anthers.
If the ovary is attached to the receptacle above the level at which the filaments are
attached (as in the drawing) it is said to be superior. If it is attached below the filaments it
is inferior.
Petals are modified leaves that surround and partly enclose the male and female
organs. The ring of petals is called the corolla. In some flowers the edges of the petals are
partly joined and the petals form a corolla tube. Together, the outer calyx and inner corol-
la are known as the perianth.
A flower that possesses stamens, carpel, calyx, and corolla is said to be complete. Many
flowers lack one or more of these parts and are therefore incomplete.
A flower that possesses both male and female organs is said to be perfect. A staminate
flower has only stamens and a carpellate flower has only a carpel. Staminate and carpel-
late flowers are imperfect.

petal

anther stigma

style
filament

ovary
ovule
nectary
sepal
receptacle

peduncle

The parts of a flower

species produce seeds that remain viable for several years.

These seeds remain in the ground, forming a seed bank. Paper
birch (Betula papyrifera) and yellow birch (B. alleghaniensis)
92 TEMPERATE FORESTS

seeds are able to survive in this way. Certain coniferous trees,

especially some pines (Pinus species), retain their seeds in
their cones until conditions are suitable for them to germi-
nate (see “Fire, and trees that depend on it” on pages
120–123).
A seed that germinates in a place overshadowed by full-
grown trees will not produce a young tree that grows very
large, but this may not matter. Walk through a forest and
you will see many small tree seedlings. These look like
young plants, but they may have remained at the same size
for many years. They are waiting until a nearby tree falls.
When it does they will grow rapidly to take its place. Not all
tree species produce seedlings that can survive for long in
this state of arrested development, but many common forest
trees do, including hemlock (Tsuga canadensis), beech (Fagus
grandifolia), sugar maple (Acer saccharum), and white oak
(Quercus alba).
Other trees produce seeds that disperse passively, carried
by the wind. Birches (Betula species), ashes (Fraxinus species),
elms (Ulmus species), and maples (Acer species) are among the
trees that produce winged seeds, or keys. The illustration
shows the keys of maple and birch.
A winged seed—the botanical term is samara—spins as it
falls. This slows its rate of fall. In still air the wings will have
no other effect, because the seed falls vertically, but even the
slightest breath of wind will carry the seed away from the
parent tree.
Even with wings, seeds seldom travel far, and whether or
not they germinate is very much a matter of luck. Most trees
that rely on this type of dispersal produce vast quantities of
seeds and the seeds themselves are usually small.

maple

birch
Winged seeds
 HOW TREES WORK 93

There is another way to disperse seeds, however, and that

is to recruit animals for the purpose. This is called active dis-
persal. Birds and mammals are free to move where they will,
so they can carry seeds much farther than the wind can.
Obviously, these seed carriers must be “paid,” and the way to
do that is to make the seeds edible. Squirrels are famous for
their habit of gathering tree seeds, preferring those of hicko-
ry, beech, and oak. Some birds, such as the acorn woodpeck-
er (Melanerpes formicivorus) and the Eurasian jay (Garrulus
glandarius) also collect acorns from around oak trees.
They gather seeds in order to eat them, of course, so this
may not seem to be the best way for a tree to reproduce. It
works because the seed gatherers collect food when it is plen-
tiful and store the surplus for the winter, when food is scarce.
Squirrels bury their hoards, but they are notoriously ineffi-
cient at finding them again. They search for their stores by
scent, but the seeds are odorless when the ground is dry.
Consequently, the squirrels never recover all of their seeds
and some, conveniently buried in fertile soil, are able to ger-
minate. The jay, on the other hand, has a truly remarkable
memory. It buries up to 4,000 acorns singly and remembers
where it put every one of them, but a jay seldom eats all of
them, and so some germinate. The acorn woodpecker does
not help the trees at all, however. It drills holes in dead trees
and fence posts and stores its acorns there, where they have
no chance of germinating.
Flowering plants have developed a much better alternative
to edible seeds: edible fruit. The fruit is rich in sugar and
therefore a valuable source of energy. It is usually brightly
colored, so the fruit eaters can find it easily, and it changes
color as it ripens so consumers can tell when it is ready to
eat—which is also when the seeds are ready to be dispersed.
The seeds are inside the fruit and have tough seed coats
that make them indigestible. If the seeds are small, the ani-
mal will swallow them as it eats the fruit. They will then pass
through its digestive system to be voided with the feces. By
the time it defecates, the animal will be far away from the
fruit tree and its feces will supply the seed with manure to
help the seedling. If the seed is large, like that of a cherry or
plum, the animal will eat the fruit from around it and leave
94 TEMPERATE FORESTS

the seed on the ground. Even then the seed may travel a long
way, because the animal may have carried the fruit some dis-
tance to a place where it can eat without fear of being
attacked or robbed.

How wood forms

At the tip of each twig there is a bud, made up of a dome-
shaped mass of cells, known as meristem cells, that are able to
divide to produce more tissue. Many trees also have meristem
tissue in the axils where twigs join the main branch. Growth
from these buds makes twigs longer and produces new twigs.
This is primary growth. If it were the only type of growth, the
tree would grow longer and longer twigs and a longer and
longer trunk, but none of these would grow any thicker. The
tree—if you could call it a tree—would sprawl across the
ground as a spindly plant too weak to support its own weight.
It is secondary growth that makes the trunk and branches
thicker—and that produces wood. Trees have trunks and
branches made from wood, and wood is the most important
material people obtain from trees.
Wood is a highly structured material. The grain, exposed
when wood is cut, and the rings that are clearly visible in the
cross section when a trunk or branch is severed reveal the
structure.
Forming a ring around the trunk that extends into the
branches, tracheids, vessel elements, fibers, and sieve ele-
ments (see “Differences between conifers and flowering
plants” on pages 87–89) are packed together in vascular bun-
dles, with the xylem and phloem lying side by side. Xylem is
on the inside of the ring and phloem on the outside. Pith rays
radiating from the center of the trunk or branch to the outer
bark consist of stacks of parenchyma cells—all-purpose cells—
most of which store starch, a source of energy for the plant.
Meristem cells are located among the vascular bundles and
in the pith rays. Those in the pith rays divide to extend the
rays as the trunk or branch grows thicker. The meristem cells
in the vascular bundles form a layer, in most trees no more
than one cell thick, of tissue called vascular cambium. A
meristem cell is known as an initial and the cells it produces
 HOW TREES WORK 95

are called derivatives. In the vascular cambium each initial

divides to produce two more cambium cells—initials. One of
the initials then divides to produce a derivative—a different
kind of cell. Initials nearest the center of the tree produce
xylem cells and initials nearest the outside of the tree pro-
duce phloem cells.
Outside the phloem tissue there is a second cambium
layer, called cork cambium. This cambium gives rise to the
cells composing the “skin” of the tree. Cork cambium initials
give rise to derivatives that lay down a coating of waxy mate-
rial, called suberin, in their walls and then die. The dead cells
form a layer of cork that makes the trunk or branch water-
proof and protects it against attack by insects. Unlike the vas-
cular cambium, the cork cambium is not permanent and
does not surround the entire trunk or branch. It is produced
where and when it is needed. After a few weeks its initials
turn into cork cells and die, whereupon the cork cambium
disappears. On the outside of the tree, the “skin” dries and
splits as the tree grows, and more cork cambium is produced
to fill the gaps.
As phloem cells die they turn into cork cells and add to the
cork layer, while parenchyma cells among the phloem cells
turn into meristem cells to produce new cork cambium.
Together, the phloem, cork cambium, and cork layer make
up the bark of the tree. The vascular cambium and xylem
comprise the sapwood. If the sapwood is cut, liquid—sap—
leaks from it. This is the living part of the trunk or branch. A
tree will die if bark is removed completely in a ring all around
the trunk. This is called ringbarking, and by severing the
phloem it prevents sugars from the leaves reaching the roots.
Unable to respire (see “Photosynthesis and respiration” on
pages 79–85), the roots die, ending the supply of water and
nutrients to the tree, and the entire tree dies.
Each year the vascular cambium produces more xylem and
phloem cells. These are needed because, as the tree grows
taller and its branches grow longer, its tissues need more
water and its leaves produce more sugars, and more vascular
tissue is needed to transport them. This gives rise to second-
ary growth, which happens because each new layer of xylem
is laid around the outside of the preceding layer, making the
96 TEMPERATE FORESTS

trunk or branch grow thicker and, of course, necessitating

the formation of more bark to cover it.
Meanwhile older xylem cells die. They are already
strengthened with lignin and this tough substance remains,
as does the lignin in the long fiber cells embedded in the vas-
cular bundles. The old cells often fill with waste products,
which alters their color. The cells are no longer capable of
conducting water. Instead they form a dense mass of tough,
fibrous material rich in lignin. This is heartwood, which forms
Cross section through a a central column that adds greatly to the strength of the tree.
tree trunk, showing the The illustration shows a cross section through a tree trunk,
dead heartwood, living with the layers labeled.
sapwood (xylem tissue), In temperate forests, growth does not continue through-
vascular cambium, out the year. It commences in spring, when the vascular cam-
and the phloem, cork bium produces wide xylem cells with thin walls. These form
cambium, and cork, a pale, fairly wide band around the previous year’s xylem.
which together Later in the summer the cambium produces smaller cells
compose the bark. with darker, thicker walls that pack into a narrow, dark-

heartwood

sapwood

vascular cambium

phloem

cork cambium bark

cork
 HOW TREES WORK 97

Peeling bark on paper

birch (Betula papyrifera)
(Courtesy of Jim
Brandenburg,
Minden Pictures)

colored band around the spring xylem. When growth ceases

for the winter the secondary growth remains as two bands
around the trunk or branch, one pale and the other dark.
Each year another pair of these layers is added. That is how
secondary growth produces the annual growth rings that are
clearly visible when a tree is felled.
CHAPTER 5

ECOLOGY OF
TEMPERATE FORESTS
How ecological ideas developed
One of the largest European temperate forests covers about
725 square miles (1,878 km2) on the plain that straddles the
border between Poland and Belarus. Its Polish name is the
Bia l⁄ owieża Forest and in Belarus it is known as the
Belovezhskaya Pushcha. It is a World Heritage Site.
The forest contains a variety of coniferous and broad-
leaved tree species. There are Scots pine (Pinus sylvestris),
Norway spruce (Picea abies), European common hornbeam
(Carpinus betulus), small-leaved lime (Tilia cordata, sometimes
called littleleaf linden), English oak (Quercus robur), Eurasian
sycamore (Acer pseudoplatanus), maples (Acer species), ash
(Fraxinus excelsior), two species of birch (Betula), aspen
(Populus tremula), and European black alder (Alnus glutinosa).
These are the principal species, but in addition there are
more than 900 other species of vascular plants, as well as
lichens, liverworts, and more than 1,500 species of fungi.
Obviously, a forest of this size harbors many animals.
There are 55 species of mammals, including the European
bison (Bison bonasus), gray wolf (Canis lupus), lynx (Felis
lynx), and wild boar (Sus scrofa), as well as 212 species of
birds, 11 of amphibians, and seven of reptiles. The forest has
more than 8,500 species of insects.
Listing the species present in a forest, or any other area,
gives the impression that the plants and animals are scattered
evenly, as though you could take an area anywhere in the for-
est and find all the plants and most of the animals. It is not
like that, however. There are plant associations. In some
places linden (lime) and hornbeam grow side by side; in
other places there are oak and hornbeam. In all, there are 12
of these associations in the Belarus forest and 20 on the
Polish side of the border. The smaller plants form associa-

98
 ECOLOGY OF TEMPERATE FORESTS 99

tions in the same way. Plant-eating animals prefer some

plants to others and consequently the meat eaters that hunt
them concentrate their searches in those same areas. Far from
being an even spread of species, the forest is an immensely
complex patchwork with a plant and animal composition
that varies greatly from place to place.
The patchwork results partly from chance: Trees grow
where their seeds fall, a matter over which the trees have no
control. Beyond that, however, the complexity results from
the fact that particular plant species prosper in some places

Parent birds must work

hard to bring food to
youngsters that may be
as big as themselves.
These are ravens
(Corvus corax).
(Courtesy of
Michael Quinton,
Minden Pictures)
100 TEMPERATE FORESTS

and fail in others, and whether or not they succeed depends

partly on the trees around them. In fact, all these species—
trees, shrubs, herbs, and animals, not to mention the soil
fungi and bacteria—shelter one another, feed on one anoth-
er, and compete for food, water, light, and space. They form a
community that is every bit as complicated as the human
community found in a city.
Scientists were becoming aware of these communities dur-
ing the 18th and 19th centuries. Charles Darwin (1809–82)
was well aware of them, and the relationships within natural
communities are central to his evolutionary theory. In On the
Origin of Species by Means of Natural Selection, published in
1859, Darwin argues that inherited natural variations be-
tween individuals equip some better than others to thrive
and reproduce under the conditions in which they find
themselves living. Those conditions include the influences of
other living organisms.
The German zoologist Ernst Heinrich Haeckel (1834–1919)
was an enthusiastic supporter of Darwin who sought to work
out all the implications of Darwin’s theory. In 1866 Haeckel
published a two-volume work called Generelle Morphologie
der Organismen (General morphology of organisms)—“mor-
phology” is the study of the form of organisms—in which he
discussed a kind of large-scale economy of nature in which
all species are engaged. Seeking a word to describe this idea
he took the Greek oikos, meaning “household,” and logos,
meaning “discourse” or “study,” to produce Ökologie, which
was written in English as “oecologie.” At the International
Botanical Congress in 1893 the spelling was officially
changed to “ecology.”
Haeckel had given a name to the emerging science of
natural communities. In 1923 one of the greatest of the early
ecologists, Sir Arthur Tansley (1871–1955), published a
book called Practical Plant Ecology in which he defined the
new science:

The word ECOLOGY, as is well known, is derived, like the common

word economy, from the Greek oikos, house, abode, dwelling. In
its widest meaning ecology is the study of plants and animals as
they exist in their natural homes; or better, perhaps, the study of
 ECOLOGY OF TEMPERATE FORESTS 101

their household affairs, which is actually a secondary meaning of

the Greek word. [The italics are his.]

In 1935, in a paper in the journal Ecology, Tansley intro-

duced another new term: ecosystem. He defined it fully in the
second edition of Practical Plant Ecology, which was published
in 1946 as Introduction to Plant Ecology and in which he also
explained the term biome.

The most natural conception that has been suggested is that which
regards the whole complex of organisms—both animals and
plants—naturally living together as a sociological unit which has
been called the biome, whose life must be considered and studied
as a whole.
A wider conception still is to include with the biome all the phys-
ical and chemical factors of the biome’ s environment or habitat . . .
as parts of one physical system, which we may call an ecosystem,
because it is based on the oikos or home of a particular biome.

Most of the early ecologists had started out as botanists,

and European ecologists devised ways of classifying commu-
nities of plants. They coined the name phytosociology for the
study of plant communities. American ecologists developed
the new science in a different direction, toward the concept
of climax communities (see “Succession and climax” on pages
118–120).

Food chains, webs, and pyramids

Scurrying about the forest floor, a mouse is searching for
berries that have fallen to the ground, seeds buried beneath
dry leaves, tender young seedlings, and anything else that
might make a tasty snack. Mice living in a temperate forest
eat plant material.
The mouse forages at night, when it is dark. Even a full
moon on a clear night is enough to keep it in its nest, where
it is hidden and safe. It dares to venture forth only when it is
dark. Darkness gives it some protection from its foes, but
only from those that hunt by sight. It is no use against the
owl that sits high on a branch, still, silent, and invisible from
102 TEMPERATE FORESTS

Mice, such as this deer the ground. The owl keeps watch with its ears. It can hear
mouse (Peromyscus and locate the slight rustling a mouse makes as it pushes
maniculatus), feed on aside leaves and grass. An owl can find and catch a mouse in
a variety of plant total darkness.
materials, but they also Plants provide the basic food for the entire forest commu-
provide a tasty meal for nity. This is because plants are the only members of the com-
owls and other hunters. munity with the ability to produce sugars, proteins, fats, and
Their large ears and other nutrients out of air, water, and the mineral substances
sharp eyes allow them to that are dissolved in the soil water (see “Photosynthesis and
remain alert at all times. respiration” on pages 79–85). Food underlies all the relation-
(Courtesy of Gerry Ellis, ships between species. Species compete with each other for
Minden Pictures) food. Plants that compete for light are also competing for
food, because they need light to manufacture sugars by pho-
tosynthesis. Some animal species eat plants, other animals
eat only meat, and there are also animals, including humans,
that eat both. The way food passes from organism to organ-
ism through the forest community is central to how the com-
munity functions.
 ECOLOGY OF TEMPERATE FORESTS 103

Mice eat plant material and owls eat mice. The relation-
ships between plants, mice, and owls look like this: plants
→ mice → owls. The arrow indicates that one group of organ-
isms is consumed by another. A sequence of relationships,
based on what each organism eats, is called a food chain.
There is another forest food chain up in the trees, high above
the forest floor. That one goes: leaves → caterpillars → small
birds → hawks. This food chain is slightly longer; it has four
links rather than three.
Insects and birds die, and uneaten leaves decay. Dead
material provides food for earthworms, fungi, and a series of
other organisms, all the way down to bacteria. The effect of
decomposition is to release nutrient compounds that dis-
solve in the water moving through the soil and are then
absorbed by plant roots (see the sidebar “The nitrogen cycle”
on page 80). This produces a rather more complicated food
chain, shown in the illustration.
Food chains are straightforward and provide a way of
demonstrating clearly the relationships among the compo-
nents of an ecosystem. A food chain diagram provides clues
about what may happen if some component changes. If a
disease epizootic—an epidemic that strikes nonhuman ani-
mals—kills large numbers of mice, for example, there will be
less food for owls, and so many of their young will starve.
Food chains have a major disadvantage, however, because
life is not so simple as they suggest. As well as seeds and fruit,
mice will eat insects. Many birds change their diet between
seeds and insects, depending on what is available. In other
words, the pattern of relationships among the members of
the ecosystem is more like a web than a chain. It is a food web.
A food web diagram attempts to link all the food chains. In
the web shown on page 105, for example, beetles, moths,
butterflies, and bees all feed on flowers. Vireos eat beetles and
butterflies and thrushes eat butterflies, moths, and bees.
Mosquitoes feed on plant sap, but they also feed on the blood
of mice and flycatchers. Flycatchers and mice eat mosqui-
toes. The diagram is complicated, but it is still only a very
simplified version of the feeding patterns in a real ecosystem.
Many relationships are omitted because including them
would make the diagram so complicated as to be confusing.
104 TEMPERATE FORESTS

There is an alternative way to represent ecological relation-

ships based on feeding, and that is to divide the organisms
according to the type of food they eat and then group togeth-
er all the members of each type. The Greek word trophe
means “nourishment,” and relationships based on food are
said to be trophic. Grouping organisms according to their diet
therefore produces a trophic arrangement.
The first group is composed of plants. They produce food
for all the other members of an ecosystem, so they are known
as producers. Animals are consumers. These groups can be rep-
resented graphically as bars of the same height but different
widths and stacked by trophic level with the producers at the
base. Above the producers are the herbivores—plant-eating
animals. They feed directly on the plants, so they form the
first consumer level; they are primary consumers. Carni-

leaves caterpillars small birds hawks

dead organic matter

earthworms
earthworms

fungi

springtails

mites

tree roots single-celled organisms

A food chain
 ECOLOGY OF TEMPERATE FORESTS 105

owl

shrew woodpecker vireo thrush flycatcher mouse squirrel

salamander spiders beetles butterflies bees flies chipmunk

nematodes

earthworms moths mosquitoes

snails

slugs

microorganisms

dead organic matter leaves flowers sap seeds/fruit

vores—meat-eating animals—feed on herbivores, so they A food web

form the next trophic level, as secondary consumers. Some
carnivores feed on other carnivores. These animals are some-
times called top predators, and they form the highest trophic
level, as tertiary consumers. The resulting diagram resembles
a stepped pyramid. It is known as an ecological pyramid or as
an Eltonian pyramid after Charles Sutherland Elton (1900–91),
the British ecologist who first devised it. The illustration on
page 106 shows a typical pyramid.
Pyramids count the organisms at each trophic level, but
there is a problem: Exactly what are they measuring? If the
pyramid shows the number of organisms at each level, it is a
pyramid of numbers. It shows that there are many more plants
than there are animals eating the plants, and many more her-
bivores than there are carnivores, which sounds sensible—
but is it? If the plants are forest trees, each tree can support a
large number of herbivorous animals, so the primary con-
sumers may heavily outnumber the producers. This will be
especially so if most of the primary consumers are insects.
The pyramid will look very different if the producers are grass
plants and the primary consumers are elephants. There will
be many plants, very few elephants, and probably no second-
ary or tertiary consumers because no carnivores hunt adult
106 TEMPERATE FORESTS

tertiary consumers

secondary consumers

An ecological pyramid. primary consumers

The block representing
each trophic layer producers

should be approximately
one-tenth the width of
the block below it.

elephants. Complicating matters further, in most ecosystems

there is a variety of organisms at each trophic level, making it
unclear just what the pyramid means. Pyramids of numbers
are seldom used for this reason.
It would be much better to group the organisms in trophic
levels and then find the weight of all the organisms at each
level. The weight, or more correctly the mass, of organisms is
known as the biomass, and a pyramid that shows the biomass
at each level is known as a pyramid of biomass. Biomass is
measured as the dry weight of organisms, which is the weight
after all the water is removed. This is important, because
water accounts for most of the weight, and the amount varies
greatly from species to species and from time to time. A pyra-
mid of biomass usually has a similar general shape to the one
shown in the illustration. This pyramid is much more useful,
because it makes no distinction between insects, mice, and
deer, or between wolves, bears, and owls, simply lumping all
of them together based on trophic levels.

How energy flows through a forest

Sunshine provides the energy for every ecosystem, because
the producers use sunlight as a source of energy for photo-
synthesis. The amount of food that plants produce represents
the proportion of the sunlight falling on them that photo-
synthesis captures. Photosynthesis is not very efficient;
plants capture no more than about one percent of the sun-
light to which their leaves are exposed.
 ECOLOGY OF TEMPERATE FORESTS 107

Plants use some of the captured energy to grow bigger and

some to repair damaged tissues, but most of the energy is
used for respiration (see “Photosynthesis and respiration” on
pages 79–85). Respiration supplies cells with the energy they
need to perform their ordinary functions. In other words, the
captured sunlight is used for maintenance. No more than
about 10–20 percent of the captured energy—10–20 percent

Winter in a temperate
forest. It is too cold for
plants to grow and
although the pool
remains unfrozen, the
soil is dry because the
water lies as snow above
it, rather than below the
surface, and water in the
soil may be frozen. The
trees have shed their
leaves and will remain
dormant until the
spring. (Courtesy of
Jim Brandenburg,
Minden Pictures)
108 TEMPERATE FORESTS

of that 1 percent, or 0.1–0.2 percent of the sunlight falling on

the plants—is available as food for the primary consumers.
Consumers use most of the energy derived from the food
they eat to maintain their own bodies. It is easy to see that
this is so. Once they are full-grown adults, animals eating a
healthy diet grow no larger. Their size and weight remains
constant, yet they continue to eat regularly. The food they
eat provides the energy to maintain their bodies, allowing
them to move, digest their food, repair or replace damaged or
worn tissue, heal wounds, and, most of all in the case of
mammals and birds, to maintain a constant body tempera-
ture. Not surprisingly, therefore, only about 10–20 percent of
the food that a primary consumer eats is converted into food
for a secondary consumer. Similarly, only 10–20 percent of
the food the secondary consumer eats is converted into food
for a tertiary consumer. The energy represented by the
remaining 80–90 percent is released by respiration and used
to maintain the animal’s body.
The fact that such a small proportion of captured solar
energy passes from one trophic level to the next explains
why pyramids of biomass rarely have more than four levels
and why the width of the levels decreases so rapidly. Primary
consumers need a certain geographical area within which to
find sufficient plant food for their needs. Secondary con-
sumers need a much bigger area in which to find enough ani-
mal food. A family of gray foxes (Urocyon cinereoargenteus)
needs a range of about one square mile (2.59 km2) in which
to find enough food, and a pack of about 12 gray wolves
needs 40–400 square miles (104–1,040 km2). Gray wolves
have largely disappeared from temperate forests because so
much of the original forests were cleared that the remaining
areas were simply not large enough to feed wolves.
Carnivore-hunting carnivores need such a vast area that they
must always have been rare.
It is possible to show the trophic levels in terms of the
flow of energy from one to another in a pyramid similar
to those already described. A pyramid of this kind is called a
pyramid of energy. It is by far the most useful of the ecologi-
cal pyramids, thanks mainly to the insight of Raymond
Lindeman.
 ECOLOGY OF TEMPERATE FORESTS 109

Raymond L. Lindeman (1915–42) was an American ecolo-

gist whose career was cut short by his untimely death at the
age of only 27. In 1942 Lindeman published a scientific paper
that opened up an entirely new area of ecological research.
The paper was called “The trophic-dynamic aspect of ecology”
and it appeared in the journal Ecology. In it Lindeman pro-
posed a new way to study ecosystems, by measuring the
amount of energy that passes from each trophic level to the
next. This initiated the study of ecological energetics.
As with so many good ideas, the value of studying the flow
of energy is obvious once someone else has thought of it. It is
also fairly easy, because energy can be measured. Heat a sam-
ple of biological material in an oven to remove all the water,
then burn the dry matter in an oven equipped with a
calorimeter—an instrument that measures the amount of
energy released. On average, one ounce of plant material
releases 114–128 kilocalories of energy (16.7–18.8 kJ/g) and
one ounce of animal material releases 142–156 kilocalories
(20.9–23.0 kJ/g).
Armed with this information, it is possible to measure the
efficiency with which energy is used. Ecological efficiency is
the percentage of absorbed energy that is converted into bio-
mass. Herbivorous mammals and birds have an efficiency of
0.3–1.5 percent, and carnivorous mammals and birds of
0.6–1.8 percent. Reptiles and amphibians are much more effi-
cient, because they do not use their food to maintain a con-
stant body temperature. Those that are herbivorous have an
efficiency of 9–25 percent and the carnivores are 12–25 per-
cent efficient.
Ecologists also measure energy to calculate the total
amount of new biomass produced each year by the plants in
particular ecosystems. This is called the gross primary pro-
ductivity or production (GPP). Deducting the amount the
plants use in respiration reduces the figure to the net primary
productivity (NPP), and the remainder after account has been
taken of the amount eaten by consumers is the net commu-
nity productivity (NCP).
These calculations allow scientists to compare one ecosys-
tem with another. In fact they are comparing the way differ-
ent ecosystems use the energy available to them. Ecologists
110 TEMPERATE FORESTS

who study temperate forests use ecological energetics to com-

pare broad-leaved forest ecosystems with coniferous forest
and with mixed forest containing both broad-leaved trees
and conifers. They also use this technique to compare natural
forests with plantations, and plantations growing one group
of tree species with plantations growing other species.

Decomposition and the cycling of nutrients

Down on the forest floor there is a layer of dead leaves and
twigs. Scattered about there are fallen branches and occasion-
ally fallen trees—old, almost dead trees brought down by a
recent storm. A group of feathers is all that remains to show
where an unwary bird fell victim to a predator. There may be
a shed snake skin or droppings left by a passing animal, per-
haps to mark the boundary of its territory, or even the dead
body of a bird or mammal.
Material lying on the ground is called detritus, a Latin word
that means “worn down.” It is dead and discarded, but it is
far from useless. The layer of this material never grows any
thicker: It is being discarded all the time, but it does not accu-
mulate because it provides food for an entire range of living
organisms—the decomposers.
Among the detritus, especially in the damper, more shaded
areas, live slugs, snails, wood lice, beetles, ants, millipedes,
earwigs, insect larvae, and earthworms. These animals feed
on dead plant material, so they are known as detritivores.
They shred it into tiny pieces and drag the fragments below
ground to eat them. This activity spreads the material
through the upper soil, where smaller worms, mites, spring-
tails, and similar tiny animals can reach it. The fragments
they leave are food for microscopically small organisms, such
as nematodes, rotifers, and protozoans.
It may look lifeless, but in fact the forest floor is densely
populated. Beneath one square yard of surface area there may
be more than eight million nematodes, 80,000 springtails,
80,000 mites, and 40,000 animals of other assorted species
(10 million, 100,000, and 50,000 per square meter, respec-
tively). In addition, there are fungi and bacteria. Altogether
there are likely to be almost 850 species in every square yard
 ECOLOGY OF TEMPERATE FORESTS 111

(1,000 per square meter). Not all of these organisms are feed-
ing directly on the detritus. There are also small spiders, har-
vest spiders (“daddy longlegs”), centipedes, and other carni-
vores hunting the herbivores.
The soil community forms the unseen part of the ecosys-
tem and, like the visible part of the ecosystem above ground,
its members form a hierarchy that can be described by eco-
logical pyramids. Because they live below ground, it is tempt-
ing to draw their pyramids upside down, but this would be
incorrect. They are just like above-ground pyramids, with the
detritus forming an equivalent to the producer level.
Detritus does not comprise growing plants, of course, and
although all of it is consumed, the consumers are not eating
whole leaves the way grazing mammals do, but particular
ingredients that they select from the menu. Some of those
ingredients are more easily digested than others, and the
most digestible are taken first.
Fruit is rarely found lying on the forest floor. Rich in sugars
and starches, fruits are highly nutritious and easily digested.
Many mammals, birds, and invertebrate animals are attracted
to them, and fungi and bacteria quickly colonize them. They
soon disappear. Other items are consumed more slowly, but
always in the same order, depending on their chemical ingre-
dients. Sugars disappear first, followed by starches. Next the
decomposers attack more complex sugars called hemicellulos-
es that are found in the cell walls of plants. Then pectins, car-
bohydrates, and proteins are consumed. This leaves the com-
pounds that are much tougher to digest. Cellulose is the first
of these to go. It is a complex sugar that provides most of the
structure in the cell walls of plants; it is probably the most
abundant of all the compounds found in living organisms.
Cellulose is followed by lignins (see “How wood forms” on
pages 94–97) and finally by suberins—the fatty compounds
that make the cork in tree bark waterproof. The last plant
compound to disappear is cutin, a mixture of big, very com-
plex molecules that forms part of the waxy coat found on
many leaves and the stems of nonwoody plants. Dead leaves
that form an autumn carpet on the forest floor are among the
last plant material to disappear, but in the end they, too, van-
ish to become part of the soil. The toughest leaves of all are
112 TEMPERATE FORESTS

the needles that fall from coniferous trees. Their thick,

strong, outer coat makes them much more durable than the
thin, more delicate leaves of broad-leaved plants that need to
last for only one season.
At each stage in the process the organisms responsible
leave wastes of their own. The overall effect is to break down
the once-living tissues step by step until they are finally
reduced to carbon dioxide, water, and a variety of simple
chemical compounds that dissolve in the water passing
through the soil. This is the water that plant roots absorb,
taking up the dissolved nutrients at the same time. Decom-
position is the process by which nutrients are recycled be-
neath the forest floor.

Habitats and niches

Living in a city has many advantages. Supermarkets supply
food at all hours of day and night, houses and apartments
provide shelter, medical services help people who are sick,
and there are emergency services, mass transit and a wide
range of entertainment and amenities. Provided a person is
not too poor to enjoy its benefits, the city has much to offer.
So far as people are concerned, the city is good habitat—a
Latin word that means “inhabits.”
People do not have the city entirely to themselves, of
course. Many species of birds also find it a good place to live,
as do the rats that live belowground, emerging at night to
feed on the garbage people discard, the mice that sneak into
homes, and the half-wild cats that hunt them. Spiders build
webs to catch the innumerable flying insects, while cock-
roaches and beetles eat their way through the food stores and
woodworms tunnel into old furniture. The city is good habi-
tat to all of these and many more.
It is not good habitat for every species, however. Red foxes
(Vulpes vulpes) have moved into many towns and thrive
there, but the bobcat (Felis rufus) stays well away. What is
good habitat for one species may be extremely inhospitable
for another. People sometimes describe an attractive area as
“good habitat,” but strictly speaking the statement is mean-
ingless unless it is qualified. The area may be good habitat for
 ECOLOGY OF TEMPERATE FORESTS 113

particular species or it may be good habitat for all the species

typical of a certain ecosystem—temperate forest habitat, for
example, or marshland habitat—but it cannot be good habi-
tat for every species.
Habitats can change. If a habitat is stable, changing little
over the years, the species occupying it will be adapted to a
situation in which the resources on which they depend—of
food, water, shelter, and nesting sites—are continually avail-
able. In this situation the species will tend to produce few off-
spring, but will care for them over a prolonged period so that
the young are almost fully grown by the time they become
independent. Ecologists call this reproductive strategy K-
selection. In a stable habitat it gives the young the best possi-
ble chance of surviving. Humans reproduce in this way; ours
is a K-species.
Unstable habitats are liable to change fairly suddenly and
with little warning. The K-selection strategy would not work
in such a situation, because the young would be subjected to
the change before they were old enough to cope with it. In
unstable habitats species are more likely to adopt an r-selection
strategy. R-species produce many young that enter the world
tiny and extremely vulnerable, but grow fast, eating the food
around them before it disappears. By the time the habitat
deteriorates, enough of them will be full grown to ensure
that the population survives. Most of the young will perish,
but this strategy gives them the best chance possible.
K-selection is always written with a capital K and r-selection
with a lower case r because the terms were first used in an
equation for calculating population size, called the logistic
equation. The logistic equation was published in 1838 by the
Belgian mathematician Pierre-François Verhulst (1804–49);
it is:
N(t) = K/(1 + ec–rt)

where N(t) is the population size at time t, K is the carrying

capacity of the habitat, e (approximately 2.718) is the base of
natural logarithms, c is a constant defining the value of N at
time t = 0, and rt is the rate of increase (r) to time t. The car-
rying capacity is the number of individuals the habitat can
support; if the population increases above this number it will
114 TEMPERATE FORESTS

later decrease again, and if it falls below this number it will

later increase.
The most unstable of all habitats appears only occasionally
and disappears rapidly. It is said to be ephemeral. A puddle of
water that vanishes within a few hours and the body of a
dead forest animal are examples of ephemeral habitats. They
attract organisms—mainly bacteria and insects—that multi-
ply rapidly, feed, grow, and then move away during the brief
time the habitat remains.
The gray squirrel (Sciurus carolinensis) lives in the forest. It is
a forest animal, so a temperate forest provides it with suitable
habitat. Its habitat is the place it lives—that it inhabits.
Knowing the habitat that suits a particular species tells us
nothing about how the species lives, however. It tells us where
it lives, but not what it does there. In fact, in spring and sum-
mer the squirrel spends most of its time high in the trees. The
leaves are especially nutritious then, and the squirrel is eating
them. It builds a big, untidy nest called a drey from leaves, far
out along a branch or, if it can find one, in a hollow in the
tree. It also builds flat feeding platforms on branches, where it
can eat and rest in comfort. By fall the leaves are becoming
less nutritious and the squirrel spends more time on the
ground, looking for fallen nuts and the fruiting bodies of
fungi. That is when it collects nuts and stashes them away for
the winter. In winter it spends most of its time on the ground,
eating the food in its stores. This is what the squirrel does, the
way it lives; that is its niche, a word derived from the French
nicher, meaning “to make a nest” or “to nestle.”
A niche does not exist until a species occupies it. Leaves,
nuts, fungi, trees and their branches are just those things
until a squirrel comes along and finds a use for them. To the
squirrel they represent an opportunity that it is equipped to
seize, using them to construct its niche. The organism actual-
ly creates the niche that it then occupies, and every niche is
unique. If the species should disappear, the niche it formerly
occupied will be left vacant. Other members of the same
species may return to occupy it, or another species may find
it congenial and take it over. In this case the niche exists
prior to being occupied, but only because a previous species
had already created and defined it.
 ECOLOGY OF TEMPERATE FORESTS 115

How trees are adapted to climate

Trees, like all other green plants, depend on photosynthesis
to provide the sugars and energy they need to grow. Photo-
synthesis and its opposite, respiration (see “Photosynthesis
and respiration” on pages 79–85), consist of sequences of
chemical reactions. The speed of these reactions, and of all
the other reactions that take place in a plant, varies according
to the temperature. They cease altogether if the temperature
falls below about 21°F (–6°C). When it is this cold, plants can-
not grow at all and they become dormant. The reactions
work best at temperatures higher than 41°F (5°C), and then
their speed doubles with every 18°F (10°C) rise in tempera-
ture up to about 85°F (29°C), which is the temperature at
which photosynthesis reaches its highest sustainable rate.
The rate continues to rise with temperature up to about 95°F
(35°C), but the acceleration is brief and the rate quickly falls
back to its previous level. Photosynthesis starts slowing rap-
idly as the temperature rises past 105°F (40°C), and most
plants die if the temperature remains higher than 113°F
(45°C) for very long.
Prolonged high temperatures pose no threat to temperate
forest plants, because the weather in temperate regions is
never that hot for long enough to harm them. Winter tem-
peratures usually fall below 41°F (5°C), however, and they
often remain below 21°F (–6°C) for weeks at a time. There are
times, therefore, when plant growth ceases.
A period of dormancy presents no problem for trees, pro-
vided it does not continue for too long. In order to repro-
duce, broad-leaved trees must produce flowers, fruits, and
seeds, and this takes time, during which the temperature
must remain higher than 41°F (5°C) and for part of which it
must be considerably higher. Many tree species, including
some maples (Acer species), lindens (Tilia species), and birch-
es (Betula species), produce flowers very early in spring,
before their leaves open, to take immediate advantage of the
rising temperature. This gives them time to ripen their seeds
by late summer or early fall.
Early flowering is an adaptation to the length of the grow-
ing season, but it is risky. Late frosts can postpone flowering
or, if they occur after the flowers have opened, destroy the
116 TEMPERATE FORESTS

flowers. Cool summers slow the rate at which seeds ripen. If

the flowers are destroyed the tree will produce no seed, and if
its seeds form too late in the season they may not be fully
ripe by the onset of winter.
Trees live for many years and an occasional failure to pro-
duce seeds is not serious, but a species that frequently fails to
produce seeds is at risk of dying out. As old trees die, the
shortage of seeds means there will be too few seedlings to
replace them, and other species, better at producing seeds,
will take their place.
Some species escape this trap by reproducing vegetatively
(see “How trees reproduce” on pages 85–87). Trees that repro-
duce through suckers—shoots that arise from roots some dis-
tance from the main part of the plant—do so more in higher
latitudes, where the climate is cooler, than in lower, warmer
latitudes, where they rely more on producing seed.
During the coldest part of winter, when the ground is
frozen, trees face a different problem. Their roots are unable
to absorb ice, and they have no means of thawing the ground
to release liquid water. Consequently, the winter is the equiv-
alent of a drought—even if the landscape lies beneath a deep
layer of snow. On sunny days the leaves may warm suffi-
ciently for photosynthesis to start. They will open their stom-
ata to allow carbon dioxide to enter and oxygen to leave, but
when they do so water will evaporate (see “Evaporation and
transpiration” on pages 66–69) and, with the roots embedded
in frozen ground, there will be no rising water to replace it. It
will not take many sunny winter days before the tree is seri-
ously, even fatally, dehydrated.
Trees have adapted to this in two ways. Some produce
thick leaves with a waxy outer covering that minimizes water
loss. Holly (Ilex aquifolium) and ivy (Hedera helix) are well
known examples. Others bear leaves only during that part of
the year when leaves are useful. Trees that do this are said to
be deciduous.
Deciduous trees produce thin, flimsy leaves. They are not
very durable, but this does not matter, because they need
to last for no more than five or six months. The leaves are
disposable. This may sound wasteful, but in fact it is very
efficient.
 ECOLOGY OF TEMPERATE FORESTS 117

To grow flimsy leaves, the tree does not have to expend Enough light penetrates
large amounts of materials, and it can grow them quickly. the forest from the side
The leaves open early in spring and reach their full size in a to allow plants to
matter of days. As summer draws to a close, the days grow blanket the ground, but
shorter and the temperature begins to fall. When they detect only where the sunlight
these changes, the leaves reduce their production of auxins— reaches. Elsewhere the
substances that stimulate plant growth—and increase their forest floor is in deep
production of ethylene, which inhibits cell growth. Tree shade and few plants
growth slows and then ceases. The leaves cease producing are able to grow. This
chlorophyll, and many of the nutrients in the leaves flow forest is in Bavaria,
back into the tree branch or trunk, where they are stored for Germany. (Courtesy
the winter. This process exposes the remaining pigments, of Konrad Wothe,
changing the color of leaves from green to yellow, orange, Minden Pictures)
and red. Near the base of the leaf stalk, or petiole, there is a
thin layer of parenchyma cells. This layer weakens until
leaves begin to fall, and a few windy days strip the trees bare.
118 TEMPERATE FORESTS

The tree has stored most of the useful chemical compounds

from the leaves and the remainder are released as the leaves
decompose (see “Decomposition and the cycling of nutri-
ents” on pages 110–112).

Succession and climax

To an ecologist, demolition sites are interesting places. Once
the old buildings have gone and the rubble has been carted
away, all that remains is bare earth, rough, uneven, and com-
pacted, where heavy machines have driven across it. Unless it
is the middle of winter and bitterly cold, within a few days
the first plants will have pushed above the surface. In the fol-
lowing few weeks, plants will spread until they cover the
entire site. Then they will flower and produce copious
amounts of seed—often the type of tiny seed that floats
through the air on a “parachute” of fine hairs called a pappus.
Ecologists are especially interested in what happens next.
Other plants have seeds that are slower to germinate. They
take longer to appear, but when they do they grow strongly,
shading out the first arrivals, called the pioneers, but feeding
on nutrients released by their roots. Grasses appear, only to
die away when the first woody shrubs rob them of the direct
sunlight they need. Leave the site for long enough—though
usually the builders start developing the area before this
happens—and tree seedlings may begin to rise above the
other plants. Eventually the site might turn into a tiny patch
of forest.
This series of changes in the composition of the plant com-
munity is called a sere or succession, and each community
that appears and is then replaced is called a stage or seral
stage. The final community that becomes more or less perma-
nently established is called the climax. Over much of the
northeastern United States, the first Europeans to arrive in
North America found forests dominated by various combina-
tions of beech, birch, maple, and oak trees, together with the
shrubs and herbs associated with them. These plant commu-
nities formed the climax vegetation. Oak and hazel dominat-
ed the primeval forest that once covered much of lowland
Britain and therefore constituted the British climax.
 ECOLOGY OF TEMPERATE FORESTS 119

Scientists began studying the way natural communities de-

velop and change over time toward the end of the 19th cen-
tury, and the idea of a sere producing a climax became very
influential by the middle of the 20th century. Unfortunately,
it can be misleading.
In the first place, one stage may or may not prepare the
way for the next—for example, by releasing nutrients.
Sometimes this is what happens, but there seems to be no
way to predict it and it is far from inevitable. Nor is it possi-
ble to predict what the climax will be. If an area of forest is
cleared and then left to regenerate, it will usually develop as
forest once more, but not necessarily with the same trees in
the same proportions. Which seeds germinate first and
which trees manage to crowd out their rivals and become
established is a matter of chance.
The idea of the climax suggests a community that remains
unaltered for as long as it remains undisturbed. This seems to
suggest a “balance of nature” in which every species has its
place and function, but natural communities are not like
that. There can be no “balance of nature” because communi-
ties are constantly changing, developing, and responding to
disturbances that occur naturally and often.
Nowadays ecologists recognize that the relationships with-
in plant communities, such as forests, are much more com-
plicated than they were once thought to be. It is inaccurate
to think of a regular series of stages through which bare
ground is transformed into a type of vegetation that defines
the area—as forest or grassland, for instance. Nevertheless, it
is still true that plants will appear to colonize an area of bare
ground. Pioneer plants that produce large quantities of rapid-
ly germinating seed will emerge first. These will give way to a
greater variety of stronger plants, and little by little the bare
ground will be transformed. This will happen, but there is no
way to predict the order in which species will appear or the
community that will finally result, and there is no way of
knowing when the process has reached a conclusion—if it
ever does.
This realization has one major implication. In the days
when people believed that a forest, essentially identical to
the original natural forest, would develop on land from
120 TEMPERATE FORESTS

which forest had been cleared, it was enough to protect the

site and allow the natural succession to proceed. Scientists
know now that this will not guarantee the re-creation of the
original forest or even of forest at all. The succession might
terminate in impenetrable scrub, and if a forest did emerge, it
would include species that have been introduced and
become invasive. The only way to create a forest resembling
the original forest is to plant it and then manage it carefully.

Fire, and trees that depend on it

A forest fire is a terrifying sight. The smoke is visible from
miles away and the fire advances as a roaring wall of flame,
leaping from treetop to treetop, devouring everything in its
path. At least, that is the way the most sensational wildfires
behave—the ones that approach the outskirts of cities, burn-
ing homes and forcing local residents to evacuate.
There have been many damaging wildfires in recent years,
but they are not new. In 1902 a fire at Yacholt, Washington,
burned about 250,000 acres (100,000 ha) of forest, and a sim-
ilar area burned in 1933 at Tillamook, Oregon. The fires that
struck southern California in October 2003 were much big-
ger. Driven by Santa Ana winds that blow from Arizona and
Nevada toward the Pacific, accelerating as they are funneled
through the Santa Ana Canyon, the fires burned about
740,000 acres (300,000 ha), killing two people in Mexico and
20 in California. More than 100,000 people were driven from
their homes.
Not all wildfires are like that and, far from being harmful,
most serve a useful purpose in rapidly converting dead plant
matter into fertilizer. Some wildfires are started deliberately by
arsonists, but lightning is responsible for igniting most. It is
the litter that burns—the accumulation of dead leaves, twigs,
and branches that have dried out during the summer—and
the fire moves rapidly but remains close to the ground. It
destroys shrubs and herbs and scorches tree trunks, and it will
kill some tree seedlings and saplings, but otherwise it does no
harm. The temperature among the burning litter is usually
about 194–248°F (90–120°C). A few inches below ground the
temperature is only slightly higher than normal. The fire
 ECOLOGY OF TEMPERATE FORESTS 121

spreads over a fairly small area and then dies down. Ground
fires burn belowground, consuming fuel more slowly than
surface fires and burning for longer. They consume root sys-
tems, producing no flames, but most ground fires die for lack
of oxygen before they have traveled very far.
A surface fire becomes much more dangerous if it ignites
the crowns of the trees. It is then a crown fire. Hot air rising
from a burning tree crown lifts burning cinders high into the
air. Carried by the wind, the cinders fall into the crowns of
other trees, igniting them. In this way the fire can advance
very rapidly. Really intense forest fires can generate fire-
storms (see “Forest fires” on pages 72–73).
Fires are rare in broad-leaved deciduous forest. Material
lying on the ground is usually too wet to burn, and the trees
are not very flammable. Their leaves contain a high propor-
tion of water and their wood is also fairly moist. Coniferous
trees have drier leaves and their wood contains resins, which
are flammable, so fires are more common in coniferous
forests growing in areas where summers are warm and dry.
Fire is an entirely natural phenomenon in forests of this type
and the forest trees are adapted to it. The fire clears away the
dead leaves and wood that have collected on the forest floor,
leaving behind a layer of ash, rich in plant nutrients.
Indeed, they do more. Some plants exude chemical sub-
stances that inhibit bacterial activity nearby. The fire burns
some of these substances and produces charcoal, which
absorbs others, allowing the bacteria to flourish—and the
bacteria include species that “fix” atmospheric nitrogen (see
the sidebar “The nitrogen cycle” on page 80). After the fire,
the ground surface is blackened. A black surface is better than
a pale surface at absorbing the warmth of the Sun, so the soil
is warm. This stimulates microbial activity, seed germination,
and the growth of young plants.
Attempts to prevent fires can result in uncontrollable wild-
fires. Extinguishing fires as soon as they are detected protects
buildings in and close to the forest, but it also allows surface
litter to accumulate. When eventually the litter ignites, the
amount of fuel is so large that the fire quickly rages out of
control. This is what happened in Yellowstone National Park
in 1988 (see the sidebar on page 122).
122 TEMPERATE FORESTS

Trees that grow in areas prone to fire have adapted in vari-

ous ways. Some, such as quaking aspen (Populus tremuloides),
balsam poplar (P. balsamifera), and paper birch (Betula papyri-
fera), produce copious quantities of seed that disperse widely.
When fire has cleared and fertilized the ground, seeds of these
trees germinate and the trees grow so rapidly they are able to
produce a fresh crop of seeds before the bigger but slower
conifers have had time to grow tall enough to shade them.
Despite being conifers, redwoods such as the coast red-
wood (Sequoia sempervirens) and Sierra redwood (Sequoia-

Yellowstone and coping with fire

In summer 1988 fire swept through Yellowstone National Park in the western United
States. All attempts at control failed and the fires did not die down until the first substan-
tial falls of winter snow extinguished them. By that time the fires had burned about 1 mil-
lion acres (400,000 ha) of the park, amounting to about 45 percent of the total area. The
fires damaged the vegetation, but in most places without killing all of it. Over about
20,000 acres (8,000 ha), however, the vegetation was almost totally destroyed.
When at last the smoke cleared, the landscape was one of utter devastation and it was
difficult to believe that the park could recover. Yet in spring 1989 plants began to emerge
from the ashes, and within a few years not only had the burned area recovered, its natural
communities had been invigorated. Ninety percent of Yellowstone is forested and 90 per-
cent of the trees are lodgepole pine (Pinus contorta var. latifolia). This is a species of trees
that store their seeds in the canopy, often for years, until hot air rising from a fire stimu-
lates their release. The forest is adapted to fire.
Fire occurs naturally in Yellowstone every few years, but for some years prior to the
1988 fire the policy had been to suppress fires as quickly as possible. This allowed dry,
dead plant material to accumulate. When it was ignited, probably by lightning, the result-
ing fire was uncontrollable.
A new management scheme was adopted in 1992, dividing the park into three zones.
In one zone, comprising about 10 large areas, fires are ordinarily suppressed, but con-
trolled fires are started deliberately if too much flammable material accumulates. In anoth-
er zone, covering most of the park, fires are allowed to burn but are controlled. In the
third zone, comprising a strip 1.5 miles (2.4 km) wide just inside the park boundary, fires
are permitted under even more strictly controlled conditions.
 ECOLOGY OF TEMPERATE FORESTS 123

dendron giganteum) are difficult to burn. They have spongy

bark that does not catch fire readily, and the bark is thick
enough to protect the living tissue beneath from all but the
hottest fires. Even then, if the fire is hot enough to burn the
tree, new growth often sprouts from the stump. Other trees,
including pitch pine (Pinus rigida), also recover by producing
new growth from the trunk of a dead standing tree.
These are trees that can survive fire. Others actually
depend on it. Longleaf pine (P. palustris), lodgepole pine (P.
contorta var. latifolia), and jack pine (P. banksiana) are seroti-
nous. This means that they delay releasing their seeds, keep-
ing their cones tightly closed until they are warmed by hot
air rising from a fire on the ground below. Then the cones
open to release seeds that germinate in the warm ashes.
Most forest animals also survive fire. Birds and the larger
mammals see or smell the fire long before it reaches them
and simply move away. This can be a dangerous strategy for
mammals, however, because just ahead of the fire there are
predators looking out for escapees. Other animals shelter
beneath logs or in burrows. During the 1988 Yellowstone fire
about 250 elk (Cervus elephas) died from smoke inhalation,
out of an elk population of about 31,000.
Forests where fire occurs fairly regularly at intervals of a
few years come to be dominated by tree species that are
either tolerant of fire or dependent on it. There are also
species like aspen that are able to exploit the opportunity to
grow rapidly and produce seed on the sites left temporarily
empty by fire. Consequently, all of the trees in the regenerat-
ed forest tend to be the same age. Most fires affect only a lim-
ited area of the forest, however, so these forests often consist
of a patchwork of areas, each patch containing trees that are
all of the same age, but with the trees in one patch of a dif-
ferent age from those in another. This type of forest commu-
nity is known as a fire climax or pyroclimax (from the Greek
pyr, which means “fire”).

Structure of a temperate forest

Trees expose their leaves to the sunlight, and in order to max-
imize that exposure their branches spread as widely as possi-
124 TEMPERATE FORESTS

ble. Many coniferous trees bear their longest branches near

the bottom and the branches are progressively shorter at
higher levels. This produces a conical shape. In contrast, the
leafy branches of broad-leaved trees are mainly near the top,
where they form a crown. In a forest the crowns of adjacent
broad-leaved trees or the longest branches of coniferous trees
Tall, slender aspen trees touch or overlap, producing a closed canopy. If the trees are
allow ample light to more widely separated than this, so there is an open canopy,
reach the ground, so the area is known as woodland. A closed canopy shades all of
the forest floor is the ground; the open canopy of woodland usually shades
densely carpeted with about 40 percent of the ground area.
vegetation. (Courtesy Although a forest is defined as an area with a closed
of Fogstock) canopy, this is not literally true. There are gaps, even in conif-
 ECOLOGY OF TEMPERATE FORESTS 125

erous forests where the trees retain their leaves throughout

the year. Old trees die and fall, creating a gap in the canopy
that remains open until a young tree grows up to close it
again. In a broad-leaved deciduous forest the trees are bare in
winter, so there is a time, from late fall until early spring,
when there is no canopy.
Where the canopy is open, direct sunlight reaches the
ground. This allows tall herbs, called forbs, to grow. These
plants cannot survive in the deep shade of a closed canopy.
The herbs in a deciduous forest take advantage of that brief
interval between the time when the temperature is warm
enough for photosynthesis and plant growth and the time
when the leaf buds open on all the trees (see “How trees are
adapted to climate” on pages 115–118). The herbs produce
flowers, which are small, because there is not enough time to
produce bigger ones and often brightly colored, to attract
insects, then set seed and die down as the canopy closes and
the light level falls. These herbs, such as primulas, snow-
drops, narcissi, and crocuses, are the spring flowers that bring
beauty to the forests, and they are widely grown in gardens.
Coniferous forests shade the ground all the time and so they
do not harbor spring flowers.
Among the herbs are also climbers, such as ivy, and shrubs
and young trees at various stages of growth. The forest con-
sists of mature trees, but it contains much more than tall
trees, and its plants form distinct horizontal layers. These
layers are sometimes called stories, like the stories of a tall
building.
Mature, full-grown trees form the canopy. These trees are
known as the dominants. Most of the dominants are of simi-
lar height. Any that grow taller and protrude above the
canopy are emergents. A tree that grows taller than its neigh-
bors is able to capture more sunlight, but it is also more
exposed to the drying effect of the wind above the canopy.
The wind can kill leaf buds, so trees must pay a high price to
win that extra light.
Among the dominants there are slightly smaller trees,
known as codominants, and below those are still smaller trees,
called subdominants. The dominants, codominants, and sub-
dominants all belong to the same species. When a dominant
126 TEMPERATE FORESTS

sapling of dominant sub-dominant co-dominant dominant co-dominant sub-dominant dominant

dominant
species

shrub shrub climber shrub

(e.g., ivy)

Forest structure. The dies, the nearest codominant grows taller to take its place and
plants in a temperate a subdominant takes the place of the codominant. Together,
forest form layers. the dominants, codominants, and subdominants compose
The tallest trees the uppermost story, called the canopy layer.
(dominants and Beneath the canopy layer there are smaller trees, still of the
codominants) form same species. The growth of these saplings is suppressed by
the canopy. Saplings the lack of light, but if the canopy should open they will
of these species form grow up to join it. There are also smaller trees that, although
a lower layer, with they are their full size, are too short to reach the canopy.
shrubs below them These saplings and small trees comprise the understory. The
and herbs growing diagram shows a typical forest arrangement.
close to the ground. Shrubs that are between three feet (1 m) and 33 feet (10 m)
tall make up the shrub layer. Woody plants such as gooseberry
(Ribes species) and witch hazel (Hamamelis virginiana) that are
less than three feet (1 m) tall form a dwarf shrub layer.
The tall herbs—nonwoody plants—form a story below the
dwarf shrub layer known as the tall herb layer or forb layer.
Forbs include such plants as the yellow trout lily (Erythronium
americanum), large-flowered trillium (Trillium grandiflorum),
 ECOLOGY OF TEMPERATE FORESTS 127

and lupins (Lupinus species). The smaller herbs, such as prim-

roses (Primula species), comprise the herb layer or field layer.
The lowest story of all is the ground layer, comprising mosses,
lichens, and other plants that grow close to the ground.
Not every temperate forest has all of these stories, and the
number of stories often varies from one part to another of
the same forest. Many forests lack an understory, for exam-
ple, but most have a rich field layer.

Forest communities
Unless one keeps to well-marked paths it is easy to become
lost in a forest. The pattern of trees and shrubs looks the
same in every direction, giving the impression that the com-
position of the forest is the same everywhere, with its plant
species distributed uniformly.
It is true that except in clearings the structure of the forest
has the same principal components everywhere (see “Struct-
ure of a temperate forest” on pages 123–127), but the compo-
sition of an undisturbed forest varies greatly from place to
place. Trees compete with each other for light, water, and
nutrients, and they vary in their susceptibility to attack from
animals and disease. These factors produce changes in the
local composition of the forest, as particular species thrive in
some areas but are less healthy in others. Animals that feed
on the vegetation also vary from place to place because they
are drawn to their preferred food plants. Consequently the
predators of those animals also occur in some places but not
others.
Far from being a uniform expanse of plant and animal
species, a forest is more like a series of small compartments,
each a little different from those around it. The resulting pat-
tern resembles a mosaic or an elaborately tiled floor.
Forest compartments are not quite like tiles, however,
because they lack sharp edges. Consequently, compartments
overlap each other, and where they overlap, adjacent com-
partments share certain species. The diagram on page 128
shows two adjacent compartments. For sake of simplicity,
each compartment contains only three species, colored blue,
yellow, and red. One compartment contains squares and the
128 TEMPERATE FORESTS

other circles. Where the compartments overlap, there are

blue, yellow, and red circles and squares. This combination of
species produces different ecological conditions in which
species can thrive that do not occur in either of the compart-
ments. These are shown as blue, yellow, and red triangles.
Consequently, while each compartment contains three
species, the overlap area contains nine. This richness result-
ing from the overlap is known as the edge effect.
The diagram gives the impression that the edge effect is
beneficial, and so it would be if species were distributed even-
ly throughout each forest compartment. But they are not.
Some species need the protection of the species around them,
so they grow only near the center of a compartment and fail
to thrive near the edge. Dividing the forest into smaller and
smaller compartments, for example, by driving roads
through it or clearing areas, increases the overall length of
forest edge, but it also confines the interior species into
smaller and smaller areas until eventually they may disap-
pear.
Edges tend to increase in length naturally. More light pen-
etrates at the outer edge of a forest and the climate is warmer
and drier there than it is deep inside. This favors plants that
require light and warmth and suppresses those preferring
cool shade. The edge climate is also windier, however, and

The edge effect. Where

two ecosystems overlap,
the overlap area
supports species from
both, plus species
peculiar to the overlap.
 ECOLOGY OF TEMPERATE FORESTS 129

Summer is a time of
abundant food, when
many birds and
mammals raise their
young. But sometimes
there are accidents.
This four-week-old
great gray owl (Strix
nebulosa) has jumped
out of its nest in an
Idaho forest. (Courtesy
of Michael Quinton,
Minden Pictures)

trees are more likely to be blown down, often bringing others

down with them. Light penetrates and this extends the edge
a little farther into the interior. It is possible for a large forest
to be reduced to nothing but edge: Local clearances and roads
divide the entire forest into patches so small that the interior
species are no longer present. Edge forests are sometimes
called by an English version of the Hindi word jangal, which
describes the type of vegetation often seen beside roads: They
are jungle.
Forests are often identified by their dominant species. The
northern hardwood forest that once covered most of eastern
Canada and the United States to the east of Minnesota
consisted of white pine (Pinus strobus), sugar maple (Acer sac-
charum), beech (Fagus grandifolia), yellow birch (Betula allegh-
aniensis), and varying amounts of hemlock (Tsuga canadensis).
Farther south in Illinois and Wisconsin the forest comprised
130 TEMPERATE FORESTS

sugar maple and basswood (Tilia americana), sometimes with

red oak (Quercus rubra). From the southern edge of this forest
as far as Texas the forest was mainly black oak (Q. velutina),
white oak (Q. alba), pignut hickory (Carya glabra), and shag-
bark hickory (C. ovata), with an understory of flowering dog-
wood (Cornus florida). In the Gulf states the forest, extending
to southern Massachusetts on its eastern side, was of white
oak, loblolly pine (P. taeda), and shortleaf pine (P. echinata). To
the east of that forest there was red oak, chestnut oak (Q. pri-
nus), and tulip tree (Liriodendron tulipifera).
Most European forests contain beech (Fagus sylvatica) as
the most widespread tree. This grows with sessile oak (Q.
petraea) and pedunculate oak (Q. robur), field maple (A.
campestre), ash (Fraxinus excelsior), and in some places small-
leaved lime, also called littleleaf linden (Tilia cordata).

Animals that attack trees

White-tailed deer (Odocoileus virginianus) are highly adapt-
able animals that thrive everywhere from the edge of the
North American tundra to northern Peru and Brazil, and they

Deer are woodland

animals that feed on
ground plants and the
leaves of shrubs and
low trees. (Courtesy
of Fogstock)
 ECOLOGY OF TEMPERATE FORESTS 131

have been introduced in Scandinavia and New Zealand.

Their success is due to their ability to eat a wide variety of
plants. Like all other deer, they are browsers, which means
that they eat the leaves of trees and shrubs, a diet that
attracts them to forests. Elk (Cervus elephas), known in Britain
as red deer, and the European roe deer (Capreolus capreolus)
also seek food inside forests.
Deer cannot reach very high, however, so they are able to
feed only on low-growing plants and on leaves from the
lower branches of trees. Their food is most plentiful around
the edges of forests, and they seldom penetrate deep inside
the forest, except when fresh young plant growth is abun-
dant in an area cleared by forest fire (see “Fire, and trees that
depend on it” on pages 120–123).
As well as feeding on leaves, stags (adult male deer) also
rub their antlers against tree trunks to remove the fur-
covered skin called velvet that covers them when they first
appear and supplies them with blood while they are forming.
This can strip away patches of bark. The loss of bark is not
directly serious because it does not damage the vascular cam-
bium, but exposure of the living tissue allows infection to
enter and the injured tree may become diseased as a result.
Deer are not the only mammals to enjoy the regeneration
that follows fires. Berries begin to appear about two years
after a fire and they remain abundant for about 20 years.
Black bears (Ursus americanus) enter the forest in search of the
berries. Moose (Alces alces), known in Europe as elk, feed on
sprouting trees and begin to arrive about three years after a
fire. Snowshoe hares (Lepus americanus) feed in the forest five
to 35 years after a fire, nibbling the emerging plants. Once
the young trees are big enough to build dams and lodges,
beavers (Castor canadensis) arrive in parts of the forest close to
rivers. Many deer and moose shelter in forests even when
they are not feeding in them.
Wolves (Canis lupus) do not eat leaves, but they hunt deer
and follow them into the forest. A forest comprising open
spaces and compartments of different ages following fires is
likely to provide food for wolves.
American bison (Bison bison) do not enter woodland, but
European bison (Bison bonasus) are forest animals. Bison eat
132 TEMPERATE FORESTS

forest plants, but they also trample and uproot plants they do
not eat. If there were large herds, bison might cause serious
damage, but these animals survive only in the Bia l⁄ owieża
Forest (Belovezhskaya Pushcha), where there are too few of
them to be harmful (see “How ecological ideas developed” on
pages 98–101). Wild boar (Sus scrofa) are common in main-
land Europe. They eat a variety of foods, but especially roots
and tubers that they dig up from the forest floor. In doing so
they also destroy seedlings and other plants.
Large mammals feed mainly in and around the edges of
clearings. They have no effect on the interior of the forest,
and ordinarily trees quickly recover from any damage the
animals cause. There are circumstances, however, in which
the damage can be excessive, and deer have caused consider-
able harm in some North American and European forests.
The problem arises when roads and isolated areas that
have been clear-felled divide the forest into fragments, there-
by extending the total length of forest edge (see “Forest com-
munities” on pages 127–130). Opening up the edge allows
deer and other large animals to penetrate more deeply into
what was formerly the forest interior, and by increasing the
area of suitable habitat it allows their populations to increase
to a level where the damage they do becomes serious. The
extended forest edge also attracts predators, such as foxes and
domestic dogs and cats. They destroy the eggs of ground-
nesting birds, kill fledglings, and even attack adult birds that
once nested securely deep in the interior of the forest.

Diseases and parasites of trees

Like any other living organism, a tree can fall sick, and from
time to time there are devastating outbreaks of disease that
destroy countless millions of trees. Two of the most serious
outbreaks in recent times occurred in North America and in
Europe.
Chestnut trees (Castanea dentata) were once common
throughout the forests of eastern North America. Then, in
about 1904, a consignment of chestnut trees arrived at New
York from China. These trees were infected with a fungus,
Endothia parasitica. No one knew about the infection.
 ECOLOGY OF TEMPERATE FORESTS 133

Probably they would have taken no action even if they had

known, because in China Endothia parasitica produces only
very mild symptoms. When the fungus infected chestnut
trees on Long Island, New York, however, the effect was very
different. It killed the American chestnut trees and also
European sweet chestnut trees (C. sativa) that were being
grown in orchards. The disease spread rapidly. There was no
way to control it and attacks were invariably fatal. It was
called chestnut blight and by about 1940 it had destroyed
almost all the chestnut trees in North America.
The European outbreak was due to the resurgence of a dis-
ease first described in the 19th century. In 1838 John
Claudius Loudon (1783–1843), an English gardening writer,
reported a disease that was afflicting elm trees (Ulmus
species). The disease broke out again in France in 1918, and
in 1919 Dutch scientists identified the responsible fungus.
Because the fungus was first identified in the Netherlands,
the disease it causes came to be called, perhaps unfairly,
Dutch elm disease.
Infection begins when tiny elm bark beetles (mainly
Scolytus scolytus and S. multistriatus), no more than 0.1–0.2
inch (3–5 mm) long, burrow into the bark and carve out nup-
tial chambers where they mate and lay eggs. Their larvae feed
on the wood and adults feed on sap. The beetles attack in
vast numbers and are capable of killing a tree without help.
They often have help, however, in the form of the fungus
Ophiostoma ulmi, spores of which they carry into the tree.
The fungus then grows in the vascular tissue (see “How wood
forms” on pages 94–97) and blocks the xylem vessels. That is
what kills the tree.
A serious outbreak of Dutch elm disease began in Europe in
1920 and spread to North America by 1930, to where the
infection was carried by S. multistriatus but sustained by an
American bark beetle, Hylurgopinus rufipes. By 1940 the dis-
ease had died down in Europe, but it continued for longer in
North America. In the 1960s the disease returned to Europe,
this time caused by O. novo-ulmi, a much more virulent strain
of the fungus. The 1960s outbreak killed approximately 25
million of the 30 million elm trees then growing in Britain.
Dutch elm disease has since spread over most of the temper-
134 TEMPERATE FORESTS

ate world. It is found wherever there are elm trees, although

not all species of elms are affected equally severely.
Elm bark beetles are related to a group known as ambrosia
beetles. These beetles cannot survive without ambrosia fungi,
so called because in Greek mythology ambrosia was the food
of the gods (their drink was nectar). Ambrosia was sweeter
than honey and anyone eating it was thought to become
immortal; it was also a medicine that was believed to heal
any wound. There are several species of ambrosia fungi. Each
beetle species has its own preferred species of fungus and
every beetle has a small pocket on its front legs in which it
carries the spores. When the beetle invades a tree, the fungus
feeds on the wood and the beetle feeds on the fungus, tend-
ing it carefully and removing waste matter. Ambrosia beetles
bore tunnels deep into the wood, and ambrosia fungi can
cause serious damage to the tree.
Woolly aphids are tiny insects that also transmit a serious
disease of trees. The insects feed on plant sap. There are many
species, most of them specializing in one or a few species of
trees. The felted beech coccus (Cryptococcus fagisuga) prefers
American and European beech trees (Fagus grandifolia and F.
sylvatica, respectively). The insect’s name refers to the waxy,
feltlike substance with which it coats large areas of the trunk.
The “felt” contains spores of the fungus Nectria coccinea,
which infects the tree with beech bark disease. The disease
slows the growth of the tree and a severe attack can kill it.
Not all fungi require insects to carry them into a tree.
Spores of tinder fungus (Fomes fomentarius), also called hoof
fungus and touchwood, enter wounds in the bark of broad-
leaved trees and the fungus grows inside the tree. It hollows
out the trunk until this splits and the branches fall. The fun-
gus produces fruiting bodies in the form of brackets that proj-
ect from the sides of the trunk. These have a leathery texture.
In the past dentists used the fungus to clean and dry tooth
cavities prior to filling them, and since Roman times it has
been used to cauterize wounds. After treatment, the material
of the fruiting bodies becomes highly flammable and is easily
ignited by a spark from a flint; it is the original tinder fungus.
Artist’s conk, also called artist’s fungus, picture fungus, and
false tinder fungus (Ganoderma applanatum) is similar. It is
 ECOLOGY OF TEMPERATE FORESTS 135

common in forests and sometimes kills beech trees. Its com-

mon name refers to the fact that a picture carved on its
smooth surface will remain there permanently.
Honey fungus, also known as bootlace fungus (Armillaria
species), is the most serious of all fungal parasites. It begins
by infecting the stump of a fallen tree and spreads outward
from there through the soil as black strands called rhi-
zomorphs that resemble black bootlaces. These enter trees—
usually young ones—through any tiny crack in a root or even
by piercing the bark. The fungus then spreads throughout
the tree, rotting and finally killing it.
Caterpillars—moth larvae—can also cause serious damage.
They eat leaves, and an infestation of thousands of them can
defoliate a tree, causing severe injury. Some of the moths
responsible are named for the time of year when the adults
emerge to lay eggs. The winter moth (Operophthera brumata),
for example, emerges between October and February. Its
caterpillars feed on several trees, including oak and spruce,
and they are serious pests in fruit orchards. Caterpillars of the
oak leaf roller moth (Tortrix viridana) live inside leaves that
they roll up to make a compartment, and they often defoliate
oak trees.

Differences between a natural forest

and a plantation
A forest that has grown up naturally over the centuries con-
tains a variety of tree species. These are not distributed even-
ly (see “Forest communities” on pages 127–130). Rather,
there are patches of forest, each with its own collection of
species, and each tree species forms part of its own smaller
community of shrubs and herbs. When a tree falls it opens
up the canopy. As the fallen tree slowly decomposes, provid-
ing food for a range of organisms and supplying nutrients to
the living plants, a young tree, perhaps of a different species,
grows up to take its place. Gradually the composition of the
community changes, while the forest remains as a patchwork
of communities.
Logging removes trees, thereby preventing them from
dying naturally and decomposing. The nutrients they con-
136 TEMPERATE FORESTS

tain are no longer recycled, and although fertilizers can be

applied to replace the nutrients that are lost, repeated log-
ging of commercially valuable trees eventually alters the
character of the forest.
Plantation forestry is the alternative to logging in natural
forest. A plantation consists of trees that have been planted.
It looks like a forest, but there are important differences.
A natural forest contains those species that arrived by
chance and survived under the prevailing conditions. A plan-
tation contains trees that will thrive on the chosen site and
yield lumber for which there is a commercial market. It will
exclude “valueless” trees that are naturally small or that fail
to grow straight. Most plantations include large blocks of a
single species. These are often coniferous trees, such as Sitka
spruce (Picea sitchensis), Norway spruce (P. abies), Douglas or
Oregon fir (Pseudotsuga menziesii), Japanese larch (Larix
kaempferi), and lodgepole pine (Pinus contorta). All the trees in
the block are planted at the same time and consequently all
the trees are the same age. At one time this produced monot-
onous, straight-edged, unattractive forests. Nowadays other,
usually native trees are planted around the edges of the
stands, and the edges are less straight than they used to be in
order to provide more visual interest.
When the trees reach marketable size they are felled.
Again, a complete block is clear-felled, leaving the site bare.
Shrubs and herbs thrive for a time, but then a new crop of
seedlings is planted and the cycle continues. Plantation
forestry is exactly like arable farming. Lumber is the crop. To
produce it, trees are planted, thinned when they reach a cer-
tain age, grown to the required size, and then harvested. The
only difference is that instead of taking one season to ripen,
the tree crop takes about 50 years.
Obviously, there is a considerable difference between a nat-
ural forest and a plantation, but this difference should not be
exaggerated. Plantations tend to support less wildlife than
natural forests, but this may be due partly to the fact that
they are often grown on land that was not formerly forested
and is far from the nearest natural forest. It takes many years
for forest species to arrive and forest communities to become
established, but this will happen in time. A plantation con-
 ECOLOGY OF TEMPERATE FORESTS 137

sists of blocks of trees all the same age, but natural forest
compartments are not very different, because they grow up
after trees have fallen, opening the canopy, and a compart-
ment often contains only one or two dominant species.
Within a plantation there are several—often many—blocks
of trees at different ages, resembling the compartments of a
natural forest. Together the blocks of the planted forest pro-
vide a variety of habitats for forest plants and animals, and
animals are able to move freely from one block to another.
Plantations that have existed for several 50-year harvest
cycles support a rich wildlife, but in many areas the planta-
tions are not yet that old.
Most important, though, is the fact that plantations pro-
duce lumber as a crop, which means they are sustainable
indefinitely, and in doing so they render logging in natural
forest unnecessary.
CHAPTER 6

BIODIVERSITY AND
TEMPERATE FORESTS
What does biodiversity mean?
On October 28, 1982, the General Assembly of the United
Nations adopted the World Charter for Nature. The charter is
not a law but a set of ideas and principles that are intended to
guide governments as they formulate policies that affect liv-
ing organisms, and governments are expected to enshrine
these ideas and principles in relevant national legislation.
The charter calls on governments not to waste natural
resources, to avoid pollution, and to protect natural habitats.
It begins with a statement of general principles, of which the
first three are:

1. Nature shall be respected and its essential processes shall not be

impaired.
2. The genetic viability on the earth shall not be compromised;
the population levels of all life forms, wild and domesticated, must
be at least sufficient for their survival, and to this end necessary
habitats shall be safeguarded.
3. All areas of the earth, both land and sea, shall be subject to
these principles of conservation; special protection shall be given to
unique areas, to representative samples of all the different types of
ecosystems and to the habitats of rare or endangered species.

The second of these principles refers to “genetic viability”

and the need to protect habitats in order to maintain popula-
tions of plants and animals. That idea was developed in the
years that followed, leading to the Convention on Biological
Diversity. A convention is an agreement between govern-
ments to take certain kinds of action or to behave in certain
ways. It is less formal than a treaty, although once it comes
into force it becomes part of international law. “Biological
diversity” is often shortened to “biodiversity” and the con-

138
 BIODIVERSITY AND TEMPERATE FORESTS 139

vention is often called the “Biodiversity Convention” (see

the sidebar on page 143).
Not all governments have signed the Biodiversity Con-
vention, but that is because they disagree with its approach
rather than with its principles. Every government accepts
that it is desirable to preserve as much biodiversity as
possible, both globally and within its national borders.
Before any government can take practical steps to imple-
ment the convention, however, it must decide just what it
is trying to preserve. What does “biological diversity”
mean?
The Biodiversity Convention defines biological diversity as
“the variability among living organisms from all sources,
including, inter alia, terrestrial, marine and other aquatic
ecosystems and the ecological complexes of which they are
part. This includes diversity within species, between species,
and of ecosystems.”
Diversity therefore exists at three levels. Diversity within
species refers to genetic diversity—the naturally inherited
variation between individuals of the same species. For exam-
ple, some people have dark skin, some light skin, some have
blue eyes and some have brown eyes, and there are several
blood types or groups so that the composition of the blood
varies from one person to another. These are examples of
within-species diversity among people.
Conserving genetic diversity presents a difficulty. Since
each individual is unique, the only way to preserve this
amount of diversity is to prevent any individual from dying,
which is clearly absurd. In practice it is taken to mean
that genetic variations among populations of a species
should be noted and, so far as possible, the populations
preserved from extinction. Local populations of a species
are sometimes classified as subspecies or varieties, but very
often they are not given a distinguishing scientific name,
although they may have a common one. For instance, the
Florida panther is a local population of the puma (Felis con-
color).
Species diversity refers to the differences between species
and the overall number of species. The concept of a species
provides the basis for the system of biological classification.
140 TEMPERATE FORESTS

This system was devised in the 18th century by the Swedish

botanist Carl Linnaeus (see the sidebar) and it assumes that
species are quite distinct from one another. That is how it
seemed until biologists began examining species at the
genetic level. They discovered that different species share a
substantial number of genes and that one species shades
into another rather than being sharply distinct from it.
Genetically, there may be more difference between two indi-
viduals of the same species than there is between the average
genetic constitution of their species and that of a related
species. As a result the concept of the species has grown

Carolus Linnaeus and

the classifying of plants and animals
People have always needed some way to describe the plants and animals they saw, but
without a formal system of classification this is more difficult than it might seem. Everyone
knows what the name wolf means, but suppose that word did not exist because no one
had ever seen a wolf, until one day someone encountered one. How might that person
describe it? Perhaps they might call it “a big, lean, mainly gray dog.” Plants were even
more difficult. In 1623, a famous Swiss scientist called Caspar Bauhin (1550–1624)
described a plant as: Ranunculus alpinus humilis rotundifolius minore (“low-growing, round-
leaved, small-flowered, alpine buttercup”). This was accurate enough as a description, but
it was difficult to remember, and anyone interested in botany had to master hundreds of
descriptions like this one.
The first comprehensive system to classify plants and animals—and also minerals—
using simple, memorable names appeared in 1735, in a book with the title Systema
Naturae, written by Carolus Linnaeus (1707–78).
Carl Linnaeus—Carolus was the fashionably Latinized version of his name that he used
in his publications—was born in Råshult, in southern Sweden, the son of a clergyman who
was also an avid amateur botanist. Linnaeus studied medicine at the University of Uppsala.
The university had a botanical garden where he spent much time; in those days medical
schools maintained botanical gardens as a source of medicinal herbs. In 1730 Linnaeus
became a lecturer in botany at Uppsala and he qualified in medicine in 1735, the year he
published Systema Naturae. He was the first president of the Royal Swedish Academy of
Sciences and physician to the Swedish Admiralty, and in 1742 he became professor of
 BIODIVERSITY AND TEMPERATE FORESTS 141

rather more vague. Nevertheless, it is convenient and useful

to group similar organisms into species.
Distinguishing between ecosystems is less controversial,
because an ecosystem is defined as a distinctive area with
boundaries that can be identified (see “How ecological ideas
developed” on pages 98–101). The convention does cover
marine ecosystems, however, which are often more difficult
to define. The level of protection is also drawn widely: It is
not only the diversity among plants, animals, and fungi that
is to be protected, but also that among microorganisms of all
kinds, including bacteria.

botany at the University of Uppsala. He was made a nobleman in 1741, and changed his
name to Carl von Linné. He died on January 10, 1778, following a stroke.
Systema Naturae contained detailed drawings of plants, animals, and minerals, group-
ing them according to their similarities. His classification of minerals was abandoned, but
his system for classifying plants and animals formed the basis of the system used today.
Linnaeus was a dedicated and very popular teacher, and devised his system partly to help
his students.
In the case of plants, the similarities arose from the number of flower parts. Linnaeus
believed this “sexual system” identified distinctive characteristics. Scientists now know
that the number of flower parts can vary considerably among closely related species, so,
despite its title, Linnaeus’s system did not reflect natural relationships and was artificial.
Despite this, the 10th edition of Systema Naturae, published in 1758, is the starting point
for the modern classification of animals, nowadays regulated by the International Code of
Zoological Nomenclature. Linnaeus’s most important botanical work was Species
Plantarum, published in 1753. This is the starting point for modern plant classification,
regulated by the International Code of Botanical Nomenclature.
Linnaeus grouped similar organisms into species, similar species into genera (singular
genus), genera into classes, and classes into orders. The French zoologist Georges Cuvier
(1769–1832) later extended the hierarchy by grouping orders into phyla (singular phy-
lum). Linnaeus found it possible to name any organism by using only two words. The first
word, conventionally spelled with an initial capital letter, is the name of the genus and the
second, or trivial, name is the name of the species. This is sometimes called the binomial
system of classification. The buttercup that Bauhin described in five words is now known
as Ranunculus alpestris L. The “L” is for Linnaeus, to identify the scientist who first named
the plant. Human beings are named in the same way, as Homo sapiens L.
142 TEMPERATE FORESTS

Why biodiversity matters

When a forest is cleared, the land that is exposed can be used
for many purposes. Originally, forest was cleared to provide
farmland, to raise livestock and grow food. This is obviously
important. Or the land might provide space to build homes,
schools, shops, hospitals, factories, and offices—places where
people can live and work. People all need to live somewhere,
they need schools, shops to buy things, and to work, and
when people fall sick they need medical care. These, surely,
are more important uses for the land than just leaving it to
grow trees, aren’t they? People cannot eat trees, and even if
their homes are built from wood, some forest must be cleared
away to make space to construct them.
There is no shortage of arguments to justify clearing forests
and other natural areas. These arguments are usually very
persuasive. Sites really are needed for perfectly sensible uses;
greed is not the only reason people clear forests. So what
arguments can conservationists advance to counter these?
They might point out that when the forest is cleared, it is
not only the trees that will be removed. So will all the herbs,
fungi, animals, insects, and other members of the forest com-
munity. They could mention, for example, that digitalis is an
important heart drug that was first isolated from the foxglove
(Digitalis purpurea), a plant that grows in forest clearings.
Perhaps they could mention an area of 270 acres (109 ha) of
forest that was scheduled to be sold in 1996. A graduate stu-
dent from Cornell University discovered on that site a tiny
mold fungus from which cyclosporin is obtained. This drug is
used to suppress the immune system to help patients avoid
rejecting grafted tissue. These are not isolated cases. More
than 10 percent of the major drugs prescribed by physicians
in the United States were derived from plants, and more than
25 percent of our most common medicines contain at least
some ingredients originally isolated from plants. The National
Cancer Institute has tested 35,000 species of vascular plants in
its search for anticancer drugs and has found that many of
them contain effective substances that have been extracted
and analyzed. Plants may, and probably do, contain sub-
stances that have not yet been identified but that might be
effective in treating a wide variety of serious illnesses. The
 BIODIVERSITY AND TEMPERATE FORESTS 143

The Biodiversity Convention

In June 1972 the United Nations sponsored the largest international conference held until
that date. Called the UN Conference on the Human Environment, it was held in
Stockholm, Sweden, and was known informally as the Stockholm Conference. Delegates
to the conference resolved to establish a new United Nations agency, to be called the UN
Environment Program (UNEP). UNEP came into being in 1973. Its tasks are to collect and
circulate information about the state of the global environment and to encourage and
coordinate international efforts to reduce pollution and protect wildlife.
UNEP sponsored several major conferences over the years, and in 1992, 20 years after
the Stockholm Conference, it organized the UN Conference on Environment and
Development, also known as the Earth Summit and the Rio Summit because it took place
in Rio de Janeiro, Brazil. It was the largest meeting of world leaders ever held. The aim of
the 1992 conference was to relate environmental protection to economic development,
and to this end the delegates agreed on the provisions that were set down in the
Convention on Climate Change and the Convention on Biological Diversity—also known
as the Biodiversity Convention.
A convention is a binding agreement between governments. Government representa-
tives sign the convention, and when their own legislatures have accepted it, the govern-
ments ratify it by signing it again, confirming their willingness to abide by its terms. The
lawmakers must then translate those terms into national law. When a majority of signato-
ry governments have ratified the convention, it becomes part of international law and is
known as a treaty. By the summer of 2005 157 countries had ratified the Biodiversity
Convention.
The Biodiversity Convention reminds governments that natural resources are not infi-
nite and promotes the principle of using resources in sustainable ways that ensure future
generations will also be able to enjoy them. The convention requires governments to
develop national strategies and plans of action to measure, conserve, and promote the
sustainable use of natural resources. National plans for environmental protection and
economic development should incorporate these strategies and plans, especially in
respect of forestry, agriculture, fisheries, energy, transportation, and city planning. As
well as protecting existing areas of high biodiversity, governments should restore
degraded areas. The convention strongly emphasizes the need to involve local com-
munities in its projects and to raise public awareness of the value of a diverse natural
environment.
Many countries have now taken positive steps to implement the Biodiversity
Convention.
144 TEMPERATE FORESTS

plants have not yet been identified, but if we destroy the

communities of which they are part we may lose them alto-
gether. Forests may contain chemical compounds with the
power to improve human lives and to save them.
These are selfish reasons, however. There are other reasons
for retaining areas of natural forest, reasons that have noth-
ing directly to do with curing diseases and improving life for
human beings, their farm livestock, or their household pets.
Wild plants contain genetic material that one day may be of
value to other plants. Genetic engineering makes it possible
to transfer genes that confer desirable qualities from one
plant to another. The loss of wild plant populations implies
the loss not only of the source of those genes but also the
natural genetic pattern against which scientists can measure
the changes they introduce.
Scientists and conservationists recognized long ago the
importance of preserving natural communities of plants and
animals as far as possible in a state where they remained
undisturbed by human activities. The variety of species such
communities contain, and the genetic variety this represents,
came to be known as “biological diversity” and abbreviated
to “biodiversity.” As evidence emerged of the rate at which
biological diversity was being lost, concern over the situation
increased until governments felt compelled to take action.
When the heads of the governments of 178 nations gath-
ered at the United Nations Conference on Environment and
Development held in Rio de Janeiro, Brazil, in June 1992,
preserving biological diversity was high on their agenda.
That conference—nicknamed the Earth Summit—was the
largest summit meeting of world leaders ever held, and one
of its major achievements was the Convention on Biological
Diversity—the Biodiversity Convention (see the sidebar).

Typical trees of temperate forests

Temperate forests occur naturally over such a vast area that it
is not surprising they contain a wide variety of tree species.
Botanists group related species into genera and related genera
into families (see the sidebar “Carolus Linnaeus and the clas-
sifying of plants and animals” on pages 140–141), and many
 BIODIVERSITY AND TEMPERATE FORESTS 145

of the most widespread trees of temperate forests belong to

just two families: Fagaceae and Betulaceae. The Fagaceae, or
beech family, includes the oaks, beeches, southern beeches,
and sweet chestnuts. These trees grow naturally throughout
the temperate and tropical regions and are the dominant
species in many temperate forests. The Betulaceae, or birch
family, comprises the birches, alders, hornbeams, and hazels,
which grow throughout the temperate regions of the
Northern Hemisphere as far north as southern Greenland,
and at higher elevations in the Andes of South America.
Oaks (Quercus species) are probably the most common
trees of all, but this is partly due to the fact that there are
approximately 600 species. North America, with 85 native
species, has a greater variety of oaks than any other conti-
nent. Many oaks produce valuable timber. White oak (Q.
alba), found on the eastern side of the continent, is the most
commercially important North American oak. It is a huge
tree, growing up to 160 feet (49 m) tall and with a trunk that
is up to five feet (1.5 m) across. Its timber was once used for
shipbuilding and to make railroad ties. It is still used to make
furniture and barrels. The most durable oak timber comes
not from white oak, however, but from live oak (Q. virgini-
ana), native to the southern states and Mexico.
Pedunculate oak (Q. robur), also known as common oak
and English oak, is the most famous of the European species.
It grows in the same areas as sessile oak (Q. petraea), also
called durmast oak, and the two trees look similar. The
species can be distinguished by the way their acorns grow.
Pedunculate oak bears its acorns attached by long stalks, or
peduncles, while those of sessile oak sit directly on the twig.
Most oak trees are deciduous, shedding their leaves in the
fall, but those growing in warmer climates are evergreen.
Evergreen oaks are known as live oaks in North America. In
addition to Q. virginiana, there are encina, also known as
California live oak and coast live oak (Q. agrifolia), canyon
live oak or maul oak (Q. chrysolepsis), and interior live oak
(Q. wislizeni). Holm oak or holly oak (Q. ilex), which has
leaves resembling those of holly, is the most widespread
European evergreen species. Bamboo-leaved (or -leaf) oak,
also known as Chinese evergreen oak and Japanese evergreen
146 TEMPERATE FORESTS

oak (Q. myrsinifolia), found only in eastern Asia, is an ever-

green species with narrow, smooth-edged leaves. There are
also semievergreen oaks, such as laurel oak (Q. laurifolia) and
willow oak (Q. phellos). Semievergreen trees retain some but
not all of their leaves throughout the year.
One evergreen oak produces cork, which is one of the most
familiar commodities in the world. The cork oak (Q. suber)
grows in southern Europe and North Africa. Cork oak pro-
duces a very thick layer of cork (see “How wood forms” on
pages 94–97), and removing cork encourages the living tree
to lay down more. Cork oaks live to a great age.
Beeches (Fagus species) have a much more restricted distri-
bution. There are only 10 species, and only one of these,
American beech (F. grandifolia) is native to North America.
European beech (F. sylvatica) and oriental beech (F. orientalis)
are the equivalent species in western and eastern Europe
respectively. Japanese beech (F. crenata) is an important forest
tree in Japan.
Southern beeches (Nothofagus species) are the second most
important timber trees of the Southern Hemisphere, after the
mainly tropical and subtropical eucalypts (Eucalyptus
species). Some are deciduous and others evergreen. There are
35 species of southern beeches, occurring naturally in New
Guinea, New Caledonia, temperate Australia and New Zea-
land, and in Chile and southern Argentina. Wood from myr-
tle beech (N. cunninghamii), native to southeastern Australia,
is used for flooring and to make furniture. Roble (N. obliqua),
ruil (N. alessandri), and Dombey’s southern beech or coigue
(N. dombeyi) are commercially important in South America.
Southern beeches have been planted extensively in Europe to
replace elms lost to Dutch elm disease (see “Diseases and par-
asites of trees” on pages 132–135).
Sweet chestnuts (Castanea species) are deciduous trees that
occur naturally only in the Northern Hemisphere. There are
12 species. American chestnut trees (C. dentata) were once
widespread in North American forests, yielding high-quality
timber and the best of all chestnuts, but chestnut blight has
rendered the species almost extinct. The European sweet
chestnut (C. sativa) grows naturally in southern Europe. The
Romans introduced it to Britain, where it is sometimes
 BIODIVERSITY AND TEMPERATE FORESTS 147

known as the Spanish chestnut, perhaps—although no one

can be sure—because until then chestnuts were imported
from Spain. Other chestnut species grow in China and Japan.
Alders (Alnus species), birches (Betula species), hornbeams
(Carpinus species), and hazels (Corylus species) all belong to
the family Betulaceae—the birch family. Alders are 35 species
of small deciduous trees that grow on wet ground, beside
rivers and lakes. Their male flowers are catkins and their
woody fruits look like small pinecones. Native Americans
used to make canoes by hollowing out the trunks of red or
Oregon alder (A. oregona), and the wood chippings were used
to smoke fish. The most widespread North American species
are American green alder (A. crispa), which grows in the East,
and Sitka alder (A. sinuata), which grows in the West, from
Alaska to northern California. Wood from black alder (A.
glutinosa), also called common alder and European alder, is
used for carving and was once used to make clogs. It may also
have been used in making Stradivarius violins.
Birches are small, hardy deciduous trees that grow best in
northern temperate regions. There are about 60 species. They
also bear catkins, but unlike alder catkins, birch catkins shat-
ter when they are ready to release their windborne pollen.
Many birches have attractive bark. The waterproof bark of
the paper birch or canoe birch (Betula papyrifera) peels off like
sheets of paper. Native Americans formerly used it to cover
canoes. This is the most widespread North American birch.
Silver birch (B. pendula) is one of the most common European
species. “Silver” refers to the color of its bark and “pendula” to
the way the tips of its branches hang down. Downy birch (B.
pubescens) is also widespread. Oil obtained from the bark of
downy birch is used in Russia to tan leather, giving Russian
leather a characteristic smell.
American hornbeam (Carpinus caroliniana) is the only one
of the 35 species of hornbeams that is native to North
America. It is grown as an ornamental tree because of its fall
colors, and it occurs naturally on the eastern side of the conti-
nent. Hornbeams are deciduous trees that produce very hard,
dense wood. Wood from the common European hornbeam
(C. betulus) is still used to make the levers between the keys
and hammers in pianos, as well as butchers’ blocks, pulleys,
148 TEMPERATE FORESTS

and similar items that must withstand hard wear. It was once
used to make the cogwheels in mills and yokes for oxen.
Hazels are 10 species of deciduous shrubs or small trees
that often occur in the understory of temperate forests in the
Northern Hemisphere. They are widely cultivated for orna-
ment or for their nuts, and also for poles produced by cop-
picing (see the sidebar “Coppicing and pollarding” on page
195). American hazel or filbert (Corylus americana), a shrub
up to 10 feet (3 m) tall, is the most popular ornamental hazel.
Nuts from cultivated hazels are called filberts. In Europe C.
maxima, a tree native to southeastern Europe, yields the best
filberts. In Britain, large filberts grown mainly in Kent are
sometimes sold as “cobnuts.” The native British and north-
ern European hazel is C. avellana, which grows to about 20
feet (6 m) tall. Turkish hazel (C. colurna), native to southeast-
ern Europe and southwestern Asia, is a much bigger tree,
reaching about 80 feet (25 m). Farther east there are Chinese
hazel (C. chinensis), which is very similar to Turkish hazel,
and Tibetan (C. tibetica) and Japanese hazel (C. sieboldiana),
both of which grow to 15–25 feet (4.5–7.5 m).

Typical animals of temperate forests

Many traditional folktales are set in the vast, seemingly end-
less forests that once blanketed northern Europe (see “Forests
in folklore” on pages 167–169). The animals that feature in
those stories are forest animals and, of all those animals,
bears are the fiercest.
Brown bears (Ursus arctos) once inhabited the Eurasian
forests from Scandinavia eastward to the Pacific, patches of
forest in southern Europe and western Asia, and the forests of
northwestern North America. Despite this, encounters
between people and bears were never common. A brown bear
is a big animal that needs a large area of forest in which to
forage for food, so bears were always widely scattered. In
addition, most bears avoid contact with humans, except
where they are attracted to food that has been discarded
around human settlements. In these places bears may have
learned to associate the smell of humans with food, making
encounters more likely. This is undesirable, because an
 BIODIVERSITY AND TEMPERATE FORESTS 149

urgent need to defend their cubs and repel rival adults makes
all bears aggressive. Combined with poor eyesight that makes
it easy for them to mistake a human for a juvenile bear in an
upright, challenging posture, their inclination to attack
makes them unpredictable and therefore dangerous. The
resulting persecution, together with changes in land use that
have divided what was once continuous forest into patches
too small to support bears, has greatly reduced the bear pop-
ulation. The brown bear is still the most widely distributed of
all bears, but large populations now survive only in north-
western North America and Russia.
There are several subspecies of brown bears. Grizzlies (U. a.
horribilis) and Kodiak bears (U. a. middendorffi) are the
biggest. An adult male grizzly weighs up to 800 pounds (365
kg) and eats up to 30 pounds (14 kg) of food a day. Male
Eurasian brown bears (U. a. arctos) usually weigh less than
600 pounds (272 kg). Brown bears feed on roots and burrow-
ing animals, which they dig up with their long claws, and on
leaves, berries, honey, fish, deer, and sometimes farm live-
stock. Despite their name, not all brown bears are brown.
Their fur ranges in color from cream through many shades of
brown to black.
Black bears (Ursus americanus) occur only in North
American forests—and they may be white or brown rather
than black. They are smaller than brown bears and eat a sim-
ilar diet.
Bears breed very slowly. Female grizzlies raise no more
than six or eight cubs in their whole lives, and black bears
raise no more than about 12. Consequently, killing bears
quickly reduces the population. Since bears need such a large
area to find food, it is easy to see why they have vanished
from most of the territory they once occupied.
Wild boar (Sus scrofa) still roams through most of the forests
of continental Europe and Asia, but it is not found in North
America. It became extinct in Britain in the 17th century; in
modern times, however, a market has developed for its meat,
leading to the introduction of farmed boars. Some of these
animals escaped, and they have now reestablished themselves
in the wild in parts of southern England. Wild boars feed
mainly on roots, nuts, and other plant material, but will also
150 TEMPERATE FORESTS

eat earthworms and other small animals. Adult males are

often solitary. Groups of wild boar consist either of bachelor
males or of sows with their young. Ordinarily, wild boars live
an unhurried existence, walking slowly across the forest floor
as they search for food and resting frequently. Despite their
relaxed appearance, anyone encountering wild boar should
give them a wide berth. They are highly aggressive if dis-
turbed, can run fast when they need to, and they are capable
of inflicting serious wounds with their powerful tusks.
Rivers that flow through forests, and the ponds and lakes
associated with them, are the home of the beaver. Because of
demand for their fur, North American beavers (Castor
canadensis) suffered greatly from hunting—wars were fought
over access to the areas where they were trapped—but they
are now protected and have been reintroduced where they
had become extinct. Today they are found throughout North
American forests, and they have been introduced in parts of
Europe and Asia. Steps are also being taken to reintroduce the
European beaver (C. fiber) in the forests it once inhabited.
Formerly found throughout Europe and western Asia north of
the Caspian Sea, the European beaver survives today only in
isolated parts of France, Germany, Scandinavia, and Russia.
Beavers mate for life and devote a great deal of care to the
young, called kits, which remain with their parents for up to
two years. Each family builds a dam across a stream, using
sticks, bigger branches, and stones, all held together with
mud. The beavers then tunnel into the bank of the stream
and extend the tunnel upward as the water level rises behind
the dam. When the tunnel breaks the surface, the beavers
cover it with sticks and mud. As the water level continues ris-
ing, the beavers raise the dam wall and also the roof of the
chamber in which they sleep and where the kits are born. All
the family except for the kits work to build and maintain the
dam and dwelling, called a lodge. The rising water usually
surrounds the lodge, so eventually it becomes an artificial
island in the middle of the pond. The drawing shows a
beaver lodge and dam.
During spring and summer the beaver family feeds on
plants around the pond, but in the fall they start to store
woody plant material to feed them through the winter. The
 BIODIVERSITY AND TEMPERATE FORESTS 151

dam water level lodge water level

food store is situated on the bottom of the pond, within easy Beaver lodge and dam.
underwater access from the entrance to the lodge. In summer Inside their lodge the
beavers often dig canals to link ponds and feeding areas. beavers live above the
Beavers are rodents that live in and beside water. Above water, but the entrance
them are rodents that live in the trees, leaping from branch to the lodge is below
to branch and running effortlessly up trunks and headfirst the surface.
down them. Squirrels are marvelously agile, and the most
widespread squirrel of temperate forests is the gray squirrel
(Sciurus carolinensis), a North American species that was intro-
duced to Britain on several occasions between 1876 and
1930. It is now common in most of England and Wales, but
occurs only in the central part of Scotland. It is also found in
parts of eastern Ireland. Gray squirrels inhabit broad-leaved
forests, especially where oak, hickory, and walnut are the
dominant trees. They eat nuts of all kinds, as well as tree
leaves, buds, pollen, and fungi.
Foresters regard squirrels as pests because they strip bark
from trees, causing considerable damage. The squirrels strip
bark to find the sweet sap flowing through the vascular tissue
just below the outer bark (see “How wood forms” on pages
94–97). They invariably choose the trees with the thickest
sap—those that the forester considers the best trees—and
ignore the others. The eastern chipmunk (Tamias striatus) is
152 TEMPERATE FORESTS

Chipmunks are lively,

inquisitive forest animals.
They feed on nuts,
berries, and all kinds of
seeds. This is an eastern
American chipmunk
(Tamias striatus).
(Courtesy of Gerry Ellis,
Minden Pictures)

also a squirrel, and very familiar because it is naturally in-

quisitive and is not afraid of humans. It does not damage
trees.
Deer also strip bark from trees in order to eat it. They also
eat buds, shoots, leaves, and flowers of a wide variety of
plants. Most deer are forest animals. They are seen in open
country, but they seldom stray far from the forest edge. The
biggest deer of the temperate forest is the red deer (Cervus ela-
phus), standing about four feet (1.2 m) tall at the shoulder,
which is found throughout Europe, Asia, and North Africa.
Some zoologists consider the red deer to be the same species
 BIODIVERSITY AND TEMPERATE FORESTS 153

as the North American wapiti or elk, but others regard the

wapiti as a separate species, C. canadensis. The white-tailed
deer (Odocoileus virginianus) is smaller, up to 3.3 feet (1 m) tall
at the shoulder, and highly adaptable. It occurs in forests,
brushland, and swampy areas from southern Canada to
Brazil and Peru. The roe deer (Capreolus capreolus) is very sim-
ilar but stands only about three feet (0.9 m) tall. It occurs
throughout Europe and Asia.
CHAPTER 7

HISTORY OF
TEMPERATE FORESTS
Forest clearance in prehistory
When Europeans first began to explore North America it was
natural for them to assume that the landscapes they saw had
existed since the dawn of time. In the east there were forests,
and beyond the forests the interior of the continent was a sea
of tall grasses and bright flowers. The French called it a mead-
ow, for which the French word is prairie.
The tallgrass prairie was vast, but it had not existed since
the dawn of time. Sediments taken from the beds of lakes on
the prairie contain tree pollen. Until about 5,000 years ago
there were groves of trees set in the grassland, making it an
open forest.
Scientists are not sure why the landscape changed, but the
most likely explanation begins with a change in the climate.
Between 5,000 and 6,000 years ago the weather became
much drier. Trees need moisture, and as the rainfall decreased
they became more scattered. Grasses are more tolerant of dry
conditions, so they thrived. The prairies became established.
That was not the end of the story, however, because about
4,000 years ago the climate changed again. This time it
became cooler and moister. Trees returned and the boundary
between the forests and the prairies moved westward. Open
forest should have become established over much of the
prairies, and it would have if humans had not intervened.
Grassland fires have always been common. Grass withers
and dries during dry weather, and then it ignites easily.
Storms were as common then as they are now, and light-
ning started fires that swept before the wind. Herds of graz-
ing animals moved ahead of the fire, and this made them
easier for skilled hunters to kill. Native Americans aided the
process by starting fires deliberately to drive game into
ambushes.

154
 HISTORY OF TEMPERATE FORESTS 155

When the fires died down, fresh grass and other non-
woody plants flourished in the ashes, improving the pasture.
There was more food for the grazing animals, so their num-
bers increased, and that meant there was more food for the
hunters. The herds became huge and the people lived well,
but the forest never returned. Fire and trampling by livestock
destroyed tree seedlings until the soil’s stock of tree seeds was
exhausted and there were no mature trees to replenish it.
More recently, the prairie has been converted to farmland.
It was not only the North American prairies that were
formed in this way. Many scientists suspect that large parts of
the South American pampa and Eurasian steppe also resulted
from this combination of fire and trampling by game. To this
day, people inhabiting the African savanna use fire to main-
tain the grassland for the benefit of the livestock on which
they depend.
Forest is the natural type of vegetation over most of the
lowlands of northwestern Europe, but it is not the vegetation
that travelers see today. Europe is farmed intensively and
fields have replaced most of the original forests.
Clearance of the British forest began about 5,500 years ago
and it was farmers, not hunters, who were responsible.
Knowledge of farming—the cultivation of plant crops—
spread across Europe from what is now Turkey. By about
6,000 years ago it had reached the Atlantic and Channel
coasts and soon afterward it entered Britain, carried by peo-
ple who had to cross the open sea to reach British shores. At
the end of the Wisconsinian (Devensian) ice age (see the side-
bar “Holocene, Pleistocene, and late Pliocene ice ages and in-
terglacials” on page 13), melting ice sheets and glaciers
released so much water that the sea level rose rapidly. About
8,000 years ago water flooded low-lying land between what
are now England and France, creating the English Channel.
The early farmers grew wheat, barley, and other plant crops
and kept pigs, cattle, and possibly sheep and goats as well.
They probably practiced swidden farming. This involves clear-
ing an area of ground to grow crops. After a year or two,
when the soil becomes depleted of nutrients, weeds prolifer-
ate out of control, and crop yields decline, the farming fami-
ly moves to a new area, clears the ground, and starts over.
156 TEMPERATE FORESTS

Swidden farmers revisit the same areas, working in a cycle,

usually of five to 10 years.
Farmers would not have allowed the animals onto their
cultivated ground, where they would have destroyed the
crops. Pigs are forest animals and would have foraged for
themselves in the forest. Cattle would have grazed in the for-
est clearings, but they fed mainly by browsing on the tree
leaves and shoots within their reach. Their herders would
have augmented this food supply by climbing into the trees
and cutting down browse from higher branches. As they fed,
the cattle trampled tree seedlings and damaged bark, some-
times severely enough to kill trees, thus enlarging their clear-
ings. If the farmers kept sheep they would have needed to
clear forest to provide pasture for them, because sheep are
grazers, not browsers.
About 5,000 years ago a sharp reduction in the number of
elm trees (Ulmus species) occurred over most of northwestern
Europe, in an episode called the elm decline. No one knows
just why this happened. Disease or a severe pest infestation
might have caused it (see “Diseases and parasites of trees” on
pages 132–135), but alternatively it might have resulted from
a change in farming technique. Farmers might have started
keeping their cattle in pens to protect them from predators
such as wolves and to prevent them from wandering into the
forest and disappearing. That would have made their farming
more efficient, but it would also mean the animals were no
longer able to find their own food. The farmers would have
had to gather food and bring it to them, as is done for cattle
raised in beef lots. Farmers would have collected browse, per-
haps seeking out elm twigs and leaves because cattle find
these especially palatable. Removing foliage before the trees
flowered would have greatly reduced the amount of pollen
they produced—and scientists know about the elm decline
because of the reduction in the amount of pollen they find in
the soil (see “Using pollen and beetles to study the past” on
pages 157–161). It may be, therefore, that the number of
trees remained fairly constant and it was only their output of
pollen that decreased.
What is certain is that it was farmers who cleared the
European forests. This sounds like a formidable task, but it is
 HISTORY OF TEMPERATE FORESTS 157

surprisingly easy. European farmers used axes with blades

made of chert—the stone called flint when it is found embed-
ded in chalk—fixed to ashwood handles. Some years ago,
Danish scientists fixed a wooden handle to a genuine 4,000-
year-old ax blade of this type and found they could cut down
100 trees before the blade needed sharpening. On another
occasion it took three men using axes with blades of polished
stone just four hours to clear about 718 square yards (600 m2)
of birch forest, helped by the fact that as one tree fell it often
brought down others.

Using pollen and beetles to study the past

Flower pollen consists of microscopically small grains pro-
duced by the anthers (see the sidebar “What is a flower?” on
page 90). Borne by the wind or clinging to the bodies of ani-
mals, pollen travels from the male organs of one flower to
the female organs of another. It is a hazardous journey, and
flowers produce copious amounts of pollen to allow for the
fact that most never reaches its intended destination.
Pollen grains are extremely tough, however. Each one is
encased in a coat called an exine that is almost indestructi-
ble. Of all the countless trillions of pollen grains that set off
each year in search of a female flower to fertilize, most fall by
the wayside, onto the ground, into rivers and lakes, or into
the sea, and are lost. But a few grains fall onto wet mud or
sink to the bed of a lake or pond, where the acid conditions
preserve them. The cells inside the pollen grain survive for
only a short time, but the exine can remain unaltered for
thousands of years.
The exine has two layers. The inner layer, called the nex-
ine, is smooth, but the outer layer, called the sexine, is sculp-
tured, with patterns of pores, grooves, or spines. Pollen grains
also vary in shape and size. Some are spherical, others oval,
and they range in size from less than 10 µm (0.0004 inch) to
250 µm (0.01 inch) across. The shape and size of the pollen
grain and the markings on its surface are unique to each
plant family and, in some cases, to particular species.
Scientists are able to extract pollen grains from soils and
sediments and identify the type of plants that produced
158 TEMPERATE FORESTS

them. Pollen from grasses and other herbs can usually be

identified to the family level, trees and shrubs to the level of
genus, and a few plants to the species level.

Radiocarbon dating
Most carbon atoms have nuclei composed of six protons (particles carrying a positive
charge) and six neutrons (particles carrying no charge); this is written as 126C or simply
12C, the “12” referring to the mass of the nucleus. Nitrogen nuclei have seven protons and

a mass of 14: 147N. Chemical elements are defined by the number of protons in the nuclei
of their atoms.
Cosmic radiation—high-energy particles from space—strike air molecules, and this
bombardment releases neutrons. Occasionally a neutron strikes a nitrogen atomic nucleus
and replaces one of its protons. The nitrogen atom now has six protons, but a nucleus
with six protons is not nitrogen but carbon: 147N has been transformed into 146C. This is
carbon-14, or 14C, and because it is radioactive it is also known as radiocarbon.
Plants use both 12C and 14C in photosynthesis, so both forms of carbon are present in
all living tissue. Only 12C is stable, however. Being radioactive, 14C decays back to 14N at a
slow and very precise rate. After 5,700–5,760 years, half of the 14C in a sample will have
decayed.
While an organism is alive the amount of 14C in its tissues is constantly replenished, so
the ratio of 12C to 14C in its tissues is the same as the ratio in the atmosphere. After it dies,
however, the organism no longer absorbs 14C, and the 14C in its tissues steadily decays,
altering the ratio of 12C to 14C. Scientists can measure this ratio in small samples of mate-
rial and use it to calculate the amount of 14C that has decayed and therefore the age of the
sample. The calculated age is reported as so many radiocarbon years BP (“BP” means
“before present,” “present” being taken as 1950).
Although cosmic radiation constantly produces 14C in the atmosphere, the rate at
which it does so varies slightly. Scientists have been able to check radiocarbon dates
against dates measured by counting the annual growth rings in very ancient trees, to
allow them to convert radiocarbon years into calendar years.
Radiocarbon dating measures the age of material up to about 70,000 years old. If the
sample is older than that, so much of the 14C has decayed that it is impossible to measure
the 12C:14C ratio accurately. The method can be used only with samples from material that
was once alive. As well as the remains of plants and animals, this also includes wood and
cloth.
 HISTORY OF TEMPERATE FORESTS 159

The operation begins by drilling out a core of soil or sedi-

ment, then laying out the material on a bench. The ages of
tiny fragments of organic material present in the core are cal-
culated from the amount of radioactive carbon-14 present in
their cells (see the sidebar), allowing the core to be labeled
with the age at each level from the surface downward.
Scientists then know when the pollen at each level was pro-
duced.
By studying the pollen, scientists reconstruct the type of
plants that were growing at different times: the vegetational
history of the site. This is displayed graphically in the form of
a pollen diagram, like the example in the illustration. A pollen
diagram devotes one column to each of the most abundant
species in the sample. Each column is graduated vertically
with the depth, in this case starting 39 inches (100 cm) below
ground level. Horizontally it shows the amount of pollen for
that species as the percentage of the total pollen present,
known as the relative pollen frequency (RPF). Some pollen dia-
grams show the absolute pollen frequency (APF), which is the

depth pine birch oak elm alder

(cm) Pinus Betula Quercus Ulmus Alnus
100

125

150

175

A pollen diagram,
showing the percentage
200 of the total pollen
contributed by five tree
species at each depth.
The diagram shows how
0 0 20 0 20 0 20 0 20 40
percentage total tree pollen the vegetation has
changed over time.
160 TEMPERATE FORESTS

number of pollen grains present. The shading makes it is easy

to see the proportion of plants at each level. In the drawing,
for example, at a depth of 60 inches (150 cm), the forest was
40 percent alder (Alnus species), 30 percent oak (Quercus
species), 25 percent birch (Betula species), 10 percent elm
(Ulmus species), and 5 percent pine (Pinus species).
Alder grows on riverbanks and on wet ground and birch is
a species found in northern forests, where the climate is cool.
The preponderance of alder suggests that at the time corre-
sponding to soil at a depth of 60 inches the site was wet, per-
haps marshy, and the climate was cool and fairly rainy.
Pollen diagrams must be interpreted with care, however, and
the obvious interpretation may not be accurate. In this case it
is important to know that alder and birch trees produce more
than twice as much pollen as the other species growing at the
site; adjust the count for those species and the picture
changes. Probably the forest was mainly oak with birch and
elm and occasional pine trees, and alder growing where the
ground was wet.
Paleoecologists and paleoclimatologists—scientists who
study past environments and past climates, respectively—
have another tool to help them: beetles. Beetles are every-
where, and there are at least 350,000 species. Of all the ani-
mals living in a forest, approximately three-quarters are bee-
tles. Individual beetles do not travel very far, and most
species survive only within a narrow environmental range. If
the climate is too warm, too cool, too wet, or too dry for
them, they disappear, so if the climate changes the beetle
community will also change. Some species will die out and
others will take their place. A representative sample of the
beetle population in a particular place is therefore an excel-
lent guide to the climate.
The ancestors of beetles had two pairs of wings, but in bee-
tles the front pair have become modified and are now elytra—
tough wing cases that open to allow the wings to be used and
that protect the delicate hind wings when the insect is not
flying. Even insects that have lost their hind wings in the
course of their later evolution still retain elytra. Like pollen
grains, beetle elytra are very tough and are often preserved in
soils and lakebed sediments.
 HISTORY OF TEMPERATE FORESTS 161

Beetle elytra are more useful indicators than pollen, be-

cause they are easier to interpret. Elytra are unlikely to be car-
ried far from where the insect died, so they really do repre-
sent the beetles living at the site. Pollen, on the other hand,
is blown by the wind, and pollen from some trees travels far-
ther than pollen from others. Consequently, unlike beetle
elytra, the pollen preserved at a site may not give an entirely
true picture of the plants that once grew there.

Forests in the days of

ancient Greece and Rome
In about 2000 B.C.E., people using tools and weapons made of
bronze, an alloy of copper and tin, entered Greece from the
north. They were called the Hellenes, or Greeks, and they
mingled and intermarried with the original inhabitants of
Greece, who used stone tools. Their migration took place in
three stages, the last in about 1100 B.C.E., and the culture the
immigrants established developed into the civilization of
ancient Greece.
The migrants brought with them their stories of gods, god-
desses, and heroes. Many of those stories are set in forests or
forest clearings, places where mortals might encounter gods
and goddesses, as well as lesser woodland spirits such as pans
and satyrs (later called fauns by the Romans)—beings who
were half human and half goat. Forests must have been
familiar places to the people who told and listened to such
stories, and who worshipped gods who lived in the forest. In
fact, in those days forests covered 84 percent of the land area
of Greece.
The forest was not to last. Farmers needed land to grow
food, and as agriculture expanded the area of forest
decreased. Mountainsides were cleared first. The land was fer-
tile and food was plentiful, but as immigration continued
and the population increased, farming spread into the low-
lands, where more trees were felled to make way for crop-
land. The success of agriculture made farmland valuable.
Neighboring communities quarrelled over boundaries and
outsiders attacked, attempting to seize land. Repeated con-
flicts drove people out of northern Greece, and many of these
162 TEMPERATE FORESTS

migrants headed south to find new ways to earn a living in

the city of Athens. But the soils of southern Greece were
poorer than the soils of the north and they were unable to
support the expanding population. The Athenians had no
choice but to establish colonies elsewhere to accommodate
people their own farms could not feed.
Athenian expansion resulted in war with neighboring
states. There was an age of wars that reached its climax dur-
ing the lifetime of Pericles (c. 495–429 B.C.E.), an Athenian
statesman who was largely responsible for the establishment
of the Greek Empire and the development of Greek democra-
cy. Greece comprises a peninsula and many islands, and in
order to wage war the Athenians needed a navy. Their ships
were built from wood, and by the time of Pericles the timber
for shipbuilding had to be imported. The forests of southern
Greece had been cleared and the northern mountainsides
were bare. Soil eroded from the mountains, washed away by
the rains, and erosion created the dramatic, harsh landscapes
of modern Greece.
More than half of the original Greek forest has gone. Today
forests cover almost 14,000 square miles (36,000 km2),
amounting to 27 percent of the total land area of Greece, and
97 percent of the forest is natural, rather than plantation (see
“Plantation forestry” on pages 199–202). The Greek forests
are now expanding. Their area increased by 9 percent
between 1990 and 2000.
Greek civilization spread throughout the lands surround-
ing the eastern Mediterranean. The mightiest empire of the
ancient world was not Greek, however, but Roman.
Immigrants from central Europe had begun settling in Italy
in about 1600 B.C.E., and at its peak in 400 C.E. the Roman
Empire stretched from Jordan in the east to the Atlantic
Ocean in the west, and from North Africa and Egypt in the
south to Britain in the north.
Roman power expanded by military conquest and by
trade, and the expansion made many Romans extremely
wealthy. Roman aristocrats invested their money in land,
establishing large rural estates, and as land prices rose, specu-
lators bought and sold farms in the hope of making a quick
profit. Small farmers found life difficult, much as they do
 HISTORY OF TEMPERATE FORESTS 163

today in the face of competition from large agricultural cor-

porations. The wealthy landowners seldom farmed the land
themselves. They employed bailiffs to manage their estates
and there were slaves to do the hard work.
The first immigrants found forests covering 94 percent of
the land area of Italy and, being farmers, they set about clear-
ing areas to grow food. The process continued, until eventu-
ally almost two-thirds of the original forest had disappeared.
Today forests cover approximately 39,000 square miles
(101,000 km2), which is 33 percent of the land area of Italy.
Almost all of the forest is natural rather than plantation. As
in Greece, the forested area is increasing. Between 1990 and
2000 it expanded by 3 percent.

How North America nearly lost its forests

Forests cover 33 percent of the land area of the United States
and 45 percent of Canada. The U.S. forests represent 5 per-
cent of the world’s forests and the Canadian forests amount
to 10 percent, so 15 percent of the world’s forests are located
in North America. The Canadian forests cover 1.6 million
square miles (4.18 million km2) and those of the United
States occupy 1.2 million square miles (3 million km2).
It is a vast area, but it used to be even greater. European set-
tlers cleared parts of the forests to make space for farming
and they used the seemingly inexhaustible supply of timber
for construction and fuel—fuel for industrial as well as
domestic use. Wood was also a source of charcoal (see the
sidebar “Charcoal and how it is made” on page 182) and of
certain industrial raw materials, such as potash. Potash, also
called pearlash, is potassium carbonate (K2CO3). It is used in
the manufacture of glass and soap and in the processing of
wool. Nowadays there is an industrial process for making
potassium carbonate, but in the 18th century it was obtained
from wood ash and in the l790s potash from burned forest
timber was one of Vermont’s most important exports.
Forests began to disappear during the 17th century, but the
process accelerated during the 19th. The rate of clearance was
dramatic. A good example of this is Cadiz, a small town in
Wisconsin with a population of 863 and a land area of 36.4
164 TEMPERATE FORESTS

George Perkins Marsh

George Perkins Marsh (1801–82) was one of the founders of the conservation movement
and the author of Man and Nature, one of the most influential books on the subject ever
published.
Marsh was born in Woodstock, Vermont, on March 15, 1801, the son of a prominent
attorney and politician. A frail child, with eyesight so poor that it often prevented him
from reading, Marsh had an exceptional memory and much of his early education con-
sisted of remembering information that was read to him. He entered Dartmouth College
to study languages and had mastered several by the time he graduated in 1820. Marsh
specialized in Scandinavian languages and later wrote a grammar of Icelandic, but he also
learned other European languages. By the time he was 30 he was able to speak 20 lan-
guages. After graduating he spent a year teaching Greek and Latin at a military academy,
but he found he disliked teaching and left it to enter his father’s office and study law. He
was admitted to the bar in 1825 and established a practice in Burlington, Vermont, while
continuing to study languages in his free time. In 1828 he married Harriet Buell; she died
in 1833. Marsh’s second wife, whom he married in 1839, was Caroline Crane, a writer and
German translator.
Marsh entered politics in 1835, as a member of the Vermont executive council—the
equivalent of the state senate. He entered Congress in 1843 as a Whig. In 1849
President Zachary Taylor appointed him to be minister resident in Turkey, where he
served until 1854, when he returned to Vermont. While in Turkey he led a diplomatic

square miles (94.3 km2). In 1881 the town had 33.7 square
miles (87.2 km2) of forest. By the following year, 1882, this
had decreased to 10 square miles (25.8 km2), by 1902 to 3.2
square miles (8.4 km2), by 1935 to 1.6 square miles (4.2 km2),
and by 1950 to 1.2 square miles (3.2 km2). Only 3.6 percent
of the original forest remained by the l950s, and the surviv-
ing forest was divided into small patches.
In the United States as a whole, approximately 177,680
square miles (460,300 km2) of the original forest had been
cleared by 1850. The rate of clearance then accelerated and a
further 297,670 square miles (770,900 km2) of forest had dis-
appeared by 1910. Between 1600 and 1920 the United States
 HISTORY OF TEMPERATE FORESTS 165

mission to Greece in 1852–53. He served on the Vermont state railway commission

from 1857 to 1859 and lectured in philology—the scientific study of languages—at
Columbia University in 1858 and 1859 and at the Lowell Institute, Boston, in 1859 and
1860.
Marsh joined the recently formed Republican Party and in 1861 President Lincoln
appointed him as the first U.S. ambassador to Italy. That was the year Italy united into a
single kingdom, rather than a number of small states. Marsh remained in this post until
his death at Vallambrosa, near Florence, Italy, on July 23, 1882.
During his years in the lands bordering the Mediterranean, Marsh saw for himself the
erosion that had resulted from deforestation centuries earlier. He also studied some of the
numerous texts that had already been written on the environmental effects of human
activities. Marsh determined to write a general account of the ways in which people had
altered the natural environment over the centuries and to issue a warning of the dangers
that might lie ahead. His book Man and Nature was published in 1864. It aroused little
interest, so he revised it and in 1874 published a second edition with a new title, The
Earth as Modified by Human Action: Man and Nature. Some of his warnings sound very
modern. Marsh wrote: “But we are, even now, breaking up the floor and wainscoting and
doors and window frames of our dwelling, for fuel to warm our bodies and seethe our
pottage, and the world cannot afford to wait till the slow and sure progress of exact sci-
ence has taught it a better economy.”
His book set out the case for conservation, but it did much more. It presented a deep
insight into history and the way societies and civilizations develop.

lost 29 percent of its original forest, almost all by conversion

to farmland.
The settlers believed that by clearing away the forest they
were improving the land. They were “taming the wilderness”
by transforming it into productive fields and orchards. Few
people thought this could be anything but beneficial.
One person who questioned the wholesale alteration of
landscapes was George Perkins Marsh (1801–82), the stu-
dious, thoughtful son of a Vermont attorney and congress-
man (see the sidebar). The Marsh family lived in a large
house on the slopes of Mount Tom, in Woodstock. Mount
Tom had once been forested, as had an estimated 95 percent
166 TEMPERATE FORESTS

of Vermont, but almost all of the trees on Mount Tom had

been removed by the time George Marsh was old enough to
play on the hillside. When he was older, he noticed that
clearing the trees from Mount Tom had allowed the rain and
wind to remove topsoil, which had been washed down the
slope and into the rivers, making them shallower and
destroying the habitat for many of the fish. At the same time,
the loss of topsoil reduced the fertility of the hillside fields. In
a speech he gave to a Vermont agricultural society in 1847
Marsh warned of the dangers of deforesting steep hillsides
and described the way forests were being managed more sus-
tainably in Europe.
George Perkins Marsh was one of the founders of the con-
servation movement, and his ideas became very influential.
He was not alone in observing the harmful effects of defor-
estation. As early as the 18th century, French colonial admin-
istrators sought to prevent further deforestation in Mauritius,
and the British established forest reserves in Tobago in 1764
and St. Vincent in 1791. The British began to draw up plans
for managing the forests of India in 1847. Marsh’s book Man
and Nature provided scientists and early conservationists with
powerful support for their own campaigns. The foresters
managing the Indian forests were known as “conservators,”
and that title is still used in Britain.
In his own country, Marsh’s work contributed to the argu-
ments favoring the establishment of national parks, starting
with Yellowstone, which opened on March 1, 1872. In 1877
the secretary of the interior, Carl Schurz, proposed that per-
manent forests be retained in public ownership. The system
of U.S. forest reserves was authorized in 1891.
Eventually the rapid deforestation of the United States was
checked, and since 1920 the forested area has increased. It
was not concern for the forests that halted the clearance,
however, but changes in farming. Until the early 20th centu-
ry, approximately one-quarter of all farmland was being used
to feed the horses, mules, asses, and oxen that provided
transport and powered plows and other farm machines. With
the introduction of automobiles, tractors, and other farm
machinery, draft animals were no longer required, freeing the
land to grow crops for human consumption. In effect, a 25
 HISTORY OF TEMPERATE FORESTS 167

percent increase in the available land was achieved without

plowing an additional acre or felling a single tree. More
recently, agricultural advances have allowed food production
to increase faster than population growth, making it possible
to take some land out of agricultural production.
Large areas of Canadian forests were also cleared to provide
farmland, but the clearance was on a much smaller scale than
that in the United States. Canada is sparsely populated and
much of its land is unsuitable for farming. Consequently
only 6 percent of the original Canadian forest area has been
converted to farmland.

Forests in folklore
Little Red Riding Hood set out one day to deliver meat and
wine to her grandmother, who lived half an hour’s walk
away, deep in the forest. When she arrived, a wolf had taken
grandmother’s place. The story is not really about little girls,
grandmothers, and wolves, of course. It has much deeper
meanings. But it all takes place in a forest—in fact, a German
forest, and the little girl’s name was Rotkäppchen, “Little Red
Cap,” in the version written down by the brothers Grimm.
Forests are portrayed in folktales as mysterious and poten-
tially dangerous. It is easy to become lost—the forest plays
tricks on travelers—and when shelter appears, it may not be
all that it seems. Remember Hansel and Gretel, who found
shelter with a witch who used candy as bait to trap children
that she then cooked and ate. Wolves and bears prowled the
forest, and murderous robbers might be hiding behind every
tree.
In the stories, a journey through a forest symbolizes the
journey of life. It is difficult to find the true path, there are
countless dangers and distractions, and, all too often, appar-
ent solutions to life’s problems turn out to be traps that lead
the traveler into still greater hazard. Truth, love, courage,
ingenuity, and loyalty will guide and protect the traveler,
however. Hansel and Gretel are saved because they protect
each other and are able to outwit the witch, and Beauty in
“Beauty and the Beast” is able to see that the fearsome
appearance of the Beast hides a true and noble heart. When
168 TEMPERATE FORESTS

she falls in love with him she sees the handsome prince that
he really is. Beauty and her Beast lived in a French forest; La
Belle et la Bête is a French story.
There were also honest people living in the forest, but
these were invariably poor. Hansel and Gretel were the chil-
dren of a woodcutter so poor that he could barely put food
on the table when times were good. People lived in the forest
because they had no choice. They had no land to grow food,
no trade to practice in the city, and therefore no means of
earning a living. Forest dwellers might cut wood for sale or
make charcoal (see page 182), or they might try to sell furni-
ture or other wooden items they made by hand, but it was a
poor way to earn a living. Some were outlaws, unable or
unwilling to pay heavy fines or avoiding justice for crimes
they had committed. There were also gangs of criminals who
robbed the wealthy—but not to give to the poor.
Villagers gathered fuel and timber from the woodlands
nearby, but the remoter parts of the forest were of no value to
them. There was no food to be had there—game such as deer
was the property of rich landowners and the punishment for
stealing it was severe.
Poverty drove people to abandon children. Hansel and
Gretel were left in the forest to perish from hunger, by their
stepmother who could think of no other way to save her hus-
band and herself from starvation. Snow White was another
child abandoned by her stepmother. “Babes in the Wood” is a
story based on “The Children in the Wood,” a ballad that was
probably written in about 1595. It is one of several similar
stories that appeared at about that time. All of them tell of a
man who dies, leaving his property to his infant son and
daughter and the children to the care of his brother, their
uncle. The uncle hires assassins to take the children into the
forest and kill them so he can acquire their inheritance. One
of the assassins relents, killing his companion and leaving
the children to their fate. The infants die and are buried in
the leaves by a robin redbreast. The assassin is later caught
and confesses.
Fear and mistrust of forests was widespread in medieval
Europe and it persisted for a long time, but the stories also
allow for hope. A stroke of good fortune can end poverty. A
 HISTORY OF TEMPERATE FORESTS 169

passing prince may fall in love with the woodcutter’s beauti-

ful daughter and marry her, transporting the entire family to
live in the palace. A poor man may save the life of a rich
man, and be rewarded. A fairy or magical animal may show
the way to endless wealth.
Russian folktales describe the forest differently. The Rus-
sian forest is so vast and familiar that people regard it as an
obstacle to the traveler, but not as mysterious or dangerous.
Russian stories are set in lands far away, beyond the edges of
the forest. They have plenty of magic, in the form of birds
and mammals that talk and involve themselves in human
affairs, and although there are evil monsters in the forest,
they do not lie in wait to leap upon passing travelers.
In the West, however, some of the dangers were genuine.
In the 10th century someone called Acehorn built a refuge
near Filey in Yorkshire, England, where travelers could find
shelter if they were being attacked by wolves. In 1712
Viscount Weymouth cleared part of Selwood Forest in
Somerset, in the west of England, and built a church on the
land in order to drive out the criminal gangs that had been
operating there. The English word savage is derived from the
French sauvage, which comes originally from the Latin silvati-
cus, meaning “woodland,” so a savage is someone who dwells
in the forest.
Europeans who settled in North America took with them
the images they had acquired from these ancient children’s
tales. When they encountered the forest that stretched end-
lessly before them, what they saw was a desolate wilderness
haunted by wild beasts and wild men. They set about clear-
ing it to make a land where civilized people could live in
safety.

History of forestry
John Evelyn (1620–1706) was a man who loved trees and
who had studied the cultivation and uses of plants in Paris as
part of his education. The author of about 30 books, from the
age of 11 Evelyn also kept a diary that gives us a detailed
description of life in 17th-century London. John Evelyn was
a founding member of the Royal Society of London—today
170 TEMPERATE FORESTS

Britain’s national scientific society and equivalent to the U.S.

National Academy of Sciences. Although he only occasional-
ly held an office at the royal court, Evelyn was often in the
company of the king and other members of the royal family.
At the time England was on the verge of war with the
Netherlands (war actually broke out in 1665), and King
Charles II was very worried about the state of the British
navy. In 1662 the commissioners of the navy asked the Royal
Society to examine and comment on the extent of the deple-
tion of English timber suitable for shipbuilding over the past
20 years. The task was passed to John Evelyn and three other
fellows of the society. Evelyn compiled and expanded their
comments. In his diary entry for October 15, 1662, he wrote:

I this day delivered my Discourse concerning Forest-Trees to the

Society, upon occasion of certain queries sent us by the Com-
missioners of his Majesty’s Navy, being the first book that was
printed by order of the Society.

He continued to expand on the ideas in his discourse, and

the resulting book, published in 1664, was called Sylva, or a
Discourse of Forest-Trees, and the Propagation of Timber in His
Majesties Dominions. Two other works, called Pomona and
Kalendarium Hortense, were included with it. Pomona gave
instructions for growing apple trees to supply cider apples,
and Kalendarium Hortense was a horticultural almanac listing
the tasks to be undertaken in the garden each month. Sylva
itself was written as a manual for the owners of large estates
and gave instructions for growing a variety of trees and uti-
lizing their timber.
Sylva was the first textbook on forestry, and its publication
marked the start of commercial forestry. Evelyn’s book
remained the standard work on forestry for the next 150
years. An enlarged second edition of Sylva was published in
1670, and a third edition, enlarged still further, was pub-
lished in 1679. The 1679 edition also included “Terra,” a sec-
tion on different soils and their treatment. A fourth edition
appeared in 1706, shortly after Evelyn’s death, with what was
now called Silva enlarged again and another section added,
called “Acetaria,” on plants suitable for use in salads. There
 HISTORY OF TEMPERATE FORESTS 171

was a fifth edition in 1729, and in 1776 an edition of just

Silva appeared, edited and revised to make it much more sci-
entific. That version of Silva, with “Terra” added, went
through several more editions, the last appearing in 1825.
Landowners had always managed their forests, encourag-
ing trees to grow from seed to replace those that were felled
and planting trees to fill gaps, and some had even planted
blocks of trees all of the same species. The first such block was
planted in 1580 when Lord Burghley (1520–98), first minister
to Queen Elizabeth I, planted 13 acres (5.26 ha) of oaks in
Windsor Forest, near London. These were to replenish timber
used to build ships during the war with Spain that lasted
from 1585 until 1603, and to provide for the future.
Evelyn was proposing something quite new, however, at
least for England. His idea was to plant entire forests on land
where no forest stood previously. Trees were to be grown as a
crop and their timber sold for profit. Evelyn and his support-
ers calculated the amount by which planting forests would
increase the value of estates.
Despite Evelyn’s enthusiasm, however, many English
landowners continued to believe that forests were wasteland
and that clearing trees improved the land. British landowners
planted at least 50 million trees between 1760 and 1835, but
most of these were ornamental species grown to adorn the
landscaped parklands that were fashionable among the
extremely wealthy. The Germans, on the other hand, began
planting forests of coniferous trees much earlier, the first near
Nuremberg in 1368. When in 1895 the British prime minis-
ter, William Ewart Gladstone (1809–98), visited the German
chancellor, Prince Otto von Bismarck (1815–98), Bismarck
presented him with an oak sapling to plant back at home.
It was not until World War I, when Britain had to import
timber through a partial blockade of its seaports, that
Evelyn’s lesson was really learned. The Forestry Commission
was established in 1919 as a British government agency
charged with acquiring land and planting forests to create a
state forest large enough to provide a strategic reserve of tim-
ber (see “Plantation forestry” on pages 199–202).
The first forest reserves were established in the United
States at the end of the 19th century (see “How North
172 TEMPERATE FORESTS

British geography
The names England, Britain, Great Britain, United Kingdom, and British Isles are often used as
alternatives for the same country, but in fact they have quite different meanings.
Britain, a name first used in about 325 B.C.E., is the island comprising the Kingdom of
England, the Kingdom of Scotland, and the Principality of Wales.
England and Wales were joined politically in 1301, when Edward I made his son prince
of Wales, a title held ever since by the eldest son of the reigning sovereign.
In 1604 King James VI of Scotland inherited the crown of England, becoming James I of
England and often known as James I and VI. This united the crowns of England and
Scotland; the two parliaments were united in 1707. The country was then called the
Kingdom of Great Britain or the United Kingdom.
The Kingdom of Ireland was joined to Britain by the Act of Union in 1801, and the
name became the United Kingdom of Great Britain and Ireland. In 1922, 26 counties of
Ireland separated from the United Kingdom to become the Irish Free State; the name was
later changed to Éire (Gaelic) or Ireland (English), but the country is often called the
Republic of Ireland.
Great Britain is the island of Britain together with its more remote offshore islands,
including the Outer Hebrides, off western Scotland, and the Orkney and Shetland Isles, to
the north of Scotland. The name refers to the fact that the island contains two kingdoms
(and is therefore “great,” that is, “large”). It also distinguishes Britain the island from
Brittany (meaning “little Britain”) in France; in French Bretagne is Brittany and Grande
Bretagne is Great Britain.
The United Kingdom consists of Great Britain and the province of Northern Ireland; the
full name is the United Kingdom of Great Britain and Northern Ireland. This is the official
name of the country, often abbreviated to U.K., but in certain situations, such as sports,
the country is called Great Britain, and car licence plates carry the national identifier “GB”
(although many Scottish drivers use “Sco” or “Ecosse”).
The Channel Islands, lying close to the Cherbourg Peninsula, France, and the Isle of
Man—the large island between Northern Ireland, Scotland, and England—are self-gov-
erning British dependencies. They are not parts of Great Britain or of the United Kingdom,
and they are not members of the European Union.
The British Isles is the geographic name for the islands of Britain, Ireland, the Channel
Islands, the Isle of Man, and all the smaller associated islands.
 HISTORY OF TEMPERATE FORESTS 173

0 miles 200

0 km 250

SCOTLAND

N. IRELAND

ÉIRE

ENGLAND

WALES

Great Britain and the United Kingdom. Éire is also known as the Republic of Ireland.
174 TEMPERATE FORESTS

America nearly lost its forests” on pages 163–167), although

the main purpose of the reserves was to protect watersheds
from erosion, rather than to supply timber. Careful manage-
ment, with controls on logging and time allowed for natural
regeneration, could be relied on to sustain the timber supply.
Today plantations account for only 7 percent of the total
forested area of the United States, although two-thirds of the
total area is managed as commercial forest. More than half of
all Canadian forests are managed commercially, but there are
only a few experimental plantations.
European plantations are more extensive. They account for
30 percent of the total forest area in Europe as a whole and
for almost 70 percent of the forest area of the United
Kingdom (see the sidebar “British geography” on page 172
for an explanation of the different names applied to Britain).
 CHAPTER 8

USES OF
TEMPERATE FORESTS
Renewable and nonrenewable resources
A resource is anything people can use to provide something
needed or desired. It could be money deposited in a bank.
When a person plans a vacation, she may start by examining
her bank account to see whether she has enough money, or
resources, to pay for it. A builder preparing to erect a group of
houses will need to ensure that the necessary materials—tim-
ber, cement, bricks, slates, and so forth—will be available as
they are required. These materials are also resources.
Slates are made from a particular type of rock obtained
from quarries. Slate is plentiful, but eventually slate quarries
are exhausted. This does not mean that all the slate has been
removed, but only that the best and most accessible slate—
cheapest to remove and most valuable to sell—has gone. It is
possible to imagine a time when all the slate quarries in a par-
ticular part of the world, or even in the entire world, are
exhausted. Slate will then exist only on the roofs of buildings
and in the other items, such as pool table tops, made from
slate. These can be recycled, but as they break and have to be
discarded the amount of usable slate steadily decreases until
finally the world runs out of slate. It is a highly improbable
scenario, but not altogether impossible, because although
slate is forming now, it is forming much more slowly than
the rate at which it is being used. True slate, as opposed to
flagstones and other materials used for roofing that are often
called “slate,” forms when a mixture of mud and volcanic ash
is heated and compressed over millions of years by move-
ments of the rocks forming the Earth’s crust. It is a process
that is going on all the time, but it happens very slowly.
Consequently slate is being used much faster than it is being
replaced.

175
176 TEMPERATE FORESTS

A material such as slate that is being used faster than it can

be replaced is said to be a nonrenewable resource. Petroleum,
gas, and coal—the so-called fossil fuels—are nonrenewable
resources; they are forming constantly, but much more slow-
ly than they are being burned. Metals are obtained from the
ground. A few, such as gold and some copper, occur in the
pure or “native” form, but most are chemically bound in
rocks called “ores.” Chromium, for example, which is used to
make certain types of high-quality steel, is obtained mainly
from the mineral chromite, FeCr2O4, which is a compound of
iron (chemical symbol Fe), chromium (Cr), and oxygen (O).
Iron is obtained mainly from the mineral hematite, Fe2O3.
Native copper is very rare, and most copper is obtained from
ores such as chalcocite, also called copper glance (copper sul-
fide, Cu2S), and chalcopyrite or copper pyrites (CuFeS2).
Metal ores are still forming, but not at anything like the rate
at which they are mined and used. They are therefore nonre-
newable.
Mining companies know how much of the fuel or mineral
their mines contain, so they can calculate how soon they will
need to abandon each mine and move elsewhere. In prepara-
tion for that day they explore and identify the location of
their next generation of mines. The measured quantity of the
resource in existing and next mines is called a “proven
reserve.” If the amount is calculated on the basis of the type
and structure of the rocks, but not measured directly, it is an
“inferred reserve.” If the presence of the resource is suspected
because of the rock formations, it is called a “potential
reserve.”
The fact that a resource is nonrenewable does not mean it
is likely to run out any time soon. If it should be in short
supply, its price will rise and this will provide an incentive
for companies to explore more vigorously and to consider
reopening mines that are not exhausted but had become
too expensive to work. Increased demand for a resource usu-
ally increases the size of the reserves. Due to increased
demand between 1950 and 1970, for example, the reserves
of ores for tin increased by 10 percent, for copper by 179 per-
cent, for aluminum by 279 percent, and for chromium by
675 percent.
 USES OF TEMPERATE FORESTS 177

Nevertheless, it is possible in principle that one day a non-

renewable resource will be completely exhausted and the
world will have to learn to live without it.
Trees are different, because when one tree is felled another
can be planted to replace it. With proper management, a for-
est can continue supplying timber and smaller cuts of wood
for thousands of years without even diminishing in size.
Wood is a renewable resource, as are cotton, silk, wool, and any
type of food.
Many people suggest that modern industrial societies
should be much thriftier in their use of nonrenewable
resources to ensure that enough remains to satisfy the needs
of future generations. This sounds like common sense, but
unfortunately there is no way of predicting what the needs of
future generations will actually be. Huge amounts of copper
were formerly used to make telephone landlines and subma-
rine cables, and at one time many people feared that the
world would run out of copper. Instead, fiber optics are
replacing copper wires, cell phones are replacing landline
telephones, and intercontinental communications are often
transmitted by radio via satellites, reducing the demand for
copper. Similarly, a century ago coal was being mined in such
large amounts that, vast though the reserves were, it surely
must have seemed that all the mines would be exhausted
before many more years had passed. What actually happened
was that cleaner and cheaper oil and gas became the more
important fuels, fuel efficiency increased, and many coal
mines were closed—not because they were exhausted, but
because they were no longer required. As technology
changes, so do the resources on which society depends, and
since technological change is unpredictable, there is no way
to say what resources will be needed decades from now.
Renewable resources will last forever, but the distinction
between renewable and nonrenewable resources is less
straightforward than it seems. Forests renew themselves, but
if we decide to increase our use of forest products we will
need to plant more forests. That requires land and the supply
of land is limited.
Rather than emphasizing the relative merits of renewable
and nonrenewable resources, the more sensible approach is
178 TEMPERATE FORESTS

to use whichever is most appropriate for the task in hand, but

to use it carefully, with the minimum waste.

Forests for fuel

Today most energy is obtained from fossil fuels—coal, gas,
and oil. Nuclear power provides smaller amounts, hydroelec-
tric power is important in certain countries, and there are
parts of the world where people still burn peat in their homes
and in power stations to generate electricity. Peat consists of
partly decomposed plant material; it has to be cut from the
ground and dried before it can be burned.
At one time wood was the most important fuel. Wood fires
provided the only heating inside buildings. They cooked all
the meals and were the only means of heating water.
Although coal became popular in Europe in the Middle Ages,
it was available only in the cities and there were attempts to
ban it because smoke from coal fires polluted the air. Country
people burned wood, as they had always done. Coal, gas, and
oil did not become economically important until the 19th
and 20th centuries: At the start of the Industrial Revolution,
in the 18th century, water mills powered the machines.
Despite the rise of alternative fuels, wood has not been
completely displaced. In the world as a whole, people burn
approximately 61.4 billion cubic feet (1.74 billion m3) of
wood each year. Wood is a more important fuel in less indus-
trialized countries than it is in the industrialized countries,
but it makes a significant contribution even in the most
advanced countries. The table shows the amount of fuelwood
burned each year in a number of industrialized countries and
the amount burned per person. The highest rate of consump-
tion is found in Russia and the United States. Canadians and
Russians, with the world’s biggest forests, burn more wood
per person than do people in any other industrial country.
Throughout history, wood has always been easy to obtain
and it is easy to burn. It is a convenient fuel, but if the wood
fire is to burn cheerfully and give out enough heat to warm
the room, the wood must be chosen carefully. Live oak,
Pacific madrone or madroña (Arbutus menziesii), and walnut
are among the best trees for fuel. Each of these delivers more
 USES OF TEMPERATE FORESTS 179

Fuelwood consumption per year

Total consumption Consumption per person

Country cubic feet cubic meters cubic feet cubic meters

United States 2.6 billion 74.6 million 8.92 0.26

Russia 1.8 billion 52.3 million 12.42 0.36
France 389.7 million 11 million 6.52 0.18
Italy 201 million 5.7 million 3.52 0.10
Germany 90.4 million 2.5 million 1.09 0.03
United Kingdom 8.2 million 232,000 0.14 0.004
Japan 9.4 million 266,000 0.07 0.002
Canada 4.4 million 12.6 million 14.1 0.4
Figures include wood burned for all purposes, including power generation, and in all forms, including charcoal.
Sources: Encyclopaedia Britannica Book of the Year 2004; Earthtrends, 2003.

than 156,000 Btu of energy per cubic foot (6 GJ/m3; 1 GJ = 1

billion joules). Pine, poplar, and willow deliver only about
94,000 Btu per cubic foot (3.5 GJ/m3).
Wood is a fibrous material that absorbs water. Freshly cut
wood, called green wood, is at least 50 percent water by
weight and can consist of two parts of water for one part of
wood fiber. Drying reduces the water content to between 15
percent and 20 percent. Even when dried, however, wood is a
bulky fuel. One ton of wood, dried to about 20 percent mois-
ture content, has a volume of about 124 cubic feet (3.2
m3/tonne). One ton of coal has a volume of about 10 cubic
feet (0.25 m3 per tonne). Its bulk means that fuelwood is
expensive to transport. Consequently, most fuelwood is
burned close to its source.
Chemically, wood fiber is 48.5 percent carbon, 43.5 per-
cent oxygen, and 6.0 percent hydrogen, with 0.5 percent
nitrogen and 1.5 percent noncombustible solids. Combus-
tion is the oxidation of carbon and hydrogen, producing car-
bon dioxide (CO2) and water (H2O), respectively. This is the
reaction that releases energy. Fully oxidizing one pound of
carbon releases approximately 14,500 Btu (33.7 MJ/kg; 1 MJ =
1 million joules) and oxidizing one pound of hydrogen
releases about 62,000 Btu (164 MJ/kg). The oxygen that wood
contains is released when the wood burns, but it contributes
180 TEMPERATE FORESTS

nothing to the release of energy; effectively it dilutes the real

fuel, which is carbon. One ton (2,000 pounds) of wood con-
tains 970 pounds (485 kg/tonne) of carbon, but one ton of
black coal contains 1,540 pounds (770 kg/tonne). This dilu-
tion of its carbon means that wood has a much lower energy
density than coal.
When wood is set alight, at first the fire gives off almost no
warmth. This is because the heat is used to vaporize water,
and vaporization absorbs latent heat (see “Evaporation and
transpiration” on pages 66–69), lowering the temperature of
the wood. When the temperature of the wood rises above
about 500°F (260°C) it begins to break down chemically,
releasing a number of gases. These include carbon monoxide
(CO), carbon dioxide (CO2), ethanoic acid (also called acetic
acid, CH3COOH), and methanoic acid (also called formic
acid, (H)COOH), together with creosote, which is a complex
mixture of compounds. Because the gases vaporize at quite
low temperatures they are known as volatiles. They do not
ignite, however, until the temperature rises above about
540°F (280°C). Then the volatiles start to burn, and that is
when the fire gives off most of its heat. As it burns, the tem-
perature of the fire continues to rise. It typically reaches
between 900°F (480°C) and 1,200°F (650°C) in the white-hot
center of the fire, but the temperature in the flames is only
200°F (93°C) to 400°F (204°C). This is cooler than the igni-
tion temperature because the gases ignite closer to the hot
center of the fire and cool as they rise. This is called primary
combustion.
As the fire burns it also generates more volatiles, known as
secondary gases. These include methane (CH4) and methanol
(CH3OH). The secondary gases do not burn, either because
the temperature of the fire is too low to ignite them, or
because in the heart of the fire there is too little oxygen avail-
able for oxidation. The secondary gases will burn only when
the temperature is higher than 1,100°F (593°C) and there is
just enough air to support combustion, but not too much.
Air is a mixture of 78 percent nitrogen and 21 percent oxy-
gen, with small amounts of other gases. Oxygen supports
combustion—oxidation—but nitrogen does not. If there is
too much air at the heart of the fire, the nitrogen absorbs a
 USES OF TEMPERATE FORESTS 181

large proportion of the heat, lowering the temperature of the

mixture of oxygen and secondary gases.
It is possible to design a furnace that will burn all of the
volatiles and achieve temperatures higher than 1,100°F
(593°C), but a wood fire cannot provide the temperatures
needed to melt iron. Copper melts at 1,982°F (1,083°C), so
copper can be melted from its ore in a skillfully prepared and
managed wood furnace. Tin melts at 450°F (232°C). These
low temperatures allowed people to alloy copper and tin to
make bronze. Wood was the only fuel needed during the
Bronze Age. Iron presented a more difficult problem, howev-
er, because it melts at 2,795°F (1,535°C). Smelting and forg-
ing iron called for temperatures higher than could be
achieved in a wood-fired furnace. A new fuel was required.
The new fuel was charcoal. It is made by heating wood in
airless conditions (see the sidebar), a process industrial
chemists call destructive distillation. The high temperature
drives off all the moisture from the wood and also most of
the oxygen, leaving the carbon and remaining hydrogen
much more concentrated and thereby increasing the energy
density of the fuel. Charcoal is an impure form of carbon,
and burning it releases at least 50 percent more heat than
burning a similar volume of wood. It ignites easily and burns
cleanly, and if a bellows is used to force air through the fire it
will burn at up to 3,000°F (2,980°C). Charcoal made metal-
lurgy possible, and it is still used in some places as a fuel for
blast furnaces. Today, though, charcoal is used mainly as a
fuel for barbecues.

Logging and timber

Wood taken from a forest is known as roundwood, and
data for roundwood production include wood that is of
every kind and destined for every use. Tall, thick tree trunks
and branches are included, and so are the stumps and
roots of felled trees if these are ripped from the ground. The
wood may retain its bark or have the bark removed, and
when it leaves the forest the wood may be round in shape,
split into half-round lengths, or squared. It is all called
roundwood.
182 TEMPERATE FORESTS

Charcoal and how it is made

Charcoal is made from small sticks and branches by the process of destructive distillation, or
heating in the absence of air. Oak makes the best charcoal, but the bark must be removed
first because oak bark contains large amounts of sulfur and produces choking fumes when
it is heated. Today charcoal is manufactured industrially, but traditionally it was made in
clearings in the forest.
Short pieces of wood would be stacked around the bottom of a pole about 6.5 feet (2
m) tall that stood in the center of a shallow, circular pit approximately 14.75 feet (4.5 m)
in diameter. Longer sticks were piled around the short ones, with all the sticks sloping
inward. More sticks were added, almost to the top of the central pole, until the stack was
dome-shaped. The stack was then covered with bracken, leaves, and finally turf, and the
pile tightly sealed with a coat of mud made from earth mixed with ash and water.
Once the mud had dried, the central pole would be withdrawn and a small amount of
charcoal dropped down the hole through the stack. Sufficient burning charcoal was then
dropped into the hole to ignite the charcoal and wood in and surrounding the base of the
hole. As soon as the person igniting the stack saw flames, the top of the hole was sealed
with mud.
White smoke soon started to appear from the stack. The emerging smoke revealed
cracks in the mud sealing the stack; these had to be repaired. From time to time the char-
coal maker would remove the mud sealing the central hole to check conditions inside. It
was important that the wood inside did not catch fire; if flames were seen, the worker
poured water into the hole to extinguish them.
After one or more days the smoke emerging from the stack turned blue, indicating that
the process was complete. The stack was then left to cool. Once it was cool the charcoal
was ready for use.
While one stack was smoldering, the next stack was being built and charcoal from the
preceding stack was being bagged for sale.
Making charcoal was hard, dirty work, and charcoal makers, who lived in the forest,
were always filthy and disheveled. They were also desperately poor. Village people often
feared and despised them, but metallurgy would have been impossible without the fuel
they produced.

(opposite page) Malagasy women making charcoal in Madagascar. (Courtesy of Frans

Lanting, Minden Pictures)
USES OF TEMPERATE FORESTS 183
184 TEMPERATE FORESTS

Once removed from the forest, the wood may be sent to

factories to be made into paper and other industrial products.
It may be sold to builders for use in construction. Companies
or individual craftspersons may buy it to make furniture, cab-
inets, and ornaments. The wood may be burned as fuel or
made into charcoal (see “Forests for fuel” on pages 178–181).
Regardless of the way it is used, it is all roundwood. It is not
bought and sold as roundwood, however, but as timber, lum-
ber, or wood.
The terms timber, lumber, and wood are often used inter-
changeably, but in fact each word has a distinct meaning;
they are not synonyms. Timber is a very ancient word.
Originally it meant a building and this meaning expanded to
include any kind of building material, including material
used to build ships—hence the old sailors’ exclamation:
“Shiver my timbers!” In English law, timber also means the
live trees more than 20 years old growing on an area of land,
especially oak, ash, and elm. Sawmills define timber as wood
cut into sections that are not less than 5 × 5 inches (12.7 ×
12.7 cm) in cross section. Wood consists of material used to
make furniture, ornaments, and other small items. Lumber
consists of felled trees that have been stripped of their
branches and cut into logs of a length that can be transport-
ed conveniently.
At the time it is cut, wood contains a large amount of
water. This accounts for at least half of its weight, and if the
timber is left lying on the ground in wet weather it can con-
sist of one part of wood to two parts of water. Also, different
trees produce wood of widely differing densities. Conse-
quently, roundwood is measured and sold by volume rather
than weight, and it is dried before it is sold to the final user.
Drying, or seasoning, takes place at the sawmill after the tim-
ber has been cut into planks or boards.
The international unit of volume is the cubic meter (m3),
but in the United States lumber is often sold by the board
foot (bd. ft.). The units cubic foot (ft3), standard, and cord are
also used. A board foot is a length of wood one inch thick,
12 inches wide, and 12 inches long (2.5 cm × 2.5 cm × 30
cm); 424 board feet = 1 m3. A standard is equal to 1,980
board feet. A cord (so called because it was originally meas-
 USES OF TEMPERATE FORESTS 185

Roundwood production, 2000

Cubic meters Cubic feet Board feet
Rank Country (thousands) (thousands) (thousands)

1 United States 500,434 17,670,324 212,184,000

2 China 291,330 10,286,862 123,523,000
3 Canada 185,659 6,555,619 78,719,416
4 Russia 158,100 5,582,511 67,034,400
5 Sweden 61,800 2,182,158 26,203,200
6 Finland 54,263 1,916,026 23,007,512
7 France 50,170 1,771,503 21,272,080
8 Poland 25,652 905,772 10,876,448
9 Germany 21,907 773,536 9,288,568
10 Japan 19,031 671,985 8,069,144
11 New Zealand 17,953 633,920 7,612,072
12 Turkey 17,767 627,353 7,533,208
13 Latvia 14,488 511,571 6,142,912
14 Spain 14,810 522,941 6,279,440
15 Czech Republic 14,441 509,912 6,122,984
16 Switzerland 10,428 368,213 4,421,472
17 Ukraine 10,008 35,338 4,243,392
18 Portugal 9,878 348,792 4,188,272
19 Norway 8,173 288,589 3,465,352
20 United Kingdom 7,451 263,095 3,159,224
Includes both hardwood and softwood species.
Source: Encyclopaedia Britannica Book of the Year 2004.

ured using a length of cord) is a pile of wood 8 feet × 4 feet

and four feet high (2.4 m × 1.2 m × 1.2 m); 1 cord = 128
cubic feet = 3.6 m3.
The table lists the annual production of roundwood for the
20 temperate countries with the largest output, ranked in
order of their production. It shows that the United States is
by far the world’s largest producer.

Chipboard, fiberboard, and paper

As well as felled timber in the form of tree trunks and large
branches, the term roundwood includes smaller branches
and even the stumps and roots of felled trees. It is not possible
186 TEMPERATE FORESTS

to cut this type of material into pieces suitable for use in con-
struction, but it is not wasted. Along with waste wood from
sawmills—small ends of logs and planks, split and broken
pieces, shavings, and even sawdust—it can be made into
board.
Chipboard, also called particle board, has many uses.
Cabinet interiors are made from it, with a door of solid wood.
It is used to make upholstered furniture, for interior walls in
buildings, and for cladding to protect exterior walls.
As the name suggests, chipboard is made from wood
chips. These are dried and then sorted according to size.
Chips of similar size are mixed with heat-setting glue, and
the mixture is spread in molds. A second layer, comprising
chips of a different size, is laid on top of the first layer, and
the mold is filled with three or more layers, each compris-
ing chips of a different type. The top and bottom surface
layers are usually thinner than the inside layers and are
made from smaller chips. When the mold is full, the mate-
rial is pressed into sheets, heated to set the glue, and
allowed to cool. The sheets are then sanded and their edges
trimmed, after which they are cut to the required size and
shape.
Fiberboard is made from waste material from sawmills and
factories that make wooden products. It is used mainly for
surfacing interior walls, insulation, and draftproofing. Its
manufacture begins by pounding and grinding the raw mate-
rial until it becomes a mass of fibers. These are mixed with
water and glue and spread continuously onto a mesh drum.
The continuous sheet of fiber is known as a “web.” The drum
rotates against a solid roller, pressing the mixture to squeeze
out excess water. The web then passes between drying rollers
to remove more water, after which it is cut into sheets. The
sheets are either dried completely or pressed and heat-treat-
ed, depending on the quality required. Hardboard is fiber-
board with a density of more than 50 pounds per cubic foot
(800 kg/m3). Boards with densities between 19 pounds per
cubic foot and 50 pounds per cubic foot (300–800 kg/m3) are
classed as semi-hard. The density of soft board is less than 22
pounds per cubic foot (350 kg/m3).
 USES OF TEMPERATE FORESTS 187

Each year the world production of chipboard and fiber-

board is approximately 5.3 billion cubic feet (151.4 million
m3). Almost one-third of the world total—1.8 billion cubic
feet (51 million m3)—is produced in North America, the
United States manufacturing about 1.4 billion cubic feet (40
million m3). Europe produces almost as much as North
America—1.7 billion cubic feet (48 million m3).
Paper is made in very much the same way as fiberboard.
Most paper is made from softwood—wood from coniferous
trees—but ash, beech, and poplar are also used. The wood is
chopped into small chips, which are ground to fiber, and the
fiber is mixed with water to form pulp. Bleach and whiteners
such as kaolin (china clay) are added to whiten the pulp, and
dyes are added if the paper is to be tinted. The pulp is sprayed
continuously onto a rotating mesh, forming a web. After it
leaves the mesh, the web passes between rollers that press
and dry it.
All the wood destined for pulping in Europe and North
America is grown in plantations (see “Plantation forestry” on
pages 199–202), using fast-growing tree species, especially
Sitka spruce (Picea sitchensis). Paper manufacture threatens
no forests, and although economizing in the use of paper
may reduce the pollution caused by paper manufacture, it
will not help to conserve forests.
The annual world production of paper and paperboard,
such as cardboard, amounts to approximately 344 million
tons (313 million tonnes). Of that total the United States pro-
duces 96 million tons (87 million tonnes), Canada 22 million
tons (20 million tonnes), and Europe 104 million tons (94
million tonnes).
Paper is extensively recycled. Ordinary households are
learning to leave their old newspapers and magazines at
collecting centers, but most recycling takes place in offices
and commercial premises where large amounts of paper are
used. Printing factories, for example, trim sheets after they
have been printed, and the trimmings are collected and recy-
cled. In the world as a whole, about 139 million tons (126
million tonnes) of paper are recycled every year. The United
States recycles 45 million tons (41 million tonnes), Canada
188 TEMPERATE FORESTS

1.6 million tons (1.5 million tonnes), and Europe 43 million

tons (39 million tonnes).

Veneers and plywood

The most beautiful wood is also the rarest and most expen-
sive. Only the wealthiest people were ever able to afford fur-
niture made from it, but many centuries ago a way was found
around this difficulty, allowing cabinetmakers to produce
furniture that looked exactly like furniture made from the
costliest materials. The solution was called veneer and it was
practiced as early as Roman times. The technique fell into
disuse during the Middle Ages but was revived by French
craftsmen in the 17th century and quickly spread through-
out Europe.
Veneer is the craft of cutting a very thin slice of expensive
wood and using it to cover the surface of an article made from
inferior wood. Nowadays veneer is often used to hide defects
in the cheap pine from which some furniture is made, and the
veneer itself may be plastic rather than made from wood.
Any attractive wood can be used as veneer. The French fur-
niture makers of the 17th century liked to use ebony and
were known as ébénistes, but by the 18th century mahogany,
walnut, box, almond, pear, and cherry were among the
woods being used. Veneer makes economical use of expen-
sive material and greatly improves the appearance of the arti-
cles it covers.
The process begins by slicing a sheet from a block of wood.
The slice may be cut by saw, but it is often taken from a
round log that turns in a lathe, the slice being removed in a
single sheet and the cut spiraling toward the center of the
log. After sanding, the sheet is usually 0.03–0.06 inch
(0.8–1.6 mm) thick. The log is soaked in water before being
cut to make it easier to work and reduce the risk of splinter-
ing, and the wet sheet of veneer is easy to flatten. When dry,
the sheet is cut to the required shape, glued to the surface it is
intended to cover, and placed in a press until the glue has set.
If the surface is molded, the veneer is pressed onto it using
sandbags that take the shape of the molding. Pressing and
drying take only a few minutes.
 USES OF TEMPERATE FORESTS 189

Veneer is also used to make plywood. Plywood is light, but

very strong; compared with solid wood it is much less prone
to splitting when sawn, drilled, or nailed; and it is produced
in large sheets of very consistent quality. It is used to make
cabinets, chests of drawers, and closets. Floors and partition
walls that are required to bear no structural load are often
made from plywood, as are the walls of many trailers and
mobile homes. Until fiberglass displaced it, plywood was
used to make the hulls and decks of small boats as well as
parts of the fuselage and wings of airplanes. The Mosquito, a
famous twin-engined British fighter-bomber of World War II,
had an airframe made from wood and covered with plywood.
Plywood veneer is usually 0.03–0.1 inch (1–3 mm) thick,
and the sheets are glued together with thermosetting resins
to make laminated boards that are pressed, dried, and cut to
the required size. If the plywood is to be used outdoors,
water-resistant resins such as phenol-formaldehyde are used
to hold the veneers together; urea-formaldehyde resin is used
for indoor plywood. Phenol-formaldehyde resists cold water,
hot and even boiling water, seawater, and dry heat. It is the
adhesive used in making marine plywood for boats.
The board may consist entirely of sheets of veneer, or of
veneer on the upper and lower surfaces with a thicker layer of
sawn lumber between them; this is known as lumber-core ply-
wood. Sandwich board consists of two layers of plywood sepa-
rated by a thicker layer of a light material, such as foam plas-
tic, balsa wood, or paper honeycomb. The plywood bears the
load and the sheet is very light, and extremely strong for its
weight. Most plywood is of all-veneer construction.
Some applications, such as boatbuilding and furniture-
making, require the plywood to be shaped. This is done by
bending and gluing the veneers in a single operation.
Each plywood veneer is laid with its wood grain running at
right angles to the veneer beneath it. This makes the con-
struction very strong and it also eliminates almost complete-
ly the problems that arise with the shrinking and swelling of
wood as its moisture content changes. The grain usually runs
in the same direction on both the outer veneers, so plywood
usually has an odd number of layers, or plies. Three-ply ply-
wood is probably the most widely used, but five-ply is much
190 TEMPERATE FORESTS

stronger and more rigid. Finished plywood sheets are from

0.12 inch (3 mm) to 1.2 inches (30 mm) thick.

Small wood and its uses

In the days when sailing ships were made from wood, oak
(Quercus species) was considered the best source of shipbuild-
ing timber. Oak is strong, and once the timber had been thor-
oughly soaked to make it swell and seal all the joints, a ship
built from oak was completely watertight. The wood did not
deteriorate after prolonged immersion in water.
Shipbuilding and construction once consumed large
amounts of timber, but forests yielded small pieces of wood
as well as whole trunks, and small wood has always had
many uses. Even today, every modern home contains furni-
ture, ornaments, and everyday utensils such as wooden
spoons and chopping boards that are made from wood.
Oak was not used only for shipbuilding. The durability of
oak wood makes it suitable for other outdoor uses. It makes
excellent fence posts and gates, and it is the best wood for
making the staves of wooden barrels. Barrels are used to store
liquids, so they are kept permanently wet.
Oak also has indoor uses, of course. Its attractive grain and
the ease with which it bends make it a good material for mak-
ing furniture.
Elm is another wood that survives immersion in water,
especially if it remains immersed all the time, rather than
being alternately wet and dry. Water pipes were once made
from elm and some of them remained in use for more than
two centuries. Waterwheels and the wooden piles supporting
jetties were often made from elm.
Beech wood does not last long outdoors, but it is strong
and resists compression. Both American (Fagus grandifolia)
and European (F. sylvatica) beech are used to make wood
blocks for floors, furniture, and high heels for shoes. Kitchen
utensils, such as wooden spoons and breadboards, are often
made from beech, as are the stocks for rifles and shotguns.
Bentwood chairs are made from beech.
Nowadays the slats of most venetian blinds are made from
plastic, but before plastic was introduced they were made
 USES OF TEMPERATE FORESTS 191

from American basswood (Tilia americana), which has a long,

straight grain. This is also the wood used to make piano keys,
topped with plastic today but formerly with ivory from ele-
phant tusks.
Quaking aspen (Populus tremuloides) of North America and
European aspen (P. tremula) also have wood with a straight
grain. In medieval times arrows were made from aspen.
Because of their straight grain, wooden matches are made
from various Populus species, known as poplars, cotton-
woods, and aspens—despite the fact that the wood is non-
flammable. Matchsticks are impregnated with paraffin wax
to make them burn. Its nonflammability made it the pre-
ferred wood for brake blocks on horse-drawn vehicles
because friction with the wheel did not make it smolder, far
less burst into flames. Populus wood does not splinter, which
makes it suitable for boxes and crates, and also for the pallets
on which goods are stacked to be carried by forklift trucks.
Tool handles, oars, and sports equipment such as hockey
sticks and baseball bats (now often made from aluminum)
must withstand shock. Ash (Fraxinus species) is often used,
but hickory (Carya species) is the most popular wood for
making tool handles. Cricket bats, however, are made from a
variety of white willow (Salix alba caerulea) known, not sur-
prisingly, as “cricket-bat willow.” Hornbeam (Carpinus
species) is even tougher than ash and hickory; it used to be
called “hard-beam.” Mallet heads and the wooden pulley-
blocks and cogwheels used in windmills and watermills were
made from hornbeam.
Rulers, set squares, and similar mathematical instruments
must be straight-edged and hard enough to withstand repeat-
ed friction from pencils and pens drawn along their edges.
Modern instruments are made from metal or plastic, but they
used to be made from boxwood, the wood from box trees
(Buxus species). Boxwood is very hard and, as its name sug-
gests, the very finest boxes are made from it. The wood is an
attractive yellow color and can be carved with delicate
designs. Woodblocks once used in printing were usually
made from boxwood.
Stringed instruments, such as guitars, violins, violas, cellos,
and basses, have backs and bellies made from maple (Acer
192 TEMPERATE FORESTS

species), usually either sugar maple (A. saccharum) or syca-

more maple, also known as European sycamore (A. pseudopla-
tanus). Sheets of maple wood, planed to a very precise thick-
ness and bent to produce exactly the right shape, make the
best resonating chamber, but maple has other uses. Sugar
maple is pale pink in color and often has an attractive grain.
Sports equipment and furniture are made from it. European
sycamore is used to make kitchen utensils and chopping
boards.
 CHAPTER 9

MANAGING
TEMPERATE FORESTS
Traditional management
Paper, chipboard, fiberboard, plywood, timber for building
houses, wood for making furniture, tools, ornaments, and
countless other items all come from forests. Even today,
when most Americans and Europeans live in towns and
cities, we still need forests. Throughout most of history, peo-
ple lived in small, rural communities where they were even
more dependent on forest products. As well as raw materials,
forests supplied them with fuel and even with some of their
food. Villagers collected wild berries and nuts and allowed
their pigs to forage in the forest for fallen fruit, nuts, and
acorns. Where the forest canopy was fairly open (see
“Structure of a temperate forest” on pages 123–127) and suf-
ficient light reached the forest floor, people gathered grass
and bracken for use as bedding for themselves and their
livestock.
Forests have always been used, and when a tree was felled
no part of it was wasted. Leaves were fed to livestock, bark
was used to tan leather, galls were dried and ground to make
ink, and small sticks were used for kindling. In Europe, most
forests were privately owned, but local people were allowed
to enter them for specific purposes, such as gathering fuel,
cutting vegetation for bedding, or allowing pigs to forage.
Some forests, called wasteland, were owned and used by the
whole community.
Such an important resource had to be protected. Each area
of forest was enclosed by a strong fence or by a deep ditch
and high bank. The barrier kept out livestock that might
trample seedlings or, worse, might kill trees by nibbling the
bark all around them. This is called ringbarking or girdling,
and it is invariably fatal to the tree. It kills the tree by sever-
ing the xylem and phloem vessels that lie just beneath the

193
194 TEMPERATE FORESTS

bark (see “Differences between conifers and flowering plants”

on pages 87–89). Ringbarking prevents water and nutrients
from reaching the part of the tree above the cut and sugars
from reaching the parts below it.
The stock of trees had to be sustained. It was essential that
whenever a tree was felled another tree replaced it. This usu-
ally happened naturally, as a sapling previously shaded by
the mature tree grew rapidly to close the gap in the canopy.
If there was no suitable sapling, however, one had to be
planted.
Nevertheless, there is a limit to the amount of timber a
community needs. Buildings, bridges, and other large pieces
of civil engineering last a long time. People have a much
greater need for small wood to make tools, furniture, lath to
repair ceilings and walls, fences, and especially for fuel. A
way of producing regular crops of small poles was discovered
at least 6,000 years ago: Traces of the method have been
found in peat of that age in Somerset, England. The tech-
nique is called coppicing (see the sidebar), and it was in wide-
spread use in England by the time of the Norman invasion in
1066.
Beginning in the late 18th century, the rapid expansion of
manufacturing based on the factory system greatly increased
the demand for iron and steel. More and more iron foundries
were built and the ironmasters who owned them needed a
regular, reliable supply of fuel. Coal had not yet become the
predominant fuel. In the first decades of the Industrial
Revolution, the ironmasters contracted with local landown-
ers to supply charcoal (see the sidebar “Charcoal and how it
is made” on page 182). Coppicing was the only management
system capable of supplying charcoal regularly in sufficient
amounts, and the technique spread throughout Britain,
eventually reaching the Highlands of Scotland.
Coniferous trees die when they are felled, and therefore
cannot be coppiced, but most broad-leaved species are suit-
able. Hazel (Corylus avellana) was popular, especially for mak-
ing fences. Sweet chestnut (Castanea sativa) was also cop-
piced for fence poles called pales and was often used to make
the fences enclosing forests—hence the expression “beyond
the pale.” Willow (Salix species) was coppiced for basket-
 MANAGING TEMPERATE FORESTS 195

Coppicing and pollarding

Cutting a tree close to the ground will not necessarily kill it. Indeed, many tree species are
invigorated by this treatment. Called coppicing, it is a traditional technique used in forest
management.
A tree trunk consists of the dead heartwood surrounded by the sapwood, which forms
a layer just inside the bark. The sapwood is the only part of the trunk that is alive, and if the
trunk is sliced through, the sapwood will continue to grow. The trunk will not regenerate,
because the heartwood is dead, but many small shoots will arise from the sapwood
around the edge of the stump—called the stool. These shoots will grow into straight
poles, called underwood.
The rate of coppice growth varies according to the species, but many species produce
a dense crop of poles within about five years. When they reach a suitable size, the poles
are cut at the point where they emerge from the stool. More poles grow to take their
place, and the coppiced trees are cropped in a rotation, with an average gap of 12–15
years between cuts, although coppice rotations can be as little as four or as much as 30
years. A coppiced stool will continue to produce underwood for many years; coppicing
often makes trees live longer than they would otherwise.
Deer and cattle will eat tender young coppice shoots, so a different technique is need-
ed in places where these animals are present. That technique is called pollarding. It is iden-
tical to coppicing, except that the tree is cut about 6.5 feet (2 m) above ground level. The
remaining part of the trunk is called a bolling. Shoots grow upward from the top of the
bolling, just as they do from a coppice stool, but they are out of reach of animals that
might damage them. Like coppicing, pollarding extends the life of the tree. Trees lining
city streets are often pollarded to prevent them from growing too large.

making. Oak (Quercus species), alder (Alnus species), field

maple (Acer campestre), and hornbeam (Carpinus betulus) are
among the other species that were frequently coppiced.
Although small wood from coppiced woodland may have
been the most important forest product, it was not the only
one. Large lumber was still needed and its production was
often combined with coppicing in a system called “coppicing
with standards.” Individual trees selected for their high qual-
ity—especially the length and straightness of their trunks—
196 TEMPERATE FORESTS

were allowed to grow to their full size, while the trees sur-
rounding them were coppiced. The tall trees, known as the
standards, benefited from not being shaded by surrounding
trees. They were felled as they were needed and the forest was
managed to ensure that the supply of standards was always
sufficient for local needs.
From about 1800, landowners began to alter the composi-
tion of their coppiced forests by removing some species and
planting others to replace them. Chestnut and hazel came to
predominate in the coppiced forests of southern and eastern
England, and oak coppice in the north.
Coppicing produces a highly artificial forest, but the sys-
tem greatly encourages biodiversity (see “What does biodi-
versity mean?” on pages 138–141). Coppiced trees never
grow big enough for form a canopy, and consequently the
forest floor is well lit in some places and shaded in others,
and the clumps of coppice poles provide shelter from the
wind. This is an ideal habitat for many plant species, and
coppice woodland supports a rich variety of herbs and small
shrubs.
During the middle decades of the 20th century it was fash-
ionable in Britain to clear broad-leaved forest and replant
with fast-growing coniferous species. Conservationists
responded by protecting as much broad-leaved woodland as
they could and by reintroducing coppicing, which had
almost died out. As coppice poles became available there was
a revival in crafts using small wood, and coppicing is now
fairly widespread. Nowadays it is practiced mainly for conser-
vation reasons, however, rather than as a system of forest
management.

Park woodland
Most cities have parks where people can walk, play games, or
just sit and enjoy the view. In fine weather many workers
spend their lunch breaks in a nearby park. Parks have open
areas covered with grass, borders growing shrubs and flowers,
paths, and trees. Parks have many trees. Parks have so many
trees, in fact, that they are considered a type of woodland,
called park woodland.
 MANAGING TEMPERATE FORESTS 197

Parks have a long history. Thousands of years ago, the first

farmers in Northern Europe developed an “in-field, out-field”
system. The in-fields were small, irregular in shape, and sur- Orchards imitate
rounded the farm buildings. They were cultivated all the woodland, which is more
time. The out-fields formed an outer circle beyond the in- open than closed forest.
fields. Out-fields were larger than in-fields and were cultivat- This is an olive grove,
ed in a rotation, growing cereals, root crops, and from time to where the olive trees
time being left fallow. Both in-fields and out-fields were (Olea europaea) are
enclosed by walls, fences, or hedges, and each out-field was widely separated,
called a “park.” A park was an enclosed area of ground some allowing other plants to
distance from the farmhouse. grow between them.
Wealthy landowners later extended this meaning of “park” (Courtesy of Cisca
by applying it to the land some distance from the house Castellijns, Foto Natura,
where owner’s family and friends could ride, walk, or rest in Minden Pictures)
198 TEMPERATE FORESTS

the shade of a tree. The park was managed but not used to
grow crops. It was open, with scattered trees, and it was
enclosed to define the boundary of the property and to keep
out livestock. By the 18th century this type of park was be-
coming fashionable, and creating one was a way to increase
the value of an estate. Entirely artificial landscapes were
made by shifting earth, making lakes, planting grass, and
establishing small groves of trees, often with cattle and sheep
grazing between the groves. The farm that supplied food for
those living on the estate and the gardens supplying flowers,
fruits, and vegetables were usually not visible from the best
rooms in the owner’s house. The view from the house looked
across formal gardens to the park beyond.
Not all parks were used for recreation, however. Parks—
enclosed areas—were also created to keep animals such as fal-
low deer (Dama dama), red deer or wapiti (Cervus elaphus),
wild pigs (Sus scrofa), and cattle. These parks had to be sur-
rounded by tall, dense hedges or fences to prevent the ani-
mals escaping. Fallow deer are sometimes called “park deer.”
Livestock will destroy tree seedlings and damage mature
trees. In some parks the trees were pollarded to protect them
(see the sidebar “Coppicing and pollarding” on page 195).
Alternatively, the trees could be confined to small areas of
closed forest, protected by fences and separated by areas of
open grassland.
Not all farmers could afford anything so grand as a park,
but they could and did plant trees for entirely practical rea-
sons. Trees provide shelter from the wind, rain, and snow in
winter and they give shade in summer. They were planted in
the open countryside for the benefit of cattle and sheep, and
also around houses and churches. Trees appeared in villages
and in the cities, and by the middle of the 18th century there
were tree nurseries to supply almost every town and large vil-
lage in England. As well as native trees, the nurseries sold
species imported from other parts of the world, such as
European sycamore (Acer pseudoplatanus), holm oak (Quercus
ilex), London plane (Platanus acerifolia), and horse chestnut
(Aesculus hippocastanum). The popularity of attractive import-
ed trees grew steadily in England from the 16th century until
the early 19th.
 MANAGING TEMPERATE FORESTS 199

Trees were planted in groves as components in carefully

designed but artificial landscapes. Farmers planted trees
because it was fashionable to do so, but also to provide shade
and shelter. Exotic trees were often planted singly, to show
them off to their best advantage. Villagers planted trees
around their cottages. In towns and cities, trees appeared in
private gardens and lining streets.
By the 19th century most European towns had parks for
public recreation and most rural estates had parks for private
recreation. The parks consisted of expanses of grassland, with
trees growing singly or as groups—park woodland.

Plantation forestry
Park woodland could not supply significant quantities of
timber, and British landowners began planting forests of tim-
ber trees. Few planted native broad-leaved species, however,
because these grow slowly and would not repay the cost of
planting within the lifetime of the person making the capital
investment. They preferred fast-growing species, mainly
conifers. The output was still insufficient for the nation’s
needs, however, and the major expansion in plantation fo-
restry in Britain began in 1919 with the establishment of the
Forestry Commission, a government agency charged with
creating and then managing state-owned forests (see
“History of forestry” on pages 169–174).
Today there are approximately 7,440 square miles (19,280
km2) of plantations growing timber in the United Kingdom,
compared with 3,340 square miles (8,660 km2) of natural for-
est. With its much bigger area of natural forest, the United
States is less reliant on plantations. It has 62,700 square miles
(162,380 km2) of plantations and 810,000 square miles (2.1
million km2) of natural forest.
Plantations produce trees as a crop, so plantation forestry
is like farming, but with a much longer period between plant-
ing and harvesting. A plantation forest must produce crops of
trees regularly and reliably. To achieve this, the forest is divid-
ed into compartments bounded by permanent features such
as roads or rivers. Each compartment contains several stands
of trees. All the trees in a particular stand are the same age,
200 TEMPERATE FORESTS

but each stand is a different age. Once the plantation is fully

mature, each year there will be some stands ready to harvest.
Timber is measured by volume (see “Logging and timber”
on pages 181–185) and the yield of a plantation can be esti-
mated in advance, before a single tree is planted, because the
average rate of growth is known for different species. The
amount by which a tree increases in volume each year is
known as the mean annual increment (MAI). The MAI rises
rapidly at first, then slows, and finally reaches a maximum
value. The maximum MAI for a stand of trees is known as the
yield class for that stand. A stand of coniferous trees, for exam-
ple, may reach a maximum MAI of 30 cubic meters per hectare
(429 cubic feet per acre), giving the stand a yield class of 30.
Some broad-leaved trees have a yield class as low as four.
Knowing the yield class helps foresters decide which
species to grow. When they have decided on the species, the
foresters will also know how soon the trees will be ready to
harvest and the annual sustainable yield the forest will deliv-
er once harvesting commences. Armed with that informa-
tion, managers can calculate the capital investment the forest
will need and the rate of return it will yield. They will also
know how many workers they will need for each stage in the
operation and when they will need them, and what
machines will be used and when. Operations can be planned
many years in advance.
The planning of a new plantation begins with a detailed
survey of the site. The survey will produce a map that marks
roads, railroads, power lines, rivers, ponds, and other perma-
nent features as well as contours showing hills and valleys.
There will also be aerial photographs to accompany the map;
these are often stereoscopic, providing three-dimensional
views. Once the map is completed, the boundaries of the for-
est compartments can be marked on it, as well as the roads
and tracks that will provide access for machinery.
Firebreaks must also be planned. In the event of fire, fire-
breaks help prevent the fire leaping from one stand to anoth-
er. They must be at least 33 feet (10 m) wide and contain
nothing that will burn readily.
Climate is important, and the survey includes details of
the average amount and distribution of rain and snow, aver-
 MANAGING TEMPERATE FORESTS 201

age temperatures through the year, and average wind direc-

tions and speed. Cotton flags are sometimes used to measure
the wind speed. Distributed at various points around the
site, flags, all made from the same thickness and quality of
cloth, will become increasingly tattered as they flap in the
wind. The rate at which the wind destroys them identifies
the places that are sheltered and those that are exposed to
the strongest winds, where trees may be blown down. The
shape of the forest edge and the species planted there can be
adjusted to reduce the risk of wind damage in the exposed
areas.
Plantations produce timber, but they are also communi-
ties of plants and animals, and they should be managed in
such a way as to maximize their value as wildlife habitat.
Trees should not be planted on riverbanks, for example, but
the riverbanks left to be colonized naturally. Stands of trees
native to the area should be planted among the commercial
stands of “exotic” species. It is desirable to provide nesting
boxes for birds and bats. Forests also attract visitors, who
need roads and paths suitable for walking, biking, and
horseback riding; toilets; and information boards. The loca-
tions for these must be decided and marked on the site
map.
Once planning is complete, the foresters can leave the
office and start work. The site may need to be cleared and the
land drained—an operation that involves huge machines
cutting deep trenches. Then the first blocks can be planted
with tree seedlings that have spent one or two years in
seedbeds and a similar time in orchards. These will be the
first trees to mature and be harvested. Once they are growing
well, the next set of blocks is planted, and then more. It will
take several years to plant the entire area. As the stands grow,
dead trees must be replaced, woody undergrowth cleared
away, and some branches pruned to improve the quality of
the timber.
Finally, up to 60 years after planting, the trees in the
first stand are ready to harvest. First, though, the routes
the machines will follow into the stand are clearly iden-
tified and marked on the ground, and the stand is divided
into sections. One or two workers are assigned to fell all
202 TEMPERATE FORESTS

the trees in each section and the operation is planned so the

distance between felling teams is not less than double the
height of the tallest tree. The first trees are cut so they all
fall in the same direction and the next trees to be cut
fall at right angles across them. This arrangement makes it
easier to remove the branches—a task called “snedding”—
and lift the trunks. Machines called forwarders lift whole
trunks and load them onto trailers, or skidders—machines
that hold one end of the trunk and drag it along the
ground.
The lumber is taken away to the sawmill, the immense
machines rumble away to another part of the forest, and the
forest block is left bare. Soon the foresters will return to pre-
pare the ground for the next crop.

What are lichens?

Some look like tiny shrubs, others resemble flattened leaves, and there are those that
look like a smear of paint. All of them are lichens. They grow on rocks, the roofs of
buildings, and the trunks and branches of trees. In some forests they festoon tree
branches.
People often suppose that lichens are a type of moss, but they are not. Strictly speaking
they are not plants at all. A patch of lichen consists of millions of single-celled organisms
living inside a fungus. This kind of close relationship, from which both partners benefit, is
known as mutualism.
Mushrooms, toadstools, and molds are all fungi, but the part of the fungus that you see
is the fruiting body—the structure the fungus produces when it is time for it to release
spores for reproduction. The main part of the fungus exists as minute threads called
hyphae (singular hypha) that form a network called a mycelium. The fungus absorbs nutri-
ents through its hyphae.
In a lichen, photosynthesizing cells are lodged among the hyphae, just below the
lichen’s upper surface. Depending on the species of lichen, these cells are either algae or
cyanobacteria, and they may live as separate cells or be joined together to form fine fil-
aments. The fungus is the main part of the lichen, and the scientific name of the lichen
is taken from that of the fungal partner.
 MANAGING TEMPERATE FORESTS 203

Pest control
Trees are plants, and like all plants they provide food for ani-
mals. Because they are so large and support so many ani-
mals—as well as mosses, lichens, and even ferns that grow in
moist hollows where large branches emerge from the trunk—
trees are complete ecosystems in their own right. The older
the tree, the more time there is for species to arrive and
become established, and some trees live to a great age. Oak
trees (Quercus species) can live for more than 1,000 years,
which is time enough to accumulate a bewildering diversity
of living things.
Windsor Forest, near London, England, contains oak and
beech (Fagus sylvatica) woods in which the trees are 300–1,000
years old. These woods support more than 2,000 species of

The fungus protects its photosynthesizing partner. Its hyphae retain water and dissolved
minerals and produce pigments that shade the photosynthesizing cells from intense sun-
light. The hyphae also secrete acids that assist in the uptake of minerals, and in some
species the hyphae secrete poisons that prevent the lichen from being eaten. The algal or
cyanobacterial partner produces sugars by photosynthesis, some of which are used by the
fungus. As well as reproducing by means of fungal fruiting bodies, lichens also release sore-
dia, structures comprising a few fungal cells together with a few algal or cyanobacterial
cells that will grow if they fall onto a suitable surface. Releasing soredia is a type of asexual
reproduction, meaning that it does not involve the fertilization of an egg by sperm.
Lichens are extremely tough organisms. They can survive prolonged drought, and
when water becomes available they can absorb more than 10 times their own weight and
resume their growth immediately. Some individual lichens are thousands of years old.
Most species of lichens cannot tolerate sulfur dioxide, however, and disappear in areas
where the air is polluted. This sensitivity makes them useful indicators of polluted air.
There are more than 25,000 species of lichens. These can be divided into three groups
on the basis of their appearance. Foliose lichens are loosely attached to the tree or rock
surface and have a flattened, leaflike appearance. Fruticose lichens are attached to the sur-
face at only one point; some are upright and shrublike, others hang down like tassels.
Crustose lichens are firmly attached to the surface and look as though they have been
smeared onto it.
204 TEMPERATE FORESTS

invertebrate animals and approximately 1,000 species of

fungi. An individual oak tree in Windsor Forest may shelter
more than 400 species of invertebrates—including the larvae
of 200 species of butterflies and moths—and more than 300
species of lichens (see the sidebar). A single beech tree may
harbor more than 300 species of invertebrates. The Windsor
oak trees are English or pedunculate (Q. robur) and durmast or
sessile oaks (Q. petraea). These are typical of temperate-forest
oaks. A much younger, but nevertheless mature specimen of
Oregon white oak (Q. garryana) supports nearly 200 species of
invertebrates, more than 100 species of lichens, and approxi-
mately 30 species of mosses and liverworts.
Some of the invertebrate animals feed on the tree itself,
eating leaves, fruits, seeds, or sap they obtain by burrowing
beneath crevices in the bark. Other invertebrates hunt the
plant-eaters, and there are birds that hunt the insects.
A tree is able to resist most of the attacks from the plants,
animals, and fungi living on its surface. It can even survive
the manipulation of its chemistry that takes place when cer-
tain insects and mites inject its leaves a substance that
induces the leaf to produce new tissue in the form of a gall. A
gall is a growth inside which the insect or mite lays its eggs
and the larvae develop, feeding on the gall tissue. Many
species of insects induce gall formation, but the most abun-
dant are parasitic wasps, none more than about one-quarter
of an inch (8 mm) long and most much smaller. There are
hundreds of species of gall mites, all belonging to the family
Eriophyidae, and they are less than 0.01 inch (0.25 mm)
long. (Mites are not insects but arachnids, related to spiders
and scorpions.) Oaks are especially attractive to gall wasps,
and a single oak leaf may carry more than 100 galls. But galls
do not harm the tree.
Some attacks are much more serious, however (see “Dis-
eases and parasites of trees” on pages 132–135), and although
trees are able to defend themselves and recover from injury,
their ability to do so is not limitless. There are moths and
sawflies that are capable of stripping a tree of all its leaves,
and certain fungi that will destroy a tree from the inside.
Gypsy moths (Lymantria dispar) will defoliate oak trees, for
example.
 MANAGING TEMPERATE FORESTS 205

Insects also transmit diseases by transporting fungal and

bacterial spores and viruses into the plant tissue. For
instance, the tiny beech scale insect (Cryptococcus fagisuga),
up to 0.04 inch (1 mm) long, lays its eggs in cracks on the
bark of beech trees (Fagus species). When the eggs hatch, the
larvae feed on the bark. Beech scale insects themselves may
weaken the tree, but more seriously they introduce a fungus
(Nectria species) that spreads around the tree. The fungal
infection can kill it. Other insects and fungi then invade the
dead tree and they, too, can spread disease to nearby trees.
Healthy trees are better able to withstand disease and insect
attack than are trees with dead or dying tissues and wounds
where dead limbs have fallen away. Wounds and patches of
decaying tissue provide sites for infection, and once a tree is
sick it can infect its neighbors.
Old-growth forests suffer more from disease and pest
attacks than do plantation forests. This is because plantation
forests consist entirely of young trees that are felled before
they are old enough to succumb. Old-growth forests contain
many old and dying trees that infect others. This does not
mean that the forest as a whole is sick. It is entirely natural in
any community that old individuals die from diseases or
insect infestations that may infect others, injuring or even
killing them. Trees die, but in dying they make space for
young trees and the life of the forest continues. Even the loss
of an entire species, such as elms through Dutch elm disease
and chestnuts through chestnut blight, does not destroy the
forest. Other species fill the gaps left by the one that has
gone. In a natural forest, insects, fungi, bacteria, and viruses
that feed on trees are not pests, but simply part of the com-
munity. The damage that they do becomes important only
when it reduces the value of the harvest that people are able
to take from the forest.
Because insects transmit many tree diseases, if an insect
infestation becomes widespread in a commercial forest—nat-
ural or plantation—it must be treated. The only practicable
treatment is to kill the insects, and the only way to achieve
that is likely to be by spraying insecticide from the air. This is
expensive, and even when the operation is conducted with
great care there is an inevitable risk to harmless insects.
206 TEMPERATE FORESTS

Modern insecticides do not persist for long in the environ-

ment and are only mildly poisonous to wildlife other than
the species against which they are directed. Nevertheless,
spraying is a last resort, and of all the pesticide used in tem-
perate regions only about 2 percent is sprayed over forests.
Modern disease control is achieved more cheaply as well as
more safely through hygiene, which involves felling serious-
ly infected trees, removing them from the area, and burning
them. Although spraying to kill the insect or fungus may
save single trees grown for ornament, fungal damage will
have reduced the value of the wood, so this treatment is sel-
dom feasible in a forest. Effective pest control relies on a
detailed knowledge of the life cycle of the pest, which allows
pesticide to be applied at the stage in its life when the target
insect is exposed and vulnerable.
In some cases it is possible to use predators to control pest
populations. There is, for example, a species of ladybug,
Chilocorus stigma, that feeds on the beech scale insect, and a
fungus, Nematogonum ferrugineum (also called Gonatorrho-
diella highlei), that parasitizes the Nectria fungi that cause
beech bark disease. This type of biological control is effective
only in certain cases, however, and it can go wrong. The
introduced predator may simply die out, or, much more seri-
ously, it may attack beneficial species, becoming a pest in its
own right.
An alternative method of biological control uses scent
attractants. The females of many species of flying insects
release chemical substances called pheromones into the air to
attract males for mating. The males are extremely sensitive to
them—males of some species can detect them up to one mile
(1.6 km) away, and male gypsy moths can detect them at
concentrations as low as one part in 100 quadrillion (1017).
Scientists have been able to analyze some insect pheromones
and make them in large enough quantities for use in pest
control. Released into the air at the time when the pest
insects are preparing to mate, they will lure males into traps.
Trapped males may be killed or exposed to radiation that
sterilizes them without otherwise harming them and then
released. The sterile males mate normally, but their female
partners produce no fertile eggs. Known as the sterile-male
 MANAGING TEMPERATE FORESTS 207

technique, this works with insect species in which the females

mate only once each season.
Pests and diseases must be controlled in commercial forests
and plantations. Pesticides will remain in use for the foresee-
able future, but a combination of hygiene to prevent infec-
tion and more advanced methods of regulating insect popu-
lations mean that foresters are becoming less reliant on
them.

Modern forestry
Forests grow on land that someone or some organization
owns, and the landowner usually seeks to sell enough lumber
and small wood at least to pay for the management of the
forest. Conservation groups also own forests and manage
them in ways that enhance their value for wildlife. In both
cases, outsiders used to be regarded as intruders. They were
not welcome, either because they might damage the forest—
by lighting fires, for example, or trampling rare herbs—or dis-
turb the wild animals.
Commercial forestry is less profitable than agriculture, so
where trees and farm crops compete for the same land the
trees usually lose. This means that commercial forest planta-
tions are sited on marginal land—land that will not grow farm
crops. Plantations are set on hillsides and in the uplands.
Forest trees can survive in such places, but conditions are
harsh and trees grow slowly. Profits are small and the planta-
tions are unpopular for several reasons: Their exposed loca-
tions makes them visually intrusive; planting in rectangular
blocks makes them ugly; when a compartment is harvested
the area is left as a wilderness of tree stumps and bare earth;
and plantations support less wildlife than permanent forests.
Over several decades of the 20th century, objections to
these aspects of plantation forestry and demands for access to
privately owned forests came from naturalists, horse riders,
and hill walkers—literally millions of members of clubs, soci-
eties, associations, and conservation groups across Europe
and North America. Landowners, including the government
agencies responsible for European state-owned forests, need-
ed an additional, noncommercial reason for maintaining the
208 TEMPERATE FORESTS

forests and the naturalists, riders, and walkers supplied

it. The landowners responded by opening the forests to the
public.
Visitors to the forests needed certain facilities. There had to
be adequate paths and maps, for example. Providing these
was straightforward, but people also wished to see some of
the plants and animals living in the forest, and that called for
modifications to the way the forests were managed. Little by
little, “forestry” transformed itself into “silviculture.”
Silviculture is the management of a forest for the benefit of
the plants and animals living in it, regardless of whether or
not all or part of the forest is being exploited commercially.
As well as being a source of timber, the forest in the silvicul-
tural approach becomes a public amenity.
Planting methods change. Instead of planting endless rows
of closely spaced trees, all of the same species and the same
age, the planting is less regular, more widely spaced, and
includes a greater variety of tree species. The resulting forest
is more interesting to look at, and more light penetrates
through it, encouraging a more varied community of herbs
on the forest floor.
All forests have paths and tracks to allow access for forestry
vehicles. When subcompartments are thinned, the trees
growing along the edges of some tracks can be removed and
not replaced. This produces a wide strip of ground on either
side of the track that can be left for wild plants to colonize.
Increasing the variety of plants and allowing more light to
penetrate to the forest floor make the forest more attractive
to wildlife. More insects arrive, including colorful butterflies,
and birds arrive to hunt them. Mammals find food and shel-
ter inside the forest. Nesting boxes for birds and roosting
boxes for bats, attached to trees, help increase the bird and
bat populations.
Surveyors identify areas that are especially important for
wildlife, such as small ponds, marshy areas, and sheltered
hollows, during the very first stages of planning for a forest
on a new site. Those areas, as well as sites of geological or
archaeological importance, are marked on the maps and
managed differently from the remainder of the forest in
order to preserve them.
 MANAGING TEMPERATE FORESTS 209

When trees are felled for their timber, foresters nowadays

prefer to allow natural regeneration to fill the gaps. They
plant tree seedlings only where there are no suitable young
trees growing from seeds dropped by the mature trees. This
practice maintains the forest while allowing it to develop a
more natural composition.
Forests are managed in this way even in remote areas that
visitors never reach. The new approach accepts that, regard-
less of their location, forests are more than sources of timber.
Their importance as wildlife habitat is fully recognized, and
forest workers have welcomed the change. After all, they
spend most of their working lives inside the forest. They are
familiar with and care about the forest’s inhabitants. Certain
species must be excluded or their numbers limited in order to
protect the trees (see “Pest control” on pages 203–207), but
most plants, fungi, and animals cause no harm and are wel-
come.
Modern forest management combines timber production
with conservation and the provision of recreational facilities
for the visiting public.
CHAPTER 10

HEALTH OF
TEMPERATE FORESTS
Are the temperate forests disappearing?
If farmers are unable to make a profit from the sale of their
produce they will go out of business and farms will close.
With fewer farms, food production will fall, and as the
amount of food reaching the market decreases the price of
food will increase. Higher prices will make farming more eco-
nomically attractive and more people will be tempted to
become farmers. There will be an endless cycle of “boom and
bust.”
Governments cannot allow this to happen, because if food
prices rise, working people will demand higher wages, but
they will not accept lower wages when food prices fall. Large
fluctuations in food prices destabilize the economy, so gov-
ernments try to keep prices constant, with only minor and
predictable seasonal variations. They achieve this by subsidiz-
ing agricultural production. Subsidies to farmers take many
forms, but they all have the same goal: to ensure that farmers
keep on producing the commodities people want to buy.
Throughout most of the 20th century, while governments
were introducing subsidies to stabilize food prices and main-
tain the profitability of farming, agriculture itself was chang-
ing. New machinery was introduced to increase the efficien-
cy of cultivation and harvesting. The new machines—espe-
cially tractors—performed work that was previously done by
horses and, in some regions, oxen. Land that had formerly
grown food for draft animals was released to produce food for
people. Agricultural chemicals boosted crop growth and
reduced losses to pests, weeds, and fungal diseases. Output
increased dramatically during the second half of the 20th
century, and the subsidies guaranteed that increasing output
did not lead to falling prices. Throughout the temperate
regions of Europe and North America, agriculture became so

210
 HEALTH OF TEMPERATE FORESTS 211

productive that farmers were producing more food than peo-

ple could eat. There were surpluses.
Agricultural subsidies keep food prices artificially low. This
is a major source of friction between rich countries and poor
countries that cannot afford to subsidize their farmers to the
same extent. But the subsidies also have another effect: By
encouraging overproduction they reduce the demand for
more agricultural land and make it possible to take land out
of farming altogether. Land that was once farmed can be con-
verted to other uses—such as forests.

Newcastle upon Tyne

Sunderland

Middlesborough
The Great North Forest

The Tees Forest

Manchester
Red Rose Forest Sheffield
Liverpool

South Yorkshire Forest

The Greenwood
The Mersey Forest

Forest of Mercia Nottingham

Milton Keynes
Birmingham
Forest of Marston Vale

Bristol Watling Chase

Great Western
Thames Chase

Forest of Avon Swindon

London

English Community
Forests. Community
0 miles 100 Forests are planned to
serve the people living in
0 km 150
major cities.
212 TEMPERATE FORESTS

The rise in agricultural productivity has coincided with the

recognition that forests are havens for wildlife and that they
can provide valuable recreational opportunities to city
dwellers, becoming the “lungs of the city.” Forests were once
seen as obstacles in the way of farming. Today they are high-
ly valued, and the area of temperate forest is expanding.
In England, government agencies, local government, and
volunteer organizations are collaborating in a program to
establish 12 Community Forests. Each forest will serve the
people of a major city; the map shows their locations. The
forests provide opportunities for leisure activities and are an
educational resource for city schools. At the same time, they
transform the landscape, in many places revitalizing aban-
doned industrial sites. In Scotland (see the sidebar “British
geography” on page 172) natural forests, such as the Argyll
Forest in the west of the country, are protected as tourist
attractions, as local amenities, and for their value as wildlife
habitat. Between 1990 and 2000 the area of natural forests in
the United Kingdom increased by 16 percent.
Forests are also expanding in the United States. During the
1990s the area of natural forest increased by 1 percent and
the area of plantations also by 1 percent. The United States
now has approximately the same area of forest that it had in
1920. This means that the rapid deforestation of the 19th
and early 20th centuries has been arrested.
According to figures collected by the UN Food and Agri-
culture Organization (FAO), during the 1990s the total area
of all the world’s forests decreased by approximately 386,000
square miles (1 million km2), from 15.1 million square miles
(39 million km2) to 14.7 million square miles (38 million
km2). That is a decrease of 9.4 percent, and equivalent to a
loss of 0.22 percent a year.
The loss was not spread evenly across the world, however.
While tropical forests diminished in size, forests outside the
Tropics—including the northern coniferous forests (called
boreal forest or taiga) as well as temperate forests—expanded.
Globally, 0.9 percent of the area of natural forest outside the
Tropics was cleared during the 1990s and the land converted
to other uses, but the forests also expanded onto unforested
land by an area equal to 2.6 percent. That amounts to an
 HEALTH OF TEMPERATE FORESTS 213

increase in area of approximately 6,600 square miles (17,000

km2) each year between 1990 and 2000. The expansion was
due to new planting, in projects such as the Community
Forests in England, and the regeneration of forest on land
that had been taken out of agriculture. Plantation forests also
expanded during the 1990s, by 12,000 square miles (31,000
km2). This added 1.2 percent to the Earth’s total forest area.
When the increases in area of natural and plantation forests
are combined, the temperate and boreal forests expanded by
2.9 percent during this period. The table shows how the area
of forest changed between 1990 and 2000 in each continent,
with the change in the most extensively forested temperate
countries shown in parentheses.
Far from disappearing, temperate forests are expanding,
and in coming years they are likely to continue expanding.
Improvements in agricultural productivity have released land
that was once needed to grow food, providing space for
forests. This change has coincided with the growing public
appreciation of forests. Forests support a wide variety of plant
and animal species (see “Why biodiversity matters” on pages
142–144) and they attract visitors to walk, bike, and ride

Forest area changes, 1990–2000

Total forest, Total forest,
1990 2000
Rate of change Forest as % of
Region 000 miles2 000 km2 000 miles2 000 km2 (% per year) land area

Africa 2,712 7,025 2,509 6,498 –0.8 22

Asia 2,130 5,514 2,115 5,478 –1.0 49
China 561 1,454 631 1,635 +1.2 17
Oceania* 777 2,013 763 1,976 –0.2 23
Europe 3,979 10,305 4,012 10,392 +0.1 46
Russia 3,282 8,500 3,287 8,513 0 50
N. America & 1,802 4,667 1,817 4,706 +0.1 26
Caribbean
Canada 944 2,446 944 2,446 0 27
Central America 341 883 304 787 –1.1 43
South America 3,563 9,227 3,419 8,856 –0.4 50
*Oceania comprises Australia, New Zealand, and the small islands of the South Pacific Ocean.
Source: FAO Global Forest Assessment, 2000.
214 TEMPERATE FORESTS

horseback through them. Forests are more popular than they

have ever been, and in the temperate regions of the world
they are highly treasured. They will not vanish.

Acid rain
In the late 1960s and early 1970s, Swedish and Norwegian
scientists noticed that the rain falling on their countries was
becoming increasingly acidic. Scientists in the eastern United
States and Canada observed the same phenomenon at about
the same time.
At first the problem centered on damage to buildings, stat-
ues, and public monuments made from limestone and sand-
stone. Acid was eroding the surfaces, leaving them pitted.
Then lakes began to suffer. Their waters became clear as the
microscopic organisms disappeared, and fish were dying.
Finally, forest trees were affected. German environmentalists
called it neuartige Waldsterben (“new form of forest death”)
and suggested that up to 10 percent of the country’s forests
were sick or dying. Forests were also suffering in parts of
Central Europe, Britain, and North America.
The damage was blamed on “acid rain.” In fact, it was
rather more complicated. Industrial furnaces, especially in
power plants, were burning coal that contained relatively
large amounts of sulfur. The sulfur entered the air as sulfur
dioxide (SO2), which then drifted with the wind. The acid
reaching Scandinavia came from Britain, Germany, Poland,
and Russia. Most German acid rain was produced in
Germany. In North America the culprits were power plants
and factories in the U.S. Midwest. Depending on the atmos-
pheric conditions, the airborne SO2 reacted with other sub-
stances in various ways, but all of the reactions led to the
conversion of the SO2 to sulfuric acid (H2SO4). This was the
source of most of the acid.
Very high temperatures allow atmospheric nitrogen to be
oxidized to nitric oxide (NO) and nitrogen dioxide (NO2).
Some industrial furnaces generate temperatures high enough
for this reaction, but most of the NO and NO2 are produced
in high-compression gasoline engines—the engines in mod-
ern cars. Fertilizer manufacture also releases some NO2. Both
 HEALTH OF TEMPERATE FORESTS 215

of these nitrogen oxides react with other atmospheric gases.

Several reactions take place and all of them lead to the pro-
duction of nitric acid (HNO3).
Acid can damage plants directly when it is deposited on
their leaves through dry deposition (when droplets of acid in
dry air collide with a leaf surface), acid mist, and acid snow. All
of these are more harmful than acid rain, which usually falls
on only the upper surfaces of leaves and then runs off them
onto the ground. Acid mist coats both upper and lower leaf
surfaces, and snow lies on plant surfaces until it melts.
Although acid rain gets the blame, it is the least harmful;
“acid precipitation” is a better term because it is more com-
prehensive.
Acid precipitation also affects soil chemistry in two ways.
Increasing soil acidity slows the rate at which organic matter
decomposes. Decomposition releases nutrients, so slowing
their release deprives tree roots of the nutrients they need,
thus harming the trees. By a process called mass action, acids
introduce hydrogen ions carrying positive charge (H+) that
displace magnesium (Mg+) and calcium (Ca+), which are
essential plant nutrients. This is a serious problem in soils
derived from hard rocks such as granite. (It is much less of a
problem for soils developed from sedimentary rocks, such as
chalk or limestone. Most sedimentary rocks are derived from
the shells of marine organisms made from magnesium and
calcium carbonates, so sedimentary soils usually contain
enough of these elements to prevent the soil being over-
whelmed by hydrogen ions; this is called buffering.) Mass
action can also liberate aluminum ions (Al+) that react with
soil water, releasing more hydrogen ions and further increas-
ing the soil acidity. The overall effect is to starve plant roots
of essential nutrients. This is much more harmful to trees
than direct damage to their leaves. It slows growth rates and
makes trees less able to combat disease and pest attacks.
Nitrogen also reaches the ground in acid precipitation.
Nitrogen is an essential plant nutrient and stimulates plant
growth, but in a forest this can be harmful. Many forests
grow on land that is poor in plant nutrients. Adding nitrogen
makes plants grow more vigorously, but because the other
nutrients they need are in short supply, the plants’ “diet” is
216 TEMPERATE FORESTS

unbalanced. The nutritional imbalance interferes with the

processes by which plants prepare for dry conditions. This
makes them less able to survive summer drought and winter
cold, when the soil water is frozen and plant roots cannot
absorb it.
Although the problem was serious, it was exaggerated dur-
ing the 1980s due to an error in the way the effect was esti-
mated. Damage was first observed in Norway spruce (Picea
abies), and trees were believed to be suffering if they lost
more than 10 percent of their needles. Since almost one-third
of spruces had lost more than 10 percent of their needles, the
damage appeared to be severe and very widespread. Foresters
then pointed out that perfectly healthy trees often lose more
than 10 percent of their needles. When the assessment was
revised, the damage was seen to be less serious. Rather than
killing entire German forests, as the name Waldsterben (“for-
est death”) suggests, a survey in 1986 found that 1.6 percent
of German forest trees of all species were severely affected
and 17.3 percent were moderately badly affected. In 1999,
approximately 25 percent of all European forest trees had lost
more than 25 percent of their foliage and therefore were suf-
fering moderate damage or worse.
Environmental groups in the United States, Canada, and
Europe campaigned for measures to address the problem in
the only way practicable: by reducing acid emissions. In 1979
governments signed the Geneva Convention on Long-Range
Transboundary Air Pollution, and the convention came into
force in 1983. It lays down the general principles and goals
that allow nations to cooperate in reducing the emission of
air pollutants that drift across international frontiers. It was
followed by eight more specific agreements, called “proto-
cols,” to reduce air pollution. The convention has been high-
ly successful.
Emissions of sulfur dioxide in the United States fell by 37.5
percent between 1980 and 2000, from 26.4 million tons (24
million tonnes) to 16.5 million tons (15 million tonnes).
They are predicted to reach 15.4 million tons (14 million
tonnes) by 2010. European emissions fell 56 percent, from 65
million tons (59 million tonnes) to 28.6 million tons (26 mil-
lion tonnes), over the same period and are expected to reach
 HEALTH OF TEMPERATE FORESTS 217

20 million tons (18 million tonnes) by 2010. In fact, sulfur

dioxide emissions had already been falling for some time
before acid rain damage began to cause concern, so the steps
taken to reduce emissions reinforced an existing downward
trend. In contrast, Asian emissions, which were 16.5 million
tons (15 million tonnes) in 1980, are expected to rise to 87
million tons (79 million tonnes) by 2010.
Nitrogen oxide emissions are also rising, due mainly to
increasing use of nitrogen fertilizer on farms. The rising num-
ber of automobiles also contributes to nitrogen oxide emis-
sions, but to a lesser extent, because vehicles are now fitted
with catalytic converters—devices that remove most pollutants
from exhaust emissions.
Forests are recovering from the harm caused by acid pre-
cipitation, but the recovery is slow. Slow progress is inevi-
table because most of the damage arose from changes in the
chemistry of water moving through the soil, and this takes
time to correct. Meanwhile there is a danger that unless
Asian emissions are curbed, damage to forests will increase in
that part of the world.

Forest clearance and climate

Farmers plant trees to shelter their fields from the wind. A
shelter belt—a row of trees planted at right angles to the direc-
tion of the prevailing wind—absorbs some of the energy of
the wind as the air expends its energy pushing against the
trees. The diagrams on page 218 show how this works. In the
upper picture the trees are planted close together, forming a
dense barrier. Some of the wind flows back from the barrier as
though reflected, forming an eddy, and more of the wind
forms a second eddy on the downwind side of the trees, but
most of the wind blows over the top of the barrier. There is
very little wind immediately behind the trees because the
wind cannot blow through the barrier, but the wind soon
recovers its strength. By a distance equal to 10–15 times the
height of the barrier it has regained about 90 percent of its
force. If the trees are more widely spaced, as shown in the
lower drawing, the wind can penetrate the barrier, but is
slowed by it. Although the wind speed on the downwind side
218 TEMPERATE FORESTS

high density

low density

How a
shelter belt works

of the trees is greater than it would be behind a denser barri-

er, the effect continues for longer. The wind does not recover
to 90 percent of its original strength for a distance equal to
15–20 times the height of the barrier. If the shelter belt is nei-
ther too dense nor too open it is possible to reduce the wind
speed for a distance up to 40 times the height of the trees.
A shelter belt is a barrier only a few trees deep, and a forest
has a much greater effect on the wind. Suppose the wind out-
side the forest is blowing at 20 MPH (32 km/h). Inside the
forest, 100 feet (30 m) from the edge, the wind speed will be
12–16 MPH (19–26 km/h), and 400 feet (120 m) from the
edge it will be about 1.2 MPH (2 km/h). In the calm center of
the forest, the trees have absorbed almost the whole of the
wind’s energy. The wind also blows around and over the top
of the forest, but it is much weaker on the downwind side
than on the exposed side. Forests shelter the land downwind
of them as well as the ground they occupy.
Not surprisingly, when a forest is cleared the local climate
becomes windier. This increases the risk of soil erosion (see
“Forest clearance and soil erosion” on pages 225–227). More
 HEALTH OF TEMPERATE FORESTS 219

surprisingly, perhaps, clearing a forest also makes the climate

foggier and dustier. Fog consists of droplets of water that are
so small they fall very slowly, and most of them are lifted by
air eddies before they can reach the ground. The droplets
remain airborne and the fog drifts slowly on the wind. Dust
particles behave in the same way—until they enter the forest.
A forest acts as a filter. As the wind carries water droplets and
dust particles through a forest, the trees trap many of them
and the reduction in the wind speed holds them inside the
forest long enough for them to fall to the ground. Conditions
are often foggy or dusty upwind of a forest, but the air is clear
and fresh on the downwind side. Remove the forest and the
fog and dust will affect a larger area.
Forests also influence the air and ground temperatures. At
night the ground radiates the heat it absorbed during the day
and its temperature falls. On cloudy nights some of this radi-
ated heat is reflected by the clouds, so the air remains warm,
but on clear nights the radiation escapes and the ground
becomes much colder. If the air is still, a boundary layer of air
next to the ground becomes cold enough for its moisture to
condense or freeze, producing dew or frost. Condensation
and freezing release latent heat (see “Dew and frost” on pages
65–66), which the ground surface absorbs, warming it a little.
Inside and on the sheltered downwind side of a forest, the air
is still, so these are places where dew and frost are likely to
form. But remove the forest and the wind increases. Close to
the ground the moving air is turbulent, rolling and eddying
as it crosses the uneven surface, and constantly drawing air
from higher levels to mix with the air next to the ground,
preventing a layer of still air from forming. Dew and frost
cannot form under these conditions, so removing the forest
makes dew and frost less common. If there is no dew or frost,
however, there is no release of latent heat to warm the
ground. In winter ground frosts will occur earlier and they
will be harder.
Forests shade the ground. On a hot, sunny day, the air
inside a forest will feel cool. If the forest is cleared, the shade
will be lost, so summer days will likely be hotter. At night and
in winter, trees absorb much of the heat that is radiated from
the ground. Without the trees, that effect will be lost and the
220 TEMPERATE FORESTS

ground temperature will fall faster and farther. Summer days

may grow warmer, but the nights may be cooler and the cold
of winter will last for longer.
Clouds often form above forests. This is because trees move
large quantities of water from the ground to the air by tran-
spiration (see “Evaporation and transpiration” on pages
66–69) and by the evaporation of rainwater from leaves and
bark. The air inside a forest is more humid than the air out-
side; the difference is greatest in summer. The clouds that
form above the forest often release rain, so the forest causes
some of the precipitation that falls on it. If the forest is
cleared, this effect will disappear. Air moves horizontally, car-
rying moisture and clouds with it, so clearing a forest may
have little effect on the amount of rain and snow that fall,
but if the forest is cleared over a very large area it is likely that
the climate will become drier.
Without the evaporation of rainwater from tree surfaces,
more of the rain that falls will reach the ground, and without
transpiration less soil water will be returned to the air. If the
ground is well drained, the overall effect is likely to make the
ground drier, but if the drainage is poor the ground may
become wetter.
If temperate forests are cleared from a large area, the cli-
mate of that area will change. Weather will be windier and
probably drier, and there will be a bigger contrast between
summer and winter temperatures and between day and night
temperatures. Fogs may be more frequent. Depending on the
drainage, the ground may be drier or wetter. If it is drier there
may be dust storms.

How will forests respond to global warming?

Since about 1880 the average air temperature over the whole
world has risen by about 1.25°F (0.7°C). Approximately half
of this warming occurred between 1880 and 1940. It was
entirely natural and was probably the final part of the recov-
ery from a cold period called the Little Ice Age that had lasted
since the 16th century. From 1940 until the middle of the
1970s the average temperature fell very slightly. Then in
1976–77 there was a sudden jump in temperature of about
 HEALTH OF TEMPERATE FORESTS 221

0.5°F (0.3°C), after which the temperature remained steady

until the early 1990s, when it began rising again.
At present Earth’s average temperature is increasing at a
rate of about 3°F (1.7°C) per century. The warming is not dis-
tributed evenly. It is strongest in the northern latitudes of the
Northern Hemisphere, and especially in northwestern North
America—Alaska and the Yukon—and northeastern Siberia.
There is very little warming in the Southern Hemisphere,
except for the Antarctic Peninsula, which has been growing
warmer for several decades, most probably due to changes in
the circulation of water in the Southern Ocean. Where it has
occurred, the warming has had the effect of raising winter
and nighttime temperatures much more than summer and
daytime temperatures, so the difference in temperature
between summer and winter and daytime and nighttime is
growing smaller.
All of the rise in temperature that has happened since the
late 19th century can be called global warming, but it is the
warming since the 1970s that worries many scientists. Part of
the warming is natural, but it is also highly likely that
humans have been contributing to it by releasing into the air
gases, known as greenhouse gases, that absorb long-wave radi-
ation. This is known as the greenhouse effect or, more cor-
rectly, the enhanced greenhouse effect (see the sidebar).
Climate scientists estimate the way temperatures may
respond to the continued accumulation of greenhouse gases
by means of extremely complex mathematical models they
construct on the fastest and most powerful supercomputers
in the world. There are many of these models and they are
improving all the time, but they are still unable to predict
what the consequences of the change in climate will be for
any particular region. It is not yet possible to calculate how
the change may affect an area the size of the United States,
far less any smaller region. The scientists hope they may
achieve this within the next few years.
Still, there are a few clues. At present the air contains less
carbon dioxide than plants are able to use. Many commercial
growers increase the carbon dioxide concentration in the air
inside their greenhouses in order to stimulate plant growth.
Plants (outside greenhouses) are responding to the increasing
222 TEMPERATE FORESTS

atmospheric concentration of carbon dioxide by growing

bigger. At the same time, milder winters and warmer nights
mean the first frosts of autumn occur later than they used to
do, and the last frosts of spring happen earlier. This has
extended the growing season. In North America the start of
spring, marked by the date of the last frost, is now four to 12
days earlier than it was in about 1980, and autumn weather,
marked by the date of the first frost, begins one to seven days
later. In Europe spring begins four to eight days earlier and
autumn begins 14–22 days later.
Warmer weather makes water evaporate faster, a higher
rate of evaporation makes more cloud form, and more cloud

The greenhouse effect

When any object is warmer than its surroundings, it emits electromagnetic radiation, such
as light or heat. The wavelength of that radiation is inversely proportional to the tempera-
ture of the object emitting it: The hotter the body, the shorter the wavelength of its radia-
tion. This is because the amount of energy carried by electromagnetic radiation is greatest
when the wavelength is shortest, meaning that more wave crests and troughs pass a sta-
tionary point in the same interval of time. Astronomical bodies such as stars and planets
are surrounded by space, which is very cold. The Sun is hot and radiates most intensely at
short wavelengths. Its radiation warms the surface of the Earth, which then emits radiation
at a much longer wavelength because it is relatively cool.
Some of the sunshine falling on the Earth is reflected into space by clouds and pale-
colored surfaces such as snow and desert sand. Some is scattered by particles and tiny
droplets in the air and returns to space without reaching the surface. The surface of land
and sea absorbs approximately 51 percent of the solar radiation reaching Earth. The
absorbed energy warms the material that absorbs it.
During the day the Earth absorbs solar radiation—sunshine—faster than it loses heat by
radiating it away, and so its temperature rises. By late afternoon the surface is radiating
heat at about the same rate as it is absorbing it; during the night it continues to radiate,
but because the Sun is no longer shining, the Earth’s surface temperature falls. It continues
to fall until about one hour before dawn, when the first light appears in the sky.
While air is transparent to incoming short-wave solar radiation, it is less so to outgoing
long-wave radiation because certain gases—principally water vapor, carbon dioxide, and
 HEALTH OF TEMPERATE FORESTS 223

means more rain and snow. In other words, a warmer climate

is also a wetter climate, and the weather today is becoming
cloudier and wetter. In eastern North America, for example,
the temperature has not risen, but precipitation has
increased. More snow falls in winter, and because the snow is
deeper it takes longer to melt in spring.
Warmer and wetter weather does not necessarily benefit
plants, however. If the climate becomes a great deal warmer,
water will evaporate from the soil faster than it falls as rain,
making the soil grow steadily drier, and once the soil is thor-
oughly dry the rainfall will decrease. So a very much warmer
climate is a drier climate. Provided the climate warms by less

methane—absorb radiation at these wavelengths. This absorption of energy warms the

air. It is called the greenhouse effect because, in a similar fashion, the glass of a green-
house allows the sunshine to enter but the warm air inside the greenhouse is unable to
escape, so the air inside the greenhouse becomes much warmer than the air outside.
The greenhouse effect is entirely natural. The absorbed radiation warms the air, which
radiates at longer wavelengths, and energy escapes into space each time it is radiated at
a wavelength no gas absorbs, 8.5–13.0 mm, known as the atmospheric window. The
departure of the outgoing energy is delayed but not prevented. Without the greenhouse
effect the average air temperature at ground level would be approximately 34°F (1°C); in
fact, the average surface temperature is 59°F (15°C).
At present, human activities are enhancing the greenhouse effect by releasing certain
“greenhouse gases,” especially carbon dioxide, into the air. We release carbon dioxide
(CO2) whenever we burn a carbon-based fuel such as coal, oil, or gas. Combustion (burn-
ing) is a chemical reaction in which carbon (C) is oxidized (combined with oxygen, O),
releasing energy and creating carbon dioxide as a by-product:

C + O2 → CO2 + heat

Most climate scientists agree that the accumulation of greenhouse gases is producing
a rise in the average temperature over many parts of the world. They disagree over the
extent to which this presents a serious problem. At present the average temperature is
increasing at about 2.3°F–3.2°F (1.3°C–1.8°C) per century. It seems probable that the
average temperature in the early 22nd century will be 1.8°F–3.6°F (1°C–2°C) higher than
it is today.
224 TEMPERATE FORESTS

than about 3.6°F (2°C) conditions will become wetter; if the

rise in temperature exceeds this threshold they will become
drier. If the temperature continues to increase at its present
rate of about 3°F (1.7°C) per century, the weather in the com-
ing decades will become wetter, but some time in the 22nd
century the increase may exceed the threshold. Of course,
the present rise in temperature may not be sustained.
This is an estimate for the whole world, however, and the
world is not warming evenly. The rise in temperature affects
mainly the temperate and high-latitude Northern Hemi-
sphere, where the threshold may be reached before the end
of the 21st century.
As northern latitudes grow warmer and probably wetter,
the change will favor temperate forests. If nature were left
to take its course, the broad-leaved, deciduous forests typi-
cal of the northeastern United States and northwestern
Europe would expand their range northward, replacing the
southern part of the boreal forest. The boreal forest would
also expand northward. The belt of tundra across northern
Canada and Eurasia would become narrower—it cannot
expand northward, because the Arctic Ocean lies to the
north.
If the climate should become warmer but drier, the broad-
leaved, deciduous forests would not fare so well. They prefer
moist summers and wet winters. As they retreat, coniferous
forests will replace them. Coniferous trees are more tolerant
of dry conditions. They flourish in the north, where the
ground is frozen for most of the long winter and their roots
cannot absorb water. But they also flourish around the
Mediterranean and in parts of the United States that have a
dry summer, such as the mountains of Colorado, Arizona,
and New Mexico.
Many scientists believe that a continued rise in tempera-
ture will sooner or later bring harmful consequences.
Humans may adapt to the changes or seek to avert them by
reducing emissions of greenhouse gases. That is the aim of
the Kyoto Protocol, an international agreement drawn up in
1997 that commits the governments that have signed it to
taking steps to reduce emissions by target amounts set for
each country.
 HEALTH OF TEMPERATE FORESTS 225

Forest clearance and soil erosion

Tree roots push their way through the soil, reaching far in
search of water and nutrients. The roots of the herbs growing
between the trees are engaged in the same pursuit. All these
roots form an intricate network belowground. When the
plants die, their roots decompose, leaving behind the tunnels
they made and once occupied. Small animals are able to
move through the larger of these tunnels, and air penetrates
all of them. Meanwhile the decayed material of the roots
becomes part of the soil, along with the decomposed remains
of leaves, twigs, and wood.
Plants supply the materials that sustain the animals, fungi,
bacteria, and other microbes living in the soil. These organ-
isms feed on the plant material, working it over and breaking
it down. Together the plants, animals, and microorganisms
make the soil, and they continue to make it for as long as
their community endures.
If the forest is cleared, the supply of plant material ceases.
This may not matter. The trees may soon be replaced, or the
land may be converted to fields to grow crops. But there are
dangers facing land that is left bare for too long.
Trees move large amounts of water from the soil and into
the air (see “Evaporation and transpiration” on pages 66–68).
When the trees are removed, that movement ceases. Rain
and melted snow drain downward into the soil and join the
groundwater. Groundwater eventually transports the water
to a river, but the movement is slow (see “How water moves
through a temperate forest” on pages 29–33). If a forest grows
on fairly level ground in a region of high precipitation,
removing the trees may mean the water drains vertically
downward faster than the horizontal movement of the
groundwater can carry it away. Under these circumstances
the water table will rise, and after a time the ground will be
waterlogged. This may harm the next crop of trees by making
the soil cold and airless at the level the roots try to penetrate.
The remedy—which is expensive—is to plow the land and
install drains to remove the surplus water.
On sloping ground the soil may start to erode. Soil is natu-
rally irregular in texture, and soil particles stick together to
form lumps. Raindrops falling onto bare soil knock particles
226 TEMPERATE FORESTS

free from the lumps; repeated impacts wash fine soil grains
into tiny hollows and crevices in the surface, eventually fill-
ing them and sealing the soil beneath an impermeable cap.
Rain then no longer penetrates the soil. Instead it washes over
the surface, washing away the surface layer little by little but
resealing the surface with small particles as it does so. The
flowing water finds its way into natural depressions, carrying
soil with it, widening and deepening the depressions until
they form rills—small channels. Further enlargement by water
turns rills into gullies—channels the size of small stream-
beds—that flow with water only after heavy rain, carrying soil
with them. This downhill transport of soil is called “erosion.”
Washing away surface soil is a gradual process, but under
certain circumstances erosion can happen faster, carrying all
of the topsoil rather than wearing away the surface layer. It is
then called “mass wasting.”
Water draining downward through soil eventually meets a
layer of rock, clay, or some other impermeable material.
Immediately above the impermeable layer the accumulating
water may turn the soil to mud. If the impermeable layer is
fairly close to the surface and if the ground slopes steeply, the
mud may start to flow downhill, carrying the overlying soil
with it. Often the movement is quite slow, the soil shifting a
little farther with each heavy fall of rain in a process called
“soil creep,” but, left unchecked, eventually it will strip away
all of the soil, exposing bare rock.
Creep is one form of mass wasting, but there are others
that are much more dramatic and dangerous. The layer of liq-
uid mud may detach the entire layer of topsoil from the
impermeable material on which it lies. All of the topsoil will
then slide downhill, riding on the lubricating layer of mud. If
the mud layer is close to the surface, the slide will carry all of
the surface soil, vegetation, and debris of all kinds, accelerat-
ing as it gathers momentum. This is a landslide. If the soil is
extremely sodden, all of it may slide as a mudflow. Both land-
slides and mudflows are catastrophic events that can block or
carry away roads and demolish homes.
Even without mudslides, erosion due to deforestation can
remove large amounts of soil. A study in England found that
sandy soil eroded at a rate of eight tons per acre (17.7
 HEALTH OF TEMPERATE FORESTS 227

tonnes/ha) every year from a hillside left bare following the

clearance of a forest.
Foresters understand the risks of erosion on hillsides and
seek to prevent it. The first step is to plan the harvesting
operation in such a way as to leave as small an area of bare
ground as possible. As soon as the lumber has been removed,
machines dig trenches at right angles to the slope. The
trenches are not meant to transport water, but only to collect
water flowing downhill and hold it long enough for the
water to soak into the ground. Work to prepare the ground
for the next tree crop begins with scarification. This operation
breaks up the soil using machines that chop up and scatter
the brash—small branches and foliage—left after harvesting.
Scarification leaves a very uneven surface that water pene-
trates easily. Finally, except on very well drained ground,
machines dig soil from hollows to make mounds on which
the new tree seedlings are planted. This also breaks up the
surface and improves drainage.
Forest clearance and agriculture are the two principal caus-
es of soil erosion. Some erosion is unavoidable, because this
is a natural geological process that becomes serious only
when soil is removed faster than new soil can form. Good
land management reduces the risk of unacceptably high rates
of erosion.

Sustainable forest management

During the 1960s many people began to fear that the
resources on which society depends were being used up at a
rate that would soon cause some of them, particularly oil and
metal ores, to be exhausted. Also, those resources that renew
themselves naturally (see “Renewable and nonrenewable
resources” on pages 175–178) were being consumed faster
than they were being replenished.
The fear was not new. In the middle of the 19th century,
mining geologists were warning that the supply of coal was
not infinite, although the known reserves were so large that
most people could not conceive of them ever being exhaust-
ed. In 1864 George Perkins Marsh (see the sidebar on page
164) warned of the dangers of clearing forests.
228 TEMPERATE FORESTS

Eventually the concerns about the depletion of minerals

and fuels—the nonrenewable resources—and the degradation
of renewable resources such as farmland led to the idea of
sustainability. The term first appeared prominently in
World Conservation Strategy: Living Resource Conservation for
Sustainable Development (1980). This was a document pub-
lished jointly by the International Union for Conservation,
Nature, and Natural Resources (IUCN), the World Wildlife
Fund (WWF), the UN Environment Program (UNEP), the UN
Food and Agriculture Organization (FAO), and the UN
Educational, Scientific and Cultural Organization (UNESCO).
World Conservation Strategy called on governments to devise
resource conservation strategies for their own countries. It
had little effect, but the word sustainable caught the imagina-
tion of environmentalists and, through them, of politicians.
Sustainability was defined in another document, Our
Common Future, the report of the World Commission on
Environment and Development. (The United Nations estab-
lished this independent body in 1983 under Gro Harlem
Brundtland, then the prime minister of Norway, and the
commission prepared the way for the UN Conference on
Environment and Development—the Earth Summit—held in
Rio de Janeiro in 1992. Our Common Future outlined the mat-
ters to be debated at the conference.) It said: “Humanity has
the ability to make development sustainable—to ensure that
it meets the needs of the present without compromising the
ability of future generations to meet their own needs.” (Our
Common Future, p. 8.)
A commercial forester might take this to mean that a forest
should produce a crop of trees each year and continue doing
so indefinitely. But that is how forests have been managed in
Europe since medieval times, so it is hardly original. A more
imaginative interpretation follows from a different definition
of forest. If the forest is seen as a community of living organ-
isms rather than simply as so much standing timber, then to
be sustainable its management must preserve the entire com-
munity—the biodiversity of the forest. Such management
does not preclude commercial exploitation. Trees and small
wood can still be taken from the forest, but in ways that
cause the least possible disturbance to other organisms.
 HEALTH OF TEMPERATE FORESTS 229

Plantation forests (see pages 199–202) supply timber and

relieve the pressure on natural forests. Today they, too, are
managed in ways that are more consistent with the aims of
conservation. British plantations once consisted almost
exclusively of coniferous trees that do not grow naturally in
Britain. It takes time for those trees to mature and many
coniferous plantation still remain, but as the old plantations
are cleared, stands of more varied species are planted to
replace them.
Plantations are less popular in countries with much larger
expanses of natural forest. There, timber is cut from the natu-
ral forest, but the forests are then encouraged to regenerate
naturally from seeds shed from the forest trees. Where that is
impractical, plantations replace cleared forest. Areas of old-
growth forest—managed or unmanaged forest that has been
standing for more than 200 years—are being identified and
marked for protection. In Britain, ancient woodland—grow-
ing on land that is known from documentary evidence to
have been forested since before 1600—is also protected.
Insensitive logging still occurs, some of it illegal, but atti-
tudes are changing. Temperate forests are now regarded as
much more than “timber factories.” They are popular for
recreation, education, and conservation. Any plan to clear a
forested area is certain to meet strong local opposition, and
if the forest is especially beautiful, extensive, or rich in
wildlife, the opposition will be national or even internation-
al. It may be too early to say that every temperate forest is
managed in a sustainable fashion, but the trend is clearly in
that direction.
CONCLUSION:
WHAT FUTURE FOR
TEMPERATE FORESTS?
Long, long ago forests of oak, elm, beech, chestnut, maple,
ash, poplar, linden, birch, and many other species blanketed
most of the lowlands of Europe. Similar forests covered
much of the northeastern United States. Many of those
forests have gone, cleared by long-dead settlers to make way
for farms.
Farms were needed, of course. People must be fed—and so
must the draft animals that helped the farmers till the land.
But forests have always been needed. Most furniture, house
frames, floors, and everyday tools are made wholly or partly
from wood. This book is made from paper manufactured
from what were once trees growing in a forest. Wood was
once the only fuel that heated every home, from the king’s
palace and the baron’s castle to the villager’s cottage. It
cooked everyone’s meals, and the only way to obtain hot
water for washing was to place a cauldron over a wood fire.
Many people burn wood to keep warm even today. We may
boast of living in the “Information Age,” but the truth is that
we live in the Wooden Age, and we always have.
Nowadays a new use has been found for our forests, based
on a deeper understanding and appreciation of the way they
function. Forests are seen not simply as sources of raw mate-
rials and fuel, but as living communities of plants, animals,
fungi, and organisms so small they can be seen only with the
aid of a microscope. The forest reveals itself as a wonderland
of diversity, a community of living organisms so vast and
complex as to be barely comprehensible.
Most North Americans and Europeans now live in cities.
The countryside and its forests are remote from the side-
walks, subways, city blocks, and suburban sprawl, and city
parks are pale imitations of genuine countryside. This makes
the forests still more special—and valuable. The forest is a

230
 CONCLUSION: WHAT FUTURE FOR TEMPERATE FORESTS? 231

place to visit. It is where families can walk, children can play,

and naturalists can observe what is around them. It is where
scientists can study the planet we all share, and where stu-
dents can learn.

As well as supplying raw

materials, forests within
reach of cities are also
amenities, places where
people can relax, play,
or be refreshed.
(Courtesy of Fogstock)
232 TEMPERATE FORESTS

Throughout much of the Tropics, forests are being cleared,

although strenuous efforts are being made to halt the clear-
ances. In temperate regions, on the other hand, the forests
are expanding. Forests that have stood for centuries, known
as old-growth forests in most of the world and as ancient
woodland in Britain, are protected. The protection is not
always effective and valued forests are sometimes lost, often
through illegal logging, but conservation bodies are increas-
ingly vigilant and the chainsaws can no longer enter the for-
est unchallenged.
It seems certain, therefore, that the temperate forests will
survive long into the future. Changes in the climate may
alter their composition, but forests are not static; their com-
position is changing all the time. Trees come and go, but the
forest will remain. Of that we may be certain.
 SI UNITS
AND CONVERSIONS
UNIT QUANTITY SYMBOL CONVERSION

Base units
meter length m 1 m = 3.2808 feet
kilogram mass kg 1 kg = 2.205
pounds
second time s
ampere electric current A
kelvin thermodynamic K 1 K = 1°C =
temperature 1.8°F
candela luminous
intensity
mole amount of cd mol
substance

Supplementary units
radian plane angle rad p/2 rad = 90°
steradian solid angle sr

Derived units
coulomb quantity of C
electricity
cubic meter volume m3 1 m3 = 1.308
yards3
farad capacitance F
henry inductance H
hertz frequency Hz
joule energy J 1 J = 0.2389 calories
kilogram per density kg m–3 1 kg m–3 =
cubic meter 0.0624 lb. ft.–3
lumen luminous flux lm
lux illuminance lx

(continues)

233
234 TEMPERATE FORESTS

(continued)

UNIT QUANTITY SYMBOL CONVERSION

meter per second speed m s–1 1 m s–1 = 3.281
ft s–1
meter per second acceleration m s–2
squared
mole per cubic concentration mol m–3
meter
newton force N 1 N = 7.218 lb.
force
ohm electric resistance V
pascal pressure Pa 1 Pa = 0.145 lb.
in–2
radian per second angular velocity rad s–1
radian per second angular acceleration rad s–2
squared
square meter area m2 1 m2 = 1.196 yards2
tesla magnetic flux T
density
volt electromotive force V
watt power W 1W = 3.412 Btu h–1
weber magnetic flux Wb

Prefixes used with SI units

PREFIX SYMBOL VALUE
atto a × 10–18
femto f × 10–15
pico p × 10–12
nano n × 10–9
micro m × 10–6
milli m × 10–3
centi c × 10–2
deci d × 10–1
deca da × 10
hecto h × 102
kilo k × 103
mega M × 106
 SI UNITS AND CONVERSIONS 235

PREFIX SYMBOL VALUE

giga G × 109
tera T × 1012

Prefixes attached to SI units alter their value.

 SOIL CLASSIFICATION:
ORDERS OF THE SOIL
TAXONOMY
Entisols Soils with weakly developed horizons, such as dis-
turbed soils and soils developed over alluvial (river) deposits.
Vertisols Soils with more than 30 percent clay that crack
when dry.
Inceptisols Soils with a composition that changes little with
depth, such as young soils.
Aridisols Soils with large amounts of salt, such as desert
soils.
Mollisols Soils with some horizons rich in organic matter.
Spodosols Soils rich in organic matter, iron, and aluminum;
in older classifications known as a podzol.
Alfisols Basic soils in which surface constituents have moved
to a lower level.
Ultisols Acid soils in which surface constituents have moved
to a lower level.
Oxisols Soils rich in iron and aluminum oxides that have lost
most of their nutrients through weathering; old soils often
found in the humid Tropics.
Histosols Soils rich in organic matter.

Reference Soil Groups of the Food and

Agriculture Organization of the
United Nations (FAO)
Histosols Soils with a peat layer more than 15.75 inches (40
cm) deep.
Cryosols Soils with a permanently frozen layer within 39
inches (100 cm) of the surface.
Anthrosols Soils that have been strongly affected by human
activity.
Leptosols Soils with hard rock within 10 inches (25 cm) of the
surface, or more than 40 percent calcium carbonate within 10
inches (25 cm) of the surface, or less than 10 percent of fine
earth to a depth of 30 inches (75 cm) or more.

237
238 TEMPERATE FORESTS

Vertisols Soil with a layer more than 20 inches (50 cm) deep
containing more than 30 percent clay within 39 inches (100
cm) of the surface.
Fluvisols Soils formed on river (alluvial) deposits with vol-
canic deposits within 10 inches (25 cm) of the surface and
extending to a depth of more than 20 inches (50 cm).
Solonchaks Soils with a salt-rich layer more than six inches
(15 cm) thick at or just below the surface.
Gleysols Soils with a sticky, bluish gray layer (gley) within 20
inches (50 cm) of the surface.
Andosols Volcanic soils having a layer more than 12 inches
(30 cm) deep containing more than 10 percent volcanic
glass or other volcanic material within 10 inches (25 cm) of
the surface.
Podzols Pale soils with a layer containing organic material
and/or iron and aluminum that has washed down from above.
Plinthosols Soils with a layer more than six inches (15 cm)
deep containing more than 25 percent iron and aluminum
sesquioxides (oxides comprising two parts of the metal to
three parts of oxygen) within 20 inches (50 cm) of the surface
that hardens when exposed.
Ferralsols Soils with a subsurface layer more than six inches
(15 cm) deep with red mottling due to iron and aluminum.
Solonetz Soils with a sodium- and clay-rich subsurface layer
more than three inches (7.5 cm) deep.
Planosols Soils that have had stagnant water within 40 inches
(100 cm) of the surface for prolonged periods.
Chernozems Soils with a dark-colored, well-structured, basic
surface layer at least eight inches (20 cm) deep.
Kastanozems Soils resembling chernozems, but with concen-
trations of calcium compounds within 40 inches (100 cm) of
the surface.
Phaeozems All other soils with a dark-colored, well struc-
tured, basic surface layer.
Gypsisols Soils with a layer rich in gypsum (calcium sulfate)
within 40 inches (100 cm) of the surface, or more than 15 per-
cent gypsum in a layer more than 40 inches (100 cm) deep.
Durisols Soils with a layer of cemented silica within 40 inches
(100 cm) of the surface.
Calcisols Soils with concentrations of calcium carbonate
within 50 inches (125 cm) of the surface.
Albeluvisols Soils with a subsurface layer rich in clay that has
an irregular upper surface.
 SOIL CLASSIFICATION 239

Alisols Slightly acid soils containing high concentrations of

aluminum and with a clay-rich layer within 40 inches (100
cm) of the surface.
Nitisols Soils with a layer containing more than 30 percent
clay more than 12 inches (30 cm) deep and no evidence of
clay particles moving to lower levels within 40 inches (100
cm) of the surface.
Acrisols Acid soils with a clay-rich subsurface layer.
Luvisols Soils with a clay-rich subsurface layer containing clay
particles that have moved down from above.
Lixisols All other soils with a clay-rich layer within 40–80
inches (100–200 cm) of the surface.
Umbrisols Soils with a thick, dark colored, acid surface layer.
Cambisols Soils with an altered surface layer or one that is
thick and dark colored, above a subsoil that is acid in the
upper 40 inches (100 cm) and with a clay-rich or volcanic
layer beginning 10–40 inches (25–100 cm) below the surface.
Arenosols Weakly developed soils with a coarse texture.
Regosols All other soils.

Note: A basic or alkaline soil is one containing more hydroxyl

ions (OH–) than hydrogen ions (H+); an acid soil contains more
H+ ions than OH– ions. Acidity is measured on the pH scale, a
logarithmic scale expressing the activity of H+ ions in solution,
where pH 7.0 is a neutral reaction. A basic soil has a pH greater
than 7.0; an acid soil has a pH lower than 7.0.
 GEOLOGIC TIME SCALE
Eon/ Era/ Period Epoch/ Began
Eonothem Erathem Subera System Series Ma*

Quaternary Pleistogene Holocene 0.11

Pleistocene 1.81
Phanerozic Cenozoic Tertiary Neogene Pliocene 5.3
Miocene 23.03
Paleogene Oligocene 33.9
Eocene 55.8
Paleocene 65.5
Mesozoic Cretaceous Upper 99.6
Lower 145.5
Jurassic Upper 161.2
Middle 175.6
Lower 199.6
Triassic Upper 228
Middle 245
Lower 251
Paleozoic Upper Permian Lopingian 260.4
Guadalupian 270.6
Cisuralian 299
Carboniferous Pennsylvanian 318.1
Mississippian 359.2
Devonian Upper 385.3
Middle 397.5
Lower 416
Lower Silurian Pridoli 422.9
Ludlow 443.7
Wenlock 428.2
Llandovery 443.7
Ordovician Upper 460.9
Middle 471.8

(continues)

241
242 TEMPERATE FORESTS

(continued)

Eon/ Era/ Period Epoch/ Began

Eonothem Erathem Subera System Series Ma*

Lower 488.3
Cambrian Furongian 501
Middle 513
Lower 542
Proterozoic Neoproterozoic Ediacaran 600
Cryogenian 850
Tonian 1000
Mesoproterozoic Stenian 1200
Ectasian 1400
Calymmian 1600
Paleoproterozoic Statherian 1800
Orosirian 2050
Rhyacian 2300
Siderian 2500
Archean Neoarchean 2800
Mesoarchean 3200
Paleoarchean 3600
Eoarchean 3800
Hadean Swazian 3900
Basin Groups 4000
Cryptic 4567.17

Source: International Union of Geological Sciences, 2004.

Note: Hadean is an informal name. The Hadean, Archean, and Proterozoic eons cover the time formerly known as the
Precambrian. Quaternary is now an informal name and Tertiary is likely to become informal in the future, although
both continue to be widely used.

*Ma means millions of years ago.

 GLOSSARY
adiabatic a change in temperature that involves no exchange
of heat with an outside source
adventitious arising from an unusual part of the plant. Roots
that emerge from NODES are said to be adventitious
air mass a large body of air, covering most of a continent or
ocean and extending to the TROPOPAUSE, in which atmospheric
conditions are fairly constant throughout
alga an organism (a protist) that performs PHOTOSYNTHESIS; it
may be single-celled or multicelled. Seaweeds are algae
alternation of generations two stages in the life cycle of a
plant. The DIPLOID generation is called a SPOROPHYTE and the
HAPLOID generation a GAMETOPHYTE
ambrosia beetle a member of a group of beetles that feed on
ambrosia, a sweet substance produced by certain FUNGI
angiosperm a flowering plant
anther the structure in a male flower, situated at the tip of a
FILAMENT, where POLLEN is produced
anthophyte a flowering plant; flowering plants were formerly
classified in the division Anthophyta
anticyclone a region in which the atmospheric pressure is
higher than it is in the surrounding air
aquifer an underground body of permeable material (e.g.,
sand or gravel) lying above a layer of impermeable material
(e.g., rock or clay) that is capable of storing water and
through which the GROUNDWATER flows
auxin a hormone-like substance produced by a plant that pro-
motes growth by making cells at the tips of branches grow
longer
axil the angle where a leaf joins the stem or a small branch
joins a larger one
bedrock the rock lying below the ground surface over a large
area
biomass the total mass of all the living organisms present in a
given area

243
244 TEMPERATE FORESTS

biome the largest biological community recognized, compris-

ing a type of vegetation (e.g., temperate forest) together with
the other organisms associated with it, occupying a large geo-
graphical area.
blocking the situation that occurs when an ANTICYCLONE or
area of low pressure becomes stationary, obstructing the pas-
sage of weather systems which are forced to move around it
bootlace fungus see HONEY FUNGUS
boreal pertaining to the north
boundary current an ocean current that flows northward or
southward close to the coast of a continent and parallel to it.
Eastern boundary currents carry cold water southward on the
eastern side of ocean basins (not along the eastern coasts of
continents). Western boundary currents carry warm water
northward on the western side of ocean basins
boundary layer the layer of a gas or liquid adjacent to a solid
surface within which the characteristics of the gas or liquid
are strongly influenced by proximity to the surface
bract a modified leaf that forms part of a flower
brash small branches and leaves that are cut from a tree trunk
before the trunk is removed from the forest
broad-leaved describes the leaves of flowering plants, which
are broad compared with the needles or scale leaves of
CONIFERS
browse to feed on the leaves and shoots of living plants;
leaves and shoots that are cut in order to be fed to livestock
calyx the collective name for the SEPALS
canopy the forest cover, more or less shading the floor,
formed by the touching and overlapping branches and
foliage of adjacent trees
capillarity (capillary attraction) the movement of water
against gravity through a fine tube or narrow passageway
capillary attraction see CAPILLARITY
capillary fringe the region immediately above the WATER TABLE
into which water rises by CAPILLARITY
carpel the female reproductive organs of a plant, comprising
the STIGMA, STYLE, and OVARY
carrying capacity the largest population of a particular
species that an area of environment can sustain
cecidium see GALL
charcoal a form of impure carbon used as a fuel, made by
heating wood in airless conditions
 GLOSSARY 245

chestnut blight a disease of chestnut trees caused by the fun-

gus Endothia parasitica that was introduced to North America
from Asia and by 1940 had killed most American chestnut
trees
chipboard an industrial and construction material manufac-
tured in sheets from compressed wood chippings
chlorophyll the pigment present in the leaves and sometimes
stems of green plants that gives them their green color.
Chlorophyll molecules trap light, thus supplying the energy
for PHOTOSYNTHESIS
chloroplast the structure in plant cells that contains CHLORO-
PHYLL and in which PHOTOSYNTHESIS takes place
chromosome one of the threads of DNA found in the nucleus
of a cell. Each chromosome carries some of the organism’s
genes, and the full complement of chromosomes (a number
that varies according to species) contains the full set of genet-
ic instructions (the genome) for that organism
climax the final stage in a plant SUCCESSION in which the plant
community reaches a stable equilibrium with the environ-
ment
companion cell a modified PARENCHYMA cell that is linked to a
SIEVE CELL
condensation the change from gas to liquid
conifer a GYMNOSPERM plant that produces seeds in cones,
releasing them when the cone scales open
continental drift the movement of the continents in relation
to one another across the Earth’s surface
coppice the practice of cutting trees close to ground level in
order to stimulate the growth of poles; an area of woodland
that has been managed by coppicing
CorF see CORIOLIS EFFECT
Coriolis effect (CorF) the deflection due to the Earth’s rota-
tion experienced by bodies moving in relation to the Earth’s
surface; bodies are deflected to the right in the Northern
Hemisphere and to the left in the Southern Hemisphere
cork (phellem) a layer of protective tissue, comprising dead
cells, that forms immediately beneath the outer surface of the
trunk and branches of a woody plant
cork cambium see PHELLOGEN
corolla the collective name for all the PETALS of a flower
cotyledon a seed leaf; the leaf that emerges from a germinat-
ing seed
246 TEMPERATE FORESTS

crown the branches and foliage at the top of a tree

crown fire a forest fire that reaches the crowns of the trees,
leaping from tree to tree
cuticle a thin, waxy outer coat that protects the leaves and
stem of a plant
cyclone see DEPRESSION
deciduous describes parts of a plant or animal that are shed at
the same season each year
denitrification the conversion by bacteria of nitrate or nitrite
to a gas, principally nitrogen and nitrous oxide (N2O)
depression (cyclone) a region along a weather front where
the atmospheric pressure is lower than it is in the surround-
ing air
destructive distillation the process of heating organic materi-
al in the absence of air in order to drive off VOLATILES, leaving
a more concentrated carbon fuel
detritivore an organism that feeds on DETRITUS
detritus fragments of dead plant and animal material that
forms a layer on the surface of the ground
dew-point temperature the temperature at which water
vapor condenses to form dew or cloud droplets
diploid describes a cell containing two sets of CHROMOSOMES,
or an organism made up of such cells
disjunct distribution the occurrence of related species in
places separated by major geographical barriers, such as an
ocean or mountain range
dominant the species comprising the most prominent plants
in a community or the species with the most influence of the
character of the community
drip line the line surrounding a tree that is formed by water
dripping from the tips of leaves
dry adiabatic lapse rate see LAPSE RATE
Dutch elm disease a disease of elm trees caused by the fungus
Ophiostoma ulmi, carried into the tree by elm bark beetles,
mainly Scolytus scolytus and S. multistriatus
ecological energetics the study of the flow of energy through
ECOSYSTEMS
ecological pyramid a diagram representing feeding relation-
ships within an ECOSYSTEM as a series of bands, all the same
thickness but varying in width, with PRODUCERS at the base
and CONSUMERS stacked above them
ecology the scientific study of the relationships among
organisms inhabiting a specified area and between the
 GLOSSARY 247

organisms and the physical and chemical conditions in

which they live
ecosystem a clearly defined area or unit within which living
organisms and their physical and chemical surroundings
interact to form a stable system
edge effect the occurrence of more species in the area where two
ECOSYSTEMS overlap than are found in the two separate ecosys-
tems combined, because the overlap area provides conditions
suitable for species that could not survive in either ecosystem
elytra (sing. elytron) the hardened forewings of a beetle that
give the beetle its characteristic appearance when not flying
and that rise to allow the hind wings to open when the beetle
is in flight
elytron see ELYTRA
embryo a young plant contained within a plant seed, or a
young animal contained within a fertilized egg or other
reproductive structure. (In humans an embryo is called a fetus
after the first eight weeks of pregnancy)
emergent a forest tree that stands taller than those around it
ephemeral short-lived
epizootic a disease epidemic in nonhuman animals
equinox March 20–21 and September 22–23, when the noon-
day Sun is directly overhead at the equator and day and night
are of equal length everywhere in the world
ethene see ETHYLENE
ethylene (ethene) a gas (C2H4) produced by plants that func-
tions as a hormone, controlling such processes as seed germi-
nation, cell growth, fruit ripening, and aging
evaporation the change from liquid to gas
evapotranspiration EVAPORATION and TRANSPIRATION considered
together
evergreen describes a plant that bears leaves at all times of
year; although it sheds its leaves, it does not shed all of them
at the same time
exine the tough outer coat of a POLLEN grain
Fennoscandian ice sheet the ice sheet that covered northern
Europe during the last GLACIAL
Ferrel cell part of the general circulation of the atmosphere in
which air rises in about latitude 60° in both hemispheres,
moves toward the equator at high altitude, subsides in about
latitude 30°, and flows away from the equator at low level
fertilization the union of male and female GAMETES to pro-
duce an EMBRYO
248 TEMPERATE FORESTS

fiberboard material manufactured in sheets from waste mate-

rial from sawmills and factories making wood products that is
ground into a mass of fibers
filament the stalk of the STAMEN of a flower, bearing an ANTHER
at its tip
fire climax see PYROCLIMAX
firestorm a fire so intense that the air drawn in to replace hot
air rising by convection moves at gale force, carrying with it
loose material that fuels the flames
floret see INFLORESCENCE
flowering plant see ANGIOSPERM
food chain a set of feeding relationships in which each in a
sequence of organisms feeds on the preceding member
food web a diagram that shows the inhabitants of an ECOSYS-
TEM linked by lines between species and the species on which
they feed; an array of FOOD CHAINS
forb a herbaceous plant other than a grass
forest limit see TREE LINE
front the boundary between two AIR MASSES
Fungi one of the kingdoms of biological classification, com-
prising non-photosynthesizing organisms that feed by
absorbing organic substances from their surroundings and
reproduce by SPORES
gall (cecidium) a growth or swelling on the roots, stem, or
leaves of a plant caused by bacterial or fungal infection or by
attack from mites, nematodes, or insects
gamete a sex cell, i.e., a spermatozoon or ovum
gametophyte the HAPLOID stage in the life cycle of a plant. In
simple plants, such as mosses, the gametophyte is the visible
plant; in GYMNOSPERMS and ANGIOSPERMS the gametophyte is
inconspicuous
girdling see RINGBARKING
glacial a period when polar ice sheets advance; an ice age
greenhouse effect the absorption and reradiation of long-
wave radiation emitted by the Earth’s surface by molecules of
water vapor, carbon dioxide, ozone, and several other “green-
house gases,” warming the air
ground fire a fire that burns across the ground surface, affect-
ing only HERBS and the lower parts of trees and shrubs
groundwater underground water that flows through an AQUIFER
gymnosperm a seed plant in which the OVULES are carried
naked on the scales of a cone. Coniferous (i.e., cone-bearing)
trees are the most abundant gymnosperms
 GLOSSARY 249

habitat the living place of a species or community

Hadley cell the tropical part of the general circulation of the
atmosphere. Air rises over the equator, moves away from the
equator at high altitude, subsides over the subtropics, and
flows toward the equator at low altitude
haploid describes a cell nucleus that contains only one set of
CHROMOSOMES
heartwood the central part of a tree trunk, made from dead
cells
herb a small, nonwoody plant in which all the parts above
ground die back at the end of each growing season
holdfast the structure by which a seaweed or other ALGA is
attached to a solid surface
honey fungus (bootlace fungus) the most serious of all fun-
gal parasites of trees. It spreads throughout the tree, eventual-
ly killing it
hoof fungus see TINDER FUNGUS
humidity the amount of water vapor present in the air
humus decomposed plant and animal material in the soil
hypha one of the minute threads that form the main part of a
fungus
ice storm wind-driven rain that falls through air and onto sur-
faces below freezing temperature, where it freezes on contact,
forming thick layers of ice
inflorescence a mass of small but complete flowers (called flo-
rets) growing together and giving the appearance of a single
flower. Sunflower and grass “flowers” are inflorescences
interglacial a period of warmer weather between two GLACIALS
Intertropical Convergence Zone (ITCZ) the region where the
TRADE WINDS from either hemisphere meet (converge)
isobar a line drawn on a weather map to join points on a sur-
face (not necessarily the ground surface) of equal air pressure
ITCZ see INTERTROPICAL CONVERGENCE ZONE
jet stream a winding ribbon of strong wind about 5–10 miles
(8–16 km) above the surface. Jet streams are typically thou-
sands of miles long, hundreds of miles wide, and several miles
deep
key see SAMARA
krummholz stunted, gnarled, small trees that grow on moun-
tainsides between the upper limit for forest and the TREE LINE
K-species species adapted to a stable environment; they pro-
duce few offspring and devote much care to their young, and
most of the young survive
250 TEMPERATE FORESTS

lapse rate the rate at which the air temperature decreases

(lapses) with increasing altitude. In unsaturated air, the dry
ADIABATIC lapse rate is 5.38°F per thousand feet (9.8°C per km);
in saturated air the saturated adiabatic lapse rate varies, but
averages 2.75°F per thousand feet (5°C per km)
latent heat the heat energy that is absorbed or released when
a substance changes phase between solid and liquid, liquid
and gas, and solid and gas. For water at 32°F (0°C) the latent
heat of melting and freezing is 80 cal. per gram (334 J/gram);
of vaporization and condensation 600 cal. per gram (2,501
J/gram); and for SUBLIMATION and DEPOSITION 680 cal. per gram
(2,835 J/gram)
Laurentide ice sheet the ice sheet that covered northeastern
North America during the most recent GLACIAL
lichen a composite organism comprising a fungus and an
ALGA or cyanobacterium
lifting condensation level the altitude at which the air is at
the DEW-POINT TEMPERATURE and water vapor begins to con-
dense to form cloud; the lifting condensation level marks the
cloud base
lignification the process by which LIGNIN accumulates in the
cells of woody plants after the cells die
lignin a hard substance that forms in the cells of woody
plants, binding other cell components together and remain-
ing in position after the cells have died
lumber felled trees that have been stripped of their branches
and cut into logs of a length that can be transported conve-
niently
macronutrient a nutrient substance that living organisms
need in relatively large amounts
mantle that part of the Earth’s interior lying between the
outer edge of the inner core and the underside of the crust
meiosis a form of cell division that occurs in sexually repro-
ducing organisms, in which the cell divides twice, producing
four HAPLOID daughter cells
meristem plant tissue composed of cells that are capable of
dividing indefinitely
mesophyll the tissue lying just below the surface of a leaf,
where PHOTOSYNTHESIS takes place
micronutrient a nutrient substance that living organisms
need in relatively small amounts
mitochondria (sing. mitochondrion) a body (ORGANELLE)
present in large numbers in every fungal, plant, and animal
 GLOSSARY 251

cell, that is responsible for releasing energy by the process of

RESPIRATION
mitochondrion see MITOCHONDRIA
mitosis cell division in which the cell divides to produce two
identical daughter cells, both of which are DIPLOID
mycelium the mass of hyphae that make up the main part of
a fungus; see also HYPHA
mycorrhiza a close physical association between a fungus and
the roots of a plant from which both organisms benefit
nectary a plant gland that secretes nectar
niche the function an organism performs in its environment
nitrification the oxidation by bacteria of ammonia to nitrite
and/or of nitrite to nitrate
nitrogen fixation any chemical reaction incorporating
gaseous nitrogen into a compound that can be utilized by
plants
occluded front see OCCLUSION
occlusion (occluded front) the stage in the life cycle of a
frontal weather system at which advancing cold air has
pushed beneath warmer air and begun to lift the warm air
clear of the surface
organelle a structure that has a specialized function within a
cell
orographic lifting the movement of air as it rises to cross a
mountain or mountain range
ovary the female reproductive organ of a flower
ovule the structure in ANGIOSPERMS and GYMNOSPERMS that
develops into the seed following FERTILIZATION
paleoclimatology the study of ancient climates
Pangaea the SUPERCONTINENT that came into existence about
260 million years ago and began to break apart about 220 mil-
lion years ago
Panthalassa the world ocean that surrounded PANGAEA
pappus a tuft of hairs or bristles that forms a “parachute”
allowing plant seeds to be carried long distances by the
wind
parenchyma plant tissue composed of unspecialized cells
peat partly decomposed plant material forming a distinct SOIL
HORIZON; peat of suitable quality is dried and burned as fuel
pedicel the stalk attaching a flower to the plant
pedology the scientific study of soils
peduncle the stalk attaching an INFLORESCENCE to the plant
perianth the CALYX and COROLLA of a flower
252 TEMPERATE FORESTS

permafrost permanently frozen ground. To become per-

mafrost the ground must remain frozen throughout a mini-
mum of two winters and the summer between
permeability the ability of a material to allow water to flow
through it
petal a modified leaf, often brightly colored; the petals sur-
round and partly enclose the reproductive organs of a flower
petiole the stalk that attaches a leaf to the stem of a plant
phellem see CORK
phellogen (cork cambium) a layer of tissue beneath the outer
bark of a tree comprising cells that divide to produce new
cork and new bark
phloem tissue through which the products of photosynthesis
and hormones are transported from the leaves to all parts of a
vascular plant
phosphorylation a chemical reaction in which phosphate
(PO4) is added
photosynthesis the sequence of chemical reactions in which
green plants and cyanobacteria use sunlight as a source of
energy for the manufacture (synthesis) of sugars from hy-
drogen and carbon, obtained from water and carbon diox-
ide respectively. The reactions can be summarized as:
6CO2 + 6H2O + light → C6H12O6 + 6O2 ↑.
The upward arrow indicates that oxygen is released into
the air; C6H12O6 is glucose, a simple sugar.
photosynthetic pigment a chemical substance that absorbs
visible light, thereby making energy available for PHOTOSYN-
THESIS. Chlorophylls a and b are the most important photo-
synthetic pigments; carotenoids are accessory pigments that
transfer energy to chlorophyll a
pith ray structures radiating from the center to the exterior of
a tree trunk or branch that store starch
plane of the ecliptic the imaginary disk with the Sun at its center
and the Earth’s orbital path around the Sun as its circumference
plant association a type of vegetation comprising certain
species that are always present and other species that are
often present
plantation a forest consisting of trees that have been planted
to provide a crop of timber that will be harvested
plate see PLATE TECTONICS
plate tectonics the theory holding that the Earth’s crust com-
prises a number of rigid sections, or plates, that move in rela-
tion to each other
 GLOSSARY 253

plumule the part of a plant EMBRYO that will grow into a

shoot
plywood a product made from thin sheets of wood that are
glued together in layers, with the grain in each layer running
at right angles to the grain in the layers on either side
polar cell part of the general circulation of the atmosphere in
which air subsides over the North and South Poles, moves
away from the poles at low level, rises in about latitude 60°,
and flows back toward the poles at high altitude
pollard to cut off the top of a tree about six feet (1.8 m) above
ground level. This produces a crop of poles that emerge too
high for browsing animals to reach
pollen the grains containing male sex cells that are produced
in the ANTHERS of flowers
pore space the total interconnected space between the miner-
al particles in a soil
porosity the percentage of the total volume of a material that
consists of spaces between particles
pressure gradient a change in air pressure across a horizontal
distance
pressure potential see TURGOR PRESSURE
prevailing wind the direction from which the wind most fre-
quently blows in a particular location
primary growth plant growth occurring at the tips of the
stem and branches causing the plant to grow taller and its
branches longer
pyroclimax (fire climax) a CLIMAX that develops where fire
occurs at fairly regular intervals; it is dominated by plants
that survive fire or benefit from it
radicle the part of a plant EMBRYO that will grow into a root
radiocarbon carbon-14 (14C), a radioactive isotope produced
in the atmosphere by cosmic-ray bombardment of nitrogen-
14. 14C has a half-life of 5,730 ± 30 years, allowing it to be
used to date material from organisms that absorbed it and
stored it in their tissues while they were living
rain shadow the drier climate on the lee (downwind) side of a
mountain range caused by the loss of moisture as air
approaching the mountains is forced to rise, resulting in the
condensation and precipitation of its water vapor on the
windward slopes. Compression raises the temperature of the
subsiding air, further reducing its RELATIVE HUMIDITY
receptacle the part of a PEDICEL from which all parts of the
flower arise
refugia see REFUGIUM
254 TEMPERATE FORESTS

refugium (pl. refugia) an isolated area in which plants and

animals survive major climatic changes taking place else-
where
relict an organism that has survived while related species
became extinct
respiration the sequence of chemical reactions in which car-
bon in sugar is oxidized with the release of energy; the op-
posite of PHOTOSYNTHESIS. The reactions can be summarized as:

C6H12O6 + 602 → 6CO2 + 6H2O + energy.

C6H12O6 is glucose, a simple sugar
rhizoid one of the hairlike structures through which simple
plants such as mosses and liverworts absorb water and nutri-
ents
ridge a protrusion of high air pressure into a region of low air
pressure
ringbarking (girdling) making a cut all the way around the
trunk of a tree that is deep enough to penetrate the bark and
sever the VASCULAR TISSUE, thereby killing the tree by prevent-
ing the transport of water and nutrients
root plate the roots of a tree, together with the soil attached
to them, that are exposed as a flat, circular, platelike structure
when a tree is blown down by wind
roundwood wood of all kinds that is taken from a forest com-
mercially
r-species a species that is adapted to an unstable environ-
ment. It produces many offspring and devotes little attention
to them. Most of the offspring die, but enough survive to con-
tinue the species and, should conditions improve, many
more will survive and the population will increase
samara a winged fruit in which the wings carry one or two
seeds (those with two seeds are called schizocarpic samaras)
sapwood the active, living part of the trunk or branch of a
woody plant, lying immediately beneath the bark
saturated adiabatic lapse rate see LAPSE RATE
scarification breaking up the soil and BRASH and scattering the
brash in preparation for planting the next tree crop in a plan-
tation
schizocarpic samara see SAMARA
seafloor spreading the theory that the ocean floor is created
at ridges where MANTLE material rises to the surface and the
crustal rocks move away from the ridges on either side, caus-
ing the ocean basin to widen as the seafloor spreads
 GLOSSARY 255

secondary growth the thickening of the stems and branches

of a woody plant by laying down successive layers of new
cells around the central HEARTWOOD
seed the body, formed from a fertilized ovule, from which a
young plant emerges
seed bank the seeds that lie dormant in the soil, or a facility
in which plant seeds are stored
sepal modified leaves attached to the RECEPTACLE that enclose
the flower bud and surround the PETALS of a flower after it
opens
serotiny the retention of seeds by a plant until conditions
favorable for their germination trigger their release
shelter belt trees grown along a strip of land at right angles
to the PREVAILING WIND that shelter crops grown downwind
shrub a perennial woody plant less than 33 feet (10 m) tall
with several main stems arising at or close to ground level,
but with no clearly identifiable trunk
sieve element one of the long, slender, tapering cells, terminat-
ing in a perforated region called a sieve plate, that join end to
end in a sieve tube, forming part of the PHLOEM tissue
silviculture management of a forest for the benefit of the
plants and animals living in it, regardless of whether or not
all or part of the forest is being exploited commercially
skidder a device that grips one end of the trunk of a felled tree
and drags it out of the forest
snedding removing the branches from felled trees
soil horizon a horizontal layer in a SOIL PROFILE that differs in
its mineral or organic composition from the layers above and
below it, and from which it can be clearly distinguished visu-
ally
soil pores see PORE SPACE
soil profile a vertical section cut through soil from the surface
to the underlying rock
solstice one of the two dates each year when the noonday Sun
is directly overhead at one or other of the Tropics and the dif-
ference in length between the hours of daylight and darkness
is at its most extreme. The solstices occur on June 21–22 and
December 22–23
specific heat capacity the amount of heat that must be
applied to a substance in order to raise its temperature by one
degree. It is measured in calories per gram per degree Celsius
(cal/g/°C) or in the scientific units of joules per gram per
kelvin (J/g/K; 1K = 1°C = 1.8°F)
256 TEMPERATE FORESTS

spore a reproductive unit, usually consisting of a single cell,

that can develop into a new organism without fusing with
another cell
sporophyte the reproductive stage in the life cycle of a plant.
In GYMNOSPERMS and ANGIOSPERMS this is the dominant stage,
comprising the visible plant. In mosses, liverworts, and horn-
worts the sporophyte is small and inconspicuous
stadial a prolonged period of cold weather that is shorter and
milder than a GLACIAL
stamen the male reproductive organ of a flower, comprising
the FILAMENT and ANTHER
stigma part of the female reproductive structure in a flower; it
has a sticky surface that holds POLLEN grains
stomata (sing. stoma) small openings, or pores, on the sur-
face of a plant leaf through which the plant cells exchange
gases with the outside air. Stomata can be opened or closed by
the expansion or contraction of two guard cells surrounding
each stoma
stratosphere the region of the atmosphere that extends from
the TROPOPAUSE to an altitude of about 31 miles (50 km)
style the part of the female reproductive structure in a flower
that connects the STIGMA to the OVARY
suberin a waxy substance in the walls of CORK cells that make
cork waterproof
succession a sequence of changes in the composition of a
plant and animal community occupying a site that continues
until a stable equilibrium, the CLIMAX, is attained
supercontinent a landmass formed by the merging of previ-
ously separate continents as a result of CONTINENTAL DRIFT.
PANGAEA was a supercontinent comprising all the present-day
continents
swidden farming a farming method in which farmers clear
forest from an area of land and grow crops there for several
seasons before abandoning the site and moving to another,
allowing the forest to regenerate
taiga the conifer forest forming a belt across northern
America and Eurasia
taproot a plant root that is large and descends vertically
temperate rain forest forest that occurs in temperate regions
where the annual rainfall is typically 60–120 inches
(1,500–3,000 mm)
temperate zone the region of the Earth lying between the
Tropics (23.5°N and S) and the Arctic and Antarctic Circles
(66.5°N and S)
 GLOSSARY 257

tepal SEPALS and PETALS when these are so similar in appearance

as to be indistinguishable
thermal wind a wind generated by a difference in air temper-
ature. JET STREAMS are thermal winds
thylakoid membrane one of the membranes inside a CHLORO-
PLAST that hold the PHOTOSYNTHETIC PIGMENTS
timber wood that has been cut into large pieces for use main-
ly in construction
timberline see TREE LINE
tinder fungus (hoof fungus, touchwood) a parasitic fungus
that attacks trees, especially beech and birch, hollowing them
out and eventually killing them
touchwood see TINDER FUNGUS
tracheid a long, cylindrical cell with a tapering, perforated
end. Tracheids join end to end to form the XYLEM tissue in
GYMNOSPERMS
trade winds the winds that blow toward the equator in
equatorial regions, from the northeast in the Northern
Hemisphere and from the southeast in the Southern
Hemisphere
transpiration the evaporation of water through leaf STOMATA
when these are open for the exchange of gases
tree a perennial, woody plant that is more than 33 feet
(10 m) tall and has one or more than one clearly identifiable
trunk
tree line (timberline, forest limit) the elevation or latitude
beyond which the climate is too severe for trees to grow
trophic pertaining to food or feeding
tropopause the boundary separating the TROPOSPHERE from the
STRATOSPHERE. It occurs at a height of about 10 miles (16 km)
over the equator, seven miles (11 km) in middle latitudes, and
five miles (8 km) over the North and South Poles
troposphere the layer of the atmosphere that extends from
the surface to the TROPOPAUSE; it is the region where all weath-
er phenomena occur
trough a protrusion of low air pressure into a region of high
air pressure
tundra a treeless plain in the arctic or Antarctic, where the
vegetation is dominated by grasses, sedges. rushes, and wood
rushes, together with dwarf shrubs, LICHENS, and mosses
turgor rigidity of plant tissues due to water held under pres-
sure in the cells
turgor pressure (pressure potential) the pressure under
which water is held inside plant tissues
258 TEMPERATE FORESTS

understory the trees that are shorter than the trees forming
the forest CANOPY
vascular bundle a strand of PHLOEM and XYLEM tissue; groups
of vascular bundles form a continuous system extending
throughout the plant
vascular cambium a layer just beneath the outer bark of a
woody plant where dividing cells give rise to new PHLOEM and
XYLEM tissue
vascular plant a plant possessing PHLOEM and XYLEM tissue
through which water and nutrients are transported
veneer a thin layer of high-quality wood that is glued to the
surface of an inferior wood to improve its appearance
vessel element one of the cells forming the XYLEM tissue in
ANGIOSPERMS
volatile describes substances that vaporize at low tempera-
tures
water table the upper margin of the GROUNDWATER; soil is fully
saturated below the water table but unsaturated above it
xylem plant tissue through which water entering at the roots
is transported to all parts of the plant
Younger Dryas a period of cold, dry weather lasting from
about 11,000 to 10,000 years ago and affecting all of northern
Europe, but not North America
zygote the fertilized ovum (egg) of a plant or animal at the
stage where it is DIPLOID, but before it has begun to divide
 BIBLIOGRAPHY AND
FURTHER READING
Allaby, Michael. A Change in the Weather. New York: Facts On File,
2004.
———. Fog, Smog, and Poisoned Rain. New York: Facts On File, 2003.
———. Temperate Forests. New York: Facts On File, 1999.
Ashman, M. R., and G. Pun. Essential Soil Science. Maiden, Md.:
Blackwell Science, 2002.
Brewer, Richard. The Science of Ecology. 2nd ed. Fort Worth, Tex.:
Saunders College Publishing. l988.
Burroughs, William James. Climate Change: A Multidisciplinary
Approach. Cambridge: Cambridge University Press, 2001.
Foth, H. D. Fundamentals of Soil Science. 8th ed. New York: John
Wiley, 1991.
Heywood, V. H., consultant editor. Flowering Plants of the World.
Updated edition. New York: Oxford University Press, 1993.
Peterken, George F. Natural Woodland. Cambridge: Cambridge
University Press, 1996.
Rackham, Oliver. Trees and Woodland in the British Landscape. 2nd
ed. London: Weidenfeld and Nicolson, 2001.
Roberts, Neil. The Holocene: An Environmental History. New York:
Basil Blackwell, 1989.
Tansley, A. G. Practical Plant Ecology. London: George Allen and
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———. Introduction to Plant Ecology. London: George Allen and
Unwin Ltd., 1946.
Williams, Joseph H., and William E. Friedman. “Identification of
Diploid Endosperm in an Early Angiosperm Lineage.” Nature 415
(January 31, 2002): 522–526.
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Common Future. Oxford: Oxford University Press, 1987.

Web sites
Arroyo, Mary T. Kahn, and Adriana E. Hoffmann. “Temperate Rain
Forest of Chile.” Available online. URL: www.nmnh.si.edu/
botany/projects/cpd/sa/sa45.htm. Accessed May 8, 2003.
Cranshaw, W. S., and D. Leatherman. “Insect and Mite Galls.”
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259
260 TEMPERATE FORESTS

online. URL: www.ext.colostate.edu/pubs/insect/05557.html.

Accessed October 5, 2004.
Forschungsstelle für Paläeobotanik. “A History of Palaeozoic
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uni-muenster.de/GeoPalaeontologie/Palaeo/Palbot/ewald0.htm.
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Gates, Matthew. “Hemidactylium scutatum: Four-Toed Salamander.”
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ummz.umich.edu/accounts/hemidactylium/h._scutatum$
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cessed May 8, 2003.
———. “The General Sherman Tree.” Available online. URL:
www.nps.gov/seki/shrm_pic.htm. Accessed May 8, 2003.
Swiecki, Tedmund J., and Elizabeth Bernhardt. “A Delicate
Balance: Impacts of Diseases and Insects on the Health of
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publications/Oakdiseaseinsect.htm. Updated December 20,
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———. “Air Pollution and Air Quality.” GEO: Global Environment
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U.S. Geological Survey. “Major Tectonic Plates of the World.” Eastern
Publications Group Web Team. Available online. URL: geology.er.
usgs.gov/eastern/plates.html. Last updated June 27, 2001.
Viau, Elizabeth Anne. “Temperate Rain Forests.” Available online.
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temp_rain/temprain.html. Accessed May 8, 2003.
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palomar.edu/trmar98c.htm. Accessed May 15, 2003.
 INDEX
Note: Italic page numbers refer to illustrations.

A temperature of 42, 219 asexual reproduction 85

warm 45 ash 191
absolute humidity 56
air frost 66 aspen trees 85, 122, 124, 191
absolutely stable air 71
air mass 41–45 associations, of plants 98–99
absolutely unstable air 71
air pressure 45–46, 57 Athens, Greece 162
absolute pollen frequency
alder trees 147, 160 atmosphere. See also air
(APF) 159–160
algae 74, 75, 202–203 general circulation of 37,
acid mist 215
alternation of generations 86 38–39, 39
acid rain/acid precipitation
altocumulus clouds 61 atmospheric window 223
214–217 altostratus clouds 61 ATP 82–83, 84
acid snow 215 aluminum 176 automobile emissions 217
acorn woodpecker 93 ambrosia beetles 134 autumn 34
active dispersal 93 ambrosia fungus 134 auxins 117
adaptation, of trees ammonia 80–81 ax 157
to climate 115–118 amphibians 109 axils 94
to fire 122–123 ancient forests 14–16 azonal soils 28
adenosine diphosphate (ADP) angiosperms 7, 88, 89. See Azore Islands 53
82–83, 84 also flowering plants
adenosine triphosphate (ATP) animals 148–153
82–83, 84 attacking of trees by B
adiabatic cooling 58 130–132 “Babes in the Wood” 168
adiabatic warming 58 forest fires and 123 bark 95, 96, 97
ADP 82–83, 84 nutrition in 108 stripping of 131
adventitious shoots 85 plantation forestry and basswood 191
agriculture. See farming 201 Bauhin, Caspar 140
air and seed dispersal 93–94 bears 131, 148–149
boundary layer of 67, 219 in trophic system 104 “Beauty and the Beast”
and cloud formation anther 91, 91 167–168
56–61 anticyclones 51–52 beavers 131, 150–151, 151
cold 45 APF. See absolute pollen fre- bedrock 31
flow of 46 quency beech bark disease 134
general circulation of 37, aphids 134 beech scale insects 205
38–39, 39 aquifer 32 beech trees 1–2, 145, 146
maritime 52 Argyll Forest 212 in ancient forests 15
rising 56–60 Aristotle 4 diseases of 134
stability of 69, 70–71 artist’s fungus 134–135 timber from 146, 190

261
262 TEMPERATE FORESTS

beetles 133, 134, 160–161 British Isles 172 circulation

Belarus, forests of 98–99 broad-leaved trees, v. conifers general 37, 38–39, 39
Belovezhskaya Pushcha 98, 6, 87–89 three-cell model of 39
132 brown bear 148–149 cirrocumulus clouds 61
Bergen School 43 browsers 131 cirrostratus clouds 61
Bergeron, Tor 43 buffering 215 cirrus clouds 61
Betulaceae 145 cities, habitat of 112
Bia lowieża
⁄ Forest 98, 132 classification
binomial system of classifica-
C binomial system of 141
tion 141 Cadiz, Wisconsin, deforesta- of climate 54–56
biodiversity 138–153 tion in 163–164 of clouds 60–61
coppicing and 196 calyx 90 of soil 28, 29
definition of 138–141 cambium 94, 95, 96 climate 34–73
importance of 142–144 Canada adaptation to 115–118
Biodiversity Convention deforestation in 163–167 air masses and 41–45
138–139, 143, 144 forestry in 174 and ancient forests 14–16
biological pest control Cancer, tropic of 4, 5, 38 blocking and 49–52, 50
206–207 Canellaceae 24, 25 classification of 54–56
biomass 106 canopy 124–125 and clouds 56–61
biome xviii, 101 capillarity 31–32 continental 52–54
birch trees 145, 147, 160 capillary fringe 31 depressions and 45–49
bark of 97 Capricorn, tropic of 4, 5, 38 forest clearance and
fire and 122 carbon dating 158 217–220
seeds of 91–92, 122 carbon dioxide, and green- and forest formation
birds, parenting 99 house effect 221–222, 223 xv–xviii
Bismarck, Otto von 171 Caricaceae 24, 25 fronts and 43, 44, 44–45
bison 131–132 carnivores 104–105, 109 during ice ages 10–14
Bjerknes, Jakob 43 carotenoids 82 jet streams and 45–49
Bjerknes, Vilhelm 42, 43 carpel 90 maritime 1, 52–54
black bear 131, 149 carpellate flowers 91 oceans and 37–41
black frost 66 carrying capacity 113–114 of past 17–18
black water 40 catalytic converters 217 and plantation forestry
blanket weed 75 caterpillars 135 200–201
blocking, and climate 49–52, cattle 156 and soil 27–29
50 cells, in reproduction 85–86 and thunderstorms
blocking highs 52 cellulose 83, 111 69–72
blocking lows 52 Channel Islands 172 climax 118–120
boar 132, 149–150 charcoal 121, 181, 182, 183 closed canopy 124
bolling 195 chestnut blight 133 clouds 69, 220
bootlace fungus 135 chestnut trees 132–133, classification of 60–61
boreal forest xvi, 6, 224 146–147 droplets of 61–62
boundaries, of forests 6 “Children in the Wood, The” formation of 56–61
boundary layer, of air 67, 168 and thunderstorms
219 chipboard 186, 187 69–71
boxwood 191 chipmunks 151–152, 152 coal 177
bracts 90 chlorophyll 80–82, 117 Coast Ranges 53
brash 227 chloroplasts 82–83 codominants 125–126, 126
Britain 169–171, 172 chromium 176 cold air 45
British geography 172, 173 chromosomes 85–86 cold fronts 44, 44–45
 INDEX 263

cold lightning 70 cuticle 75 dominant species 129–130

combustion, primary 180 cutin 111 dormancy 115
community 100 cyanobacteria 202–203 drainage 32, 32–33
of forests 127–130, 128 drey 114
succession in 118–119 drip line 77
Community Forests 211, 212
D dry adiabatic lapse rate
companion cells 89 DALR. See dry adiabatic lapse (DALR) 70, 71
complete flowers 91 rate dry deposition 215
condensation 3, 57, 66, 69, dams, beaver 150, 151 dust, forest clearance and
70 Darwin, Charles 100 219
conditionally unstable air 69, daughter cells 85 Dutch elm disease 133–134
71 deciduous forests 1 dwarf shrub layer 126
coniferous forests fire in 72, 121
fire in 72, 121 global warming and 224
structure of 124–125 structure of 124–125 E
coniferous trees xvi, xviii deciduous trees xvi–xvii, Earth
growth conditions for 6 xviii, 1 mantle of 21
shape of 124 growth conditions for 6 rotation of
v. flowering plants leaves of 116–117 and climate 35, 35–37
87–89 decomposers 110–112 and jet stream 51
Conrad index 54, 54 decomposition 103, 110–112, Earth as Modified by Human
consumers 104, 108 225 Action: Man and Nature, The
continental climate 52–54 deer 130, 130–131, 152–153 (Marsh) 165
continental drift 19–23, 20 deer mouse 102 Earth Summit 143, 144
convective cloud 60 deforestation 212–213, 213 eastern boundary currents
Convention on Biological and climate 217–220 41
Diversity 138–139, 143, history of 154–157 ebony 188
144 in North America ecliptic, plane of 35
Convention on Climate 163–167 ecological efficiency 109
Change 143 and soil erosion 225–227 ecological energetics 109
cooling, adiabatic 58 demolition sites 118–120 ecological pyramids 105–106,
copper 176, 177, 181 denitrification 80–81 106, 111
coppicing 194–196 depressions 48, 49, 51–52 ecology 98–137
Coriolis effect (CorF) 51 derivatives 95 economics, of farming 210
cork 95, 96, 146 destructive distillation 181, ecosystems 141
cork cambium 95, 96 182 edge effect 128, 128–129
corolla 91 detritivores 110 edge forests 128–129, 132
corolla tube 91 detritus 110, 111 egg cells 86, 87
cosmic radiation 158 dew 65–66, 219 electron-transport chain 82
cottonwood 191 dew-point temperature 66, elk 131
cotyledons 87 70 elm trees 133, 156, 160, 190
Cretaceous period 7–8 Dietz, Robert Sinclair 21 ELR. See environmental lapse
crown, of trees 124 digitalis 142 rate
crown fire 73 diploid cells 85 Elton, Charles Sutherland
crustose lichen 203 diseases, of trees 132–135 105
cumulonimbus clouds 61, 69, disjunct distribution 23–25, Eltonian pyramid 105–106,
71 35 106
cumulus clouds 60, 71 dispersal, of seeds 89–94 elytra 160–161
currents, ocean 37–41, 40 dominants 125–126, 126 embryo 87
264 TEMPERATE FORESTS

emergents 125 Ferrel, William 38 frontal zone 44

energy, flow through forest Ferrel cells 38, 39 frost 66, 219
106–110 fertilization 87 fruit 86, 93–94, 111
energy density, of wood 180 fiber 88, 96 fruiting body 202, 203
England 172, 173 fiberboard 186–187 fruticose lichen 203
Community Forests in field layer 127 fuel
211, 212 filament 91, 91 fossil fuel 176, 178
English black walnuts 86 filbert nuts 148 wood as 178–181, 179
English yew 18 fire. See forest fire fungus 132–135, 202–203,
environmental lapse rate firebreaks 200 205
(ELR) 70–71 fire climax 123
ephemeral habitats 114 firestorm 73
epizootic disease 103 flagstone 175
G
Equatorial Currents 37 flint 157 gales 69–72
equinoxes 36 florets 90 galls 204
essential elements 79 flowering plants gametes 86, 90
ethylene 117 fruit of 93 gametophytes 86–87
Eurasian jay 93 trees as 115–116 gases, secondary 180–181
Europe v. coniferous trees 87–89 General Sherman tree 2–3
changes in forests of flowers 7, 7–8, 87–89, 90–91, Generelle Morphologie der
155–157 91 Organismen (Haeckel) 100
plantation farms of 174 fog 219 genetic diversity 139
European sycamore 9 foliose lichen 203 genetic engineering 144
evaporation 66–68, 220 folklore 167–169 genetic viability 138
evapotranspiration 68 food, prices of 210–211 Geneva Convention on Long-
Evelyn, John 169–170 food chain 101–103, 104 Range Transboundary
evergreen trees xviii food web 103, 105 Pollution 216
exine, of pollen grain 157 forb 125 geography 1–18
forb layer 126–127 geology 19–33
forest, definition of xvi, 228 Germany
F forest fire 71, 72–73, acid rain in 214, 216
Fagaceae 145 120–123, 200 forestry in 171
fallstreaks 62 forest management, sustain- germination 77, 90–92
false tinder fungus 134–135 able 227–229 girdling 193–194
farming xviii forestry, history of 169–174 glacials 10–11, 13
in ancient Greece 161 Forestry Commission 171, Gladstone, William Ewart
in ancient Rome 199 171
162–163 forests, natural versus planta- global warming 220–224
and deforestation tion 135–137 glucose 83, 84
166–167 forked lightning 70 GPP. See gross primary pro-
economics of 210 fossil fuels 176, 178 ductivity
in-field, our-field system four-toed salamanders 15 grain, of wood 94
of 197 foxes 108 gray squirrel 114
and park woodland foxglove 142 Great Britain 172, 173
197–199 frigid zone 4 great maple 9
and soil erosion 227 front(s) 43, 44, 44–45 Greece, ancient, forests of
swidden 155–156 frontal cloud 60 161–162
felted beech coccus 134 frontal depressions 48, 49 greenhouse effect 221,
Fennoscandian ice sheet 14 frontal theory 43 222–223
 INDEX 265

greenhouse gases 221, 223 humidity 59, 220 L

green wood 179 hydrogen bond 67
lakes, acid rain and 214
grizzly bear 149 hyphae 202
land, specific heat capacity of
gross primary productivity
41
(GPP) 109
ground fire 72–73
I landslide 226
ice 62, 63 lapse rates 70–71
ground frost 66
ice age 10–14, 12 latent heat 67, 70
ground layer 127
ice storms 63–65 Laurentide ice sheet 10, 11,
groundwater 32–33, 225
imperfect flowers 91 12–13
growth
incomplete flowers 91 leaves
primary 94
initials 94–95 of broad-leaved trees
secondary 94
insecticides 206 87–89
gullies 226
insects colors of xvii, xvii, 34
gymnosperms 76, 88–89
and pollination 8 of deciduous trees
gypsy moths 204
and tree diseases 116–117
gyres 40, 40–41
204–207 decomposition of
interglacials 10–12, 13 111–112
H Intertropical Convergence loss of 117
habitats 112–114 Zone (ITCZ) 38, 39 new 79–80
Hadley, George 38 intrazonal soils 28 Leipzig School 43
Hadley cells 38, 39 Introduction to Plant Ecology lichens 202–203
Haeckel, Ernst Heinrich 100 (Tansley) 101 lifting condensation 3, 70
hail 63 Ireland 172, 173 light-dependent stage, of pho-
“Hansel and Gretel” 167, iron 181 tosynthesis 82–83
168 Isle of Man 172 light-independent stage, of
haploid cells 86 isobars 46–47 photosynthesis 83
hares 131 ivy 17 lightning 69–71, 80
harvesting, of plantation lignification 76
forests 201–202 lignin 76, 88, 96, 111
hazels 147, 148 J Lincoln, Abraham 165
heartwood 96, 96 jet stream 47–52, 48, 49, 50 Lindeman, Raymond L.
heat, latent 67, 70 jungle 129 108–109
heat capacity, specific 41 Linnaeus, Carolus 140–141
herb(s) 125 litter, fire of 120, 121
herbivores 104, 109 K “Little Red Riding Hood”
herb layer 127 Kalendarium Hortense (Evelyn) 167
hickory 191 170 lodgepole pines 122, 123
highs 51–52 kelp 74–75 logging 135–137, 181–185
hoar frost 66 keys 92, 92 logistic equation 113–114
holdfasts 75 Köppen, Wladimir Peter Loudon, John Claudius 133
holly 17 55–56 lows 51–52
Holocene 10–11, 13 Köppen classification system lumber 182, 201–202
honey fungus 3, 135 55–56 lumber-core plywood 189
hoof fungus 134 krummholz 4
horizons, soil 26, 27, 28 K-selection 113
hornbeam 147–148, 191 K-species 113 M
horse latitudes 38, 39 Kuroshio Current 40 macronutrients 79
hot lightning 70–71 Kyoto Protocol 224 madrone 178–179
266 TEMPERATE FORESTS

magnolia 8, 9 N oceanic climate 52–54

Magnoliaceae 24, 25 oceans
NADP 83
MAI. See mean annual incre- and climate 37–41
NCP. See net community pro-
ment currents of 37–41, 40
ductivity
management of forests old-growth forests 9, 205, 229
nectaries 90, 91
193–209 olive trees 197
net community productivity
modern 207–209 On the Origin of Species by
(NCP) 109
sustainable 227–229 net primary productivity Means of Natural Selection
traditional 193–196 (NPP) 109 (Darwin) 100
Man and Nature (Marsh) 164, nexine, of pollen grain 157 open canopy 124, 125
165, 166 niches 114 orchards 197
mantle, of Earth 21 nicotinamide adenine dinu- orographic cloud 60
maple 191–192 cleotide phosphate (NADP) Our Common Future 228
marginal land 207 83 ovary 90, 91, 91
maritime air 52 nimbostratus clouds 61 ovules 86, 90, 91
maritime climate 1, 52–54 nitric oxide 80 owls 102, 103, 129
Marsh, George Perkins nitrification 80
164–166, 227 nitrogen 215–216
mass action 215 P
nitrogen cycle 80–81, 81
McKenzie, Dan 21 nitrogen dioxide 214–215 paleoclimatologists 17–18
mean annual increment nitrogen fixation 80 Pangaea 21
(MAI) 200 nitrogen oxide 214–215, 217 paper 187
medicinal plants 142–144 nonrenewable resources paper birch 97
meiosis 86 175–178 paperboard 187–188
meristem cells 94–95 North America pappus 118
metals 176 forest loss in 163–167 parasites 132–135
meteorology 43 old-growth forests of 9 parenchyma 89, 94
micronutrients 79 refugia of 15, 16 parks 9
minerals 176 temperate rain forests of park woodland 196–199
mining 176 2 particle board 186
mistletoe 17–18 Northern Hemisphere, forests pedologists 26
mitochondria 84 of 5–6 pedology 29
mitosis 85 NPP. See net primary produc- peduncle 90, 91
mixing ratio 59 tivity perianth 91
moose 131 numbers, pyramids of 105 permafrost xv
morphology 100 nutrients, cycling of 110–112 permeability, of soil 31, 31
moths 135 nutrition, of trees 77–79 pest control 203–207
mountains petals 91, 91
forests of 4, 16 petiole 117
rainfall on 3 O pheromones, in biological
Mount Tom 165–166 oak leaf roller moth 135 pest control 206
mouse 101–102, 102, oak trees 74, 145–146, 160 phloem 88–89, 94, 95, 96
103 burning for fuel 178–179 phosphorylation 84
mudflow 226 charcoal from 182 photosynthesis 79–84,
mudslides 226 life span of 203–204 106–108
mutualism 202–203 lumber from 190 in lichens 202–203
mycelium 202 oarweed 75 pigments in 80–83
mycorrhizae 79 occlusion 45 in trees 115
 INDEX 267

phytosociology 101 predators 105, 206 respiration 84, 107–108

picture fungus 134–135 pressure gradient 46–47 RH. See relative humidity
pigs 156 pressure surface 47 rhizoids 76
pine trees 92, 122, 123, 160 prevailing winds 37 rhizomorphs 135
pioneer species 118, 119 primary combustion 180 ribulose biphosphate (RuBP)
pith rays 94 primary consumers 104, 108 83
pits 88 primary growth 94 ridges, of jet stream 48, 48
plane of the ecliptic 35 producers 104 rills 226
plant(s) pyramids, ecological ringbarking 95, 193–194
associations of 98–99 105–106, 106, 111 rings, of wood 94, 97
evolution of 74–76 of biomass 106 Rio Summit 143
in food chain 102–103 of energy 108–110 roe deer 131, 153
land versus water 74–76 of numbers 105 Rome, ancient, forests of
in trophic system 104 pyroclimax 123 162–163
vascular 75–76, 88 root(s) 77–79, 78
water uptake in 68, 88 root cap 78
woody 76 R root hairs 78
Plantaceae 24–25, 25 radicle 77, 87 root plate 77
plantation forests 174, radiocarbon dating 158 roundwood 181–185, 185
199–202, 213, 229 raindrops 62, 63, 64–65 RPF. See relative pollen fre-
management of 207 rainfall quency
versus natural forests evaporation of 220 r-selection 113
135–137 global warming and r-species 113
plates 22, 23 223–224 RuBP 83
plate tectonics 19–23, 20, 22 on islands 53 Russian folktales 169
Pleistocene 11, 13 on mountains 3
plies, of plywood 189–190 movement through forest
Pliocene 13 29–33, 77, 225–226 S
plumule 87 rain forests, temperate 1–2 salamanders 15
plywood 189–190 rain shadow 53–54 SALR. See saturated adiabatic
Poland 98–99 ravens 99 lapse rate
polar cells 38, 39, 39 receptacle 90, 91 samara 92, 92
polar front 39 recycling 187–188 sandwich board 189
pollarding 195 red deer 152–153 sap 95
pollen 157–160, 159, 161 redwood trees 2–3, 122–123 sapwood 95, 96, 195
pollen diagram 159, 159 refugia 12, 15–16, 16 saturated adiabatic lapse rate
pollen grains 86–87, 91 relative humidity (RH) 3, 56, (SALR) 70, 71
pollination 8, 89 65–66, 68 savage 169
pollution 214–217 relative pollen frequency scarification 227
Pomona (Evelyn) 170 (RPF) 159 Scotland 172, 173, 212
poplar 122, 191 relicts 15 seafloor spreading 21
pores, of soil 30 renewable resources 175–178 seasons xvi–xvii, 34, 115–117
porosity, of soil 30–31, 31 reproduction, of trees 85–87 reasons for 34–37
potash 163 reptiles 109 and temperature 42
Practical Plant Ecology reserves, of nonrenewable and tree growth 96–97
(Tansley) 100–101 resources 176 seaweed 74–75
prairie 154 resources, renewable and secondary consumers 105,
precipitation 45 nonrenewable 175–178 108
268 TEMPERATE FORESTS

secondary gases 180–181 Southern Hemisphere, forests supercooled cloud droplets

secondary growth 94, 95 of 5–6 62
seed(s) 85, 87 species sustainability 227–229
dispersal of 89–94 diversity between swidden farming 155–156
in fruit 93–94 139–141 sycamore 192
germination of 77, diversity within 139 sycamore trees 9
90–92 dominant 129–130 Sylva, or a Discourse of Forest-
seed bank 91 K-species 113 Trees, and the Propagation of
seed coat 87 pioneer 118, 119 Timber in His Majesties
seed leaves 87 r-species 113 Dominions (Evelyn)
seedlings 92 Species Plantarum (Linnaeus) 170–171
sepals 90, 91 141 Systema Naturae (Linnaeus)
seral stage, of succession 118 specific heat capacity 41 140–141
sere 118 specific humidity 59
sexine 157 sperm 86, 87
sexual reproduction 85–87 sporophytes 86 T
shade 219–220 squirrels 93, 114, 151 taiga xvi, 6
sheet lightning 70 stability, of air 70–71 tall herb layer 126–127
shelter belt 217–218, 218 stadial 14 Tansley, Arthur 100–101
shipbuilding 190 stages, in succession 118 taproots 77
shoots, adventitious 85 stamens 90, 91 taxonomy 140–141
shrub(s) 76 staminate flowers 91 Taylor, Zachary 164
shrub layer 126, 126 standards 196 technology, and farming
sieve elements 88–89 starches 111 210
sieve plates 89 sterile-male technique, in bio- temperate forests xvii, 231
sieve tubes 89 logical pest control ancient 14–16
Silva (Evelyn) 170–171 206–207 animals of 148–153
silviculture 208 stigma 90, 91 area changes of 10–14,
skidders 202 Stockholm Conference 143 212–214, 213
slate 175 stomata 68, 75 community of 127–130,
snedding 202 stories, of forest 125 128
snowflakes 62, 63 stratocumulus clouds 60 defining xv–xviii
snowshoe hare 131 stratosphere 38, 46 development of 7–10
Snow White 168 stratus clouds 60 formation of xv–xviii
soil 25–29 style 90, 91 future of 230–232
capillarity of 31 subdominants 125–126, 126 history of xv–xviii, 7–10,
classification of 28, 29 suberin 95, 111 154–174
community of 110–112 subsidies, agricultural 210–211 human effects on 9–10
erosion of 225–227 succession 118–120 locations of 4–6, 5
horizons of 26, 27, 28 suckers 116 natural versus plantation
nutrients in 103 sugars 111 135–137
permeability of 31, 31 sulfur dioxide 214, 216–217 structure of 123–127,
porosity of 30–31, 31 summer 129 126
waterlogged 30 sunlight trees of 144–148
soil creep 226 and climate 38 types of 1–4
soil profile 26, 27, 28 penetration into forest uses of 175–192
solstices 35–36 117, 129 temperate rain forests 1–2
soredia 203 and seasons 35–37, 36 temperate zone 4–6, 5
 INDEX 269

temperature respiration in 84 vascular plants 75–76, 88

of air 42, 219 roots of 77–79, 78 vegetative reproduction 85
and clouds 57–59 seed dispersal by 89–94 veneer 188–190
and dew 65 seedlings of 92 Verhulst, Pierre-François 113
dew-point 66, 70 water uptake in 68, 79, vessel elements 88
forest clearance and 219 116 virga 62
global warming and wood of 94–97 volatiles 180
220–224 tree trunk 94–97, 96
of ground 42 trophic levels 104–105,
and ice storms 64 108–110
W
lapse rates of 70–71 trophic relationships 104 Wales 172, 173
range of 52–53 tropopause 38, 46 walnut 178–179
seasons and 36–37 troposphere 38, 46 warm air 45
of wood 180 troughs, of jet stream 48, 48, warm fronts 44, 44–45, 64
tepals 8 50, 51 warming, adiabatic 58
tertiary consumers 105, 108 tulip poplar 8 water
thermal wind 47 tulip tree 8, 9 and cloud formation
three-cell model, of atmos- tundra xvi 56–61
pheric circulation 39 turgor pressure 76 evaporation of 66–67
thunderstorms 69–72 movement through forest
thylakoid membranes 82 U 29–33
timber 181–185, 182 specific heat capacity of
UN Conference on 41
timberline 4
Environment and and transpiration 68
tin 176, 181
Development 143, 144 uptake in plants 68, 88
tinder fungus 134
UN Conference on the uptake in trees 68, 79,
top predators 105
Human Environment 143 116
torrid zone 4
understory 126 in wood 179
touchwood 134
UNEP. See United Nations waterlogged soil 30
tracheids 88
Environment Program water table 31
transpiration 68
United Kingdom 172, 173 weather stations 46
trees 144–148
forests in 212 Wegener, Alfred Lothar 21
adaptation to climate of
plantation forestry in 199 western boundary currents
115–118 United Nations (UN), World
attacks by animals on 41
Charter for Nature of 138 white-tailed deer 130–131,
130–132 United Nations Environment
coniferous v. flowering 153
Program (UNEP) 143
87–89 wild boar 132, 149–150
United States
diseases of 132–135 wildfire. See forest fire
deforestation in 163–167
disjunct distribution of willow 191
forestry in 171–174
23 wind
forests of 212
evolution of 74–76 directions of 47
refugia of 15, 16
forest fire and 120–123 and evaporation 67
unstable air 69
life span of 203–204 gale 71–72
nutrition of 77–79 and ocean currents 37
parasites of 132–135 V and plantation forestry
photosynthesis in 79–84 vascular bundles 94 201
reproduction of 85–87, vascular cambium 94–95, 95, for pollination 89
116 96 prevailing 37
270 TEMPERATE FORESTS

wind (continued) definition of 182 X

and seed dispersal 92 formation of 94–97
xanthophylls 82
shelter belts and as fuel 178–181, 179
xylem 88, 94, 95, 96, 97
217–218, 218 woodland 124, 196–199
thermal 47 woody plants 76
Windsor Forest 203–204 wooly aphids 134 Y
winged seeds 92, 92 World Charter for Nature, of Yellowstone National Park
winter 66, 79, 107, 115, 116 United Nations 138 122
winter moth 135 World Conservation Strategy:
wolves 108, 131 Living Resource Conservation
wood for Sustainable Development Z
composition of 228 zonal soils 28
179–180 World Heritage Site 98 zygote 87