(Biomes of The Earth) Peter D. Moore, Richard Garratt - Tundra (Biomes of The Earth) - Chelsea House Publications (2006)

BIOMES OF THE EARTH
TUNDRA
Peter D. Moore
Illustrations by
Richard Garratt
Tundra
Copyright © 2006 by Peter D. Moore
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Library of Congress Cataloging-in-Publication Data

Moore, Peter D.
Tundra / Peter D. Moore; illustrations by Richard Garratt.
p. cm. — (Biomes of the Earth)
Includes bibliographical references (p. ) and index.
ISBN 0-8160-5325-1
1. Tundra ecology—Juvenile literature. 2. Tundras—Juvenile
literature. I. Garratt, Richard, ill. II. Title. III. Series.
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Illustrations by Richard Garratt
Photo research by Elizabeth H. Oakes
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From Peter Moore:
To Eunice, Helen, and Caroline
From Richard Garratt:

To Chantal, who has lightened my darkness
CONTENTS
Preface ix
Acknowledgments xiii
Introduction xv
CHAPTER 1
GEOGRAPHY OF THE TUNDRA 1
Where is the tundra biome found? 1
What factors determine tundra distribution? 4
The Earth’s climate patterns 7
Climate in the polar tundra 14
Albedo 15
Climate in the alpine tundra 18
Lapse rate 19
Diversity of tundra landscapes 22
Arctic tundra vegetation 23
Alpine tundra vegetation 26
Conclusions 29
CHAPTER 2
GEOLOGY OF THE TUNDRA 31
Polar geology 31
Mountain geology 33
Plate tectonics 34
Ice accumulation 35
Ice sheets, ice caps, and glaciers 36
Effects of glaciation 40
After the ice 43
Patterns on the ground 46
Soil formation in the tundra 55
Soil types of the tundra 57
Soil change in time 62
The tundra atmosphere 64
Conclusions 67
CHAPTER 3
THE TUNDRA ECOSYSTEM 69
What is an ecosystem? 69
Tundra food webs and energy flow 72
Nutrient cycling in the tundra 76
Stability of the tundra ecosystem 81
Conclusions 82
CHAPTER 4
BIODIVERSITY OF THE TUNDRA 84
Living in the freezer 84
Tundra plants 86
Evergreen or deciduous? 88
Solar-tracking flowers 91
Microbes of the tundra 94
Lichens 97
Tundra invertebrates 98
Energy and tundra insects 102
Tundra birds 102
Putting on weight in the tundra 106
Polar tundra mammals 111
A tundra heavyweight, the musk ox 117
Alpine tundra mammals 121
Adaptations in tundra mammals 126
Conclusions 128
CHAPTER 5
HISTORY OF THE TUNDRA 130
Ice ages 130
Glacial history 133
Cycles of climate change 138
Causes of glaciation 142
The Milankovitch theory 144
Biological history of tundra 145
People in the tundra 148
The woolly mammoth 150
Tundra exploration 153
Conclusions 159
CHAPTER 6
USES OF THE TUNDRA 161
Hunting and trapping 161
Mineral reserves 163
Ecotourism and recreation 168
What is ecotourism? 169
Tundra as a carbon sink 172
Conclusions 173
CHAPTER 7
THE FUTURE OF THE TUNDRA 175
Consequences of climate change 175
Ozone holes 178
Pollution 180
The Vienna Convention and the Montreal Protocol 180
Tundra conservation 182
Bringing back the wolf 185
Conclusions 186
Glossary 189
Further reading 203
Web sites 205
Index 209
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. Sci-
entists 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 TUNDRA
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
I should like to record my gratitude to the editorial staff at
Chelsea House for their untiring support, assistance, and
encouragement during the preparation of this book. Frank K.
Darmstadt, executive editor, has been a constant source of
advice and information, and Dorothy Cummings, project
editor, has edited the text with unerring skill and impeccable
care. I am grateful to you both. I should also like to thank
Richard Garratt for his excellent illustrations and Elizabeth
Oakes for her perceptive selection of photographs. I have also
greatly appreciated the help and guidance of Mike Allaby, my
fellow Chelsea House author. Thanks to my wife, who has
displayed a remarkable degree of patience and support during
the writing of this book, together with much needed critical
appraisal, and to my daughters, Helen and Caroline, who
have supplied ideas and materials that have enriched the
text. I must also acknowledge the contribution of many
generations of students in the Life Sciences Department of
the University of London, King’s College, who have been a
constant source of stimulation and who will recall (I trust)
many of the ideas contained here. Thanks are also due to my
colleagues in teaching and research, especially those who
have accompanied me on field courses and research visits to
many parts of the world. Their work underlies the science
presented in this book.
XIII
INTRODUCTION
Few words conjure up such bleak images as the word tundra.
If asked to write down the ideas that come into their heads
on hearing the word, most people would probably include:
cold climate; unproductive, treeless, bare landscapes; and lit-
tle sign of life, with a total lack of human habitation. There is
a measure of truth in these ideas, but they are far from the
whole truth. In fact, summer days in the tundra can be
extremely warm, and for short periods the vegetation can be
very productive. Although low in diversity, the tundra is rich
in living organisms. Trees are present, but they tend to grow
only to knee height, and humans manage to make a living in
these regions despite the rigors of the tundra winter.
What is the tundra?

The word tundra is derived from the language of the Sami
(Lapps) in northern Finland, Sweden, and Norway, who
inhabit this open landscape. The tundra is a biome that is
associated with the high, polar latitudes, occupying land that
is farthest from the tropical, equatorial regions. But it is also
found on the summits of the world’s high mountains, which
experience a similar climate of low average annual tempera-
ture. Tundra vegetation is low and open, for the strong winds
and icy conditions of winter are too severe for the growth of
tall trees, whose buds would be blasted by airborne ice parti-
cles if they were to extend far above the surface of the
ground. Low shrubs and dwarf trees survive, however, form-
ing a bushy cover of vegetation lying close to the ground. In
some locations, especially on exposed, rocky ridges, even this
amount of vegetation cannot survive, and bare rocks or
gravel ridges are almost devoid of plants apart from a com-
pressed cover of scale-like lichens or small cushions of mosses
XV
XVI TUNDRA
that find shelter in the crevices. Within these patches, inver-

tebrate animals such as insects and nematode worms eke out
a living and serve as food for predators, particularly the birds
that arrive in the area to breed during the summer.
Below the surface of the ground, the life-bearing soil is
shallow because the deeper layers (the permafrost) are perma-
nently frozen, and neither the roots of plants nor the animals
of the soil can penetrate into this zone. The upper layers of
the soil also freeze during the winter but defrost and become
active with life during the summer season. This is another
reason why tall trees cannot survive in tundra: Trees need
deep roots to give them stability and an assured supply of
water, but due to the permafrost the available soil is too shal-
low for tree roots to flourish.
When temperatures are low and water is frozen, plant
roots are no longer able to obtain it from the soil, so water
supply can actually be a problem in the tundra, especially the
polar tundra where precipitation (snow and rainfall) is low.
The well-drained ridges in particular suffer because they may
also become very dry in summer and plants can find diffi-
culty in obtaining enough water to survive. In the lower-
lying areas between ridges, on the other hand, water
accumulates and creates wetlands that provide breeding
grounds for invertebrates and waterbirds during the summer
months. Often these wetlands develop patterns over their
surfaces that are very apparent from the air, giving the
appearance of marble or of honeycombs on the surface of the
ground.
The wildlife of polar tundra reaches its peak of activity in
the brief summer, when the vegetation is at its most produc-
tive. The resident mammals, which may include seals, polar
bears, arctic foxes, or, in the southern polar regions, pen-
guins, enter their breeding season. Other animals arrive from
the lower latitudes where they have spent the unfavorable
winter season, among them the caribou migrating northward
from the coniferous forest zone, and birds that come to take
advantage of the abundant supply of insect life and the long
days in order to feed themselves and their hungry offspring.
Away from the poles, on the summits of high mountains
even in the tropical zone, another tundra habitat can be
INTRODUCTION XVII
found. This is called alpine tundra. The word alpine is taken

from the Alps, the high mountains of central Europe, but
geographers and ecologists use it to refer to all high moun-
tain habitats. Alpine tundra has many similarities to the
polar tundra: It is treeless, annual average temperature is low,
and the more mobile animals may migrate into the zone dur-
ing the productive summer months. Many of the plants
found in alpine tundra are also found in the polar tundra,
which shows how similar conditions are. But there are also
some important differences. Near the equator (such as on the
mountains of East Africa, latitude 0°) there is very little dif-
ference between summer and winter conditions, but the con-
trast between the seasons becomes increasingly extreme the
farther the location lies from the equator. The difference
between summer and winter becomes greater moving north-
ward through the Himalayas (30°N), Sierras (40°N), Alps
(48°N), Scotland (60°N), and Brooks Range, Alaska (70°N).
The winters are increasingly long and cold when the moun-
tains lie closer to the poles. (These mountain ranges are
shown in the illustration on page 20.) It is not only in the
seasonal climate that differences in these montane tundra
habitats are found, however; there is also a contrast in the
daily range in the temperature of alpine tundra habitats. In
tropical mountains, and even in the temperate mountains
that lie closer to the equator, such as the southern Sierra
Nevada in California, there is a great difference between day-
time and nighttime temperatures; this is less marked as one
moves toward the poles. Alpine tundra regions also tend to
have higher precipitation (particularly snowfall) than the
polar tundra, and this often accumulates, leading to the for-
mation of packed ice masses, the glaciers. These climatic dif-
ferences as well as the isolation of alpine tundra, which has
led to evolutionary changes, mean that alpine and polar tun-
dra differ from one another in their physical conditions and
consequently in many of the plants and animals that occupy
them.
Tundra, therefore, is quite a widespread and varied habitat.
It has developed in response to extreme conditions, particu-
larly the impact of intense cold. It contains wildlife that is
not as diverse as that of the lowland Tropics but is of great
XVIII TUNDRA
biological interest simply because it is capable of withstand-

ing or avoiding the stresses to which it is exposed. Highly
adapted plants and animals are found here that have evolved
distinctive mechanisms for coping with harsh conditions by
modifying their structure, their biochemistry, or their behav-
ior. The tundra is, therefore, a living laboratory in which biol-
ogists and ecologists can study the way in which nature deals
with intense challenge.
Why is tundra important?

The most frequently used argument for preserving a habitat
is the value of its biological diversity, or biodiversity. In the
range of living organisms found within a biodiverse habitat
there resides a great bank of genetic information locked
within the cells of animals and plants, and the genes they
contain may be of importance to humans in the future.
Within these genes lie the keys to food production and drug
manufacture that may one day be vital to human survival. To
lose any plant or animal because of extinction is a mistake
that is irreversible and so is best avoided. It is relatively easy
to argue for the importance of the tropical rain forest because
of its immense biodiversity. But what of the tundra? It is a
general principle of biogeography that as one moves from
the equator to the poles, there is an overall reduction in bio-
diversity. The tundra is lower in biological diversity than any
other biome, even lower than the hot deserts. There are fewer
species of plants, insects, birds, and mammals in this biome
than in any other.
Does this mean that the tundra is of little importance in
the conservation of biodiversity? The answer is no. Tundra is
important because the organisms that are found in this
biome are entirely different from those of the Tropics. There
may not be as many species, but the species that occur are
not the same ones that are found in the rain forest. More-
over, they are also species that have developed under extreme
pressures, and this means that their structure and their bio-
chemistry are very distinctive and well adapted to these con-
ditions. The genes that control these traits could well be
useful to humans in the future, so these species also need to
INTRODUCTION XIX
be conserved. If humans need to improve crops, or domesti-

cated animals, or forest trees for use in cold climates, then
the genes contained in the tundra wildlife may hold the clue
to such developments. The mechanisms by which warm-
blooded animals can hibernate beneath the snow, or by
which plants can withstand the drought or the cold of a tun-
dra winter, may one day be vital to medicine and agriculture,
respectively.
Another reason why tundra is important is the fact that it
lies in remote locations. Tundra habitats are very useful as
monitoring stations of relatively undisturbed and unpol-
luted habitats, so they can be used to check the general state
of health of the world. The discovery in the 1950s of the
pesticide DDT in Antarctic penguins, for example, provided
an early warning of the way in which this harmful material
had found its way even into the most remote parts of the
planet. The shock generated by information of this sort
helped persuade governments to ban the use of this particu-
larly persistent toxin. Likewise, the discovery of a hole in
the ozone layer of the stratosphere (a layer of the Earth’s
atmosphere) over the Antarctic and, more recently, the Arc-
tic has emphasized the importance of the polar regions in
the functioning of the entire planet. The behavior of the
polar atmosphere needs more research, and the impact of
these processes on living plants and animals is best studied
in the tundra biomes.
Besides these quantifiable benefits provided by this biome,
people also gain much pleasure from the tundra in leisure
and recreational activities. Ecotourism that takes parties into
the high Arctic and Antarctic is now both popular and prof-
itable. Parties of tourists find relaxation, education, and
pleasure in watching the calving of icebergs in Alaska, the
breeding flocks of emperor penguins in the Antarctic, or the
wandering polar bears of Hudson Bay. Many other people
who will never be able to visit these regions of the world
enjoy the images and information about them now available
from books, magazines, and television. Tourism is also associ-
ated with the alpine tundra habitats of the world. Besides the
seashore, mountains are for many the most attractive envi-
ronment for rest and relaxation, but the pressures that
XX TUNDRA
humans place upon this fragile habitat can be harmful to its

wildlife, so care is needed in the development of tourism.
If humans are to take care of the tundra, then they must be
able to understand how it works, which is the subject of this
book. There are many questions that need to be answered in
order to understand the way the tundra functions, and these
will be considered in the course of the book. Where are the
tundra habitats found in the world, and what are the envi-
ronmental factors that control the distribution of this
biome? In lands that often lie beneath ice and snow, what is
the impact of glaciers upon the geology of the landscape?
What unusual features about the tundra ecosystem enable it
to function under climatic stress? How do plants and animals
manage to make a living in such a difficult environment?
What kinds of plants and animals survive here, and what are
the adaptations that make them successful? How long has
the tundra existed on Earth, and how has its extent varied in
the past?
The tundra has a history of human interaction, and people
will continue to use the tundra, exploiting its mineral wealth
and enjoying its grandeur. But the impact of past and present
human activities on the biome must be calculated, and
future human impact must be controlled. The tundra is a
unique habitat that deserves to be actively conserved, and
the final section of the book will explore how this may be
done. The future of tundra lies in human hands.
CHAPTER 1
GEOGRAPHY
OF THE TUNDRA
Geography is the study of the Earth and includes the way in
which landforms and living things are distributed over the
surface of the Earth, what factors have caused their distribu-
tion, and how the world came to contain the forms now
found here. Geography can teach people much about the
tundra biome, including the parts of the world in which it is
found and the reasons why it has this distribution pattern.
Where is the tundra biome found?

Tundra ecosystems are found only under very cold condi-
tions and these are most typically associated with the polar
regions. Perhaps surprisingly, neither the North Pole nor the
South Pole is occupied by tundra vegetation. At the North
Pole, the reason for this is that the pole lies far from solid
land in the middle of the Arctic Ocean. In winter the sea is
largely frozen so that the pole can be reached on foot, but the
pack ice becomes unstable in the summer and partially
breaks up. The lack of land in the immediate vicinity of the
North Pole means that there is no true tundra vegetation.
The South Pole, on the other hand, does lie on a large conti-
nental landmass, but the bulk of the continent is occupied by
a permanent ice cap, so once again tundra vegetation is
absent from the pole and its surroundings. The South Pole
actually lies two miles above the land surface, perched on top
of this massive thickness of ice. In both situations, therefore,
the tundra habitat forms a ring around the poles at some dis-
tance from them. Tundra is also present at other places on
the Earth’s surface where conditions are sufficiently cold, par-
ticularly on high mountaintops often far from the polar
regions. Here also, the very highest peaks are bare of any veg-
etation and the packed ice of glaciers may occupy the valleys
1
2 TUNDRA
North Pole
extent of tundra
The extent of the Arctic
tundra. South of this
line, forest becomes
increasingly abundant.
at high altitude. Below this ice zone lies the alpine tundra,
again forming a ring between ice and forest.
The map above shows the extent of the tundra region
around the North Pole. It occupies the coastal region of Alaska
and the Aleutian Islands and the northern parts of Canada,
sweeping south to include the coast of Hudson Bay, the north-
ern tip of Newfoundland, and the whole of Greenland and Ice-
land. It includes the northern coastal strip of Norway and the
northern edge of Russia, sweeping south again in the northern
Pacific to skirt the Bering Sea. In Antarctica, the great extent of
the ice cap, much of which is situated on land below sea level,
restricts tundra to isolated fragments along the coastal regions
and islands, especially at the tip of the Antarctic Peninsula.
Alpine tundra is, of course, more scattered over the face of
the Earth, wherever high mountains are to be found, from
the Arctic to the equator. In the Americas, alpine tundra is
found most abundantly in the mountain ranges of Alaska; in

the Rockies, Cascades, and Sierra Nevada in the west of North
America; and down the Andes chain of western South Amer-
ica. In Europe, the mountains of Scandinavia and Scotland
bear tundra vegetation, as do the Alps and Pyrenees. In Asia
there are high mountains, the Caucasus, to the west of the
Caspian Sea, but the continent is dominated by the massive
Himalayas, the great mountain plateau of Tibet, and the
ranges running eastward through Mongolia to the northeast-
ern regions of Russia. In Africa the Atlas Mountains lie on the
north and the Drakensberg Mountains in the far south;
between them are several very high mountains almost on the
equator in East Africa, and also in Ethiopia. Other isolated
high mountains are scattered around the world, as in Mexico
and Japan.
The polar tundra, by definition, occurs far from the equa-
tor in what are known as the high latitudes. The high lati-
tudes are given this name because the lines of latitude that
encircle the world are numbered from the equator, which has
AT L A N T I C O C E A N
Ronne Ice Shelf
Map of the Antarctic

South Pole continent, showing the
extent of the ice sheet
West Antarctica
PACIF
and the distribution of

bedrock above sea level
IC
beneath the ice. Some of

OC
N
EA
EA
the ice sheet lies directly

OC
N
Ross Ice Shelf

on bedrock below sea
AN
DI
level. The ice shelves

IN
bedrock above sea level

ice shelf
consist of floating
permanent sea ice.
4 TUNDRA
The distribution of the a latitude of 0 degrees. A cross section through the Earth
main mountain ranges would show that the North and South Poles are at right
of the world. The two angles to the line of the equator, so these have latitudes of
major ice sheets and the 90°N and 90°S, respectively. Between the equator and the
main regions of poles, the angles of the lines of latitude lie between 0° and
mountain glaciers are 90° with the higher values being closer to the poles; hence
also shown. the polar regions are referred to as the high latitudes and the
equatorial regions as the low latitudes. Mountains occur both
at high and low latitudes, as shown in the illustration on this
page and also in the illustration on page 20.
What factors determine tundra distribution?

Most of the world’s biomes, such as rain forest, desert, and
savanna grassland, occupy distinct zones around the world,
usually lying within a particular range of latitudes. The tun-
dra biome is evidently different, being found at high lati-

tudes but also scattered through the low latitudes. There
must be certain factors in common between the polar regions
and the high mountain habitats in locations closer to the
equator. All of the major biomes, including those listed previ-
ously, are ultimately controlled by climate, so it is reasonable
to begin an investigation into tundra distribution by looking
at climatic factors.
The most obvious feature that all areas of tundra, both
polar and alpine, share is their cold climate. High latitude
and high altitude both have low average annual tempera-
tures but, as will become clear, for different reasons. In gen-
eral all tundra habitats, whether polar or alpine, have average
annual temperatures below 25°F (–4°C). Low temperature is
important in maintaining the tundra biome because it
restricts tree invasion. When the overall temperature is above
this critical limit, tree seeds, such as birch, spruce, and pine,
are able to germinate and establish themselves. Forests of
coniferous and deciduous trees usually mark the edge of the
polar tundra that lies closest to the equator, while permanent
ice or ocean determines its poleward edge. In the case of
alpine tundra, forest similarly forms a lower altitudinal
boundary to the extent of the biome. So tundra exists
because the climate is too cold to permit forest growth.
The annual average temperature, however, is not an ade-
quate index of the general climate of tundra regions. A site
with warm summers and cold winters may have the same
average temperature as a location that is cool throughout the
year, so the seasonal variation in temperature needs to be
considered. Also important for vegetation is the length of the
growing season; a very short growing season, even if warm,
may be inadequate for forest growth and therefore may
encourage the development of tundra. But even tundra vege-
tation has its limits and needs some time clear of snow in
order to photosynthesize and build up a store of energy.
There are other factors apart from cold that play a part in
the maintenance of the tundra. Among these, high winds
are particularly important. The open expanses of the polar
tundra and the high peaks and ridges of the alpine tundra
are prone to high winds, sometimes bearing particles of ice
6 TUNDRA
suspended in the fast-moving air, and this can be extremely

damaging to vegetation if it projects far above the surface of
the soil. Shrubs are constantly trimmed by ice-blasting, and
taller trees are quickly eliminated.
The soils of the tundra, both polar and alpine, may be
frozen for much of the year or buried beneath the snow. The
germination of seeds is hindered by snow burial and the
development of roots is restricted by frost, so tree invasion is
prevented. The soils of the tundra regions will be considered
later (see “Soil formation in the tundra,” pages 55–57).
Water availability may not seem a problem in lands of ice
and snow, but the cold conditions can make it difficult to
obtain. Vegetation needs a reliable supply of water because
evaporation is constantly removing water from leaves, so
when cold makes water difficult to obtain, the plant suffers.
This is particularly true of plants with large leaves, including
many trees.
So, the two main climatic factors that influence plants and
animals are temperature and water. The graph below repre-
sents the occurrence of the tundra biome with respect to
these factors as compared with the other major biomes. Here
the annual precipitation is shown on the vertical axis and the
annual temperature on the horizontal axis. Biomes that need
high rainfall and high temperature, such as tropical forest,
160
average annual precipitation (in)
120
The relationships
between the major tropical
rain forest
biomes of the Earth and 80 temperate
forest
the climatic factors of
temperature and taiga
precipitation. The 40 grassland and savanna

tundra biome is found
only in regions of low desert tundra
temperature and low 0

60 50 40 30 0 -10
precipitation. average annual temperature (°F)
are therefore restricted to the upper left region of the dia-

gram. Grasslands occur over a wide range of temperature,
depending on whether they are tropical savannas or temper-
ate prairies, but need less rainfall than forests, so they lie
below these on the graph. Deserts can also occur in hot or
cold regions of the world, but characteristically they occupy
very dry climatic regions, so they lie at the base of the graph.
Tundra is found in the bottom right-hand corner. Its climate
is both cold and dry.
Climate is clearly the key to tundra distribution, therefore,
and in order to understand the pattern of tundra over the
surface of the Earth it is necessary to consider what controls
the world’s climate patterns.
The Earth’s climate patterns

The Earth receives most of its heat from sunlight that strikes
the land surface, warming it. The ocean surfaces are also
warmed in this way, but much of the energy that falls upon
them is reflected, so that the warming effect is generally less
than on land surfaces (see the sidebar “Albedo,” page 15).
The high latitudes (that is, the regions most distant from the
equator) receive less energy from the Sun than do the low
(tropical) latitudes. There are two main reasons why this is
the case. The first is related to the angle at which the Sun’s
rays strike the Earth, as shown in the illustration on page 8.
At the equator, the Sun is almost directly overhead for much
of the year, while in the high latitudes the same amount of
solar energy is spread over almost twice the land surface area
because of the low angle that is achieved. The second factor
that has to be considered is the thickness of atmosphere that
the Sun’s rays have to pass through to arrive at the Earth’s
surface. When the light comes vertically through the atmos-
phere (as at times in the Tropics), it passes through less
atmosphere than in the polar regions. The atmosphere dissi-
pates some light energy, especially if there is dust or impurity
present, so less energy reaches the ground. This again means
that the polar regions are starved of energy in relation to the
Tropics. Exactly how much energy is lost in this way depends
upon numerous factors, such as cloud cover, high-altitude
8 TUNDRA
solar energy
atmosphere
Sun
Incident solar energy on dust caused by volcanic eruptions, and the precise angle at
the surface of the Earth which the sunlight strikes the atmosphere. In general, how-
during a Northern ever, the equatorial regions receive about six times the
Hemisphere summer. amount of incident energy from the Sun per unit area when
Energy is spread over a compared with the polar regions.
greater area near the At the same time as energy is arriving at the surface of the
North and South Poles Earth, it is also being radiated from the Earth back into space,
than at the equator. The but the overall balance varies from one part of the planet to
sunlight also passes another. In the low latitudes, more energy is arriving than is
through a greater leaving, creating an energy surplus that becomes dispersed by
thickness of atmosphere movements of the atmosphere and the ocean currents. But in
at the poles, so more the high latitudes, especially those between 75°N or 75°S and
energy is absorbed the poles, more energy is being radiated than is received. So
or reflected. these polar regions are in an energy deficit. If there were no
mechanisms by which energy was redistributed around the
world, then one would expect the equatorial regions to be
25°F (14°C) warmer than they actually are, and the poles
would be 45°F (25°C) colder. As it is, the atmosphere and the
oceans are moderating the temperature differences between
equator and poles.
Seasonal changes also contribute to the uneven distribu-
tion of energy over the surface of the Earth. Seasonal varia-
tions in climate result from the fact that the Earth is tilted,
not upright, on its axis. This angle varies and is currently

23.5 degrees. The Earth orbits the Sun once a year and the
effect of the tilt is to create the different seasons (see illustra-
tion below). When the tilt of the Earth brings the North Pole
toward the Sun, the Northern Hemisphere experiences sum-
mer conditions. Conversely, when the South Pole is pointed
toward the Sun, the Northern Hemisphere is in winter. In the
Tropics, between 23.5°N and 23.5°S, the Sun is directly over-
head twice a year and is always high in the sky at noon. But
in the high latitudes, the seasonal variation is more severe.
Regions at latitudes higher than 66.5°N or 66.5°S of the equa-
tor (that is, 90 minus 23.5) will experience certain days in the
winter when the Sun does not rise above the horizon. In the
north of Ellesmere Island in northern Canada, for example, The tilt of the Earth on
the Sun rises above the horizon for only 143 days of the year. its axis results in
For the remaining 222 days of the year the area is in darkness seasonal variation in
or dim twilight. Conversely, during the summer period, there climate. In the Northern
are nights within these polar realms when the Sun never sets. Hemisphere, days are
The latitudes of 66.5° therefore mark important limits longer when the axis
around the poles and are called the Arctic and Antarctic Cir- tilts toward the Sun
cles, respectively. (summer) and shorter
The uneven heating of the Earth’s surface creates turbu- when the tilt is away
lence in the atmosphere. Light energy striking the land sur- from the Sun (winter).
spring
winter
(Northern Hemisphere
tilts away from Sun)
23.5 °
Sun
summer
(Northern Hemisphere
tilts toward Sun)
fall
10 TUNDRA
high
low polar front
North Pole
high
equator
low ITCZ
The circulation cells of

the atmosphere
high
redistribute energy
around the Earth.
Descending air produces
high-pressure areas in
South Pole
the subtropics and in
the polar regions, while low polar front
rising air results in low
high
pressure close to the
equator and around the ITCZ = Intertropical Convergence Zone
polar front.
face is converted to heat, and the air in contact with the

heated land becomes warmed. Warm air is less dense than
cold air, so it is forced upward from the hot equatorial regions
by cooler, denser air that moves toward the equator from
higher latitudes (see illustration above). This convergence of
air masses forcing up the heated air over the equator is called
the Intertropical Convergence Zone (ITCZ). Hot air is capable

of holding more water vapor than is cold air and the rising air
over the equator is heavily laden with water. But as it is forced
upward it cools and the water vapor condenses into clouds
and rain, so the equatorial region has a wet, rainy climate.
The constant upward movement of air means that low atmos-
pheric pressure predominates in the region. As more and
more air rises over the equator, these cooling air masses are
forced north and south, but because they are now colder they
are also denser and begin to fall. The descent of the dense air
takes place in the regions of 25° to 30° latitude north and
south of the equator and results in the development of two
belts of high atmospheric pressure around the world. As it
descends, the air comes into contact with land surfaces once
more, becomes heated by them, and takes up moisture from
the environment. The outcome is that the high-pressure belts
are arid and are the location of the world’s great deserts.
Some of the descending air moves back toward the equa-
tor and the Intertropical Convergence Zone, but some heads
toward the poles, taking some of the low-latitude heat
energy into the higher latitudes and assisting in the redistri-
bution of energy over the planet. When these poleward-
moving masses of air reach latitudes of 50° to 60° north and
south of the equator, they meet cold, dense polar air heading
toward the equator, and the warmer, lighter air masses are
forced upward, creating a low-pressure zone. This low-pressure
zone is called the polar front. Once again, the upward move-
ment of warm, moist air creates wet, climatically unstable
conditions. Some of the air pushed upward in this low-pres-
sure zone moves on to the poles, where it descends once
more, creating patches of high pressure with low precipita-
tion at the North and South Poles. Climatically, the polar
regions are cold deserts. The movement of air masses over
the surface of the Earth is further complicated by the spin of
the planet on its axis. The Earth is spinning from west to
east, so air moving toward the equator (including the cold
polar air) is deflected to the west. This can affect the position
of the polar front and set in motion swirling low-pressure
systems called depressions that track from west to east over
the temperate zone.
12 TUNDRA
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
Circulation patterns in Air masses thus assist in the dispersal of heat around the
the Earth’s oceans. world, ensuring that the overall temperature difference
Note that the warm between the equator and the poles is not as great as one
waters of the Gulf would expect from studies of solar energy input. Ocean cur-
Stream penetrate into rents also have an important role in global heat transfers (see
the Arctic Ocean from illustration above). In the eastern Atlantic, warm water from
the North Atlantic, but the Caribbean region, called the Gulf Stream, moves north
there is no such warm and east, becoming the North Atlantic Drift, which warms
movement in the North the western coast of Europe. Bergen, in Norway, is at the
Pacific. As a result same latitude as the southern tip of Greenland and of Baffin
northern Alaska is Island in Canada. But Bergen lies within the coniferous forest
much colder than (or taiga) biome, well to the south of the tundra, while
northern Scandinavia. Greenland and Baffin Island lie deep in the tundra biome.
(See also the illustration Warmed by the ocean currents of the east Atlantic region, the
on page 141.) climate of Bergen is warm enough to support forest growth.
There is also a warm North Pacific Drift, but this does not
penetrate past the Aleutian Islands because of the southward-
moving cold waters from the Arctic. It is deflected southeast-

ward toward Seattle and Vancouver, where it brings abun-
dant precipitation to the temperate rain forests of the Pacific
Northwest of North America.
The movement of warm water in the eastern Atlantic has a
strong impact, therefore, on the distribution of tundra in
northwestern Europe, pushing it northward by warming the
climate of the region. In the western Atlantic Ocean, how-
ever, cold polar water also flows south from the Arctic Ocean
between Greenland and eastern Canada, cooling these
regions and encouraging a southward development of tun-
dra. The North Pacific Ocean is also chilled by cold water cur-
rents from the Arctic Ocean moving southward through the
narrow straits between Alaska and eastern Russia, chilling the
Bering Sea and the Aleutian Islands.
Water loses its heat more slowly than landmasses do when
conditions become cool, so all areas of land that are close to
the sea are kept warmer as a result, even if there are no warm
currents present. Continental landmasses gain and lose heat
much more rapidly than the coastal regions, so that they
become hotter in summer and cooler in winter. This can
affect the distribution of vegetation. The boundary between
the tundra and the forest in Alaska and the Yukon Territories,
for example, lies at about 69°N, while in eastern Canada, in
the Hudson Bay region, the boundary lies much farther from
the pole, at close to 55°N. This is due in part to the very cold
winters in the continental interior of eastern North America
that hold back the northward extension of forest and favor
the tundra. The same applies to eastern Russia; the coldest
place on Earth is the north of Siberia.
When all of these different factors are considered, it
becomes clear that the polar regions of the world are going
to be cold. They receive less energy from the Sun, and the
influence of warm currents and warm air masses from the
equatorial regions is very limited. Where warm currents do
penetrate, as in the eastern Atlantic Ocean, the climate is
correspondingly warmer. The coldness of the landmasses is
also lessened to a certain degree when the ocean lies close
by; the coldest parts of the polar regions lie away from
oceanic influence.
14 TUNDRA
Climate in the polar tundra

At the South Pole lies the great continent of Antarctica, which
is covered by the world’s largest ice sheet. The influence of the
oceans is felt only along its fringes, so the interior is incapable
of supporting tundra vegetation. Almost the entire continent
lies within the Antarctic Circle, the exception being the
Antarctic Peninsula, which extends north toward the tip of
South America. Most of the landmass, in fact, lies within the
latitude 70°S. Surrounded by oceans and confined within
these high latitudes, these lands display no transition of vege-
tation from the polar ice to temperate forests, as is the case in
the Northern Hemisphere. The tundra of the Antarctic is
therefore isolated on the coastal fringe of the continent, the
Antarctic Peninsula, and some islands of the Southern Ocean.
In the Northern Hemisphere there is no ice sheet over the
North Pole because it lies within the Arctic Ocean. The lands
fringing that ocean and the North Atlantic and North Pacific
regions bear tundra vegetation, and at its southern edge this
tundra is bounded by the boreal forests of birch, pine, spruce,
and larch. This border forms a timberline, beyond which the
growth of tall trees is severely limited and only shrub growth is
possible. This timberline corresponds quite closely with the
average July temperature of 50°F (10°C). A line joining all the
places that share a particular temperature is called an
isotherm, and this average 50°F July isotherm correlates well
with the timberline. In other words, if the average temperature
in July is less than this value, healthy tree growth becomes
very difficult. This summer temperature factor, therefore,
seems to be the maximum tolerated by the tundra biome.
Low temperature within the tundra biome is partly deter-
mined by the nature of the ground cover. One inevitable fea-
ture of the tundra landscape, as a consequence of its
generally low temperature, is the presence of areas of snow
and ice. The occurrence of water in these solid forms results
in the loss of further energy, because the sunlight is reflected
off their white surfaces and is not absorbed by the ground
beneath. The degree to which a material reflects light is
termed its albedo (see sidebar on opposite page).
Snow and ice, therefore, have a high albedo, meaning that
they reflect much of the light that reaches their surfaces, while
dark rocks, organic soils, and many types of vegetation cover
Albedo
Albedo can be defined as the proportion of incoming solar radiation that is reflected from
a surface. Pale, shiny surfaces reflect much of the light that is incident upon them, while
dark, dull surfaces absorb most of the incoming energy. The albedo value is usually
expressed as a decimal, so that an albedo of 0.9 means that 90 percent of the incoming
radiation is reflected, while an albedo of 0.1 means that only 10 percent is reflected. The
reflected energy is measured either by an instrument located a few feet above the surface
or, on a larger scale, it can be measured from a low-flying aircraft. The following table
gives some albedo values for different kinds of surfaces.
Fresh snow 0.75–0.95

Old snow 0.45–0.70
Snow-covered vegetation 0.25–0.80
Open tundra 0.15–0.20
Coniferous forest 0.05–0.15
generally have a lower albedo. The heat absorption by the

darker areas and objects in a tundra landscape can create differ-
ences in the local climate, but the generally white surface of the
tundra helps to keep the region cold. This can operate even on
a very small scale in a microclimate rather than a climate. The
soil surface absorbs sunlight and becomes warmer, often by 1°F
(0.5°C), than the air temperature above. Even small differences
can be critical for living plants and other organisms. Dark
stones in the ground can act as heat absorbers, and thin layers
of gradually melting ice can behave like miniature green-
houses, raising the local temperature near the soil.
Taking into account the average temperature through the
entire year, tundra habitats are limited to regions with an
average below about 25°F (–4°C). But average temperatures
do not give a full impression of the climate of an area. The
temperature on a single day in July, usually the warmest
month of summer, can rise as high as 86°F (30°C), especially
in continental sites, such as eastern Canada or Siberia. But
the nighttime temperatures are low, so the average for the
month remains below 50°F (10°C). The length of the summer
season is also important for tundra plants and animals
16 TUNDRA
because they have to complete their reproductive cycles in a

limited period. In Churchill, Manitoba, for example, only dur-
ing the months of June, July, August, and September does the
mean temperature rise above freezing (see graph below). At
Isachsen, in the Northwest Territories of Canada, however,
July may be the only month in the year with an average tem-
perature above freezing. This places a much greater strain on
the organisms inhabiting the region. Both of these sites are
The annual cycle of examples of polar tundra, and in the graph their climates are
temperature in polar contrasted with that of alpine tundra at Niwot Ridge, Col-
and mountain sites, orado (see “Climate in the alpine tundra,” pages 18–22).
expressed as mean The overall climate of the Arctic tundra thus consists of long,
monthly temperature. extremely cold winters and short, cool summers. The seasonal
Niwot Ridge, Colorado, range of temperature is considerable, mainly as a result of the
is an alpine tundra absence of solar energy input in winter. The diurnal (daily)
location, while Churchill, range of temperature, however, tends to be relatively small as a
Manitoba, and consequence of the low Sun angle and the constant night in
Isachsen, Northwest winter and day in summer. The Antarctic, which has extensive
Territories, are polar ice sheets and high mountains, is generally colder than the Arc-
tundra sites. tic, which is one reason why it is so poor in flowering plant life.
°C °F
20 Niwot Ridge, Colorado (mountain – 12,000 ft) 68
Churchill, Manitoba (polar)
Isachsen, Northwest Territories (polar)
10 50
0 32
-10 14
-20 -4
-30 -22
-40 -40
Jan Feb Mar Apr May Jun Jul Aug Sept Oct Nov Dec
The polar tundra of the Northern Hemisphere is found in

a zone where climate varies very considerably, from the for-
est in the south to the ice in the north. Geographers have
found it convenient to divide this into two distinct zones,
the Low Arctic and the High Arctic. The term High Arctic is
usually used for the most northerly groups of islands
between the North Pole and Canada, Russia, Scandinavia,
and Greenland, lying north of latitude 75°N. The term Low
Arctic, on the other hand, is generally used for the more
southerly mainland areas of tundra. The growing season in
the Low Arctic can be between three and five months, but in
the High Arctic it is generally less than two and a half
months. In the Low Arctic, the mean July temperature is
around 46°F to 50°F (8°C to 10°C) and in the High Arctic is
only 38°F to 43°F (3°C to 6°C). The vegetation of these two
regions varies significantly as a consequence of the climatic
differences, as will be seen later (see “Arctic tundra vegeta-
tion,” pages 23–26).
The climate of the tundra is not only cold, it is also dry,
especially in the polar regions. The annual total precipita-
tion (including rain and snow) in polar tundra does not nor-
mally exceed 10 inches (25 cm). For this reason, the tundra
regions are often referred to as “cold deserts.” Although the
low temperature is generally very obvious to any visitor to
the tundra, the dryness of the climate might not be so appar-
ent. The ground is mainly wet in summer and rich in ice and
snow through the winter, but this is due to the very low lev-
els of evaporation. What precipitation falls generally fails to
evaporate back into the atmosphere; it either freezes,
remains in pools, or moves over the surface in streams and
rivers. The polar regions of the High Arctic have even less
precipitation than the surrounding ring of Low Arctic tun-
dra. True polar desert usually receives less than 4.3 inches
(11 cm) of precipitation (usually snow) in a year. This is
comparable to the rainfall in a low-latitude hot desert. For
instance, Death Valley in California generally receives 1.5
inches (4 cm) of rainfall a year and the average for the
Mojave Desert is nearer to eight inches (20 cm). So the polar
regions really can be very dry, and the use of the term polar
desert is entirely appropriate.
18 TUNDRA
Climate in the alpine tundra

The climate of alpine tundra is more complicated than that of
the polar tundra because so many factors affect the timberline
and therefore the altitude at which tundra vegetation begins.
Altitude itself is the dominant feature because the temperature
of the atmosphere generally decreases as one goes higher. The
rate at which temperature drops with altitude is called the lapse
rate and this varies with a range of atmospheric conditions
and geographical locations (see sidebar on opposite page).
Mountains in the high latitudes (closer to the poles) will
bear tundra habitats at lower altitudes than mountains in the
Tropics, because the temperature at the bottom of the moun-
tain will be lower to start with. Even in a single range of
mountains, it is found that the timberline is higher as one
approaches the equator. In the Sierra Nevada of California,
for example, the timberline is around 1,000 feet (300 m)
higher at the southern end than at the northern end around
300 miles (500 km) away. For every one degree of latitude
closer to the equator, the timberline rises by about 360 feet
(110 m). Mountains in the Tropics, therefore, need to be very
high in order to support tundra (or, to put it another way, to
fail to support forest). The illustration on page 20 shows the
relative heights and latitudes of some major mountain ranges
and indicates the altitudes at which the timberlines are to be
found. There is a marked increase in the altitude of the tim-
berline from higher latitudes (closer to the poles) to lower lat-
itudes (closer to the equator).
The average annual temperature in alpine tundra locations
is similar to that of the polar tundra sites. Detailed climate
records are available from many mountain sites, such as
Niwot Ridge in Colorado (see the graph on page 16). Here,
the mean July temperature is 46°F (8°C) at an altitude of
12,300 feet (3,750 m), but the average in December and Janu-
ary is only 9°F (–13°C). The seasonal variation in tempera-
ture, therefore, is less extreme than that of an Arctic site, such
as Churchill, Manitoba, as shown in the diagram. In alpine
tundra habitats, this seasonal variation in temperature depends
upon latitude. High-latitude mountains, close to or within the
Arctic Circle, have the same wide range of seasonal variation as
the Arctic tundra, but sites closer to the equator, such as the
Lapse rate
When a fixed amount of air (for convenience called a “package”) is in contact with a hot
land surface, it absorbs heat and it expands. This means that the molecules within the
package of air occupy more space and the air becomes less dense. When air is less dense
than the air above it, it rises, being displaced by the denser air. As it rises, the package of
warm air is subject to less and less atmospheric pressure because it has less weight of air
pressing down upon it from above, and the lower pressure means that it expands. The
molecules in expanding gas have to push other molecules out of their way, and doing so
slows down their movements and causes them to lose energy; in other words, the gas
cools. This is called adiabatic cooling; the word adiabatic means that there is no energy
exchange between the gas and its surroundings. Adiabatic cooling resulting from gas
expansion with altitude causes a general decline (or lapse) in air temperature with height
above the ground, and the rate at which this occurs is called the dry adiabatic lapse rate.
The adiabatic cooling of the air, however, means that it cannot hold as much water
vapor, because warm air has a higher capacity for water-holding than cold air. As the air
in the package cools, therefore, its water vapor condenses into droplets. A special prop-
erty of water comes into play here: its resistance to changes of physical state. Water occurs
in three physical states—solid (ice), liquid, and gas (water vapor)—and a change of state
requires a relatively large gain or loss of energy. When water changes state it either
absorbs or releases energy in the form of heat, called latent heat. During condensation
water releases some of this energy, warming the air. The actual observed lapse rate at any
location results from a combination of the two processes of dry adiabatic cooling and the
additional input of latent heat. This will vary depending on such factors as the humidity of
the atmosphere, but a general figure is approximately 3.6°F fall in temperature for every
gain of 1,000 feet in altitude (6.5°C fall for every 1,000 m).
high Ruwenzori Mountains of Uganda or Mount Kilimanjaro

in Kenya, have little variation in their overall climate through
the year. In alpine tundra, however, the daily variation in
temperature is often extreme. On Mount Kenya in East
Africa, for example, at 13,800 feet (4,200 m) the air tempera-
ture may rise to 60°F (15°C) in the day and fall to 23°F (–5°C)
in the night, and this can present considerable problems to
the plants inhabiting such a location. This degree variation
between night and day temperatures is not found in the
polar tundra, where the Sun hardly sets in the summer and
20 TUNDRA
altitude
(feet) Himalayas
30,000
25,000
20,000 Mt. Kenya,

East Africa
European
Alps
Sierra Nevada,
15,000 California
12,000
11,000
10,000 10,000
Brooks Range, Scotland

Alaska
5,000
3,000
2,600
0
80 70 60 50 40 30 20 10 0
latitude (°N) equator
A selection of Northern hardly rises during the polar winter. The range of tempera-
Hemisphere mountains ture between day and night is thus relatively small.
bearing alpine tundra. An additional climatic factor of importance in both alpine
The closer a mountain is and polar tundra is wind action. High wind speeds remove
to the equator, the the layer of air close to the ground, which is often the
higher the altitude at warmest air when vegetation is present because of the heat
which the tundra biome absorption by dark surfaces. If this warm layer is stripped
is found. away it leaves vegetation and soil cold. The presence of ice
crystals in the wind adds to its abrasive capacity and is partic-
ularly harmful to plants that emerge. For this reason, wind is
an important factor in determining the altitude of the tim-
berline on mountains. The direction of the prevailing winds
can often be detected from the pattern of the forest/tundra
border on mountains. In the Sierra Nevada of California, for
example, tundra vegetation can be found at lower elevations
on the western side of the mountains (facing the Pacific
Ocean) than on the east. On the west, the limit of subalpine
forest is about 10,500 feet (3,200 m), while on the protected
eastern side it is at about 11,800 feet (3,600 m). High winds
may completely eliminate trees and produce a more tundra-

like vegetation in lower latitudes than one would expect on
the basis of temperature, especially in oceanic regions (that is,
areas close to the oceans and strongly influenced by them).
Such oceanic, tundralike vegetation is found in Newfound-
land and northern Scotland, both in the North Atlantic, and
in Tierra del Fuego in South America. The microclimate of
mountains is made even more complicated by the varied
topography. The steepness and the aspect (the direction a
slope is facing, north or south) influence how much sunlight
reaches any particular location, so forest can extend to higher
altitude on south-facing slopes. As a result, timberlines are
often ragged and do not run neatly along the contours of
mountains. It is the height of the timberline that determines
how low down a mountain alpine tundra is able to develop.
Arctic tundra, as has been described, is effectively a cold
desert. The descending air masses at the poles bring little pre-
cipitation (see the illustration on page 10). In mountain sites,
on the other hand, air from lowland regions or from the
oceans is often pushed by winds and forced upward over the
mountain ranges. As air cools, its water-holding capacity is
reduced and its water content condenses as cloud; this may
then precipitate as rain droplets or, if the temperature is suffi-
ciently low, snow. The side of the mountain facing the pre-
vailing winds is subject to the highest levels of precipitation.
In the sheltered lee of the mountain, air descends once again
and precipitation is less intense. The lee side of the mountain
is said to lie in a rain-shadow, producing drier conditions.
In general, precipitation increases with altitude. In the
Rocky Mountains, to the west of Boulder, Colorado, for
example, at an altitude of 5,250 feet (1,603 m) the average
annual temperature is 48°F (8.8°C) and the annual precipita-
tion is 15.5 inches (395 mm). At altitude 8,460 feet (2,580
m) temperature is 42°F (5.6°C) and precipitation 21.3 inches
(540 mm), while at altitude 12,300 feet (3,750 m), tempera-
ture is 26°F (–3.3°C) and precipitation 25.2 inches (641 mm).
By comparison, the data for Fort Yukon, Alaska, at an alti-
tude of 416 feet (127 m), show an average annual tempera-
ture of 20°F (–6.7°C) and an annual precipitation of 6.8
inches (172 mm).
22 TUNDRA
One final factor of the physical environment of alpine

habitats that needs to be considered is the low air pressure
and consequently the low availability of oxygen. This is espe-
cially important to active animals, which need oxygen for
their respiration. The oxygen density at 18,000 feet (5,500 m)
is only half that found at sea level, so animals, including
human climbers, find strenuous activity particularly difficult
there. Most mammals, even wild sheep and ibex, are largely
limited to altitudes below 19,000 feet (5,800 m). Birds and
some cold-blooded invertebrate animals, however, are able to
exist even higher (see “Tundra invertebrates,” pages 98–102,
and “Tundra birds,” pages 102–111).
Diversity of tundra landscapes

The tundra is not a uniform biome; its general form, or land-
scape topography, is very variable. In the Arctic, high moun-
tains are found in some regions, such as the Brooks Range of
Alaska, which forms the southern limit of the tundra in that
state. Between the mountains of the Brooks Range and the
Arctic Ocean lies a plateau of foothills and then a strip of flat
coastal plain. The plain gives a generally uniform impression
of extensive flat lands but has numerous local landscape fea-
tures, including lakes, bogs, and patterned lands. Much of the
tundra of northern Canada is also relatively flat and it is
crossed by some major rivers, such as the Mackenzie River.
The landscape becomes more mountainous again in the east,
where the Torngat Mountains of Labrador rise to more than
6,500 feet (2,000 m). Scattered along the north Canadian
coastline are many islands, including some of the largest
islands in the world, such as Baffin Island. Farther east lies the
massive island of Greenland, whose interior (80 percent of the
total area) is covered by a permanent ice sheet. Along its east-
ern coast are high mountains, through which the ice spills as
a series of mountain glaciers. Out in the North Atlantic lie the
islands of Jan Mayen and Svalbard. Svalbard consists of a
group of five major islands and has rugged mountain topog-
raphy and a coastline of deep fjords. The remainder of the
Arctic tundra lies within northern Scandinavia and Russia.
Much of this vast area consists of an extensive flat plain, split
from north to south by the Ural Mountains, which continue
northward in the form of the islands of Novaya Zemlya. To

the west of this divide lies the Russian Plain, and to the east is
the Siberian Plain. These flat lands have much in common
with the coastal plain of northern Alaska. Despite this great
variety in landscape, all these habitats are components of the
tundra biome and all support tundra vegetation.
In the Antarctic, the world’s largest ice sheet covers most of
the continent, so the landscape is essentially ice-dominated
apart from some projecting mountain peaks. Beneath the ice
lies a landscape that is only just being discovered as tech-
niques of coring and mapping improve. One newly discov-
ered feature is the existence of lakes of great antiquity
beneath the ice, and these will undoubtedly prove of interest
to geologists, chemists, and microbiologists. The abundance
of ice, the very cold climate, and the lack of exposed land sur-
face means that true tundra vegetation is scarce and mainly
coastal. Patches of lichen and moss, together with small
quantities of grass, are able to survive in the limited patches
of land that are free of snow and ice.
As discussed previously, the climate of the polar tundra
changes as one approaches the poles, and the nature of the
landscape and vegetation reflects this. It is possible to divide
these tundra regions into different zones. One must remem-
ber, however, that strict lines of division cannot usually be
discerned; the landscape gradually changes with increasing
latitude. In the Southern Hemisphere, the isolation of the
Antarctic continent by oceans disrupts any zonation, so there
is no transition between tundra and forest as in the Arctic
regions.
Arctic tundra vegetation

In the Arctic, at the northern edge of the forest, where tree
growth becomes difficult because of the short summer season
and the intense cold and wind blasting, is a zone called forest
tundra. This represents the southern limit of the Low Arctic,
and the trees here are scattered and stunted. In North America,
black spruce and tamarack (on wetter areas) and white spruce
(on drier soils) often form the timberline. But nonconiferous
trees can also form an abundant component of the flora,
including balsam poplar, paper birch, and alder. Pines are
24 TUNDRA
Shrub tundra in the Low rarely found in the Arctic timberline of Alaska or Canada, but
Arctic. An autumn they are widespread in Europe and Asia, particularly the Scots
landscape in Denali pine (Pinus sylvestris). Isolated trees, sometimes contorted into
National Park, Alaska twisted shapes by the wind, may grow on favorable sites, or
(Photo by sometimes groups of trees survive in sheltered hollows, pro-
Michio Hoshino) tected from the wind. Beneath the trees is a dense layer of
shrubs and herbs. This zone is really a transition between the
boreal forest or taiga (the biome that forms a zone in the lati-
tudes below the tundra) and the true tundra. Ecologists refer to
such gradual boundaries as ecotones.
Beyond the zone of true tree growth, in the Low Arctic, the
rolling landscape is dominated by low woody plants, forming
the tall shrub tundra. The shrubs are characteristically less
than three feet (1 m) in height but may occasionally grow as
high as six feet (2 m). Some dwarf trees and shrubs are con-
tained within this vegetation, largely dwarf birches and wil-
lows, together with alder. Among these are other shrubs,

mainly evergreens, including Labrador tea and several species
of cranberry and crowberry.
In more exposed sites and in the more northern regions
of the Low Arctic, dwarf shrub tundra becomes the domi-
nant vegetation type. The taller species are no longer pres-
ent and the general vegetation canopy is only around one
to two feet (20 to 40 cm) above the ground surface. The
main components are still woody plants forming a dense
thicket, and these include bearberry and crowberry,
together with cotton grasses. In some places the cotton
grasses may assume dominance and the landscape becomes
what may be termed tussock-heath tundra. This type of tun-
dra is particularly well developed in Alaska, western
Canada, and eastern Siberia.
The main difference between the High Arctic and the Low
Arctic is that the vegetation in the High Arctic is composed
largely of herbaceous plants rather than woody ones. This is
called grass-moss tundra. Here, the intensity of cold, the very
short growing season, and the constant blasting of winds that
often carry ice crystals make growth very difficult for any plant
that extends far above ground level, so the characteristic
growth form is a cushion clinging closely to the soil surface.
The wind trims the tops of these cushions and prevents them
from growing higher, especially over the tops of exposed
ridges, where summer drought may also constrain growth. Just
how long the summer growing season lasts is dependent in
part on the local topography, because the sheltered hollows
often have patches of snow that last well into the early sum-
mer, and this restricts the time for growth and reproduction.
Areas of late-remaining snow are usually dominated by mosses
rather than the grasses, sedges, or other flowering plants.
Melting snow creates an abundance of water in the sum-
mer. At the same time, the rates of evaporation are low, so
most low-lying areas become wetlands despite the low rain-
fall. The action of frost often creates complicated patterns on
the ground surface, with ridges and hollows (see “Patterns on
the ground,” pages 46–53), and the hollows develop into
pools with sedge-dominated edges. These wet tundra habitats
of the High Arctic become valuable breeding areas for wading
birds and other wildfowl during the summer.
26 TUNDRA
Away from the wetland sites, however, the High Arctic

drought has a strong impact upon the vegetation and the land-
scape. Scattered grasses, mosses, and cushion plants form an
open vegetation known as the High Arctic semidesert. Lichens
and mosses often account for half of the vegetation cover, and
a range of drought-resistant tundra plants joins the commu-
nity, including saxifrages and mountain avens. Farther north,
more of the soil becomes visible as the vegetation thins. This is
the true polar desert in which only scattered plants grow. It
eventually meets the Arctic Ocean or the permanent ice.
Alpine tundra vegetation

Alpine tundra contains a range of vegetation zones very sim-
ilar to those of the Arctic, but they are arranged in relation to
altitude rather than latitude. The environment again
stretches from the timberline at its lower end to the bare
mountain peak or the ice of a glacier. Mountain timberlines,
rather like the Arctic timberlines, are often a ragged affair of
stunted and twisted individual trees that somehow manage
to survive after a chance germination in this harsh terrain. In
sheltered valleys, the trees may extend a little higher than in
the more exposed locations, and there is sometimes an eco-
tone of dwarf trees around the limits of their growth. Ger-
man botanists coined a very expressive word for this stunted
tree zone, calling it krummholz, literally “crooked wood.” The
zone is present, however, only where the natural transition
between forest and alpine tundra vegetation has developed.
Often people have interfered with this boundary, especially
in the more heavily populated mountainous regions of the
world, where the high tundra and grasslands have been used
for sheep and cattle grazing. In many mountain regions,
domestic animals have been taken to the high regions each
summer, grazing not only on the grasses and herbs but also
on the young trees. This additional stress has usually elimi-
nated the krummholz zone and in places even lowered the
timberline, creating a grassland/tundra transition zone.
There are some locations where the altitudinal limit to tree
growth may be determined not by climate nor by human
activity but by soil factors. In the southern Appalachian
Mountains and in Montana, grassy “balds” occur on the
summits of some mountains. These areas have shallow soils

and are exposed to intense drought in summer, so tree inva-
sion is not possible and instead a kind of grass subalpine veg-
etation develops. Generally, however, it is the cold climate
that limits the forest and leaves the habitat open to alpine
tundra vegetation. Timberlines are often lower on the north-
ern aspect of mountains, where the sunshine does not pen-
etrate well. In the Brooks Range of southern Alaska, for
example, woodland may extend to around 3,000 feet (1,000 m)
on the south-facing slopes while attaining only 2,000 feet
(700 m) on the northern side.
High mountains in the Tropics, such as Mount Kenya in East
Africa, display a series of vegetation zones at different heights.
The illustration on page 28 shows this zonation with altitude.
The mountain, which is situated close to the equator, has trop-
ical savanna and forest at its base. At higher altitudes these
types of tropical vegetation give way to a more temperate forest
of bamboo, which is succeeded by a shrub zone of heath
plants. Above 12,000 feet (3,600 m) the alpine tundra biome is
found, and above this altitude lies the permanent snow.
As in the case of the Arctic tundra, it is possible to recog-
nize different vegetation types in the alpine tundra. Shrub
tundra is found in the lower and less exposed regions, as seen
on Mount Kenya, and contains several of the plants of the
heath family (Ericaceae). Some of these are also found in the
Arctic, including crowberry and bearberry. Grass and cush-
ion-plant tundra is present in the more exposed, colder, and
drier sites, and this thins out to lichen and moss vegetation
as soils become thin and rock debris is the main constituent
of the soil. Within the alpine landscape, it is also possible to
distinguish wet meadows, dry meadows, and bare rocky
regions with limited vegetation cover, called fellfields. These
vegetation types reflect the increasing degree of drought
resulting from topographic position (hollows and ridges) and
the relative depth and drainage in the soil.
In both the Arctic and the alpine tundra landscapes, snow
plays an important part. Snowflakes are formed in the air
when the temperature falls below freezing and water droplets
become crystallized before settling. On the ground, they
begin to lose their crystal form over a few days, sometimes
melting and refreezing to form a more powdery or granular
28 TUNDRA
rock hyrax
alpine tundra
12,000 ft
tree-groundsel
leopard
giant heath
heath
9,800 ft
duiker
bamboo
7,500 ft
bamboo
tree fern
rain forest
5,400 ft
forest buffalo
savanna
elephant acacia tree
The major zones of material over the course of years. This is called firn. As new
vegetation found at snow accumulates from above, firn becomes compressed and
different altitudes on compacted into ice, and in this way, over the course of about
Mount Kenya in East 10 years, the snow becomes fully incorporated into an ice
Africa. This tropical mass, or glacier. But not all snow is converted to glacial ice;
mountain lies on the some simply melts with the coming of summer and exposes
equator yet has tundra once again the land that it buried.
vegetation near “Snow patches” are temporary accumulations of snow that
its summit. may survive over a few years but never form true glaciers.
Some melt each year and leave behind a wet patch of land-
scape, often in a sheltered hollow where the wind is not too

corrosive and the Sun has limited powers of penetration. The
snow has a number of effects upon the local climatic condi-
tions. Its presence late into the spring means that the ground
is protected from sunlight for longer; in other words, the snow
patch effectively prolongs winter for the inhabitants of the soil
it covers. But the snow also serves as a protective thermal blan-
ket from the worst of the winter cold, so soils beneath the
snow blanket remain warmer than those where snow is blown
away and the frost can penetrate. Both plants and animals are
affected by these two influences of a snow patch, and these
habitats support different flora and fauna as a consequence. It
is the property of thermal insulation in compacted snow, of
course, that forms the basis for its use in the construction of an
igloo. Here human beings are taking advantage of the very
effective insulation that snow provides, for they can heat the
inside of the igloo and raise the temperature below the snow
blocks considerably above the outside air temperature, and the
snow retains the hotter air. Many animals also use the snow in
a similar way, burrowing within it to keep warm.
The general persistence of snow varies in different regions
and in different microsites. Many tundra regions, such as the
Siberian tundra, are covered by snow for 200 to 280 days in
the year (compared with about 60 to 80 days in the Russian
steppes, for instance). Its depth is also variable, but the tun-
dra regions have low precipitation, so a snow depth of
around eight to 12 inches (20 to 30 cm) is fairly typical. In
alpine sites, on the other hand, precipitation can be high, so
snow patches can be deep. The local topography, including
slope, often limits the snow depth, and where deep snow can
accumulate and persist, glaciers begin to form. Local land-
form can also be important in the tundra plains, because the
wind will sweep snow from exposed ridges and raised areas
and allow it to collect in hollows.
Conclusions
Tundra is found in those parts of the Earth where conditions
are extremely cold in their average temperature but are still
able to support life. The high latitudes bear tundra because
30 TUNDRA
the amount of energy reaching these regions from the Sun is

much lower than is found in the equatorial regions. The tilt
of the Earth on its axis also produces great variations in sea-
sonal conditions with extremely long days in the summer
and very short days in winter.
The poles are regions of descending air masses, so they
receive low precipitation. The South Pole occurs on a great
landmass, Antarctica, and lies on the world’s greatest ice
sheet. The North Pole, on the other hand, occurs in the Arc-
tic Ocean, so although there is pack ice, there is no ice sheet
present. The landmass of Greenland does support an ice
sheet. Antarctica is surrounded by oceans, so ice-free land is
very limited and there is little tundra vegetation, but the Arc-
tic Ocean is surrounded by the continents of North America
and Eurasia, so much land is available for tundra vegetation,
which grades southward into the boreal forest biome.
Alpine tundra is found scattered around the world wherever
there are high mountains. Air temperature falls with altitude,
so if the mountain is high enough, tundra ecosystems can
develop even on tropical mountains. In the low latitudes, the
seasonal differences in temperature are less than those found
in the polar regions, but the differences between day and
night are often greater. Precipitation is generally higher in the
alpine tundra than in the polar tundra.
Tundra vegetation is similar in its basic structure in both
polar and alpine habitats. Tall trees are absent, but stunted
dwarf tree and shrub species can survive, generally extending
less than two or three feet (1 m) above the ground. Grasses,
sedges, mosses, and lichens cover the open patches of the
ground. This vegetation is capable of withstanding cold,
abrasive winds and periodic burial by snow. In the southern
parts of the Arctic tundra, and at the lower boundary of
alpine tundra, this vegetation type grades into stunted trees
of birch, pine, spruce, and larch.
Tundra thus occupies the cold extremity of the Earth’s var-
ied climate. Its plants, animals, and microbes have been
selected by nature to cope with the highly stressed condi-
tions they must endure. The extremes of cold also have their
impact on the rocks and the soils of the tundra region, as will
be discussed in the next chapter.
CHAPTER 2
GEOLOGY OF THE TUNDRA

Geology is the study of rocks, their formation and their
breakdown. Both processes are at work in the tundra regions.
In the oceans adjacent to the tundra lands, and in the lakes
that occupy some polar and alpine landscapes, eroded soils
and rock fragments are constantly accumulating as sedi-
ments. These build up and eventually form the sedimentary
rocks of the future. Volcanoes are active in some tundra
regions, both polar and alpine, bringing from deep below the
Earth’s crust the molten rock that cools and hardens on the
surface, forming new landscapes that will be invaded by tun-
dra vegetation. Erosive forces are also active in the tundra,
driven by low temperatures. Precipitation may accumulate as
ice. The weight of ice upon the surface of the Earth’s crust
may depress it, and as it slides over the ground the mass of
ice will grind down the rocks to the consistency of flour. This
“rock flour” is then eroded by streams and rivers, which form
from the melting ice and snow, and is redistributed over the
landscape. Among the rock detritus left by the activity of ice,
plants begin to grow and soils begin to form; the tundra
ecosystem begins to evolve. The tundra regions are thus areas
that are strongly affected by geological activity.
Polar geology
As explained in chapter 1, most of the polar tundra habitats
on Earth are found around the edge of the Arctic Ocean. The
rocks of this region lie in three massive sections, called
shields, which occupy the three continents that surround the
Arctic Ocean. These are the Canadian Shield, which includes
Greenland; the Fennoscandian Shield, covering northern
Scandinavia and Finland in Europe; and the Angara Shield,
extending through the northern edge of Asia.
31
32 TUNDRA
The shield rocks are extremely ancient, having been formed

in Precambrian times (more than 540 million years ago), when
some of the world’s oldest rocks were produced. The Earth is
believed to have formed about 4.6 billion years ago, and
Greenland contains some of the oldest rocks yet recorded,
with an age of 3.8 billion years. The Precambrian is sometimes
called the Cryptozoic eon (cryptozoic means “hidden life”). At
this time the only life on Earth was minute, so fossils are rare.
The Cryptozoic eon includes the bulk of the Earth’s history
(the first 4 billion years) and is followed by the Phanerozoic
eon, in which all visible forms of life evolved. The Cryptozoic
is divided into three eras: the Priscoan or Hadean era (about
4.6 billion to 4 billion years ago), the Archaean era (about 4
billion to 2.5 billion years ago), and the Proterozoic era (about
2.5 billion to 500 million years ago). It was during the
Archaean era that the rocks of the great shields were formed.
The Archaean era was a time of crustal development as the
Earth cooled. Afterward, during the Proterozoic era, the
Archaean rocks were altered by intense heating and pressure,
forming a compacted rock called gneiss (pronounced “nice”).
This is the rock that dominates the northern shields. Rocks
that have been melted, crushed, and transformed by these
processes are known as metamorphic rocks. It is likely that
these rocks once formed great mountains that have eroded
away over time, leaving a much more level landscape. The
rocks are generally hard and contain little in the way of plant
nutrients, so they give rise to acidic, nutrient-poor soils. But
the fact that the rocks are being ground to a pulp by ice
action (see “Effects of glaciation,” pages 40–43) does mean
that soils are frequently replenished.
Some parts of the Arctic have evidence of past volcanic
activity, such as Jan Mayen Island and Greenland. Indeed,
volcanic activity still takes place in the region, but mainly
under the sea. The island of Surtsey appeared overnight off
the coast of Iceland as a result of the eruption of an underwa-
ter volcano in 1963. Volcanic activity is associated with the
Mid-Atlantic Ridge, a seafloor line of volcanoes running right
up the Atlantic and extending into the Arctic Ocean.
The Antarctic continent is almost totally covered by the
Antarctic Ice Sheet, so the solid geology is visible only where
mountaintops protrude through the ice or in those coastal

regions where ice does not form a complete blanket. The
underlying geology of Antarctica consists of Precambrian
rocks, similar to those of the Arctic shields, but a series of
younger rocks are also present, particularly in West Antarc-
tica. The continent of Antarctica was at one time joined with
all the other major landmasses of the Earth in a single super-
continent, linking Africa, India, and Australia. Fossils in the
rocks show that the climate in the past was very different
from that of present-day Antarctica. In Permian times, about
290 million years ago, broad-leaved deciduous forests grew
close to what is now the South Pole, so the Earth must have
been much warmer. The wood of these ancient forests has
growth rings, showing that there were strong seasonal differ-
ences in climate even though the overall conditions must
have been warmer than at present. The discovery of fossils of
these Glossopteris trees by early explorers demonstrated the
continuity of vegetation between Antarctica and Australia,
where these fossils are also found, and led geologists to
believe that the continents were once part of a single land-
mass (see the sidebar “Plate tectonics,” page 34).
A major ridge of mountains runs from the eastern edge of
the Antarctic Ross Ice Shelf and arcs close to the South Pole
before extending along the line of the Antarctic Peninsula,
west of the Ronne Ice Shelf (see the map on page 3). These
mountains, called the Transantarctic Mountains, contain
sandstone rocks that bear fossils of fish from the Devonian
period and of dinosaurs from the Jurassic period. It was dur-
ing the Cretaceous period, around 140 million years ago, that
the great continent broke up. By 40 million years ago the last
land connections with South America and Australia had been
lost, and Antarctica was on its own.
Mountain geology
Tundra habitats are found not only in the polar regions but
also upon many of the world’s mountains, even in the low,
equatorial latitudes (see the map on page 4). The mountain
regions of the world are very varied in their geology and the
mountains may be formed from a whole range of rocks, both
34 TUNDRA
Plate tectonics
The relatively cool surface of the Earth, called the lithosphere or crust, is floating upon a
molten core. Within this fluid core, convection currents occur, in which hot, less dense
materials rise and cool, while dense materials descend. These currents cause the surface
crust to strain and shift. The crust is not a uniform layer, but is divided into a series of
plates, and the study of these plates and their movements is called plate tectonics.
In some regions, two plates are being pushed apart because hot material from the
mantle breaks through the crust, forming a zone of volcanic activity. The Mid-Atlantic
Ridge is one such region. It is a north-to-south-running line where rising hot and relatively
low-density molten rock from deep in the Earth rises to the surface, causing the crust to
split and move away from the line, both to the east and the west. So the seafloor of the
Atlantic is spreading very slowly, widening between two and four inches (5 to 10 cm) per
year; America and Europe are gently drifting apart. The distance sailed by Columbus in
1492 was approximately 150 feet (50 m) shorter than he would have to sail today.
Another case of crustal spreading caused the fragmentation of the landmass that once
contained Antarctica.
Crustal movements not only take continents away from one another; they can also
result in collisions. One such collision, of India with Asia, caused the line of contact to
buckle, thrusting up the Himalaya Mountain chain. When plates collide, one may slide
over the top of the other, forcing some of the crustal material down into the molten man-
tle, and the region where this occurs is called a subduction zone.
volcanic and sedimentary. The forms of their landscape and

their vegetation are often closely related to the nature of
their rocks, especially the way in which they become eroded
and the forms that they develop. The formation of moun-
tains depends upon the movements of the Earth’s crust (see
the sidebar above).
The Alps of Europe were created by pressure of the African
plate moving north into the Eurasian plate, beginning at the
same time as the Himalayan uplift, around 5 million years
ago. Since then the Alps have been forced upward about
6,500 feet (2,000 m) and the Himalayas about 10,000 feet
(3,000 m). The Sierra Nevada of California is a younger range
than either the Alps or the Himalayas. It was created within
the last 3 million years by the crushing and movement of
adjacent plates on the western side of North America. Within

that time the Sierra Nevada have risen about 6,500 feet
(2,000 m), and the area is still active, with relatively frequent
earthquakes. In 1872, near the town of Lone Pine, California,
a strong earthquake created cliffs of more than 20 feet (6.5 m)
in height, showing that uplift of the mountains is still taking
place. All mountains begin to be leveled as soon as they are
formed, however, so even the relatively young Sierra Nevada
have much evidence of erosion. Just like the Precambrian
rocks of the northern shields of Canada and Fennoscandia,
they will one day be worn down by the action of climate.
One of the most active agents of erosion, both in the Arctic
and the alpine tundra, is ice.
Ice accumulation
Ice accumulates where there is an abundant supply of snow
and where the temperature is low enough to prevent the
snowfall from melting completely away. Snow crystals, with
their complex and intricate forms, become altered within a
matter of days after they settle. They fragment into pieces,
they may melt in the sun, or they may become compacted
by the accumulation of further snow. They become denser
and harder, forming sugarlike grains that are crushed
together, eliminating the air spaces (although some bubbles
of gas may be permanently trapped within them). If temper-
atures are low enough to permit their survival, snowflakes
will have changed their form totally within two years, to
produce what has been referred to as firn, or “old snow.”
Within another three or four years, the firn will become
compacted to glacial ice.
Although ice caps and glaciers are composed mainly of ice,
they also contain many impurities. The snow that accumu-
lates may contain a range of airborne contaminants, from dust
and pollen grains to human-made pollutants. Especially in
geologically active regions such as Iceland, where active volca-
noes are present, volcanic dusts consisting of fine glass parti-
cles called tephra may settle in layers over glaciers during
periods of eruption. These tephra layers become buried as fur-
ther ice accumulates, and their presence provides a means of
36 TUNDRA
dating particular horizons in the ice mass. Meteorologists, for

example, may take borings through the ice to determine how
fast ice has accumulated in the past, and the tephra helps in
determining dates. The chemistry of the fine tephra dust is
very distinctive, and it is possible to identify the source of the
eruption by analyzing the particles trapped in ice. The chem-
istry may even differ during separate eruptions, so that specific
eruptions of particular volcanoes in the past can be identified.
Ice sheets, ice caps, and glaciers

There are several types of ice accumulation on Earth, includ-
ing ice sheets and glaciers. Only two true ice sheets now
exist, one occupying most of the continent of Antarctica, and
the other covering much of Greenland. The Antarctic Ice
Sheet occupies an area twice the size of Australia, bigger than
the whole of Canada. About 85 percent of the world’s fresh-
water is locked up in that one mass of ice, which in places
achieves a depth of 2.5 miles (4 km). Below it is a land sur-
face, much of which lies below the current sea level, espe-
cially in the west (see the map on page 3). This is so because
the massive weight of the ice sheet actually presses the
Earth’s crust deeper into the fluid layer of mantle beneath as
their combined mass achieves equilibrium. Along the eastern
edge of Antarctica and along the Transantarctic Ridge, on the
other hand, are ranges of mountains, some of which are high
enough to project above the ice surface. These mountainous
projections are given the name nunataks, derived from the
Inuit language. In between the western and eastern sections
of the Antarctic Ice Sheet is a fairly flat area of ice, the Ross
Ice Shelf. This ice shelf floats on the surface of the sea, which
forms a bay penetrating deep into the heart of Antarctica
toward the South Pole, and its eastern edge is fringed with
high mountains.
The Greenland Ice Sheet is considerably smaller than that
of Antarctica, but it is still as big as Mexico. It contains about
a further 8 percent of the world’s freshwater. The center of
Greenland is an enormous basin surrounded by high moun-
tains, and the ice sheet fills the basin and flows through the
mountains to the North Atlantic Ocean, where it breaks up
snow accumulation melting ice (ablation) Profile through the long

axis of a valley glacier.
Snow accumulates on
the upper part of the
glacier, where it is
compressed into ice. In
bedrock time the ice moves
along the lines shown,
basal melting
in “warm” glaciers eventually reaching the
site of ice melt.
into icebergs. In places this ice sheet reaches depths of about

two miles (3 km). Scattered around the polar regions there
are several smaller ice caps, often developed on high plateaus
in mountainous regions. These are much smaller than the ice
sheets but form centers of ice accumulation, from which the
ice spreads outward in the form of glaciers. Such minor ice
caps are found, for example, in Iceland and the Arctic island
of Svalbard.
Glaciers are mobile masses of ice that radiate from ice
sheets and ice caps or may form in the valleys of the moun-
tain regions of the world. Glaciers are much smaller than ice
sheets and ice caps, usually covering only a few square miles.
Glaciers are common in the mountain regions of the far
north and the far south, as in Alaska and New Zealand, but
can also be found on high mountains nearer the equator, as
in the Alps of Europe, the Himalayas of Asia, and even the
mountains of East Africa, which lie almost upon the equator
itself (see the map on page 20).
Ice sheets, ice caps, and glaciers may grow or shrink
according to the balance of snow accumulation and ice melt.
In mountain glaciers, snow accumulates fastest in the higher
regions, so the glacier effectively grows from the top (see the
illustration above). The increasing weight of ice results in a
gradual slippage downslope, where conditions are warmer, so
the bottom end of the glacier will be the main area of melt-
ing. The Earth’s mantle, beneath the crust, is hot and molten,
so warmth may arrive at the glacier from the rocks beneath.
Ice does not transmit heat well, and its insulating properties
38 TUNDRA
mean that the underside of the glacier is warmed and is also a

zone of melting. The process of ice loss is called ablation.
Whether a particular ice mass is growing or shrinking, there-
fore, depends upon the balance between accumulation and
ablation. This in turn depends upon the climatic conditions
and the local topographic setting. Obviously, ablation is at its
peak in summertime, while accumulation tends to be greatest
in winter, so the overall growth of a glacier is the outcome of
the full year’s gains and losses. If one could follow the history
of a single molecule of water falling on the upper section of a
glacier, it would probably go something like this: The molecule
would first be compressed into the local ice, then be buried as
further snow accumulated. It would then gradually slip down-
hill through the glacial mass until eventually it emerged at the
lower end of the glacier, where it would melt and flow away.
The movement of ice in a glacier is normally very slow,
much too slow to be detected by eye. A speed of three feet (1 m)
per day is quite fast for a glacier. But occasionally a glacier
will accelerate in what is known as a surge. In 1986 one of
North America’s longest glaciers, the Hubbard Glacier in
southern Alaska, suddenly increased its rate of movement to
about 33 feet (10 m) per day. It advanced across a bay, crush-
ing the trees on an island in its path and finally colliding
with the far side of the bay, where it blocked off a large lake
on its landward side. The Hubbard Glacier’s acceleration
seems to have been caused by a sudden increase in the supply
of ice from one of its tributary glaciers as a result of high lev-
els of precipitation. The surging of glaciers is not common
but is known to have taken place in Iceland, Greenland,
Alaska, and several locations in Russia. It is very unusual in
mountain glaciers away from the Arctic, but glacial surges
have been recorded in the Andes of South America.
Ice, therefore, like water, flows downhill. The weight of new
ice accumulation at the top pushes the entire mass down-
ward. The warming effect produced by the Earth’s heat at the
junction between the ice and the underlying rock also assists
this flow, melting the basal ice and lubricating the base of the
glacier. The mass movement of ice in a glacier creates lines of
tension and strain. Surface cracking may occur, resulting in
the formation of deep cracks that run along the line of move-
ment of the glacier; these cracks are called crevasses. In the

lower parts, mass slippage can cause cracks to form at right
angles to the direction of movement, called shear lines. Melt-
ing at the sides of the glacier and at its lowest point can result
in overhangs of ice, and melting beneath the ice can result in
the emergence of streams from tunnels, called ice snouts.
When light shines through the ice in these more fragmented
parts, a vivid blue color may be apparent. This is due to the
fact that, although ice reflects much of the light that falls
upon it, the spectrum of the light that does penetrate into the
ice is mainly absorbed, only blue light being transmitted.
When glaciers or ice sheets flow directly into the ocean, Europe’s largest glacier,
they can form high cliffs of ice and their continued movement the Aletsch in central
into the sea results in spectacular collapses of these ice walls. Switzerland. As the
Whole sections of the ice may break away and be released in glacier moves gently
floating islands of ice, the icebergs. This process is known as downhill it transports a
calving because it is almost as though the glacier is giving birth load of rock detritus on
to new, small ice masses. The glaciers of Glacier Bay, Alaska, are its surface and within
best observed from ships offshore, and the region has become the ice. (Photo by
a tourist attraction because of its impressive ice cliffs. Julian Barkway)
40 TUNDRA
Effects of glaciation
The high mountain ranges in which many glaciers form are
subject to intense erosion, and the resulting rubble, soil, and
detritus often ends up on the ice surfaces of the glaciers. It
falls from cliffs and slips down slopes, piling onto the stream
of gently moving ice in the valley below. Even more eroded
material may be gained as the ice scours the ground surface at
its base. The heavy mass of ice, slowly grinding its way down-
hill, carves out rocks and breaks them up under its massive
weight. When ablation (the loss of ice by melting) occurs, all
of these suspended materials are released once again and may
be either deposited on the spot or transported farther by
water that has resulted from ice melt, known as fluvial out-
wash. Long after the ice has gone, these deposited materials,
often found far from any remaining areas of active glaciers,
provide evidence of former glaciation.
Around 5.8 million square miles (15 million km2) of the
Earth’s land surface, or about 10 percent of the total, is cur-
rently covered by ice. Only 22,000 years ago, an area about
three times as large was ice-covered. We know this because of
the direct and indirect signs the glaciers left when they
retreated. Many regions of tundra, both polar and alpine,
bear the marks of ice action in the very recent past.
When glaciers grind their way down a valley, they excavate
materials in a very characteristic manner (see illustration on
opposite page). The floor of the valley becomes smooth and
rounded, while the sides of the valley are left steep, so that in
section a glaciated valley has a distinctive U shape. The steep-
ened sides may cut back into the side valleys that empty into
the main valley, leaving them cut short, or “hanging,” often
with their streams descending as waterfalls into the main val-
ley. The head of the main valley is often carved out in a steep,
rounded, basinlike form, creating what is known as a cirque
or corrie. A mountain that has formed the focal point from
which several valley glaciers have arisen often remains as a
sharpened peak, eroded on all sides, like the famous Matter-
horn in Switzerland, while hills carved out by foothill or
“piedmont” glaciers are rounded and smooth. Rock faces
exposed to passing glaciers may bear scars in the form of par-
allel lines and scratches caused by the slow grinding of
kame terrace esker
end moraine
retreat
ice
bedrock
esker till
rocks contained in the moving ice. Central Park in New York Cross section of a
City has a number of large rocks with such marks upon them, decaying valley glacier
bearing witness to former glaciation in that area. This pro- along both its long and
vides evidence that the tundra biome was once much more short axes. Melting ice
extensive on Earth, reaching as far south as modern New at the base of the
York City. glacier produces ridges
When glaciers retreat, they leave materials behind them of detritus called eskers,
that provide further evidence of their former extent and the and the sides of the
direction of their movement. The mass of material trans- glacier retreat to form
ported by ice includes large rocks and boulders as well as lateral banks, or kame
finely ground detritus, sometimes referred to as “rock flour.” terraces. At its lowest
When this mix is deposited in an unsorted mass, it is called point, the retreating
till, or sometimes boulder clay. Much of North America and glacier leaves an
northern Europe and parts of northern Asia are covered by end moraine.
such tills derived from rocks of the far north, which were car-
ried south by large-scale glacial movements. Many tills are
derived from geologically recent glaciations (often occurring
during the past 1 million years), but there are also ancient
tills from much earlier times. These very ancient tills, usually
termed tillites, have become consolidated, cemented, and
hardened until they have the appearance of a rock. They are
42 TUNDRA
very valuable to geologists because they provide clues to the

timing and location of ancient glaciations.
Till may entirely bury the parent rocks of a region, and it
may also contain large rock passengers carried from far afield,
known as erratics. Studied in the context of the geology of
surrounding areas, these erratics can provide evidence of the
direction of flow of a glacier. Sometimes the rock fragments
within a till all point in the same direction, and their orien-
tation can indicate the direction of former ice flow. The same
is not true of tills derived from floating ice. When an iceberg
melts, it also deposits all of the detritus it contains onto the
A glacier in retreat near seabed, but there will be no orientation of stones in a sedi-
Obergurgl, Austria, has ment of this type. Sometimes such material contains some
left behind a terminal microscopic marine fossils, however, and this can help to
moraine of rock identify the materials left by sea ice.
detritus, which is being Glacial advances and retreats are determined by the climate
eroded and sorted by and its effect on the accumulation/ablation balance of the ice
the flow of water from mass. A glacier may remain static for a time, perhaps a few
melting ice. (Photo by decades or even centuries, and then undergo retreat, in which
Peter D. Moore) case it often leaves behind a distinct ridge of till marking its
terminal position. Called a moraine, this feature often has a

profound effect upon the future development of the land-
scape. If it blocks the flow of water from the melting glacier or
from the stream that succeeds it, it can result in the formation
of a lake. Many lakes in regions formerly glaciated have been
created in this way. The formation of these moraine lakes
greatly diversifies the glaciated landscape, as can be seen in
the Sierra Nevada of California, where Convict Lake is an
excellent example of a moraine glacial lake. The sediments
that accumulate within these lakes provide a record of the cli-
matic, vegetation, and other environmental changes that
took place during the course of the lake’s history.
As well as being deposited at the terminus of a glacier, till
may also build up beneath the ice, in which case the piles of
deposited material take on a distinctive, streamlined form,
aligned to the direction of glacial flow (see the illustration on
page 41). These formations, called drumlins, become apparent
when the ice melts. Isolated patches of ice left behind on
level valley floors and plateaus, called dead ice, melt to
release an amorphous, randomly scattered series of piles of
till called hummocky moraine. Finally, single blocks of ice
buried in till melt slowly to form deep, steep-sided hollows
that often fill with water to form kettle-hole lakes. The pot-
hole wetlands of the prairie regions of North America have
developed from the infilling of kettle-hole lakes.
After the ice

Clearly, when changes in climate cause glaciers to retreat,
they leave behind many clues to their former presence, marks
on the landscape that tell a geologist that ice has been pres-
ent in the past. But the impact of the ice itself is not the only
clue. Glacial melt not only deposits detritus but also releases
large volumes of water, so it is inevitable that much of the
material resulting from glacial activity is washed and sorted
by water flow. While the glacier is still intact, streams form
within the very body of the ice, and these ice-confined
streams sort the detritus into long, sinuous ridges called
eskers. These can be up to 100 feet (30 m) in height and
remain distinctive long after the ice has departed. Along the
44 TUNDRA
edges of the glacier, meltwater can create similar strips of

sorted material that form terraces on the valley side. These
are termed kame terraces. Downstream of the glacier,
deposited detritus continues to be sorted by meltwater flow,
forming “braided” streams running in a series of parallel rib-
bons, their waters often milky in appearance because of the
high content of clay resulting from glacial erosion.
Water is not the only means by which glacial detritus can
be further transported and sorted; the wind is also an active
agent. When water carries and sorts the different sizes of par-
ticles that emerge from glaciers, the particles become
deposited in a series of narrow, parallel ridges in the riverbed,
called braided ridges, and there they dry out. Once the particles
that form the ridges are dry, the strong winds that often occur
in the vicinity of a glacier pick up the finer sediments and
carry them over great distances before depositing them in
new locations. Since the rivers provide a constant source of
this material, the wind may carry large quantities and deposit
them all together, forming a silty or sandy loess soil. Loess
soils are therefore rich in fine particles. Because they are
formed from freshly ground rock, they are also rich in chemi-
cal nutrients, so they produce very fertile ground. Loess soils
are particularly frequent in Alaska, in central Europe (from
Belgium to Ukraine), and in China. The great agricultural
regions of the American Midwest are based upon loess soils.
Fossils are, not surprisingly, scarce in glacial deposits. The
degree of physical damage and abrasion precludes the survival
of most biological material. Apart from that, glacial landscapes
do not normally support large numbers of animals and plants.
Occasionally, however, the bones of large mammals survive in
glacial outwash, along with other very resistant materials, such
as the wing cases of beetles. In rare instances, pockets of peaty
soils are preserved, containing plant fragments usually of
Arctic-alpine vegetation. Caves around the glaciers are also
good sources of fossils, because animals, including humans,
often used these for shelter and remains may have accumu-
lated on the cave floor. The deposits build up over the course
of years in a sequence of layers, the deepest being the oldest.
Cave deposits can thus provide evidence of the biological his-
tory of a glaciated region over the course of time.
PACIFIC OCEAN
Cordillera
Laurentide
ARCTIC OCEAN
Siberia
Greenland
NORTH ATLANTIC OCEAN
Scandinavia
ice sheets and large ice caps
By studying these various glacial features and their marks The maximum extent of
upon the landscape, glacial geologists have been able to put ice sheets and glaciers
together detailed pictures of former glaciation events. in the Northern
Answering the seemingly simple question of how many Hemisphere during the
glaciations have taken place in recent times, however, has most recent ice age
been made difficult by the fact that the latest glaciation has
often destroyed or obscured the evidence of former glacial
events. The glacial features themselves are relatively easy to
recognize, but deciding exactly when they were produced,
and in what sequence, is much more difficult. Geologists
have mapped the extent of ice when the last ice age was at its
maximum, around 22,000 to 20,000 years ago, and the map
above shows this extent in the Northern Hemisphere. Many
of the regions covered by the ice sheets had been glaciated
46 TUNDRA
during former cold episodes over the past million years, but
much of the evidence for these earlier glaciations was
destroyed by this final ice advance.
Patterns on the ground

The region around the edge of a glacier following glacial
retreat is called the periglacial region. The conditions here are
cold but not quite cold enough to support a year-round ice
cover. Even when ice has retreated from an area, leaving bare
rock and glacial debris, ice may remain beneath the surface of
the ground. This happens especially in polar regions but also
occurs in some high parts of the world such as Tibet and
Mongolia. If the mean annual temperature remains below
the freezing point of water, then water below the ground may
remain in a frozen state. The surface water, however, is
warmed by the Sun and remains in a liquid state through the
brief summer, so a wet soil overlies permanent ice. Perma-
nently frozen subsoil is called permafrost. Regions affected in
this way are conveniently divided into two types: continuous
permafrost, in which there is an uninterrupted layer of per-
manently frozen soil beneath the ground; and discontinuous
permafrost, where frozen soil is patchy and interspersed with
areas that seasonally defrost. The map on page 47 shows the
geographical distribution of permafrost. It is in the southern,
warmer regions, usually occupied by coniferous forest rather
than tundra, that permafrost becomes sporadic. Where it is
discontinuous, the remaining patches of permafrost tend to
be confined to the more exposed ridges where protective
blankets of snow fail to accumulate in winter. The depth of
the permafrost is variable, but it can be very deep. Records
from Canada suggest that permafrost can extend to two-
thirds of a mile (1 km) in depth.
The map shows the areas of both discontinuous and con-
tinuous permafrost, and it can be seen that the true tundra
regions (shown in the map on page 2) fall mainly within the
area of continuous permafrost. The permafrost map also
shows locations of permanent offshore ice, which are really
the oceanic equivalent of permafrost, since the waters remain
frozen at all times of the year. Observe that the areas of per-
mafrost extend farthest south in areas with the most conti-

nental climates, as in Siberia. Note also the large block of dis-
continuous permafrost well to the south of the main
permafrost distribution in the highlands of Tibet. These
regions are very far from any ocean and therefore cannot be
heated by the influence of warm oceanic currents. Land-
masses also lose heat faster than the oceans, so the interiors of
continents tend to experience low winter temperatures. In the
case of the highlands of Tibet, the high altitude (most of the
land is higher than 12,000 feet [3,600 m]) also contributes to
the low temperatures. The influence of the warm ocean cur- The extent of
rents moving northward in the North Atlantic (shown in the continuous and
map on page 12) is also clear in this map. Northern Europe is discontinuous
clear of continuous permafrost because of the warming effect permafrost in the
of these ocean currents, despite the fact that it lies as far north Northern Hemisphere
150° 180° 150° 120°
20°
30°
40°
50°
120°
60°
90°
70°
80°
90°
70°
60°
50°
60°
offshore permafrost
continuous permafrost
60°
discontinuous permafrost
30° 0° 30°
48 TUNDRA
as much of Siberia and Canada. No warm currents penetrate

to these regions, so these lands are locked in permafrost.
The surface of the ground is strongly affected by the cycle
of freeze and thaw of the soils above the permafrost. This
upper layer of soil, which is defrosted each spring, is called
the active layer. It is within this layer of the soil that most
physical, chemical, and biological soil processes take place.
The plant roots are confined to this layer. Plant roots often
occupy only the very uppermost levels (about the top four
inches [10 cm]) of the soil but may extend down to about 10
inches (25 cm) in places. The depth of the active layer
depends on a number of factors, including the nature of the
soil. In flat meadow soils, only the top 12 to 17 inches (30 to
40 cm) may thaw in summer, while on drier sites the thaw
may extend to a depth of about three feet (1 m). If there is a
layer of organic litter or wet peat over the mineral soil (which
is often the case), this insulates the soil against the spring
warming and the active layer remains very shallow.
When water freezes it expands, exerting pressure on any
object in contact with it. As winter commences, the water
contained within the deeper layers of soil, which are in con-
tact with the permafrost, begins to freeze. The temperature of
the surface soil, which is exposed to the cold night air, also
falls rapidly and the water it holds also freezes. The layer of
wet, unfrozen soil sandwiched between the upper and lower
ice zones is consequently placed under pressure because the
soils above and below are expanding. The result is that bub-
bles of wet soil may burst through the surface of the ground,
releasing a mass of muddy material over the local vegetation.
Where this happens, the surface of the tundra becomes
patchy, with bare soil areas often raised above their sur-
roundings by the pressures from below.
The great pressure exerted from below by the wet soil,
expanding as it freezes, forces any stones or rock fragments in
the soil up toward the surface. This forcing of rocks to the
surface is called frost heaving. The reason why stones in par-
ticular are pushed upward through the soil is that they lose
heat more rapidly than their surrounding soil. Stones con-
duct heat well, so they quickly become hot or cold under the
right conditions. This is why stones feel hot to the touch in
the sunshine but very cold at night. As a result of this prop-

erty of the stones, the soil around them (particularly the soil
immediately below them) becomes cold faster, and this is
where water begins to freeze. The expansion that results from
this freezing pushes the stones upward through the soil and
they end up on the surface.
Larger stones are usually forced upward more effectively
than small stones, so the frost sorts the rocks into different
sizes. Moreover, when rocks and stones are brought to the
surface, they are not randomly spaced but often develop in
patterns. This results from the fact that the stones move
sideways as well as upward. On flat ground, the stones
become arranged in a kind of network called stone polygons.
Why they should form a network is a question best answered
by considering what happens when a pool of water dries out:
Even in temperate climates, the exposed mud usually
becomes caked and cracked in a distinct pattern of polygons.
These usually six-sided shapes form because of the way in
which soils dry out and contract. Similarly, in tundra habi-
tats the freezing and thawing of the ground causes the soil to
contract periodically into polygons, and the stones on their
way up through the soil move into the cracks on the surface,
forming stone polygons. On a slope, these polygons are nar-
rower, and on steep slopes they form a linear series of stone
stripes, aligned along the contours. These stone patterns on
the surface of the ground can still be detected in regions now
far from the tundra, such as New England and southern
Britain, indicating that these regions once experienced
periglacial conditions.
Another feature of periglacial regions is the formation of
ice wedges in the soil (see the illustration on page 50). When
soil freezes, it forms cracks over its surface and, as described,
these often take on the form of a network of polygons, look-
ing rather like a honeycomb from the air. Water abounds in
low-lying regions of the tundra because evaporation is low in
the cool temperatures, and it fills these cracks and then
freezes and expands in winter, pushing the cracks deeper into
the soil and widening them in the process. In cross section,
the water that freezes within these cracks looks like a wedge
that is driven into the soil, so these features are called ice
50 TUNDRA
wedges. They are widest at the surface of the ground, where

more water is added each summer, and narrowest at their
deepest tip. The resulting patterned landscape takes on the
form of ice-wedge polygons, and these cover many areas of the
tundra landscape. The center of the polygon may be raised
into a dry crest, or it may become sunken and bear a pool of
Some of the geological water during the summer, so that the center and the edges of
features found in Arctic the polygon bear different vegetation, emphasizing the pat-
tundra. Ice lenses below tern further. The wedges themselves thaw out in summer and
the ground surface may form an interconnecting web of narrow waterways, and these
expand to form large become filled with aquatic vegetation and then silt as the sur-
mounds called pingos. rounding soils become eroded into the channels.
Ice can also penetrate Ice-wedge polygons will form only under very cold condi-
the soil from the tions, usually with a mean annual temperature of 21°F (–6°C)
surface, often forming or lower. They can occur in a fossil state in lands far south of
ice wedges that their current distribution, indicating that very low tempera-
are arranged in tures prevailed in that region in the past. The size of the poly-
polygonal patterns. gons is variable, from a few feet to as much as 300 feet (100 m)
water ice polygons
ice ice
ice
permafrost
lens of ground ice growing pingo collapsed pingo ice wedges
base of permafrost unfrozen rocks

in diameter. When the perimeter becomes slightly raised

above the center because of the expansion of the ice wedge,
which forces up the sediments in the crack, then the depressed
middle of the polygon may become an open pool of water (see
the illustrations on page 52).
Patterns of stone stripes and polygons are also found in
alpine situations where the conditions are cold enough.
Even on the summits of the equatorial mountains in East
Africa, polygons can be found, but here they are much
smaller, often only four to eight inches (10 to 20 cm) across.
They are formed by the same basic mechanisms as their
polar counterparts, but the freezing and thawing is due to
the difference in day and night temperatures rather than any
difference in season. Perhaps this is why their development
is on so much smaller a scale.
In some periglacial regions, huge mounds (perhaps even
several hundred feet in height and diameter) may rise out of
an otherwise flat landscape. These are especially frequent in
the coastal plain of Alaska and in the coastal zone of eastern
Greenland. These structures, called pingos, often develop
over dried-out lake basins or along the sides of streams and
rivers. Pingos become elevated by the growth of an ice core
in the center of the mound, formed as a result of water pres-
sure from below. Usually they develop in the sites where
springs of water rise from the ground under pressure and
feed the ice core, which freezes as it nears the surface of the
ground. The upper part of the mound has a very different
microclimate from the surrounding lowlands, and the
southern slopes of the pingo mound may bear vegetation
characteristic of a warmer climate, possibly even a tree cover.
If the general climate becomes warmer, however, the ice core
in the center of the pingo melts and the whole mass col-
lapses to form a deep pool, surrounded by a circular rim or
“rampart” formed from the soil and other detritus raised up
by the mound. Circular ponds with ramparts can be found
far to the south of the present-day tundra regions, indicating
the periglacial conditions that once prevailed there (see the
illustration on opposite page).
A similar type of structure is sometimes formed in those
regions of the Arctic that are rich in peat bogs, and this is
52 TUNDRA
Profiles of Arctic
polygons, showing both
the low-centered and
the high-centered forms.
The active layer of the
permafrost
surface soil melts each active layer
spring, while the deeper water
layers (permafrost)
remain frozen. Low-
centered polygons bear
marsh vegetation, while
high-centered forms are
dry and covered only
with mosses and lichens.
termed a palsa. Palsas, like pingos, are large mounds that

are formed by the development of ice beneath the surface
of the ground. Generally smaller than pingos, palsas are
only six to 10 feet (2 to 3 m) in height and about 150 feet
(45 m) across. They often lie within wetland areas, where
they are interspersed with open pools and other palsas in
various states of development. Palsas begin their formation
as small irregularities on the surface of a sedge marsh. Even
a very small elevation of an inch or two can affect snow
accumulation. The snow is blown from the slight mound
and accumulates in the hollows between, so the more ele-
vated spots have less snow cover in the winter. They there-
fore become colder, because snow acts as an insulating
blanket, retaining some of the ground’s heat that it gained
in the summer sun. By contrast, the cold penetrates any
small raised areas of ground, causing an ice core to develop
within them (as shown in the illustrations on pages 50, 52,
and 54). As the ice forms, it expands, pushing the hum-
mock even higher so that even less snow is retained in the

high winds. Palsa growth is thus a self-propagating process:
The higher the palsa grows the colder it becomes, and the
colder it becomes the more the ice core expands and
pushes the palsa upward. The ice core survives through
the summer, so its growth continues for many years, even
centuries.
As the palsa grows in height, drainage improves and the
vegetation on its surface becomes drier. The surface of the
palsa may then develop a vegetation cover of lichens. Many
lichens are white or pale gray, and these reflect the sunlight,
keeping the mound cool in summer. In the course of time,
dwarf shrubs replace these lichens, but these have a darker
color and absorb more heat from the summer sun. This, cou-
pled with the increasing height of the dome, eventually
leads to erosion of the surface soil, revealing the peat
beneath. The dark-colored peat absorbs even more heat, and
its exposure leads to the meltdown of the ice core. The core
collapses quite quickly, leading to the formation of an open
pool of water that is available for recolonization by sedges as
the cycle begins again. Finnish scientists, working in the
1980s, demonstrated experimentally how this process oper-
ates. They spent a whole winter visiting an area of Arctic
sedge bog where, using a broom, they swept an area clear of
snow and kept it clear through the winter. The result was the
development of a permanent ice core beneath the ground,
and within a few years the swept locality was developing
into a palsa.
All of these structures associated with tundra environ-
ments lead to a diversity of landscapes and landforms. This is
very important when one considers the range of microhabi-
tats that are available for the plants and animals that inhabit
the inhospitable world of the tundra. The mounds and hol-
lows, wetlands and drylands, slopes and cliffs all offer oppor-
tunities for living things to find a home, survive, and breed.
As will become clear, the biodiversity of the tundra is not
high, but such richness as it contains is due largely to the
diversity of its landscape features. Geology, or more strictly,
geomorphology, plays an important role in the maintenance
of the tundra’s biodiversity.
54 TUNDRA
3 70–330 feet
10–17 feet
4
bare peat
water
cotton sedge dwarf shrubs lichens

unfrozen peat frozen peat unfrozen silt frozen silt glacial deposits
Soil formation in the tundra

As has been described, in areas of permafrost, the deeper
parts of soils are frozen throughout the year while the upper
layers melt during the summer. The result of this annual
drastic change is that soil conditions are also altered strongly
from one season to another. The formation of ice means that
soils are constantly being heaved, stirred, contorted, and dis-
turbed, leading to a degree of instability. On slopes, the upper
layer is likely to become highly mobile in summer as the wet
surface soil slides over the top of the frozen lower layers. Soil
slips downhill and gathers at the base in a process called
solifluction. Even on level ground, soils in effect plow them-
selves as stones are heaved upward to the surface, resulting in
a constant mixing of the upper layers. As they move, these
stones are fragmented by the activity of frost and mechanical
damage.
All soils throughout the world are produced initially by the
breakdown of rocks into small fragments. This process, called
weathering, results from the action of physical and biological
factors on the rock. In the tundra, the action of freezing and
thawing is particularly important because the penetration of
ice into rock surfaces can split them into sections so that the
rock decays under the stress. This is particularly apparent
when rocks and cliff faces are directly exposed to the air,
where the changes in temperature are strongest. Freezing and
thawing causes rock falls and generates masses of rock rubble,
known as tallus scree, on the slopes beneath the cliffs. This is
particularly obvious in mountain regions, and rock break-
down may be particularly active in summer in alpine sites.
The very hot summer of 2003 in the European Alps led to
pronounced rock falls and even led to the closure for safety
reasons of one of the favorite peaks for mountaineers, the
Matterhorn in Switzerland. In winter, snow avalanches are
often the greatest danger to humans on mountains. They are
also a major contributor to rock breakage and destruction.
(opposite page) The development of palsas. Ice forms beneath the

tundra soil surface and expands to build a mound. The elevated
surface dries, leading to changing vegetation and eventually to
erosion, when the palsa collapses to form a pool.
56 TUNDRA
Within tundra soils, rock weathering takes place primarily

in the active layer, where thawing and refreezing take place
each year. Deeper in the soil, in the permafrost, such temper-
ature changes do not occur and the underlying rocks remain
below freezing. The outcome is that physical weathering
occurs mainly above ground or within the active layer of the
soil. This may not be as spectacular as the large-scale rock
breaking that occurs on cliff faces, but it is nevertheless an
important process. Rock particles are split into smaller por-
tions by water that penetrates into cracks and freezes. Over
the course of time, the chemicals contained within the rocks
are released after being trapped in the rocks for many mil-
lions of years, and they become available to the living plants
within the ecosystem once more.
In addition to physical breakdown, weathering can also
result from biological activities, in which living organisms
assist in the breakdown of rocks. Lichens on the surface of
exposed rocks, for example, actively attack the rock by secret-
ing acidic compounds that dissolve the rock surface beneath
them. This can strip away flakes of rock in a process called
exfoliation. Algae may likewise coat the surface of the rock.
Grazing mollusks such as snails that eat algae and lichens
may then scrape the rock itself with their rasping tongues.
Plant roots, including those of dwarf willows (Salix species)
and the mountain avens (Dryas octopetala), may penetrate
cracks in rocks and enlarge them as the roots grow and
expand. The force exerted by roots extending as they forage
for water and minerals in the ground is entirely adequate for
the splitting of rocks.
Although physical weathering of rock occurs relatively
quickly in the tundra, biological and chemical weathering
take place rather slowly. The chemical reactions in the soil
that assist in the breakdown of rocks, such as the impact of
acids from the atmosphere on the soil particles, are slower in
the tundra than in the Tropics because of the low tempera-
ture, so chemical weathering is slow. The action of bacteria
and fungi in the soil produces corrosive materials that attack
both mineral and organic matter, but this process is also
slower in the tundra or restricted to those limited times in
the year when the temperature is high enough to allow bac-
terial and fungal growth. So, although freezing and thawing

have a strong impact on rock weathering, the more subtle
action of chemicals and soil organisms is slow.
The limited activity of bacteria and fungi in the soil means
that organic matter produced by the dead portions of plants
does not decay rapidly. Consequently, soils may develop
reserves of organic material. In the wetter locations this
organic matter may even accumulate as peat. Organic matter
is good at holding water, like a sponge, so it prevents many
tundra soils from becoming excessively dry in summer. It is
also efficient at holding the scarce mineral elements that
plants require for their nutrition, so that the peat forms an
important reserve of elements within the soil. Most rocks of
the Arctic are relatively poor in the chemicals needed by
plants. Phosphorus and nitrogen, in particular, are in short
supply. The low level of precipitation also means that rela-
tively little in the way of chemical input arrives from the
atmosphere (see “Nutrient cycling in the tundra,” pages
76–81).
Soil types of the tundra

It is possible to divide tundra soils into two main types,
“skeletal soils” and peaty soils. The skeletal soils, so called
because they consist mainly of rock fragments of various
sizes and very little organic matter, develop in well-drained
sites, such as on the tops of esker ridges. Here vegetation is
sparse and the limited precipitation from the atmosphere
passes quickly through the soil, moving down the slopes to
the water-accumulating regions in the valleys and on the
plains. Within these wetter locations decomposition is inhib-
ited by the abundance of water, and so organic matter does
not rot away but develops as peat over the mineral soil.
Skeletal soils, sometimes called “rankers,” are found where
the process of rock weathering has produced masses of frag-
ments but where the long-term effects of vegetation devel-
opment and the percolation of water have not taken place.
In a sense, they can be regarded as young or immature soils.
They have the basic mineral components but lack the full
range of animals, plants, and microbes that are found in a
58 TUNDRA
fully developed mature soil. As these additional components

arrive, and as the constant downward movement of water
through the soil takes place, the soil develops a distinct
series of layers, called horizons. The orderly series of horizons
within a soil is known as its profile. The profile of a soil is its
cross section, and soil scientists usually view it by digging a
pit right down to the underlying rock (see photograph on
opposite page). In the skeletal soils, the soil profile is very
simple, consisting of broken rock overlying the basal rock or
the ice of the permafrost. The underlying basal rock is often
the source of the fragments that make up the soil, so it is
called the parent material. Skeletal soils are found where gla-
ciers have recently retreated or on slopes and dry ridges,
often with little or no vegetation cover.
In the shrub tundra of the Low Arctic, where vegetation
forms a continuous cover, the soil may mature to form a
more complex profile. The surface vegetation of this region,
consisting of plants belonging to the blueberry and heather
family (Ericaceae), produces a very acid litter as the dead
leaves of these plants fall to the ground. This litter of dead
plant material usually accumulates as a layer of slowly
decomposing organic matter, called humus, on the soil sur-
face. The mineral soil below this humus consists of two dis-
tinct layers: an upper pale, bleached layer, and a darker,
reddish colored lower layer. The organic top cover and the
bleached layer are termed the A horizon and the lower red
layer is the B horizon. There is a lower subsoil called the C
horizon, which is really just decomposing rock, often lying
within the permafrost. A soil with this layered, or stratified,
profile is called a podzol. It is very common in the boreal for-
est (or taiga) zone, but also extends into the southern parts of
the tundra where there is an extensive dwarf-shrub vegeta-
tion cover. The A horizon of a podzol is usually acidic and
poor in nutrients, so the development of this type of soil pro-
file has an impact on the vegetation and consequently on the
animals that graze upon it.
The conditions needed to produce the layers of a podzol
are quite complicated. Water from melting snow or summer
rain moves downward through the soil under the influence
of gravity. As it passes through the organic litter layer, the
The profile of a soil

developing over a
bedrock of chalk. The
disintegrating rock at
the base is overlain by
smaller rock fragments
mixed with organic
matter derived from
the surface vegetation.
(Photo by Peter D.
Moore)
water picks up various organic chemicals, including some

called polyphenols, which are derived from the breakdown
of the plant tissues. These acidic compounds help to dissolve
many of the inorganic chemicals in the upper layers of the
60 TUNDRA
soil, including iron and aluminum, and they also cause fine
A podzol soil profile clay particles to become mobile and to migrate down the soil
from northern Finland. profile. All of these moving materials become deposited in
The surface layer the lower B horizon, where they form a dark reddish zone,
consists of organic litter mainly because of the color of the iron oxides present. This
derived from the type of soil profile is dependent on the downward movement
vegetation, below which of water, so it can develop only if there is relatively free
is a pale layer that has drainage of water through the soil.
been leached of its iron. When drainage is poor, as when a site is low-lying and
Lower in the profile is a receives an inflow of water from surrounding areas, the soil
deposition layer underneath is often permanently waterlogged and this
containing iron, strongly affects its chemistry and its biology. Poor drainage
aluminum, and organic also affects its general appearance. Digging a pit in such a
matter leached from waterlogged Arctic site would reveal, first, a black layer of
above. (Photo by poorly decayed organic matter, and below it, a mineral soil,
Peter D. Moore) often with a blue-gray appearance. Such a soil is called a gley
soil. The reason for its gray color is the fact that it is starved
of oxygen. Most healthy soils have air spaces within them
and the penetration of air into the soil allows oxygen to
reach the bacteria, fungi, earthworms, plant roots, and other
living organisms that dwell in the soil and depend on oxy-
gen to breathe. When a soil becomes fully saturated with
water, all air spaces and channels to the surface become
blocked by standing water and the air can no longer pene-
trate to the lower layers. Although oxygen does dissolve in
water and can move by diffusion through water, its rate of
movement when dissolved in water is about 10,000 times
slower than is possible in the air. So, in a water-soaked envi-
ronment oxygen is often in very short supply, and this
means that microbes, animals, and plant roots are all starved
of oxygen. Many plants and animals literally drown in such
a difficult habitat.
The dearth of oxygen also affects chemical processes.
Some chemical reactions in soils are dependent on oxygen,
especially reactions involving the element iron. Iron is a
very common element in all soils, and in a well-aerated soil
it is usually present in the oxidized form iron-III (ferric
iron). Iron-III is often further combined with oxygen to
form iron oxides that have a reddish (rusty) color, and this
often gives a soil its rich, dark color. When oxygen is in
short supply, however, as in gley soils, iron becomes
reduced to its iron-II form (ferrous iron). In the iron-II form,
the element takes on a blue-gray color that is very apparent
in the gley soil profile. So the waterlogged soils of the tun-
dra take on the gray character of the form of iron contained
in an oxygen-poor environment.
A careful examination of a gley soil profile will sometimes
reveal flecks or lines of reddish color. These rusty spots and
stripes are usually associated with the channels and tubes
through which roots of plants have penetrated: Old root
channels in which the root has died and decayed form
access routes for air into the soil. They create tubes along
which oxygen can travel, and its presence is displayed by the
formation of iron-III oxides with their characteristic rust
color.
62 TUNDRA
Debris from glacial Soil change in time

retreat near Lech in Soils change over the course of time, and this is particularly
Austria has been apparent where glaciers retreat and leave areas of ground-up
colonized by herbaceous rock behind them that will gradually develop into soils.
vegetation. Some shrubs Indeed, some of the most important scientific studies of soil
are beginning to invade development have been associated with glacial retreat sites,
the developing soils. especially where the timing of the glacial movements has
(Photo by been recorded and where a timescale can therefore be used as
Peter D. Moore) a framework for soil development. The process of ecosystem
development and maturation involves an array of plants and
animals, and the arrival and growth of organisms at a site
causes changes in the tundra soil (whether polar or alpine) as
it develops.
The soil deposited by a retreating glacier is essentially a
mass of rock particles ground down by ice action, perhaps
sorted by water movements, and eventually deposited in a
site where it becomes physically stable. This is a typical

ranker or skeletal soil, poor in organic matter. Any larger
rocks remaining become colonized by lichens, and these fur-
ther corrode the solid materials, producing smaller particles
that settle on the soil below. Many lichens are also able to
“fix” nitrogen from the air (see “Nutrient cycling in the tun-
dra,” pages 76–81), and these play a part in providing a plant
nutrient that is essential to all living organisms. Plants can
then begin to colonize the soil, and as they thrive and ulti-
mately die they add organic matter to the soil. This is
because when a plant dies, a proportion of the living mate-
rial it produced by photosynthesis is deposited in the soil, Dwarf shrubs colonize
where it changes the soil’s very nature. The organic matter an area of glacial
acts as a sponge and holds water, making the soil a moister retreat at the
place where more plants can grow. It also supplies an energy Kangshung Glacier, east
source for animals such as springtails and earthworms that of Mount Everest in
feed upon dead vegetable matter. The organic humus com- Tibet. (Photo by
ponent of the soil is also very effective at holding various Colin Monteach/
minerals in the soil (such as calcium and potassium) so that Minden Pictures)
64 TUNDRA
they are not lost but remain as a reservoir available to plants

for their growth.
Some mineral elements, however, are inevitably lost. As
water drains through the soil profile, it dissolves some of the
elements released from the decaying rock particles and car-
ries them away in the drainage water that leaves the soil. This
loss of elements is called leaching. Some of the chemicals dis-
solved in rainwater assist in this process, especially acids such
as carbonic acid and sulfuric acid that result from the uptake
of carbon dioxide and oxides of sulfur respectively from the
atmosphere. Plants themselves may also contribute to the
speed of leaching, as in the case of podzol development (see
“Soil types of the tundra,” pages 57–61).
The gradual development of a soil is accompanied by
changes in local vegetation and animal life, so it leads the
entire ecosystem along a track of growth. Exactly which
plants and animals manage to invade and take up residence,
depends, of course, on the climatic factors that apply in any
given location. Colonization of the soil by dwarf shrubs of
the Ericaceae family can lead to the development of podzol
soils on the better-drained sites. Where conditions are wetter
and drainage poor, then the peaty soils and gleys develop. So
the tundra region bears a variety of soils and these change
both in space and in time.
The tundra atmosphere

All soil processes, from the weathering of rocks to the forma-
tion of peat, are affected by the air above the surface and
within the pores of the soil structure. The soil has an atmos-
phere of its own, which interacts and exchanges gases with
the atmosphere above. Precipitation passes through the
atmosphere and gathers some chemicals as it falls toward the
ground, so the atmosphere can supply elements to the soil in
this way. The vegetation that grows upon the surface of the
soil and adds organic matter to its structure derives the ele-
ment of carbon from the atmosphere in photosynthesis. To
understand the chemistry of tundra environments, therefore,
it is necessary to know something of the atmosphere above
the ground.
The Earth is surrounded by the atmosphere, a layer of gases

held close to the planet by its gravitational pull. The most
abundant gas is nitrogen, composing about 79 percent of the
atmosphere. This is a relatively nonreactive gas, and
although nitrogen is vital to all living organisms as a compo-
nent of proteins, the gaseous nitrogen of the atmosphere
can be tapped only by a small number of microbes. Of the
remaining 21 percent of the atmosphere, the bulk is made
up of oxygen, which is, of course, vital for respiration.
Together with most animals and plants, people take in oxy-
gen and use it in a controlled combustion in their cells to
release energy; this is called respiration. Atmospheric oxygen
used in respiration is replenished by green plants, which
produce oxygen as a waste product of their activities in pho-
tosynthesis. A very small proportion of the atmosphere is
occupied by carbon dioxide (less than 0.04 percent), water
vapor, and various trace gases. These smaller components of
the atmosphere are important because some of them have
the ability to absorb and retain heat energy. They create a
kind of thermal blanket around the Earth that absorbs the
heat radiated from the Earth and maintains a fairly stable set
of conditions at the Earth’s surface. Human disturbance of
this balance of scarcer gases and the way in which such dis-
turbance affects the tundra are problems that are becoming
increasingly urgent (see “Tundra as a carbon sink,” pages
172–173).
The atmosphere is densest close to the surface of the
Earth because of the pressure of gases above it. But Earth’s
atmosphere is considerably less dense than that of some
other planets, such as Venus, where a high concentration of
carbon dioxide produces an atmospheric pressure roughly
90 times that of the Earth. The atmosphere becomes thinner
with increasing height above the surface of the Earth, and it
is convenient to divide it into various layers according to its
composition and properties. The lowermost layer, which
extends up to a height of about seven miles (11 km), is the
troposphere. Most meteorological activity takes place here;
clouds, even the highest cirrus ice clouds, are mainly
located within this layer. As explained earlier, temperature
decreases with increasing height within the troposphere
66 TUNDRA
(see the sidebar “Lapse rate,” page 19), but at the very top of
this layer a change occurs at a point called the tropopause.
Just above the tropopause is the stratosphere, and this layer
extends up to a height of about 31 miles (50 km). The strat-
osphere contains significant quantities of ozone, and this
gas has properties that are of particular importance to the
polar tundra regions. Ozone absorbs solar radiation in the
ultraviolet (UV) part of the spectrum. As a result, parts of
the stratosphere actually become warmer than the upper
layers of the troposphere, sometimes reaching as high as
50°F (10°C). Most of the light that is visible at the surface of
the Earth and that plants use in photosynthesis passes
through the stratosphere, but the very short wavelength
energy of UV is absorbed by the stratospheric ozone and fil-
tered out. This is important to all the living things on the
land surfaces of the Earth because high-energy UV radiation
is potentially damaging to cell structure. It causes break-
down of various cell components, including the DNA,
which carries genetic information, and this damage can
lead to cell malfunction and even cancer, especially of the
skin. So the ozone of the stratosphere is critical for the pro-
tection of life on Earth. In the tundra regions, especially of
the Antarctic, stratospheric ozone destruction has been
noted in recent years and this is a cause of concern (see
“Ozone holes,” pages 178–180). What happens in the polar
atmosphere can provide an early warning for those parts of
the world where human populations are denser and may be
in danger.
Another essential aspect of the polar atmosphere relates
to the layers that lie above the stratosphere. From a height
of 31 miles (50 km) to about 50 miles (80 km) lies a layer
called the mesosphere, at the top of which the temperature
may be around –180°F (–120°C). The density of the atmos-
phere here is very low, but it still contains enough oxygen
to burn up the meteorites that head toward the Earth, creat-
ing shooting stars. Above this, up to around 250 miles (400
km), lies the thermosphere, where the atmosphere gradu-
ally thins to the emptiness of space. Satellites are positioned
at around this altitude. It is also in this layer that a remark-
able polar phenomenon, the aurora, takes place. This con-

sists of flickering sheets of light that pass across the night
sky in the very high latitudes in both the Northern and
Southern Hemispheres. In the north this is known as the
aurora borealis (Boreas is Greek for “north wind”), or
“northern lights,” and in the south the aurora australis
(Auster is Latin for “south wind”). These spectacular displays
occur when the molecules of the thin upper atmosphere are
bombarded by charged particles emitted by the Sun, known
as the solar wind. The reason why auroras are restricted to
the polar regions is that the magnetic poles attract and con-
centrate the charged particles of the solar wind. The inten-
sity of particles emitted from the Sun varies with the
activity of the Sun in the form of sunspots. When a solar
flare takes place on the surface of the Sun, spectacular dis-
plays of the aurora occur about 24 hours later. Sometimes
these polar displays can be seen as far south as the northern
United States or in England. Their occurrence makes the
atmosphere above the polar tundra of particular interest to
atmospheric physicists.
Conclusions
The cold conditions of the regions surrounding the North
and the South Poles have encouraged the accumulation of
permanent ice sheets. Mobile masses of ice in the form of
glaciers have also developed both in polar and some moun-
tainous regions of the world, and the presence of ice has had
a major impact on the physical environment of the tundra.
Over long periods of time ice can grind away mountains and
create valleys. When ice melts it deposits detritus that cre-
ates a variety of distinctive landforms. Ice in the soil, the
permafrost, is also responsible for the development of pat-
terned ground and strange geological blisters called pingos
and palsas. Soils in the tundra are often unstable because the
freezing and thawing processes place them under great
stress, but in time tundra soils mature to produce distinc-
tively layered profiles, and this maturation is accompanied
by the invasion and stabilization of tundra vegetation.
68 TUNDRA
Above the Earth, the polar atmosphere is the location of one

of the most spectacular light displays to be found on the
planet. It is hardly surprising, then, that the tundra environ-
ment has attracted close attention from both geologists and
atmospheric physicists.
CHAPTER 3
THE TUNDRA ECOSYSTEM

For thousands of years, naturalists have studied the structure
of different plants and animals and the ways in which they
resemble one another. They have observed how living things
grow, move, and reproduce and how they are distributed over
the surface of the Earth. As a result of these studies, scientists
in the 19th century, particularly Charles Darwin but also
many others, began to ask questions about how species inter-
act with one another and how they respond to the physical
environment, including the climate and the soil. The study of
the ways in which organisms relate to their surroundings,
both biological and physical, is called ecology. Central to the
science of ecology is the concept of the ecosystem.
What is an ecosystem?
Living organisms can be looked upon in many different
ways. The simplest approach is to study the individual: its
structure, its biochemistry, or its behavior. But individuals
usually occur in groups, and to understand the individual it
is sensible to study it in the context of the other members of
its species. A group of individuals of the same species is called
a population, and individuals usually modify their behavior
to fit in with the other members of the population. In nature,
however, pure populations of a single species are unusual;
more frequently different species are found mixed up
together. A collection of individuals belonging to a number
of different species is called a community. When different
species are mixed together in this way, they interact. One
species may eat or may act as a parasite upon another. Some
species demand the same resources from their surroundings
and find themselves in competition with one another. Some
species may even inadvertently aid one another, as when a
69
70 TUNDRA
fern grows in the shade and protection of a tree, or when a

bee, as it collects nectar, accidentally carries pollen from one
flower to another. All kinds of interaction are possible
between species in a community.
One further component is involved in the ecosystem, and
that is the physical environment. The ecosystem includes all
of the living organisms found in a location (the community)
plus all of the nonliving materials and factors that form their
background (including the chemistry of the atmosphere, the
soil, and the rocks that underlie the habitat). One could draw
an imaginary line around any patch of the natural world and
regard it as an integrated ecosystem, containing individuals
of different species interacting with each other, all being
affected by the physical and chemical environment and also
modifying that environment by their own presence. A single
tree, for example, changes the light intensity, the humidity,
and even the chemistry of the space beneath its canopy,
affecting what species can grow there.
Certain processes take place within the ecosystem that can
be studied only when all of the living and nonliving compo-
nents of the system are considered together. Energy, for
example, is taken into the ecosystem, usually when photo-
synthetic plants trap the energy of the Sun and use it to con-
vert carbon dioxide gas from the atmosphere into sugars and
then other organic molecules. This process is called primary
production because it represents the first step in the move-
ment of energy into the ecosystem in a form that can be used
by organisms in general. Once energy has been trapped in
this way the plant can use it to help itself perform work, such
as collecting elements from the soil and concentrating them
inside its cells. To perform this energy-demanding task, the
plant needs to release some of the energy trapped by photo-
synthesis, and it does this by respiration, a biochemical
process that resembles a slow, controlled combustion of the
energy-rich sugar within the cells.
The energy that plants bring into the ecosystem fuels not
just the plants but also the animals. Animals are unable to fix
their own energy from the Sun, so they rely on plants as an
energy resource. Herbivores eat the plants; carnivores eat the
animals that have eaten the plants. So there are different lev-
els of consumption within the ecosystem, called trophic levels.

Plants are the first trophic level, herbivores the second, pred-
ators that eat herbivores the third, predators that eat other
predators the fourth, and so on. Few ecosystems have more
than four or five trophic levels. Grass → caterpillar → shrew
→ kestrel is an example of a food chain involving four trophic
levels. In fact, energy rarely moves in simple lines through
the ecosystem but may pass into a range of tracks, forming a
kind of web of different options. This set of relationships is
called a food web (see “Tundra food webs and energy flow,”
pages 72–76).
Not all plants and animals are consumed by predators.
Some die a natural death or, in the case of plants, various
parts—including leaves, stems, flowers, and seeds—may fall
to the ground uneaten. The dead remains of organisms pro-
vide an energy resource for the decomposers of the ecosys-
tem. These are bacteria and fungi, which, like animals, are
unable to fix their own energy but rely on plants for their
supplies. Energy from the plants, therefore, can move either
into the consumer food web or into the decomposer system.
All of these consumers and decomposers, however, use up
the energy they acquire as they work and grow. The energy,
which is made available through the process of respiration,
dissipates in the form of heat as the organisms respire it
away. Once energy is trapped by plants it flows through the
ecosystem, becoming dissipated and dispersed as it flows.
Chemical elements are also involved in food webs, includ-
ing the carbon taken by plants from the atmosphere and the
nitrogen and phosphorus absorbed by plants from the soil.
Plants incorporate these elements into their own structure;
nitrogen, for example, is an essential component of all pro-
teins. These elements, too, move through the food webs as
one organism consumes another, but, unlike energy, these
atoms are not dissipated and lost; they cycle within the
ecosystem. When decomposers finally dispose of dead
remains, the elements contained are released into the envi-
ronment, usually the soil, and they are available to plants so
that they can be reused and recycled within the ecosystem.
Energy flows through the ecosystem from high intensity
(sunlight) to low intensity (dissipated heat), while chemical
72 TUNDRA
elements cycle within the ecosystem. These two processes,

energy flow and nutrient cycling, are fundamental to the
functioning of an ecosystem.
The ecosystem concept can be applied at any scale. It is
often used when scientists study a particular habitat, such as
a pond, an area of woodland, or grassland. But it can be
applied at a much smaller scale, perhaps to a decaying log, a
clump of grass, or even, in the case of medical studies, to the
human gut. Larger units can also be studied using the ecosys-
tem concept, such as a desert, an ocean, or even an entire
planet.
Tundra food webs and energy flow

As in most ecosystems, energy enters the tundra ecosystem in
the form of sunlight. The polar tundra is unusual, however,
in the way sunlight is distributed over the course of time. In
the polar tundra summer, the days become very long and the
nights very short. Between the Arctic and Antarctic Circles
and the poles there is no night at all for part of the year. In
the polar winter, on the other hand, there is little daylight or
none at all for part of the time. Day length in alpine tundra
varies seasonally according to how far north or south of the
equator it is found. Close to the equator there is very little
seasonal variation in day length.
The duration of daylight affects the amount of photosyn-
thesis that green plants can achieve. They continue to respire
during the hours of darkness, so if day length becomes very
short, plants may lose more energy in respiration than they
are producing by photosynthesis, in which case they have to
draw upon stored food reserves to sustain life. Many plants,
therefore, enter a period of dormancy throughout the polar
winter so that they can reduce respiration losses of energy to
a minimum. Even in alpine tundra, plants may respond to
low temperature and snow cover with a winter shutdown
rather than try to maintain low levels of productivity.
The short growing season of the tundra ecosystem means
that the total amount of primary production (the overall
carbon-fixation by green plants) occurring in the course of a
year is relatively small compared to that of other ecosystems.
It is not easy to measure primary production accurately for a

number of reasons. Some of the energy that accumulates is in
the form of roots, and root growth is not easily measured.
Parts of plants become detached and decomposed, so ecolo-
gists have to make allowances for the losses of litter and the
death of roots. Animals, including invertebrates, are con-
stantly consuming plants, so energy is moving along the
food webs all the time and is not left to accumulate in the
growing plants. Despite all these problems, ecologists have
now built up many separate estimates of primary productiv-
ity from the various biomes of the world. Generally these
estimates are expressed in terms of how much dry weight of
plant material accumulates in a specific area of an ecosystem
in the course of a year. Dry weights are used because different
plants contain different amounts of water, regardless of the
amount of energy present. Strictly speaking, it would be bet-
ter to express productivity data in the form of energy accu-
mulated rather than weight, and sometimes this is done, but
weight of plant material is normally used because the varia-
tion in energy content of different plant materials is not nor-
mally a source of great error.
Tundra vegetation, already described, is very variable (see
“Diversity of tundra landscapes,” pages 22–23). There are
locations where vegetation is unable to survive, and these
areas will have no productivity at all. The amount of annual
dry matter productivity found within vegetated Arctic tundra
habitats ranges from 0.25 to 0.42 pounds per square yard
(0.14 to 0.23 kg/m2). Alpine tundra productivity is similar,
usually below a value of 0.5 pounds per square yard (0.3
kg/m2). This is very low productivity compared with a tropi-
cal rain forest, which has an annual productivity of about 5.4
pounds per square yard (3 kg/m2), or even a temperate decid-
uous forest yielding 1.8 pounds per square yard (1 kg/m2). It
is very similar, however, to the value for annual productivity
found in the hot dry deserts of the world. From a productiv-
ity point of view, therefore, the tundra ecosystem is equiva-
lent to a desert.
Ecosystems with low primary productivity generally sup-
port a limited amount of animal life and have a relatively
poor biodiversity, and this is true of the tundra. The diagram
74 TUNDRA
predatory invertebrates Arctic foxes, owls, wolves

(spiders, beetles, etc.) jaegers, gyrfalcons
wading birds invertebrate microbes

detritivores
A food web for a herbivorous invertebrates lemmings dead plant ducks, geese caribou
(leaf hoppers, etc.) litter
tundra community. All
organisms, when they
die, provide food for the
primary productivity by plants
detritivores and (sedges, grasses, willows, dwarf birch, etc.)
the microbes.
illustrates the type of food web that is found within tundra

ecosystems. New plant material is produced as soon as the
light and temperature conditions become favorable in the
spring, and invertebrate grazers, including caterpillars, leaf-
hoppers, plant-feeding bugs, weevils, and grasshoppers,
begin to take advantage of the warm conditions and the
availability of fresh food. Predatory invertebrates, including
carnivorous beetles, spiders, and harvestmen, then con-
sume the increasing populations of invertebrate grazers.
There are also larger animals that take advantage of the
growing numbers of insects, including birds. Some birds,
such as the rock ptarmigan (Lagopus mutus) and willow
ptarmigan (Lagopus lagopus), are residents in the tundra and
feed upon vegetation for most of the year but, in the spring
when their young hatch, they also turn to tundra insects as
a source of food rich in protein for their growing chicks.
Other birds arrive in the Arctic tundra from their winter
locations farther south, including a number of wading
birds, such as plovers (Charadrius species), godwits (Limosa
species), and sandpipers (Calidris species). These take advan-
tage of the long summer days and the rich invertebrate har-
vest to increase their breeding success. Birds that normally
eat seeds, including snow bunting (Plectrophenax nivalis),
Lapland longspur (Calcarius lapponicus), and horned lark
(Eremophila alpestris), also resort to insectivory at this time
of year in the tundra, as they boost their protein intake

prior to egg laying.
Ducks and geese arrive in the Arctic tundra as the days
lengthen, and they feed directly on vegetation but, like the
buntings, also supplement their diet with invertebrate pro-
teins. Caribou (Rangifer tarandus) herds move in from the
south to their preferred breeding locations and consume the
nutrient-rich new growth of vegetation, and small herbivo-
rous mammals, including lemmings (Clethrionomys species),
take the increased supply of fresh food as a stimulus to begin
their breeding cycle.
The increasing numbers of vertebrates provide a food
resource for carnivores, including both birds and mammals.
Predatory birds, such as gyrfalcons (Falco rusticolus), snowy
owls (Nyctaea scandiaca), and rough-legged hawks (Buteo lago-
pus), hunt their prey over the open tundra. Gyrfalcons feed
mainly upon other birds, especially ptarmigan, while the
snowy owl and rough-legged hawk concentrate on the small
mammals, as do arctic foxes (Alopex lagopus). The large graz-
ing mammals, especially caribou, are preyed upon by wolves
(Canis lupus).
This food web is essentially land-based, but there is also a
flow of energy from the marine waters that surround the
polar tundra regions. Primary productivity in the oceans is
mainly due to the growth of microscopic plants that float
in the upper layers of the ocean, called phytoplankton. These
are food for tiny swimming animals, especially crustaceans,
which in turn form the diet of fish and of some of the great
whales. Fish are eaten by seals and also by many seabirds,
and both of these predatory organisms leave the ocean for
breeding and take to the land, either open tundra, cliffs,
and beaches or the pack ice. Here, seals may fall prey to
one of the top predators of the Arctic tundra, the polar
bear (Ursus maritimus), and seabirds provide an additional
source of food for arctic foxes and predatory birds. One
group of birds called jaegers (Stercorarius species) is not truly
predatory, in the sense that they kill their victims, but
they do harass fish-catching seabirds and cause them to
disgorge their stomach contents, which these food thieves
then consume.
76 TUNDRA
Any of these organisms, from the plants to the polar bears,

may escape being consumed by an enemy and may die a nat-
ural or an accidental death. If this happens, then the dead
remains, together with the excreted material of the living
animals, enters the soil and is consumed by detritivores and
decomposers. Earthworms, the larvae of beetles and crane
flies, and tiny springtails all act as detritivores, devouring all
kinds of dead material that still contains residual energy and
making a living from its breakdown. Finally, the fungi and
bacteria in the soil attack whatever remains and extract the
last remnants of energy, converting it into heat.
The low primary productivity of the tundra means that
only a limited amount of energy enters the ecosystem to sup-
port consumers, so food webs tend to be relatively simple
compared with those of biomes that have high productivity,
such as tropical forests. Food webs also expand and contract
according to the season and the availability of energy, so that
the winter effectively means a shutdown of the ecosystem.
Even the decomposers are unable to remain permanently
active because of intense cold. The different living members
of the tundra ecosystem either become dormant or leave the
area when the winter sets in.
Nutrient cycling in the tundra

Chemical elements are durable and they can be constantly
used and reused by living organisms. The element carbon, for
example, which is the most abundant element in living
things, is taken out of the atmosphere by photosynthetic
plants and is constantly being returned to the atmosphere in
the respiration of plants, animals, and microbes. There is an
endless cycle of exchange between the living and the nonliv-
ing components of the ecosystem. The same is true of all the
elements that are found in living bodies.
The atmosphere consists of approximately 80 percent
nitrogen gas and 20 percent oxygen. Small traces of other
gases are present, the most abundant of which is carbon
dioxide, the raw material of photosynthesis, but this com-
poses only about 0.04 percent by volume of the atmosphere,
so it is a very tiny component. Most organisms use oxygen
for their respiration, and there is no shortage of this material

in the atmosphere. Nitrogen is likewise a very important ele-
ment for all life because it is an essential component of all
proteins. But nitrogen gas is highly inert. This means that it is
extremely stable and does not easily enter into reactions with
other chemicals. So, although the atmosphere is an enormous
reservoir of nitrogen, it is effectively unavailable to living
things. Fortunately, certain bacteria have developed biochem-
ical techniques for trapping nitrogen gas and converting it
into the building blocks of protein. These “nitrogen-fixing”
bacteria occur either free-living in the soil or associated with
other organisms, such as higher plants (which house them in
their roots) or with fungi (which combine with the bacteria to
form lichens). The presence of nitrogen-fixing organisms in a
community is extremely important because it enables the
whole ecosystem to avail itself of the nitrogen reservoir in the
atmosphere.
The atmosphere is thus a source of certain vital elements
for the sustenance of life. There are two other major reser-
voirs of chemical elements in ecosystems: One is the soil, and
the other is the mass of living things, the biomass.
Soils in tundra regions are formed by the breakdown of
underlying rock, so much of their chemical composition is
determined by the nature of that rock (see “Soil formation in
the tundra,” pages 55–57). Many of the rocks underlying the
herbivores predators seabirds The cycling of elements

in the tundra ecosystem.
Nutrients move
dung and dead dung and dead feces at nest
bodies bodies and roost sites constantly between soil,
vegetation, grazers, and
vegetation
predators. Excretion,
death, and decay
drainage (conducted by
soil ocean
water detritivores and
decomposers) ensure
rock
rain and that elements can be
snowfall
recycled repeatedly.
78 TUNDRA
A breeding colony of polar tundra are poor in chemical elements, but the fact that
thick-billed murre (Uria they are constantly being shattered by freezing and thawing
lomvia) on a cliff face of means that there is a constant supply of new material into
the High Arctic island of the soil. The dead remains of plants and animals also serve as
Spitsbergen, Svalbard. a source of elements in the soil. As decomposers extract
These birds feed on energy from the organic components of dead material, they
marine fish and then also leave behind residual elements, such as nitrogen, phos-
deposit nutrients in their phorus, calcium, potassium, magnesium, and many others
excreta on the tundra that were once part of the living organism. As they are
landscape at their released, these elements enter the fabric of the soil.
breeding sites. (Photo by Animals deposit some materials into the soil even before
Patricio Robles Gil/Sierra death in their excreta. Urine is a waste product that enables
Madre/Minden Pictures) an animal to rid itself of toxic elements and also those mate-
rials that are in excess supply, especially nitrogen. Predators,
in particular, take in more protein than they need, so their
bodies remove the nitrogen component and convert the
remainder into carbohydrates or fats. The nitrogen then
passes out of the body in the urine. Mammals excrete their

nitrogen in the form of urea, while birds and reptiles void it
as uric acid, but both provide a source of nitrogen to the soil.
Fecal material is more complex because it consists of partially
digested organic materials of all kinds. Like urine, however, it
is rapidly colonized by soil fungi and bacteria, which liberate
many of its component elements and release them into the
soil. Some parts of the tundra ecosystem may be strongly
influenced by the input of urine and feces, particularly those
areas where seabirds roost or breed. In such locations, nutri-
ent elements derived from marine food webs are brought to
the land and deposited in great abundance around nests and
roosting sites, greatly enhancing the nitrogen and phospho-
rus content of local soils.
A further source of chemical elements to the soil arrives in
the form of rain and snow. As precipitation falls through the
atmosphere, it collects dust and other suspended materials,
including crystals of salt or droplets derived from the ocean.
Precipitation thus brings sodium and chlorine from the sea,
together with calcium and magnesium, also present in sea-
water. In tundra regions lying close to the ocean, the input of
these elements can be extremely high, but more isolated and
continental areas receive little deposition of this kind.
Indeed, as noted previously, the quantity of precipitation
over the polar tundra is very low, so nutrient input from this
source will also be small (see “Climate in the polar tundra,”
pages 14–18). Industrial pollution of the atmosphere, how-
ever, penetrates far into the uninhabited regions of the world
and may bring other chemical elements to the tundra.
Radioactive cesium was carried into the tundra regions of
northern Europe and Asia following the nuclear accident in
the Soviet power station at Chernobyl in 1986. More general
in their dispersal are the oxides of nitrogen produced by
industrial processes and the combustion of fossil fuels.
Clearly, then, tundra soils can gain nutrient elements in a
number of different ways, but these elements can also be lost
from soils. Plant roots absorb many of them, expending
energy in order to concentrate the required elements from
the soil and assemble them into new forms within the plant
body. The water that passes through the soil may also take
80 TUNDRA
away some of the elements and convey them into neighbor-

ing aquatic ecosystems in the process of leaching.
The living bodies of plants and animals form the biomass of
the ecosystem, and in most terrestrial ecosystems the vast bulk
of the biomass is in the form of vegetation. Vegetation, there-
fore, represents a major reservoir of chemical elements in most
ecosystems. But the tundra does not support bulky vegetation.
Even at its most luxuriant, where the ground is clothed in
dwarf shrubs of birch and willow, the total amount of dry mat-
ter in vegetation does not often exceed one pound per square
yard (0.6 kg/m2). This compares with 55 pounds per square
yard (30 kg/m2) in a temperate deciduous forest and 75
pounds per square yard (45 kg/m2) in a tropical rain forest. So,
although the vegetation of the tundra is an important reser-
voir of nutrient elements, it is still not a very large one.
The vegetation is grazed by animals, which obtain both
energy and chemical elements from this source of food, and
these elements pass through the food web in the same pat-
terns as the energy. The main difference is that none of the
chemicals are lost to the ecosystem in the process but are
recycled through excretion, death, and decay. The decom-
posers, therefore, play a very important part in the nutrient
cycle; if it were not for their activities chemical elements
would remain locked up in dead organic matter and would
no longer be available for reuse by plants. Decomposition is
particularly important in the tundra ecosystem because the
total quantity of nutrient elements present is low, so recy-
cling is critical. Decomposition, however, is often limited by
two important environmental factors, low temperature and
an abundance of soil water.
In well-drained sites during the warmth of the summer,
decomposition of organic matter in the soil proceeds rapidly,
but in wet sites decomposition is slow because dissolved oxy-
gen is soon consumed. Oxygen diffuses through water 10,000
times more slowly than it does in the atmosphere, so it
becomes scarce, and therefore microbial respiration becomes
slow under waterlogged conditions. Microbial activity is also
affected by low temperature. Just as food lasts longer in a
refrigerator, so organic debris remains intact longer in a cold
soil, because the microbes fail to break it down. The outcome
is that decomposition can be slow at certain times and in

some localities in the tundra, leading to the accumulation of
organic matter as peat in the soil, and locking some nutrients
away from the recycling processes of the ecosystem.
Some ecologists believe that nutrient cycling patterns
influence the cycles of population in tundra animals. Popula-
tion explosions in herbivorous animals, such as lemmings
and ptarmigans, could be related to changes in the availabil-
ity of a vital and limiting element like phosphorus. When
phosphorus becomes available as a result of faster decompo-
sition, vegetation grows more rapidly and produces a richer
source of nutrition, resulting in faster population growth in
the herbivore. This is just one of many suggestions that could
account for population cycles in the Arctic.
Stability of the tundra ecosystem

Some ecosystems are fragile, easily disrupted by disturbance
and slow to return to their original condition. A stable
ecosystem is one that is not easily perturbed by disruptive
forces and can rapidly heal itself if it is damaged. An impor-
tant question concerning the tundra ecosystem, therefore, is
whether it is robust or fragile. Is it a stable ecosystem? There
are two ways of measuring stability: First, one can determine
how much disturbance must be applied to the ecosystem to
unsettle its inertia; and second, one can observe how quickly
the ecosystem recovers from disturbance (called resilience).
Most scientists agree that tundra ecosystems are fragile on
both counts.
The number of different species of plants, animals, and
microbes found in the tundra is relatively low, and ecosys-
tems with few species often lack inertia. The loss of one
species as a result of disturbance, such as a pollution episode,
could have an impact on many other species, especially any
predators that were dependent on the lost species for food. In
a more biodiverse ecosystem there would be other species
that a predator could turn to, but prey-switching options are
limited when the ecosystem is low in diversity.
The soils of the tundra are unstable because of the constant
freezing and thawing that they undergo (see “Patterns on the
82 TUNDRA
ground,” pages 46–53). Soils on slopes are constantly on the

move, and any additional pressures on the environment,
such as the impact of human trampling or, even worse, the
pressure of vehicle wheels, can cause mass instability.
The low biomass and poor nutrient cycling in the tundra
ecosystem can also result in fragility. In general, ecosystems
with large amounts of biomass (with massive reservoirs of
nutrient elements stored in the plant tissues, as in the case of
a forest) are less easily disrupted by minor disturbance events
than ecosystems with low biomass. It is rather like the world
of high finance. A person who has a sum of money to invest
in a bank is likely to be concerned about the size of the assets
of that bank. If it is a large bank with assets of billions of dol-
lars, then the investment is more likely to be safe than it is in
a bank with very limited assets. Similarly with ecosystems:
An ecosystem with large nutrient reserves is less likely to
become chemically bankrupt than one with small reserves.
Tundra ecosystems have very small reserves, so their nutrient
capital is always at risk. The fragility of tundra ecosystems is a
matter of concern to conservationists (see “Tundra conserva-
tion,” pages 182–186).
Conclusions
The tundra can be viewed as an ecosystem because the ani-
mals, plants, and microbes that it contains function as an
integrated unit, interacting with their chemical and physical
setting. Using the ecosystem approach to the study of tundra,
it is possible to trace the processes of energy flow and nutri-
ent cycling that link the living and nonliving components of
the system.
The energy input from the Sun is relatively low in tundra
ecosystems when compared with those of all other biomes,
and it is strongly seasonal in its distribution. Polar sites
undergo a prolonged winter in which energy input is very
low or completely absent. Low temperature in the winter sea-
son reduces the primary productivity in both polar and
alpine tundra. As a result, energy flow in the tundra ecosys-
tem is limited and consequently there are relatively few links
in food chains and food webs. The outcome of these energy
limitations is that the number of different types of living

organisms that can exist in tundra is restricted.
Nutrient cycling between vegetation and soil is limited by
the rate of decomposition, and this is affected by temperature
and wetness. Under conditions of low temperature or water-
logging there is reduced microbial activity and hence low
decomposition. As a result, some elements are locked up in
dead organic matter in the ecosystem and are not released for
recycling. This slows down the process of nutrient cycling
and may restrict plant growth and consequently herbivore
abundance.
The low biodiversity, soil instability, and restricted nutrient
cycling combine to make the tundra ecosystem relatively frag-
ile. It lacks inertia, so it is easily damaged, and it is not very
resilient, so it does not recover quickly from disturbance.
CHAPTER 4
BIODIVERSITY
OF THE TUNDRA
Life in the tundra is both harsh and bleak. It is not surprising,
therefore, that it contains a relatively low diversity of plants,
animals, and microbes. Few organisms are able to survive in
the difficult conditions that prevail in the tundra, but those
that do so are of considerable ecological and evolutionary
interest because of their high degree of specialization. Tundra
plants and animals have to be tough to endure the problems
that face them and they have developed some unusual fea-
tures that help them to survive.
Living in the freezer

The cold is the most obvious problem in the tundra. It is not
a constant cold; indeed, the temperature in summer on the
ground can rise to more than 80°F (26°C), and the high angle
of the Sun over tropical mountains can take the daytime tem-
perature well above these levels. But the average temperature
is low, and all chemical reactions take place more slowly at
low temperatures. This means that the enzymes, the chemi-
cals within all cells that control all operations, are not able to
work as fast as they could at higher temperatures. In plants
and in animals that are “cold-blooded” (that is, animals that
are unable to control their own temperature and simply
adopt the temperature of their environment), all activities
slow down when the temperature of their surroundings is
low. They grow more slowly, they reproduce more slowly,
and, in the case of mobile animals, they move more slowly.
All of these factors reduce the ability of an organism to com-
pete with its neighbors or to evade attack by a predator. So
cold makes the tundra a hard place in which to survive.
In the case of the polar tundra, the habitat is actually dark
for a large portion of the year because the Sun never rises.
84
This is one of the major reasons why average annual temper-

atures are low, but it provides an additional problem for
plants, which depend upon sunlight for the energy needed in
photosynthesis. Without light, the green plant is unable to
continue its growth. The tundra plant, therefore, must be
able to survive in a dormant form in this darkness. Animals
that require light to find their food will similarly be disad-
vantaged by darkness. They can either copy the plants and
shut down for the polar winter or adopt the alternative strat-
egy of migrating away from the harsh conditions until things
improve in the spring.
Winds can be strong in polar and alpine regions, and they
can add to the problems of cold by removing the warm air
layers that may cover the surface of an organism. Wind chill
further reduces the temperature of both cold-blooded and
warm-blooded organisms by stripping these insulating layers.
The development of hair on both plants and mammals helps
to protect this precious layer of air from being disturbed by
the wind. When snow has settled on the ground and has
undergone changes by melting and refreezing, it produces
small ice particles that can become airborne in the strong
winds. The result is that suspended particles of ice are swept
along by the moving currents of air, and these have a biting
and penetrating impact on any living things, from shrubs to
musk oxen. The ice-blasting of these winds can strip away
leaves and skin, leaving plants and animals damaged and sus-
ceptible to infection.
Burial by snow can be either helpful or harmful. It adds to
the effects of darkness, so green plants buried beneath snow
patches have a shorter period in summer when they can grow
and reproduce. Relatively few plant species are able to cope
with this kind of stress, some mosses being among the most
tolerant. On the other hand, a snow cover insulates buried
organisms from the worst effects of the cold because heat
transfer from the ground to the cold air above is slower when
there is a snow layer. Hibernating animals and dormant
plants are protected from extremely low temperatures and
from the impact of ice-blasting while they remain buried.
Drought may also be a problem in polar and alpine tundra
habitats. Where the soils are rocky and free draining, water
86 TUNDRA
passes quickly through the soil, so plants and any inverte-

brate animals with low mobility (such as snails and earth-
worms) can be subjected to desiccation, especially if the
rainfall or snowfall is low. In montane areas, precipitation is
usually high, but the polar regions have low precipitation
and the soil can easily become dry. Tundra plants, therefore,
often exhibit adaptations to drought conditions, including
evergreen leaves with tough, waxy coverings. The cold can
also make water difficult to obtain, especially when it is
frozen in the soil. Plants then experience what is known as
physiological drought, meaning that there is plenty of water
about, but it is in the wrong form or cannot be easily
accessed in the soil. Again, the plant is subjected to water
stress.
All of these problems add together to make the tundra one
of the most difficult environments for plants and animals to
live in. The result is that tundra has low overall biodiversity,
but the species that do manage to cope are often highly
adapted and specialized in their structure and physiology.
They are often restricted to the tundra habitat simply because
they have become so specialized that they cannot compete
with the organisms belonging to any other type of habitat.
So the tundra plants and animals represent a distinct group,
most of which are found in none of the other biomes of the
world. In this way they contribute a very distinctive aspect to
the world’s biodiversity.
Tundra plants
The impact of cold and wind blasting has led to the evolu-
tion of a distinctive plant life-form in the tundra. Most plants
are of a type called chamaephytes. These are small shrubs and
perennial herbs that grow to a height of just a foot (25 cm) or
so and often take the form of a cushion or blanket over the
surface of the ground. Thus they hold their buds above the
soil surface, yet close to the ground. The advantage of this
structure is that the wind moves over the top of the low veg-
etation. Any shoot that emerges above the general level is
damaged and trimmed by the icy blast so the growth is held
back. Within the vegetation blanket, the temperature is
maintained at a higher level than either above the plant

cover in the cold air, or below in the frozen soil of the per-
mafrost. The microhabitat created by this life-form is of great
importance to invertebrate animals that can find shelter
beneath the plant (and sometimes snow) canopy at the sur-
face of the litter and soil.
Many species of saxifrage provide good examples of cush-
ion-forming chamaephytes. They produce a dense hemi-
sphere of evergreen leaves, and the individual plants often
coalesce into hummocky carpets. Beneath the dense surface
layer of green leaves is a mass of fibrous stems and dead
leaves that provide shelter for a host of invertebrate animals.
Dwarf shrubs, such as the arctic heather (Cassiope tetragona)
and the mountain cranberry (Vaccinium vitis-idaea), also form
dense hummocks and carpets over the surface of the ground,
and even the small Arctic trees, such as the dwarf birch
(Betula nana) and willows (Salix species), rarely raise their
heads far above the general canopy.
Just as the aboveground parts of plants do not extend far
from the soil surface, so the roots do not penetrate deeply
into the ground. Often the soil is poorly structured and rocky
(see “Soil types of the tundra,” pages 57–61), hindering root
penetration, but even where peaty soils provide easier access,
the permanently frozen layers prevent deep root develop-
ment. So most of the activity of roots in the soil, including
water absorption and the uptake of nutrient elements, takes
place in the uppermost few inches. Root growth begins in the
spring as soon as the soil thaws, and continues throughout
the growing season. Most of the sideways growth of roots
takes place late in the season. These lateral roots allow the
plant to forage for nutrients and water in new areas and may
also allow the plant to spread.
Protection against drought, either true drought or the phys-
iological variety (that is, lack of water availability because of
intense cold), demands certain modifications to plant form.
One of the most conspicuous parts of a plant in the tundra is
the leaf. Evergreen leaves predominate among tundra plants,
so these must provide some advantages in the tundra habitat
(see sidebar on page 88). The deciduous habitat is an alterna-
tive approach to the problem of survival during drought, for
88 TUNDRA
Evergreen or deciduous?
Most Arctic and alpine plants are perennial and evergreen. The saxifrages, cranberries,
heathers, and willows are all perennial plants. The likely reason for this is that the growing
season is so short. As soon as the Sun appears in the spring and the temperature begins to
rise, the plants need to start their growth. Being evergreen is an advantage because the
leaves that perform the photosynthesis are already present and able to begin their work.
Some plants, including the willows and birches, are deciduous, and they need to develop
new leaves as soon as conditions improve in the spring. They do this very rapidly but still
lag a little behind their evergreen competitors. Where they gain some advantage is in their
smaller bulk during the harsh winter. Stems without leaves are less likely to be damaged by
wind blasting than those that still bear a leafy cover. So there is a kind of trade-off between
keeping leaves through the winter and losing them, each strategy having certain advan-
tages and disadvantages. On the whole, however, the evergreen habitat probably proves
the more effective and successful. Evergreen leaves have one additional advantage: Con-
structing them costs less. When a plant builds a new leaf it has to invest some of its food
reserves, including the sugars produced by photosynthesis and the nutrient elements, such
as nitrogen, phosphorus, and potassium, taken out of the soil by the roots. Evergreen
leaves last longer, perhaps two or three years, so the investment is more worthwhile.
the presence of fewer leaves during fall and winter means that
the plant requires less in the way of water supply. This is one
of the great advantages of being deciduous. Evergreen leaves
could result in water loss to the plant through the whole year,
so they need to be constructed in such a way that water loss is
reduced to a minimum. The long-lasting evergreen leaves are
usually thick, rounded, and glossy with wax, and they may
bear a cover of hairs. All of these adaptations serve to reduce
water loss by cutting down the evaporation of water (transpi-
ration) from the pores upon the leaf surfaces. In addition to
reducing water loss, some of these adaptations, such as silvery
hairs on the leaf surface, can also serve to reflect light and to
prevent the leaf from becoming heated. If a leaf becomes hot
it loses water faster, so keeping the leaf cool on a hot summer
day can be important even in the tundra.
One method some plants adopt in order to avoid periods
of great stress is to die. It sounds like an extreme solution to
the problem, but the plant may leave behind a reserve of

seeds in the soil that can germinate when conditions are
more favorable once more. Annual plants of this kind, how-
ever, are extremely rare in the tundra. One annual plant that
has proved successful is the Iceland purslane (Koenigia
islandica). This exceptional little plant, only two inches (5 cm)
high, is found in the Arctic and subarctic, in the mountains
of Asia and the Rocky Mountains of North America, and on
the southern tip of South America, in Tierra del Fuego. So it is
clearly a very successful plant of tundra habitats despite
being an annual. The problem with being an annual plant is
that the entire life cycle (germination, establishment,
growth, flowering, and fruiting) has to take place in a single
growing season. Where the season is very short (as in the
tundra) this can be extremely difficult, so annuals are rare.
But where there is perpetual daylight, a very fast-developing
annual can make it, and the Iceland purslane fits this bill. It
grows in open rocky areas and at the end of the summer sea-
son it dies completely, leaving only its tiny seeds in the sur-
face layers of soil. These survive the rigors of winter beneath
a blanket of snow and begin the cycle again the following
spring.
A feature of Arctic and alpine plants that has made them
extremely attractive to gardeners is that they often bear very
large flowers in comparison to their small stature. The purple
saxifrage (Saxifraga oppositifolia), for example, is a tiny cush-
ion plant that has large purple flowers, and these flowers are
larger in the most northern populations of the Arctic than
they are in more southerly regions. The brilliant blue gentians
(Gentiana species) of the European Alps and some poppies,
such as the arctic poppy (Papaver radicatum), have extremely
large and showy flowers. Since the general tendency of tundra
plants is to maintain a low profile, it may seem strange that
they often have such conspicuous flowers. The reason is prob-
ably their need to attract insect pollinators in an environment
where time is short and competition for the limited number
of appropriate insects is high. Flowers are a kind of advertise-
ment that demands the attention of passing insects and, like
the billboards along a highway, each flower is trying to out-
compete its neighbors in visual attraction. In such situations
90 TUNDRA
The spring gentian large flowers with bright colors are likely to succeed best. Some
(Gentiana verna), flowers may even turn during the course of the day, always fac-
growing on the ing the Sun. In this way they gather heat, which further
mountains of encourages insect activity (see sidebar on opposite page).
Switzerland, blooms as The high mountains of the Tropics are a home to some
soon as the winter snow remarkable plants. Many of the typical Arctic-alpine plant
has melted. As with species of the high latitudes have failed to colonize these iso-
many alpine plants, its lated patches of tundra habitat, far from the main regions of
flowers are large and the tundra biome. The outcome is that some types of plants
brightly colored in order that are not usually associated with tundra have evolved some
to attract the attention features that enable them to survive in the cold conditions
of pollinating insects. that they find there. Among these are members of the genera
(Photo by Senecio (a group that includes the ragwort) and Lobelia (a group
Peter D. Moore) that includes the cardinal flower and Indian tobacco). Both of
these plant groups are generally herbaceous (that is, they have
no woody tissue) but in the tropical tundra habitats they have
taken on a most remarkable form (see illustration above). They
Solar-tracking flowers
The flowers of many tundra plants have the capacity to track the Sun across the sky. Often
their petals have the curved shape of parabolic reflectors that can gather sunlight and
focus it upon the vital part of the flower, the fruit-producing carpels. In this way they can
ensure that the development of their reproductive structures gains as much energy as
possible. But the energy-focusing effect of the petals is also a means of making life easier
for the pollinators. Most pollinators in the tundra are cold-blooded insects, whose activity
is determined by their immediate energy intake, often largely from the Sun. So providing
the insects with a hot flower ensures that they can work faster and provide a more effi-
cient pollination. Poppies are an excellent example of this use of flower shape to encour-
age visits from cold pollinators. The visual outcome of the flower’s need to attract
pollinators is that many tundra habitats, especially the high mountains, are decorated by
a wide variety of large and colorful flowers through the summer season.
In the cold alpine climate some flowers, such as this alpine pasque flower (Pulsatilla
alpina) in Switzerland, track the Sun across the sky and gather warmth by adopting a
parabolic shape that focuses the rays on the flower’s center. (Photo by Peter D. Moore)
92 TUNDRA
The alpine zone of high produce thick, treelike trunks that are clothed with a compact
mountains in equatorial mass of dead leaf bases that wrap around the trunks like fur
regions is home to some coats. Like animal fur, this fibrous covering forms an insulat-
remarkable plants. ing layer around the living cells of the stem. Although frost
Giant senecios, such as damage is a problem that plants of the Tropics do not nor-
the species Senecio mally face, at night the air temperature at 13,000 feet (3,800
elgonensis shown here m) can fall below freezing even on the equator. The layer of
on Mount Elgon, leaf bases on the trunks of the giant Senecio and Lobelia plants
Uganda, East Africa, are ensure that even if the air temperature falls to 25°F (–4°C), the
treelike forms of a plant temperature beneath the insulating layer, where the living
group that is normally cells are found, remains at about 36°F (2°C). For a tropical
herbaceous. (Photo by plant, this is the difference between life and death.
Peter D. Moore) Coping with the cold is evidently the most serious problem
for tundra plants. Unlike mammals that can migrate or hiber-
nate beneath the snow and maintain a living body tempera-
ture by their biochemical activities, plants (together with
microbes and cold-blooded animals) assume the temperature

of their surroundings, and this can be very low. That is why
many are clothed in wrappings of dead leaves or high densi-
ties of hair. The real problem for living cells when they are
exposed to intense cold is that their high water content puts
them in danger of freezing. If an automobile is left out in
severe cold without the protection of antifreeze compounds
in its cooling system, then the water in its radiator and pipes
will freeze. The driver may not notice this until the thaw
comes, and then it becomes obvious that the pipes have burst
because all the water leaks out. The problem is, of course, that
water expands when it freezes and thus ruptures the pipes,
but this becomes evident only when the temperature rises. It
is the same with plants. The freezing of a cell involves ice crys-
tals forming inside the cell itself, and the frozen water occu-
pies a greater volume, so it may begin to damage the structure
of the cell, particularly the membranes that surround the del-
icate organelles within the cell (nucleus, chloroplasts, mito-
chondria, and so on). When the thaw comes, the damage
becomes evident and the ruptured membranes often lead to
the death of the cell.
Just as people use antifreeze to prevent damage to their
cars, plants have developed antifreeze compounds that can
help them cope with low temperature. Most chemicals, when
dissolved in water, lower its freezing point, acting as a sort of
antifreeze. Common salt, for example, is used in this way on
highways to reduce the impact of frost. This explains why the
sea freezes only when the temperature is well below that
needed to freeze freshwater. But salt is not a good material to
use as antifreeze either in automobiles or in living cells. It is
corrosive in the metal tubes of an engine, and it causes dam-
age to enzymes in a cell. In the cell, it can be accumulated in
the vacuole, which is a kind of reservoir carefully enclosed
and separated from the working parts of the cell by a mem-
brane, but it cannot be used in the cytoplasm, the living and
working part of the cell. Other compounds, however, can be
used, such as certain amino acids, which are the component
units of proteins. The amino acid proline, for example, serves
as antifreeze in some plants as well as in some fish. Addition-
ally, some plants use sugars and sugar alcohols, such as glyc-
erol, mannitol, and sorbitol. In these ways, plants can lower
94 TUNDRA
the freezing point of their cell contents and increase their

chances of survival.
Among the plants that cope best in tundra habitats are the
mosses. This may seem surprising, but despite their small
stature and delicate structure, mosses are actually very tough.
They have no waxy covering to prevent their leaves from des-
iccating, but many species are able to survive in a dry state for
years without losing their ability to recover and grow once
again. They can also continue to photosynthesize and grow at
lower temperatures than is possible for most flowering plants.
The moss Bryum argenteum, for example, can continue to pho-
tosynthesize below the freezing point of water. The success of
the mosses in tundra habitats can be judged by the fact that
the whole of Antarctica contains only two species of flowering
plants but more than 100 species of mosses. Biologists often
refer to mosses as being “primitive” in the sense that they are
structurally simple and have a low position on the evolution-
ary scale, but this does not mean that they are unable to com-
pete with more advanced plants when conditions are tough.
Microbes of the tundra

When people want to preserve foodstuffs over long periods,
they store them in a refrigerator or, better still, in a freezer.
This is to prevent bacteria and fungi consuming the food. Just
like humans and other animals, most bacteria and all fungi
obtain the energy they need for life from organic materials
that have ultimately been assembled by green plants. If food
lies unattended in warm temperatures, then microbes (bacte-
ria and fungi) soon make use of it and, from the human point
of view, it decomposes. When people are cunning enough to
put food in very cold containers, most bacteria and fungi are
unable to attack it because their biochemistry, like humans’, is
adapted to operate in warmer temperatures. For much of the
year, the tundra is like a gigantic freezer, and almost all micro-
bial activity ceases. But in the summer, the soil, where most of
the microbes live, becomes warmer, the water within it melts,
and the microbes take up their activity once more.
There is plenty of food for microbes in many tundra soils.
When the spring comes, the soil is usually rich in dead plant
material derived from the previous year’s growth, or even
from before that. Some of the organic materials in tundra soils

may date back many thousands of years and yet still lie only
partially decomposed. This is why the remains of humans
buried in the tundra during early expeditions of discovery
have remained intact and have occasionally been recovered
by more recent travelers. It is why the remains of mammoths
are still being recovered from the Siberian tundra; Russian
restaurants have even been known to serve mammoth steaks.
So the microbes are well supplied with food, but the physical
conditions of the winter prove too severe for most of the bac-
teria and fungi found in temperate, warmer soils, and only
very few species are present to begin their spring activities.
The microbial biodiversity of the tundra, in other words, is
very low when compared with that of warmer climates. But
those microbes that are present perform very valuable tasks,
enabling nutrients to cycle so that they are not locked up per-
manently in dead matter, and fixing nitrogen from the atmos-
phere, which is so vital to both the plants and animals of the
tundra (see “Nutrient cycling in the tundra,” pages 76–81).
Very little is known about how the cold-loving microbes
manage to function at low temperature. Some, of course, sur-
vive the extreme cold as resistant spores and thus avoid being
desiccated or having their cell membranes ruptured by the
formation of ice crystals. Others may avoid the problem
completely by becoming extinct in a site each winter and
then invading again by aerial dispersal every spring. The very
small size of the dustlike spores and their capacity for long-
distance movements in the air would easily allow such rein-
vasion every year. But some microbes do manage to maintain
activity even at very low temperatures, and these probably
survive the tundra winter. The enzymes (proteins in the cells
that manage most of the cell processes) of these cold-loving
microbes must be able to function at low temperature and
avoid the molecular breakdown that would occur in most
plant and animal cells under those conditions. Microbes use
antifreeze compounds in a similar way to some tundra
plants. They may accumulate sugars, or glycerol, or various
fats in their cells that lower their freezing point, and in this
way they can remain unfrozen even at temperatures of 5°F
(–15°C). Understanding how microbes operate at low temper-
ature might enable scientists to use genetic manipulation to
96 TUNDRA
construct plants and animal that would similarly grow and

function under low temperature. There could be opportuni-
ties here for developing a cold-climate agriculture that would
help to feed the world. Such an understanding could also be
of great importance in the food storage industry in trying to
prevent the destruction of frozen foods by certain microbes
and thus increasing the storage life of foods.
One very important group of microbes in the tundra is the
blue-green bacteria (often referred to as blue-green algae, which
is not strictly accurate). As their name implies, these are pig-
mented (colored) microbes, and they use their pigments to
catch the energy of sunlight and to fix carbon dioxide in pho-
tosynthesis. They therefore contribute to the energy-fixing
process in the tundra community. Because of this, just like
green plants, they are able to live only in locations that receive
light. They are found mainly on the surface of the soil, or in
association with fungi in the lichens on the rocks or on the
ground. Not only do they create organic matter from inorganic
carbon by photosynthesis, they are also capable of taking nitro-
gen from the atmosphere and incorporating it into organic
molecules, including proteins, so that it then becomes avail-
able to plants and animals. The blue-green bacteria, therefore,
play a vital role in the functioning of the tundra ecosystem.
Some blue-green bacteria associate with fungi in the formation
of lichens (see sidebar on opposite page), which are also impor-
tant components of many tundra communities.
Lichens, judging by their abundance, clearly have great
advantages in the tundra habitat. They contain photosyn-
thetic organisms (the algae or the blue-green bacteria), so
they need light to supply the energy they require. They also
have a fungal component, and the function the fungus serves
is to provide a protective structure in which the photosyn-
thetic component can operate. So, most of the crusty, leafy,
or branched parts of the thallus (the body of the lichen) are
composed of fungal tissue. A close look under a microscope
shows that the fungus consists of a mass of interweaving
strands, forming a spongy tissue, within which the green
cells of their associated species are located. The spongy fun-
gal tissue is highly efficient as protection against the ele-
ments, particularly against the impact of frost and drought,
the two chief problems that face isolated algal cells in the
Lichens
Lichens are both abundant and very apparent in the tundra regions. The lichen is a
remarkable construction that looks like a single organism (and is classified and named as
such by biologists) but is in fact a combination of two separate organisms, closely bound
in an interdependent union. The combination consists of a fungus and either a green alga
or a blue-green bacterium.
The body of the lichen (the thallus) can take on a variety of forms. It may form a crust,
almost like paint, on the surface of a rock, or it may take on a leaflike form, curling up at the
edges, or it may even have stalklike structures that stand up from the ground and may
branch like miniature trees. Even the tallest of lichens, however, are rarely more than two
inches (5 cm) in height. But since they can grow in dense colonies and can form compact
mats over the surface of the ground (especially over shallow and dry soils), they may form
the main vegetation cover in some of the polar semidesert regions of the tundra. Where
there is taller vegetation, such as dwarf shrubs, lichens may grow on the surface of the
shoots and branches of their hosts. When they adopt this way of life, they do not act as par-
asites but simply use their host as a means of support and of escape from the shade on the
ground beneath. An organism that uses a plant for support in this way is called an epiphyte.
Epiphytism is a very common way of life in tropical rain forests, where obtaining light can be
a severe problem, but it is found even in the tundra habitat if shrubs are present.
tundra. The alga (or blue-green bacterium) thus provides the

food by photosynthesis, and the fungus provides the protec-
tion in return for a share of the food. If the photosynthetic
component is a blue-green bacterium, then the lichen is also
a means of nitrogen fixation, so this is a source of nutrition
for many other tundra organisms.
Many lichens are strongly pigmented and have bright colors;
orange is particularly common. The pigments protect the deli-
cate cells from the high intensity of light, especially ultraviolet
radiation, in the clear air of the tundra. One disadvantage of
lichens is that they are very slow growing, so they are sensitive
to high levels of grazing (by animals such as caribou) and to
trampling that can crush and damage their branching struc-
tures. Despite their sensitivity to physical damage, lichens are
tough under cold conditions, as illustrated by their ability to
continue photosynthesis in very low temperatures. The lichen
Umbilicaria aprina, for example, is still photosynthetically
98 TUNDRA
active at 2°F (–17°C). So lichens exceed even the mosses when

it comes to coping with the cold.
Although microbial biodiversity in the tundra is low, the
value of these organisms for ecosystem function can be very
high. People often evaluate habitats according to their biodi-
versity. Tropical rain forests and coral reefs are held in high
regard, for example, because they are so diverse, and this is rea-
sonable. But the tundra is valuable despite its low biodiversity
because it contains such remarkably well adapted species; they
are packets of genes that are found nowhere else on Earth.
Tundra invertebrates
Invertebrate animals (animals without backbones) are
extremely numerous in almost all the habitats of the world,
and the tundra is no exception. All invertebrates, however,
have the disadvantage of being cold-blooded, which means
that their body temperature is largely determined by the tem-
perature of their surroundings. In a cold habitat, such as the
tundra, this limits the activities that invertebrate animals can
undertake, especially during the winter season. Invertebrates
are generally much smaller than vertebrates, so it is useful to
try to see a habitat from an invertebrate’s point of view. For
an animal the size of an ant, the mat of lichens and mosses
on the surface of the soil appears as a forest would appear to
a person, having a high canopy and branches and a sheltered
floor beneath that lies in its shade. Within this microhabitat,
therefore, the conditions may be very different from those in
the air up above. The temperature will fluctuate less, because
the trapped air layer beneath the canopy neither heats up so
much in the day nor cools down so rapidly at night. The
humidity also remains high, which is an important consider-
ation for many invertebrates that lose water easily through
their body surfaces.
The soil itself is also an important habitat for invertebrates
within the tundra, for the surface layers are usually warm and
moist during the summer months. Many different types of
invertebrates manage to make a living here. There are two
types of worm, for example, the nematodes (phylum Nema-
toda) and the enchytraeids (phylum Annelida). Nematodes
are among the most numerous and diverse of all animal
groups. In tundra soils as many as 5 million have been

recorded within 1.2 square yards (1 m2). They have bodies
that are not divided into segments and they have a very wide
range of lifestyles. Most are detritivores, which means that
they feed upon dead organic matter, mainly derived from the
fallen litter of plants. Some feed upon algae in the soil, or
even on soil bacteria. But there are others that prey on other
small invertebrates or are parasites upon larger animals,
including mammals, or on higher plants. Some eat fungi,
including lichens, but there are some fungi that reverse the
feeding chain and trap nematodes in ringlike snares, thus
killing them and feeding upon their decaying remains. Some
nematodes are highly tolerant of low temperature. The nem-
atode worm Panagrolaimus davidi, for example, remains
unfrozen even at a temperature of –40°F (–40°C) because of
the antifreeze chemicals in its body. The enchytraeid worms
are also largely detritivores, but they differ in having bodies
that are divided into segments, rather like the common
earthworm but very much smaller. They are particularly
abundant in organic, peaty soils.
The springtails (Collembola) are another group of detritiv-
orous invertebrates. These are tiny, wingless insects that gain
their name from a forked spine at the rear of their bodies that
can flick outward, allowing the animal to spring into the air.
This usually occurs when they are disturbed, so this trick pre-
sumably functions as a means of escaping a predator. The fact
that springtails cannot fly raises the question of how these
small creatures are so widely dispersed in the Arctic, often on
isolated islands. Recent studies have shown that they are
capable of surviving for long periods on the surface of the sea
and can be dispersed in this way. Their bodies are covered
with very small hairs and scales that make them virtually
incapable of becoming wet, so they float on the water surface
instead of sinking. This can be observed when water is added
to a potted plant, because springtails are usually present in
the soil and they often float on the surface of the water that
has been added. So springtails can disperse on ocean surfaces
to distant islands and coasts; mountain populations of
springtails with an Arctic distribution, however, are more dif-
ficult to account for. It is likely that these were left behind
after the last ice age. One species of springtail, Tetracanthella
100 TUNDRA
arctica, is found in Greenland and Iceland and also in the

Pyrenees Mountains of Spain and France. It is unlikely that it
spread from the Arctic islands to the continental mountains,
but perhaps it was once much more widely distributed and,
as the ice retreated and forests spread, this cold-loving species
of springtail became confined to the isolated peaks and the
high-latitude islands.
The tundra contains many wetland habitats, including
pools and peaty mires (see “Diversity of tundra land-
scapes,” pages 22–23). These are rich areas for many inver-
tebrates, including midge and mosquito larvae (both in
the order Diptera). Some of the midge larvae, such as the
chironomids (family Chironomidae), can also live in soils.
Any human visitor to the tundra in summer is bound to be
impressed by the abundance of biting insects; clouds of
mosquitoes arise whenever vegetation in the wet tundra is
disturbed. Only the female mosquito takes blood, and she
requires this in order to lay her eggs and complete her life
cycle. Fortunately, the species of mosquito found in the
tundra do not convey any parasitic diseases, unlike their
tropical relatives, but their bites can be extremely irritating
and can cause blisters and inflammation. Although people
tend not to appreciate this abundance of insect life in the
Arctic, many birds do. Several bird species move into the
tundra to exploit this source of food during their summer
breeding seasons (see “Tundra birds,” pages 102–111).
Plant-feeding insects are found in the tundra, including
some beetles (especially weevils; family Curculionidae) and
also the crane flies (family Tipulidae). Some plant-eating
insects, such as the sawflies (suborder Symphyta), lay their
eggs in the leaves of dwarf willows, and as the larva devel-
ops, the leaves produce extra tissues, forming grotesque
swellings, or galls. Eventually the mature insect emerges
from these, mates, and goes on to lay eggs in new leaves.
Many flies and butterflies serve important functions as polli-
nators for the flowering plants, and they obtain their food
from the supplies of nectar or surplus pollen from these
plants. There are also surprisingly many moths that con-
tinue the pollination process during the tundra night. One
of the most spectacular butterflies of the alpine tundra is the
Apollo (Parnassus apollo). It is a European species but is a
member of a group of butterflies found in North America,

Europe, and Asia, into the Himalayas. Black and white in
color, it has large spots on its hind wings that look like eyes,
and these serve to distract a predator when they are sud-
denly flashed as the insect opens its wings. Both the wings
and the body of the Apollo are densely hairy, and this is a
feature of the butterflies of alpine and Arctic habitats, help-
ing to conserve warmth. The large wings provide the Apollo
with strong flight and the ability to soar rather than flutter.
These qualities are very advantageous in the mountains,
where winds are often strong and flight for a light organism,
such as a butterfly, is risky. Indeed, many insects in the
mountains have lost the ability to fly as flight can be a
source of danger rather than an aid to dispersal.
Another butterfly family that has proved successful in both
Arctic and alpine habitats is the Satyridae, which includes
meadow browns and wood nymphs, found both in the Old
World and the New World. These butterflies are good exam-
ples of the predominance of dark colors in the tundra insects.
While this seems to go against the trend found in the birds
and mammals, which are often white or pale colored for
camouflage, it must be remembered that butterflies are active
only in the summer period, when snow is less abundant and
both the ground and the vegetation are dark. Also, dark col-
ors are more effective at absorbing heat from the Sun, and in
a cold-blooded animal this is an important source of energy.
The generally small size of tundra insects may also be related
to energy transfer (see sidebar on page 102). A small insect
has a larger amount of surface per unit of body size, so if it
basks in the sunshine it can quickly transfer heat to all its
parts. Mammals, on the other hand, have evolved very differ-
ent strategies. Because they are warm-blooded it is heat loss
that concerns them, not heat gain, so a small surface area per
unit volume is preferable and most mammals of the tundra
are large and relatively rounded in shape (see “Alpine tundra
mammals,” pages 121–126).
Once an adult insect emerges in the spring, the process of
locating a member of the opposite sex, mating, and egg lay-
ing becomes urgent, and some insects have developed a
shortcut for avoiding the risks involved in the mating
process. Blackflies, many midges, stone flies, and sucking
102 TUNDRA
Energy and tundra insects

Energy is the controlling concern of tundra insects. Even the flies that are responsible for
the consumption of dead bodies of birds and mammals lay their eggs first on the side of
the carcass that faces the Sun, for the larvae develop faster within the heated meat. But-
terflies and flies bask wherever they find a combination of sunshine and shelter from the
wind. Within the cups of flowers facing the Sun is a favorite location. Of all the insects,
only some bumblebees have mastered the art of generating a little heat of their own.
They vibrate their wings violently, causing an intense buzzing noise and building up heat
by friction. Some flowers have developed a mechanism of pollen release that relates to
this frantic buzzing, the pollen-bearing anthers rupturing in response to the resonant
vibration created by the bee and showering it with pollen.
Although insects have adapted to take advantage of any opportunity to obtain solar
energy in the tundra, the predominant temperature is low. For this reason many of the
processes of growth and development have to take place in the cold. In the case of mos-
quito larvae, for example, growth and development from egg to adult can take place in
temperatures just above freezing point. Even so, development is slow and many insects
may take two years to complete their growth and reproductive cycle instead of the usual
one. The larvae must therefore develop mechanisms for survival through the winter,
which involve the use of antifreeze chemicals and sometimes the desiccation of their bod-
ies into a torpid, dormant state.
bugs have virtually abandoned sexual reproduction. Males

are extremely rare or absent, and the females lay eggs requir-
ing no fertilization. This is obviously a simpler and faster
reproductive system, similar to the vegetative propagation of
plants. But it has the disadvantage that the offspring are all
clones of the parent, so new variations are not being intro-
duced into the population. There is a trade-off between
genetic diversity and speed of replication, and the system
could misfire if there are sudden changes in the environ-
ment, such as climate change.
Tundra birds
The long days and the abundant supplies of insects and
berries, together with the availability of seafood along the
coasts, make the Arctic a very attractive place for migrant

birds to breed. The Antarctic also has an abundance of
marine food resources, but the generally poor vegetation lim-
its its possibilities for birds that do not depend on the ocean.
Mountain tundra also provides local food supplies from
alpine vegetation, but it generally lacks the oceanic advan-
tages and certainly cannot supply the long daylight hours for
feeding the young, unless the mountains happen to lie
within the Arctic Circle. So the tundra, especially the Arctic
tundra, attracts immigrant bird populations during the sum-
mer months. Alaska has 113 species of breeding birds, while
the Canadian Arctic has 105 species and the Russian Arctic
136. These are quite high numbers considering how far north
these regions lie. But very few of these bird species remain in
the Arctic through the winter.
The willow ptarmigan (Lagopus lagopus) and the rock
ptarmigan (Lagopus mutus) are good examples of the truly
Arctic residents. These two birds are found right around the
Arctic and subarctic regions of the world and are difficult to
distinguish from each other. Both are almost pure white in
winter, with just a little dark on the tail, while in summer
they become dark brown with only the wings remaining
white. They flock together during the winter and split into
pairs for the breeding season in summer. As their names
imply, they have different habitat preferences, the willow
ptarmigan preferring dwarf shrub tundra while the rock
ptarmigan occupies mainly the open rocky regions with little
vegetation cover. Apart from their change in color with sea-
son, which has a clear advantage for avoiding predators, they
also have feathered feet, which prevents heat loss to the
snow and the cold ground. Both species are mainly plant
eaters, consuming the young growth of leaves, together with
berries; they also eat insects during the spring. The tundra’s
abundant insect supply is important in providing protein for
the young birds in the nest.
Other permanent residents of the Arctic are the snowy owl
(Nyctea scandiaca), the peregrine falcon (Falco peregrinus), and
the gyrfalcon (Falco rusticolus), all of them predators. The gyr-
falcon is a large falcon that preys mainly upon the two
ptarmigan species (88 percent of the diet in Alaska and 96
104 TUNDRA
percent in Greenland consists of ptarmigans). Other prey ani-

mals include seabirds, snow bunting (Plectrophenax nivalis),
and small mammals, such as lemmings (Dicrostonyx species).
The somewhat smaller peregrine falcon competes with the
gyrfalcon for ptarmigan in the Arctic. The peregrine falcon is
much more widely distributed throughout the Northern
Hemisphere, but it succeeds well in the tundra regions, nest-
ing on cliffs. Snowy owls are also important predators of the
tundra, but their main food source is the lemming. Like the
lemming’s, the snowy owl’s population fluctuates strongly
depending upon the availability of its prey. White-tailed
eagles (Haliäetus albicilla) are often associated with tundra
coasts, as in western Greenland, but these are mainly fish-
feeders and only occasionally take tundra birds and mam-
mals, such as the arctic fox (Alopex lagopus).
Among the smaller, perching birds, the hoary redpoll
(Carduelis hornemanni) is one of the most truly tundra
species, being resident in much of the Arctic region,
although in winter it tends to move southward into south-
ern Canada, so it should really be regarded as a partial
migrant. The common redpoll (Carduelis flammea) is mainly
a summer visitor to the tundra, preferring the shrub tundra
and willow thickets to the open tundra landscapes. Both are
mainly seed eaters. Several other perching birds visit the
tundra to breed and then move south for the winter. The
Lapland longspur (Calcarius lapponicus) is one such bird that
breeds on the open short grass habitats in the tundra of
Canada and Alaska and moves south into the United States
for the winter. It is also found in the Eurasian tundra and
again migrates into the temperate zone around the Black
and Caspian Seas to avoid the severity of the tundra winter.
The Lapland longspur is mainly brown in coloration, the
male having a black facial mask and yellow beak in summer.
Some of the tundra perching birds use white patches and
wings as a means of camouflage in sites that retain some
summer snow. The snow bunting (Plectrophenax nivalis), for
example, has a brown body and white wings with black tips.
When it flies among rocks and snow patches it becomes
very conspicuous, but it seems to disappear once it lands
and closes its wings. A predator trying to locate it would be
greatly confused by the way it appears and then vanishes so
effectively. Snow buntings, like Lapland longspurs, breed on

barren, open ground in the polar tundra, but their range
extends south in Europe on the Scandinavian and Scottish
mountains. In North America they come south only in the
winter. They are mainly seed eaters and they carry seeds in
their crops when they migrate. When a new volcanic island
arose out of the sea off the coast of Iceland, some of the first
land plants to arrive came in the guts of migrating snow
buntings. Some of these birds made a landfall and then
died, but their crops still contained living seeds that they
had eaten, possibly as far away as Scotland, and these ger-
minated within the birds’ corpses and formed the first wave
of land plant invasion on the island.
The snow bunting does not occur on more southerly
mountains except as a winter visitor in North America, but
the mountains of Europe and Asia have a very similar bird
called the snowfinch (Montifringilla nivalis) that spends its
entire life on the alpine tundra, changing its altitudinal range
a little with seasons. It occurs in the Pyrenees, Alps, Cauca-
sus, and eastward into the mountains of Tibet and China. It
is very like the snow bunting in appearance, with its black-
tipped white wings, and uses the same tricks to evade preda-
tors, but it is not very closely related. This is a good example
of how evolution can come up with the same answer to a
challenge on more than one occasion.
Many types of waterbirds spend their summers in the Arctic
and breed there in the peace and tranquility, far from human
settlements. These include wading birds and also ducks and
geese. Some of the most familiar waders of North American
winter coasts, such as the red knot (Calidris canutus), the
sanderling (Calidris alba), and the dunlin (Calidris alpina),
breed in the very far north of the Arctic and migrate over
huge distances to do so, despite their diminutive size. They
find the tundra ideal for their breeding as there is an abun-
dant supply of aquatic invertebrates in the tundra wetlands.
Most of these waders, as their name implies, have long legs in
relation to their bodies so that they can wade into shallow,
muddy pools and probe the mud with their long bills. Some
of them, like the phalaropes (Phalaropus species), have devel-
oped a behavior pattern in which they swim rapidly in circles.
This creates a vortex in the water that brings the mud and the
106 TUNDRA
Putting on weight in the tundra

Wading birds, like all migrants, need to accumulate large reserves of fat in their bodies to
supply the energy needed for the long migration flight. The sanderling, for instance, dou-
bles its weight just prior to the migration and loses all of this excess of fat during the stren-
uous journey. Some wading birds spend the winter in the southern parts of South America
and Africa. Most of the journey is carried out at night, using the stars to assist in naviga-
tion, together with the kind of magnetic compass that birds have within their brains and
that enables them to find a correct course even in cloudy weather. During the day they
rest at stopover sites where they can replenish their food supplies. In the case of waders,
many migrate along coastlines so that suitable habitats are quite frequent, but some pass
over the land, even over deserts, where isolated water sources become vital for their suc-
cessful completion of the journey.
small animals it contains to the surface, where the phalarope

can feed upon them. Their chicks are said to be nidifugous,
which means that they can walk and run within an hour of
hatching, and they leave the nest to shelter in the long vege-
tation of cotton grasses and sedges. Their parents continue
feeding them through the long summer days, but they
quickly develop the ability to feed themselves and prepare for
the long journey that lies ahead of them. Storing enough
energy in the form of fat is vital if migrant birds are to com-
plete their long journeys successfully (see sidebar above).
Ducks, geese, and swans—the waterfowl—also find an ideal
breeding ground in the Arctic tundra. The small, dark brant
goose (Branta bernicla), for example, breeds on the tundra
throughout the northern fringe of North America and Green-
land, and a closely related species breeds in northern Russia.
Like the waders, it tends to migrate along the coasts and spends
its winter in estuaries and coastal areas farther south, even as
far as Florida and Baja California. The snow goose (Chen
caerulescens), on the other hand, which also breeds in Arctic
Canada and Alaska, has several inland flyways: west down the
Sierras, through central North America along the Mississippi
River, and also down to the East Coast, cutting through Hud-
son Bay and Ontario. These flyways are also used by the white-
fronted goose (Anser albifrons). This species is more widespread

than the snow goose, however, being found in western Green-
land and through northern Russia and Siberia. There are several
subspecies of this goose, each of which has its own breeding
area, migration route, and winter habitat, as shown in the illus-
tration. The Greenland population, for example, travels to Ire-
land and western Britain to spend the winter. These separate
populations have probably established their flyway behavior
patterns since the end of the last glaciation. The separation of
these different groups of birds with their various patterns of
movement has ensured that they have begun to evolve sepa-
rately. Many different species of waterfowl use the same fly-
ways, including the aptly named tundra swan (Cygnus
columbianus), which travels to the Gulf of Mexico and other
coastal regions for the winter. This same swan breeds in the
Russian tundra and travels all the way to England in the west or
to Japan in the east for its winter season.
Seabirds are a feature of the tundra coastlines and use both
the flat, open tundra and the cliffs of the more mountainous
locations for their breeding. Some of them, such as Ross’s gull
(Rhodostethia rosea) and the ivory gull (Pagophila eburnean),
spend virtually all their lives in the cold northern oceans and
the fringing land, but others breed in the tundra and then
migrate to more southern coasts for the winter. These
migrants include eider ducks (genera Somateria and Polystica) The breeding range,
and long-tailed ducks (Clangula hyemalis), the common wintering grounds, and
murre (Uria aalge) and the pigeon guillemot (Cepphus migration routes of the
columba) of the west coast of North America and the razorbill white-fronted goose.
white-fronted goose
Anser albifrons
108 TUNDRA
arctic tern
Sterna paradisaea
The breeding range and (Alca torda) of the east coast. The Pacific loon (Gavia pacifica)
the migration routes of is another seagoing migrant that moves along the west coast
the arctic tern (Sterna in flocks of several hundred as it migrates between the tundra
paradisea), perhaps the and the ocean off Mexico, where it spends the winter. But of
best-traveled of all birds all the migrant seabirds that nest in the tundra the prize for
the most well traveled must go to the arctic tern (Sterna par-
adisaea). This small white bird, sometimes called a “sea swal-
low” because of its elegant flight and long forked tail, breeds
throughout the tundra regions of the north. It then sets off
on a journey that takes it deep into the Southern Hemi-
sphere, all the way to Antarctica and the Southern Ocean (see
illustration above). Researchers have studied its main migra-
tion routes by banding birds (placing small, harmless rings
marked with code numbers around their legs) and recording
their places of recovery when they are trapped for observa-
tion or found dead. The arctic terns from North America and
eastern Russia move from the Arctic mainly down the west
coast of America (North, Central, and South), while those

from Greenland, Europe, and western Russia travel down the
west coast of Europe and Africa. They then disperse in the
southern oceans, feeding upon the abundance of small fish
until the time comes to reverse the journey. The total dis-
tance traveled in a year by an arctic tern is thus around
22,000 miles (36,000 km). As a result of its journey, it
undoubtedly enjoys more hours of sunlight in a year than
any other creature on Earth.
The only bird that comes near to the arctic tern in the dis- The arctic tern (Sterna
tances covered in a year is the parasitic jaeger (Stercorarius par- paradisea). By migrating
asiticus), known in Europe as the arctic skua. The American annually between the
name communicates something of the behavior of this tun- Arctic and the Antarctic
dra-breeding bird, for it obtains its food by attacking other it spends much of its life
seabirds, particularly gulls and terns, and harassing them in in perpetual daylight.
the air until they regurgitate their stomach contents. The (Photo by Lauri
jaeger then leaves off its pursuit to collect the lost food, Dammert)
110 TUNDRA
Emperor penguins sometimes even catching it before it lands in the water. These
(Aptenodytes forsteri) bully birds of the tundra seldom actually attack and kill the
in a colony at Cape birds they pursue but may catch their wings and tails in their
Crozier, Antarctica. bills and cause them much distress. Like the arctic tern, they
These are the largest of breed in the open tundra of the Arctic and then migrate
the penguins, and it is southward along the coasts of all the major continents in
the male that takes sole winter. They may reach as far as New Zealand, southern Aus-
responsibility for tralia, and Tierra del Fuego, so they come close to equaling
incubating the single the terns in the distances covered. There is an equivalent
egg. (Photo by Norbert Antarctic jaeger, called the south polar skua, which is a gen-
Wu/Minden Pictures) eral predator of smaller birds, including penguins. It nests on
the bleak, boulder-covered coastal regions of Antarctica.
Perhaps the most familiar of the Antarctic birds, however,
are the albatrosses (family Diomedeidae) and penguins (fam-
ily Spheniscidae). Albatrosses are enormous seabirds with
wingspans of up to 12 feet (3.5 m). They fly by soaring and
gliding, keeping their long wings perfectly stiff and using
every updraft from the ocean waves to propel them forward.

This is a highly efficient way of flight, involving no flapping,
and they can travel great distances in this way. They may
cover 5,000 miles (8,000 km) in a week, searching for food for
their chicks. Eight different species are frequently found in
the southern oceans, but all of them nest on oceanic islands
rather than on the mainland of Antarctica.
Penguins, of which there are 17 species, do breed on the
coastal strips of the continental mainland of Antarctica. The
largest is the emperor penguin (Aptenodytes forsteri), with a
height of 45 inches (115 cm). Emperor penguins eat fish,
swimming underwater with their flipperlike wings and diving
for as long as nine minutes at a time. During these dives they
can reach depths of 1,300 feet (400 m). Their large size is a
great advantage under very cold conditions because they have
less surface area (through which body heat is lost) per unit vol-
ume than smaller penguins, such as the little penguin (Eudyp-
tula minor). The little penguin has a height of just 16 inches
(40 cm) and is not nearly as cold-tolerant, breeding no farther
south than southern New Zealand. Emperor penguins breed
on the bare coastal flats of Antarctica, the male bird being
responsible for incubating the single egg, holding it on top of
its feet while covering it with a flap of feathered fat. They
breed through the Antarctic winter, standing together in large
huddles for nine weeks in constant darkness and in tempera-
tures of –76°F (–60°C). They constantly shuffle around the
group, which may number as many as 5,000, trying to find the
warmest position, and the movement ensures that no individ-
ual is exposed on the outside of the congregation for too long.
Penguins, being unable to fly, are permanent residents of
the land of ice. They do not have the option of long-distance
migration that so many Arctic birds enjoy.
Polar tundra mammals

The tundra is not rich in mammal diversity, but it does con-
tain some very distinctive species. Like penguins, most mam-
mals are unable to fly, so the opportunities for long-distance
migration are limited. Some large mammals undertake regu-
lar seasonal movements in the Arctic regions, and bats are
exceptional mammals in having the capacity to fly, so they
112 TUNDRA
can migrate to a certain extent. But most mammals are forced

to cope with the polar winter. Alpine tundra mammals find it
easier to move up and down mountains according to the sea-
son, so they often adopt this way of life. Relatively few mam-
mals are found in both polar and alpine tundra habitats.
The animal most characteristic of the Arctic tundra is the
polar bear (Ursus maritimus). This is a massive mammal, often
growing to nine feet (2.8 m) in length and 1,500 pounds
(700 kg) in weight. It is unusual among bears in being totally
The harp seal (Phoca carnivorous, feeding mainly upon seals, waiting for them at
groenlandica) is one of their breathing holes and snatching them as they surface for
the main prey species of air. It is then very happy to consume up to 30 pounds (13 kg)
the polar bear. Harp of blubber in one sitting. The polar bear has very thick fur that
seals breed on ice floes, protects it from the cold and even allows it to spend many
as shown here in the hours swimming in the icy polar waters without suffering any
Gulf of St. Lawrence, harm. The insulating properties of the fur are so efficient that
where a pup is greeting ice forming on the outside fails to melt because the bear’s body
its mother. (Photo by heat does not penetrate. Even the soles of its feet are covered
Michio Hoshino/ with hair, preventing frostbite in its toes as it pads across the
Minden Pictures) ice. Its creamy white coloring allows it to blend perfectly into
its surroundings of snow and ice floes. Whereas most camou-

flage in animals is a means of avoiding detection by predators,
in the polar bear it is more likely to be valuable in deceiving its
prey, because it has no predators apart from humans. A white
predator is more likely to be successful in approaching a bask-
ing seal than a dark-colored one. The Inuit people hunt and
eat polar bears, but they do not consume the liver, which is
poisonous because of its exceptionally high level of vitamin A.
The hairs of the polar bear are hollow, which makes them even
better at thermal insulation than solid ones, but the hairs also
allow light energy to penetrate, right through to the black skin
beneath. Here it is absorbed and helps to maintain the bear’s
temperature. One problem encountered when polar bears are
kept in southern zoos is the growth of algae within the hollow
hairs, which turns the bear’s coat green.
Polar bears spend most of their lives close to the sea, or
even out on the floating ice. There are risks involved in living
on the ice, for it can break up into drifting sections and move
away from land. Polar bears have been recorded on ice floes
as far as 200 miles (320 km) away from land. But their power-
ful swimming ability enables them to move from one raft to
another and ensures their survival. They seem to be inces-
santly on the move, either in search of prey or, in the case of
males in the spring, in search of a mate. It is at this time of
year that the wandering males are in their most irritable and
dangerous moods and may attack people.
The female polar bear hibernates through the long dark
winter. She burrows beneath the snow, which insulates her
from the worst of the cold conditions, and there she lives off
her fat until the arrival of spring. Females that have become
pregnant as a result of a spring mating give birth to tiny, rat-
size cubs inside the winter den. Although the polar bear is
among the largest of the four-legged carnivores of the world,
its young are among the smallest in relation to the mother’s
size. Most commonly, twins are born in December or January.
They suckle on fat-rich milk until they emerge in the spring,
and the female at this stage is in a light state of dormancy
and wakes easily if danger threatens. When they emerge, the
young are weaned onto a diet of meat, usually arctic hare
rather than seal in their first year. The cubs stay with the
mother for the whole of the following summer; she mates
114 TUNDRA
0 miles 500 1,000
0 km 500 1,000 1,500
Greenland
ARCTIC OCEAN
Beaufort Sea Baffin Bay
Alaska
Yukon Northwest Territories
PACIFIC OCEAN
Hudson Bay
British
Columbia
total range Québec
Alberta
known calving grounds
Saskatchewan Manitoba Ontario
The range of caribou in only every other year. Males, as well as females with growing
the tundra of North cubs, tend not to build snow dens, but remain active through
America. The species is the winter.
distributed in a number While polar bears stay in place through the winter and
of herds, each of which endure the hardships, the caribou (Rangifer tarandus) escapes
occupies its own calving the worst of winter conditions by moving south to areas of
grounds in the spring, better food supply. The caribou, or reindeer as it is called in
migrating south in Europe and Asia, is the only deer that survives north of the
the winter. Arctic tree line throughout its life. Although there is just one
species of caribou, it is widely distributed throughout the
polar tundra of the north and is divided into several sub-
species that have different ways of life. In Siberia, for exam-
ple, and also in western North America, there are subspecies
that spend their lives in the forest rather than out on the tun-
dra. But the most common forms of this animal graze on the
open tundra vegetation. The North American caribou is gen-
erally larger than the Eurasian reindeer, a bull often growing
to a height of more than four feet (120 cm) at the shoulder
and weighing up to 600 pounds (270 kg). The cows are
smaller. North America’s caribou population occurs in a series
of separate herds, each of which has its own preferred calving

grounds, as shown in the illustration.
Like the polar bear, caribou are covered in dense hair that is
hollow and provides extra thermal insulation. These air-filled
hairs also enable the caribou to float very efficiently, and it
swims strongly with about a third of its body above the water.
In summer the caribou are brown or gray, but they become
paler, almost white, in winter, especially on some of the Arctic
islands. The hooves have spongy footpads, which effectively
spread the weight of the animal and allow it to move over
areas of soft, peaty soils without sinking into the mire. In win-
ter, the pads harden and develop bunches of hair, which serve Caribou (Rangifer
both to insulate the hooves from frost and to give additional tarandus) herd with
grip on the slippery ice and snow. With these adaptations the young at its breeding
caribou can run rapidly, achieving speeds of about 50 mph grounds in the tundra
(80 km/h) and can escape such predators as wolves in this of the Alaska National
way. But the caribou is a sprinter and cannot keep up such a Wildlife Reserve. (Photo
speed for very long, so it may fall prey to wolves if these ani- by Michio Hoshino/
mals persist in their pursuit until it weakens. Migrating herds Minden Pictures)
116 TUNDRA
of caribou are often accompanied by wolf packs that pick off

the old, the fragile, and the young as their victims.
Both male and female caribou have antlers, but those of
the male are usually larger. In the North American caribou
the antlers can grow to five feet (150 cm) in length. Males
fight one another with their antlers during the rutting
(breeding) season in the fall, and antlers also provide a means
of defense against predators, such as wolves (Canis lupus) and
wolverines (Gulo gulo). After the rut (by about December), the
males shed their antlers, but the females retain theirs
through the winter, shedding them in April when the calves
are born. During the winter, the females are able to assume a
degree of dominance because they have retained their
antlers, so they can gain better access to the richer vegetation
when they are pregnant and need the additional nutrients.
Caribou are very particular about their food. Although
they will eat the young leaves of willow and birch, they pre-
fer the new growth of grasses and sedges. They also eat
lichens, especially those of the genus Cladonia, the so-called
reindeer mosses. These small, slow-growing organisms (actu-
ally a combination of algae and fungi) are dry and crisp and
do not appear very attractive as food, but they are actually
quite nutritious. Their very slow growth of about one inch
(2.4 cm) every five years means that a grazing herd of caribou
can remove many years of growth in a very short time. Con-
sequently the caribou have to keep moving over the tundra
to seek out new resources while the grazed areas slowly
recover. The outcome is that the tundra can support only low
densities of such grazers; a density of one caribou per 150
acres (60 ha) is about as high as can be maintained.
A large proportion of the world’s caribou lives in Russia,
where almost half of the 3 million or so animals are semido-
mesticated. Pastoral peoples follow the herds on their migra-
tions, protect them from predators especially in the calving
season, and harvest them as a source of milk, meat, and skins.
It is likely that caribou were once very much more abundant
than they are now, and one estimate indicates that there were
once more than 3 million caribou between Hudson Bay and
the Mackenzie River. Human predation has undoubtedly con-
tributed to their decline, but they also undergo strong popula-
tion fluctuations. There has been much research on what
A tundra heavyweight, the musk ox

The musk ox (Ovibos moschatus) is an impressive grazer of the tundra, and this herbivore
must rank as one of the most highly adapted of Arctic mammals. It is totally confined to
the Arctic tundra, particularly the polar desert regions. Hunting has greatly reduced pop-
ulations of musk oxen, driving the species to extinction in Alaska by 1860, and the animal
has also been eliminated from the Russian Arctic, apart from one island where it has been
reintroduced from North America. Musk oxen are now found largely in the Canadian Arc-
tic and Greenland.
Actually more closely related to sheep and goats than to cattle, musk oxen are stocky
animals, standing about five feet (1.5 m) in height and weighing up to 670 pounds (300
kg). The musk ox grows a coat of dense shaggy hair, with an undercoating of very soft
hair that some Native American tribes use in the construction of cloth. It bears massive
horns that curve downward on either side of the head. The musk ox owes its name to its
strong smell, which is not in fact due to musk but to its fetid urine. Like caribou, musk
oxen are herbivores, but they are more inclined to eat the woody tissues of dwarf shrubs,
although they take grasses and sedges during the spring growth. Musk oxen are not
migratory but wander around in small groups of about 20 to as many as 100 animals con-
sisting of both males and females. When they are threatened by predators, they form a
defensive circle with their heads and formidable horns pointing outward, keeping any
young animals safely within the circle. It would take a brave or foolhardy predator to
attempt to penetrate the wall of horns that meets it.
causes caribou populations to rise and fall every 60 years or

so. Natural and human predation, together with climate and
food supply, influence population levels.
Of all the animals of the Arctic tundra, few have generated
more interest or legend than the lemming (family Microti-
dae). These small rodents have a reputation for mass suicide;
folk tradition asserts that lemmings leap from cliffs by the
hundreds during episodes of population explosion. There are
several different species of lemmings, all of them small
rodents that survive under very extreme conditions of cold.
The one that has generated most of the fabulous stories of
suicide is the Norwegian lemming (Lemmus lemmus), which
occupies the mountainous tundra of Scandinavia. It is cer-
tainly true that the populations of the Norwegian lemming
118 TUNDRA
undergo very considerable fluctuations, and there are years

in which large numbers of the animals migrate in massive
numbers, usually down the valleys from their mountain
homes. When they reach a water barrier, Norwegian lem-
mings are often undecided what to do because, although
they swim quite well, they are reluctant to take to water
unless they can clearly see the other side. The pressure of the
crowd at such times undoubtedly forces many individuals off
the edge of cliffs or riverbanks and into the water, where they
drown. The discovery of large numbers of drowned lem-
mings washed up on the shores of fjords is probably what
inspired the stories of mass suicide.
Exactly what causes the population explosions among lem-
mings is still not entirely clear. There is an approximate three-
to four-year cycle in the population fluctuations. When the
population is at its peak size, there may be as many as 160
lemmings in each acre of tundra (400/ha). The buildup in
population probably results from an abundance of high-qual-
ity food. During low lemming years the grazing pressure is
much diminished and the vegetation grows more rapidly. The
growing vegetation then becomes richer in certain essential
elements that are in short supply, such as phosphorus. The
outcome is a lemming plague that consumes the resources, so
the population crashes once again as quality food becomes
scarce. During the stage when food resources are beginning to
run low, the lemmings migrate outward from their centers of
population, giving rise to the myths associated with them. In
peak population years, the lemming is a major source of food
to a wide range of predators, including snowy owl, arctic fox,
rough-legged hawk (Buteo lagopus), and pomarine (Stercorarius
pomarinus) and long-tailed jaegers (Stercorarius longicaudus).
These predators also undergo population cycles that match
the lemming cycles, illustrating how dependent they are
upon this source of food. Snowy owls, for example, raise as
many as 10 or more young in a good lemming year but may
fail to rear any when lemming populations are low. It is
unlikely that the intensity of predation actually causes the
crash in lemming populations, however. Even heavy preda-
tion is likely to account for only about 10 percent of the lem-
ming population, which is hardly enough to cause a
population crash; food limitation is much more likely to be
the cause. A drop in the availability of food would affect the

rate of reproduction, which would have a rapid impact on the
overall population of the lemmings.
Like most other small mammals, lemmings are capable of
very rapid breeding. Unlike many of the other small rodents
of the tundra (such as voles), they have the remarkable abil-
ity to breed all year. In one study of brown lemmings (Lem-
mus sibiricus), 33 percent of females were found to be
pregnant in the December-to-April period and about 80 per-
cent in July. The arctic lemming (Dicrostonyx torquatus) has
even higher rates of breeding, with 40 percent of females
pregnant in winter and 90 percent in July.
In the Arctic summer, of course, there is no opportunity for
nocturnal activity to avoid the bird predators that mainly
hunt by daylight. As the summer wears on, however, the
lemmings tend to confine their activity to the hours of dark-
ness to avoid the attention of birds and foxes. Lemmings are
herbivores, and the different species have varying food pref-
erences, which is why they are able to coexist in the same
area. The arctic lemming, for example, prefers broad-leaved
herbs and dwarf willows, and it eats very little in the way of
grasses, sedges, mosses, and lichens. The brown lemming, on
the other hand, concentrates on these latter plants and takes
very little of the broad-leaved and woody plants. In winter
lemmings burrow beneath the snow and the breeding
females construct well-insulated nests of sedge stalks to pro-
tect themselves against the penetrating cold.
Another grazing mammal of the tundra is the arctic hare
(Lepus arcticus), which has the most northerly distribution of
any of the world’s hares and rabbits (lagomorphs). It occurs
in the tundra regions between Labrador and the Mackenzie
River, being replaced in Alaska by the Alaska hare (Lepus
othus). The arctic hare is one of the largest of the hares,
weighing around 12 pounds (5.5 kg). It is quite fussy about
its food, concentrating upon arctic willow (Salix arctica),
which means that its distribution is limited by this particular
food species. It is quite unusual in such a harsh environment
to have a large herbivore that is so strictly limited in its food
preferences, but it seems to thrive despite this specialized
diet. The arctic hare remains active through the winter and
therefore has to cope with extreme cold in the months of
120 TUNDRA
darkness, often with monthly average temperatures of –35°F

(–38°C). Unlike the lemming, which burrows below the
snow, it has to survive without any insulation apart from its
thick fur. Its breeding, however, is restricted to the summer,
and it produces a litter of five or six young in a scraped hol-
low, often in the shelter of a rock. The female arctic hare is a
more devoted mother than is the case with most lago-
morphs. She remains with the young for as long as two or
three weeks; then they form small groups or crèches and the
mothers return regularly to nurse their own offspring. After
about nine weeks the young are left to fend for themselves.
Adult arctic hares remain quite solitary during the summer,
but in the fall they form large herds. Flocks numbering into
the hundreds have been recorded on Ellesmere Island, and
this may offer them some protection against such predators
as the arctic foxes, which find it difficult to stalk a grazing
flock without being detected. The hare herds can cause inten-
sive grazing pressure on the tundra vegetation and may out-
compete some of their fellow grazers, such as musk oxen and
caribou, especially in years of high populations. Like lem-
mings, they exhibit a cyclic pattern of population variation,
but with peak populations roughly every nine years.
In Europe and Asia the mountain hare, or blue hare (Lepus
timidus), is found. This is closely related to the arctic hare but
is unusual among tundra mammals in being found also in
the more southerly mountains of Europe, including the Alps.
It is called the “blue hare” because its summer coat is a blue-
gray color, but in winter it molts and produces a white coat
with just the tips of the ears remaining black. It has a wider
range of food preferences than the arctic hare, feeding on
grasses and heathers. The change in coat color is of obvious
advantage in habitats that are dark in summer and white in
winter, but the timing of the molt is crucial in determining
how effectively this camouflage defends the animal against
predators. A combination of day length and temperature
determines the onset of the molt, but if the timing is wrong
because of early snow in fall, for example, then the outcome
can be fatal. White hares on the dark surface of the snowless
tundra, or gray hares on the surface of the snow, are easily
spotted by the foxes and eagles that prey upon them.
One group of mammals that may not immediately spring

to mind in the context of the tundra is bats (family Vesper-
tilionidae). It is certainly true that bats are scarcer and less
diverse in the tundra regions than they are in the temperate
and tropical regions, but they do occur as seasonal migrants,
taking advantage of the great abundance of insects found in
the Arctic summer. Bats are nocturnal insectivores, flying
and hunting at night to avoid the effects of daytime preda-
tion by such birds as hawks and falcons. By confining their
activity to dusk and nighttime, they also avoid direct com-
petition with insect-eating birds, such as swallows and
swifts. In the Arctic, of course, bats are faced with the prob-
lem of the absence of any real night during the summer.
Studies on bat activity times in northern Norway have
shown that they maintain a period of hunting that corre-
sponds with their perceived “night,” generally between 10
P.M. and 2 A.M. During that period the activity of bird insecti-
vores is also much lower, so the bats and the birds still seem
to divide the available time between them despite the
absence of darkness as a cue. This division is not ideal for
bats, however. Insect activity is lower at “night” even
though the Sun is still above the horizon. So the bats have
the poorest slot for their insect hunting and yet are still
exposed to attack by bird predators. It is not surprising that
bats are scarce in the tundra.
The mammals of the Antarctic are entirely marine rather
than true tundra species. Whales, dolphins, porpoises, seals,
and sea lions spend most or all of their lives in the oceans or,
in the case of the seals, on nearby beaches and ice floes. No
land mammals have been able to colonize or survive on this
extremely inhospitable continent.
Alpine tundra mammals

Relatively few mammal species are found in both polar and
alpine tundra, except where mountain chains connect the
two, as in the North American Rocky Mountains and the
mountain ridge of central Norway. The wolverine has a range
that extends from the Arctic tundra south into the United
States along the Rocky Mountain chain, and the Norwegian
122 TUNDRA
Pair of yellow-bellied lemming is found on mountains over the whole length of

marmots (Marmota Norway. The mountain hare is unusual in being found both
flaviventris) in in the tundra of the Arctic and also on isolated, more
Yellowstone National southerly mountains, such as the Alps. On the whole, the
Park, Wyoming. These mountains of the world support their own distinctive mam-
alpine tundra mammals mal communities.
are herbivores and build One of the most typical of mountain mammals is the mar-
up enough fat during mot (Marmota species). There are, in fact, many species of
the short summer to marmot and they are widely distributed around the moun-
enable them to tains of the world, including the Rockies, the Alps, and right
hibernate through the across to Siberia. They are not large mammals, generally
long winter. (Photo by about two feet (60 cm) in length, although the hoary marmot
Carolyn McKendry/ (Marmota caligata) of western Canada can grow to almost
IstockPhoto) three feet (90 cm). What they lack in length, they make up in
girth, being rotund animals with a stocky build. Marmots are
closely related to the squirrels and, like them, are herbivo-
rous, spending the summer grazing on the herbage of the
alpine tundra above the tree line. They enjoy well-drained,
sloping grassland, often with rocky outcrops and scree, where
they will sit and watch the skies for predatory eagles and give
warning to their grazing colleagues by emitting loud and
penetrating whistles when they see danger threatening.
Marmots are largely social and cooperative mammals, liv-
ing in extended family groups, but they defend their local
territory and will chase off individuals that seek to graze their
territory. Males will also wrestle with one another during the
breeding season, standing on their back legs and pushing one
another with their forelegs. They live in burrows or rock
crevices beneath the ground, and they have short, strong
limbs for burrowing in the rocky soils. Here they hibernate
through the long cold winter. Once the spring arrives and the
melting snow reveals green plants, the marmots emerge, bask
in the sunlight, and clean themselves. They are active in the
daytime and are relatively unafraid of people, so they are
easy to observe in their activities. One of their first activities
on emerging in the spring is to clear out their bedding and
replace it with clean materials. When a marmot tries to carry
too much new bedding material, it may end up rolling on its
back holding a mass of bedding with all four legs. When this
happens, its companions drag it by the tail into their burrow.
This behavior was first recorded by the Roman naturalist
Pliny (23–79 C.E.) from his observations in the European
Alps, and, like some of Pliny’s other tales, it was once
regarded as suspect. But scientists in recent times have
observed the same behavior, so Pliny’s unlikely-sounding
story has been confirmed. When their winter bedding has
been replaced, marmots turn their minds to breeding, and
the first young are born within about six weeks of mating.
The family stays together for several years before the young
are sufficiently mature to set off on their own.
In the short season of the high mountain tundra, the mar-
mot must spend much of its time feeding, especially as the
winter approaches. Marmots do not store food in their bur-
rows but build up large reserves of fat to keep them alive
through the hibernation. This, together with their dense fur,
serves as an insulating layer to retain some body warmth,
and human hunters prize both the fur and the fat. The
black-capped marmot (Marmota camschatica) of eastern Rus-
sia is one of the most proficient sleepers in the animal king-
dom. It can spend up to nine months of every year asleep.
124 TUNDRA
By mid-September it is covered with a thick layer of fat and

weighs about 11 pounds (5 kg). It descends into its burrow,
along with the other members of its family, and they curl
into a tight ball with their forepaws covering the sides of
their heads. As the external temperature drops, so does the
body temperature of the marmot, often becoming as low as
40°F (4°C) and sometimes even below freezing. At this stage,
it is very difficult to detect any sign of life in the animal. Its
heart beats just once every two or three minutes, and it
breathes only once every 10 minutes. Even this very low level
of body activity does generate some waste materials, so the
marmot warms up and wakes about every three weeks so that
it can urinate. By the time it finally wakes in May, it has lost
half of its body weight, so it is easy to appreciate its eagerness
to emerge and start eating. The marmots are a group of mam-
mals that are essentially alpine tundra rather than polar tun-
dra in distribution. The same can be said of the sheep and
goats; many of these are mammals of the mountains.
The American mountain goat (Oreamnos americanus) is
actually not a true goat but belongs somewhere between the
goats and the antelopes. It occurs above the tree line in the
mountains of Alaska, British Columbia, and the Rocky
Mountains south into Colorado, and South Dakota. Like
those of many of its relatives, its hoof is adapted for gripping,
having a hard outer rim and a softer inner part that can grip
slippery surfaces. It lives on the high, steep, and inaccessible
cliffs of the mountains, producing young in the spring that
are able to climb on the precipices virtually from birth. The
kids may fall prey to golden eagles, but their greatest danger
is death by misadventure. A slip on a mountain ledge, or a
loose rock, can easily lead to a fatal fall. The chamois (Rupi-
capra rupicapra) of the European and western Asian moun-
tains is a close relative of the American mountain goat and
has many of the same adaptations. Like the wild goat, it is
polygamous and the males compete for harems of females in
the November rutting season. The chamois has been exten-
sively hunted in Europe but is now widely protected and is
becoming a frequently seen animal of the Alps and Pyrenees.
It can live up to 22 years and is capable of survival at very
high altitudes, the highest on record being 15,430 feet (4,750
m) on Mont Blanc, Europe’s highest mountain.
Another agile rock-climbing mammal of the American

mountains is the bighorn sheep (Ovis canadensis). It occupies
the more remote areas of the Rockies and the Sierras, avoid-
ing contact with human beings. It lives in groups, consisting
of up to 15 animals (mostly females and young) in summer
and congregating into herds of up to 100 for the winter. The
males join the female groups during the rut, when battles for
access to females reach epic proportions. Competing males
charge one another head-on, and the crash of their colliding
horns can be heard a mile away. The heads of the males have
a very dense cover of bone at the front, so a direct clash rarely
causes any serious injury. These battles may last many hours
until the fitter male wins the day and has the privilege of
passing on his genes to the next generation. The lambs are
susceptible to predation, particularly from golden eagles and
mountain lions. Wolves, coyotes, bears, bobcats, and lynx
can also pose a threat to them if they wander from their pre-
cipitous habitats into lower parts of valleys. Bighorn sheep
tend to migrate from the higher regions that they occupy in
the summer, feeding on grasses and sedges, to the lower parts
of the mountain in winter, where woody materials including
willow form their major food. Like the chamois, the bighorn
has been intensively hunted in the past, but such activities
are now carefully controlled to ensure its long-term survival.
There are species of mountain sheep in Asia, but on the
whole these tend to be animals of rolling hills rather than
steep cliffs and rocky crags. The nearest equivalent to the
bighorn is the snow sheep (Ovis nivicola) of the eastern Asian
mountains, which extend into the northern highlands
beyond the Arctic Circle. In most of the Eurasian mountains,
however, the ecological equivalent to the bighorn sheep is a
type of goat, the ibex (Capra ibex). It is a large goat, up to
three feet (90 cm) in height and notable for its conspicuous,
backward-curving horns, often around 30 inches (75 cm) in
length. The horns are black and strongly ridged, and, as with
bighorn sheep, they are used in battles between males during
the rutting season. Human hunters almost drove the alpine
ibex to extinction, partly for the trophies supplied by their
enormous horns, but also for the so-called bezoar stone. The
animal’s stomach contains this hard concretion of undi-
gested material. People believed that this “stone” possessed
126 TUNDRA
great potential for healing, including the cure for various

types of poisoning, and slaughtered the animals to obtain it.
Fortunately, when the population had been reduced to just a
small herd in northern Italy, authorities imposed protection
and instigated a program of reintroduction over its former
range. The scattered populations of ibex are now growing
once again.
In general, the mammals of the world’s mountains have
suffered more from human hunting than those of the polar
tundra. Mountains are more accessible to people and conse-
quently their mammalian inhabitants more vulnerable. As
tourism and mineral harvesting develop in the polar regions,
disturbance, habitat loss, and hunting may well intensify
there too.
Adaptations in tundra mammals

The mammals of both polar and alpine tundra face similar
stresses. Periods of very low temperature are common to both
habitats, and the survival of any animal depends upon its
being adequately equipped to deal with this problem. As a
consequence, many animals of the tundra share a number of
general features:
1. Although size varies (from lemmings to musk oxen) all
of the animals tend to be compact and rounded, having
relatively small extremities, such as tails and ears. Tun-
dra animals need to maintain a blood supply to their
extremities to keep them warm and avoid frostbite, and
a rounded form is most efficient in maintaining body
temperature and avoiding heat loss. Even birds, such as
penguins, are large and somewhat rounded.
2. An abundance of hair ensures that the hot skin of mam-
mals, richly supplied with blood vessels, resists heat loss
to the surrounding cold air. This works by trapping a
layer of air close to the skin that becomes warm but is
not easily removed. Many tundra animals have hollow
hairs, which are even more efficient than solid hairs in
holding in the warmth. Hairiness is also a feature of
many tundra insects, including butterflies and bumble-
bees. In birds, of course, it is replaced by feathers.
3. Changing color with the season, usually white in winter

and a darker color in summer, has many advantages in
terms of camouflage. It also lets an animal absorb heat
from the Sun more effectively in summer and avoids
the problem of radiating energy in winter. Blue hares
and arctic foxes are examples of tundra animals that
change color seasonally.
4. Hibernation is a means of reducing the biochemical
activity of the body in times of severe cold, and this per-
mits an animal to reduce to zero its requirement for
food and allows it to survive on its accumulated fat
reserves. It leads to problems of waste disposal, but the
greatly reduced levels of body activity solve these.
5. Migration is an alternative to hibernation that allows
an animal to take advantage of the resources of the tun-
dra in its productive summer and avoid the stresses of
its unproductive winter. In the case of mountain tundra
animals, this may involve movements up and down the
mountain with the changing seasons.
6. Summer breeding is the general rule with Arctic mam-
mals, but some of the most successful species, like the
lemming, have mastered the art of year-round breeding.
This is possible only for a small animal that is able to cre-
ate its own microclimate beneath a protective layer of
snow. Winter production of young (as in polar bears and
emperor penguins) has the advantage that the young are
sufficiently grown by the time they emerge from winter
dens to be able to take full advantage of the short sum-
mer season for their feeding and further growth.
7. Many tundra species have relatively wide distributions
compared with the plants and animals in other parts of
the world. The mountain avens (Dryas octopetala), the
white-fronted goose, and the polar bear are examples of
plant and animal species that are found throughout the
Arctic regions of Russia, Alaska, Canada, Greenland,
and Scandinavia. It is important to remember, however,
that the distances between continents are not as great as
might be assumed from conventional maps. Represen-
tations of the surface of the Earth (a sphere) on a flat
piece of paper tend to exaggerate the size of the polar
128 TUNDRA
regions. The total length of latitude 70° is actually only

one-third that of the equator.
The animals and plants of the tundra regions of the world
have thus come to share a number of structural, physiologi-
cal, and biogeographical adaptations to their common prob-
lem, life in a frigid climate.
Conclusions
Intensely cold conditions cause all chemical reactions to pro-
ceed slowly, and this includes those occurring within living
cells. For organisms that cannot control their body tempera-
ture (which includes all microbes, plants, and invertebrate ani-
mals) activity in the tundra is confined to those occasions
when the temperature rises in summer. Wind, burial by snow,
and even drought can add to the stresses of life in the tundra.
Tundra plants are generally herbs or dwarf shrubs, fre-
quently take on a cushion form, and are often evergreen.
Trees are absent, and annual plants are extremely rare. There
is a short season for flowering, and pollinating insects may
not be abundant, so there is much competition to attract
their attention. Consequently, flowers are often large and
brightly colored, especially among species of alpine tundra.
Many plants have hairy surfaces and cells rich in chemical
antifreezes, which enable them to survive the winter cold.
Tundra soils are often rich in undecomposed organic matter,
so there is plenty of food available for microbes, but for much
of the year the temperature is too low for these organisms to
be active. Even in summer, many parts of the tundra become
too wet for decomposer microbes to operate efficiently.
Lichens are among the most successful groups of microbes.
The combination of photosynthetic (cyanobacteria or algae)
and structural (fungi) components allows them to grow
slowly, fix nitrogen, and avoid desiccation in summer.
Invertebrate animals abound during the summertime.
They include soil dwellers that feed on the organic debris,
herbivores that consume the growing vegetation, pollen
feeders, and predators. Tundra wetland provides a rich habi-
tat for many aquatic insects, such as mosquitoes.
Birds take advantage of the brief tundra summer for breed-

ing, many of them migrating from distant regions. In the
polar tundra the long days allow birds extended time in
which to feed their young. Seabirds, wetland birds, and wad-
ing birds are abundant, but there are also resident bird
species, including herbivorous ptarmigans and predatory fal-
cons and owls.
Few mammals are found in both polar and tundra habitats.
Mammals of the Antarctic tundra include many that are
mainly marine but come ashore to breed, including seals.
The Arctic has a much larger range of mammals including
herbivores, such as lemmings and caribou, and predators,
such as arctic fox and polar bear. The alpine tundra is the
home of several species of marmot and also of various sheep
and goat species. These are adapted to life in steep and inac-
cessible places, where they can avoid predation by humans.
The diversity of animals, plants, and microbes found in the
tundra is undoubtedly lower than in other major biomes, but
that does not reduce the scientific interest or the value of this
biome. The living organisms of the tundra face some of the
most severe conditions to be found on Earth. Only a volcanic
crater could be regarded as more inhospitable than these
frigid regions, yet life in abundance is found there. The
species that survive demand admiration, for they have
achieved something that very few humans have been pre-
pared to face, namely living in the tundra. These species rep-
resent highly tuned products of evolution that show an
extraordinary set of adaptations to their harsh environment.
In their structure, their biochemistry, and their behavior,
they have managed to develop strategies for survival, and
humans can learn much from them that would be of great
value. Low biodiversity in the tundra, therefore, does not
mean low value.
CHAPTER 5
HISTORY OF THE TUNDRA

Tundra can exist only in extremely cold conditions, so the
question “How old is the tundra?” is really asking how long
the Earth has known such cold episodes. The answer is that
coldness has a very long history on this planet; on the other
hand, it has been a relatively rare experience. Cold periods
have occurred perhaps nine or 10 times in the history of the
Earth, but these times of cold are the exception rather than
the rule.
Ice ages
The occasions in the Earth’s history when permanent ice has
been present in the form of ice sheets and glaciers are often
referred to as ice ages. They rarely last more than a few mil-
lion years, which is quite brief in relation to the 4.6 bil-
lion–year history of the Earth, and these ice ages are not
times of uniform cold but are often interrupted by relatively
warm episodes. So an ice age consists of alternating cold and
warm conditions. A time of cold, when ice sheets and glaciers
spread, is called a glacial, and a warmer episode when ice
retreats is called an interglacial.
The accompanying table shows the occasions when ice
ages have occurred on Earth. An approximate reconstruction
of the climatic history of the Earth is given in the graph at
right.
Past ice ages have left evidence behind them in the rocks.
As discussed in chapter 2, the movement of ice over the sur-
face of the ground and its subsequent melting leave behind
distinct clues (see “Effects of glaciation,” pages 40–43). Some
of these clues are the scars that they create on the landscape,
but that kind of evidence does not survive the millions of
years involved here. Glaciers also leave distinct deposits, such
130
millions of mean global temperature

years ago cold present warm
Quaternary
1.6
Pliocene
Miocene
Oligocene
Eocene
Paleocene
6.5
Cretaceous
Jurassic
Triassic
245
Permian
Carboniferous
Devonian
Silurian
The changes in global
Ordovician temperature (black line)
Cambrian over geological time,
570
determined from
1,000
chemical and biological
evidence. The wide blue
bands represent the ice
2,000 ages, the times in
Precambrian Earth’s history when
glaciers and ice sheets
3,000 and hence the tundra
biome were present.
Note that the time axis
4,000 ice ages
has been expanded for
more recent times.
as boulder clays, or tills, which can become effectively “fos-

silized.” These tills may be buried by subsequent sediments
and incorporated into rocks, and they are then called tillites
to distinguish them from more recent glacial deposits. These
tillites have been found in Precambrian rocks dating from
132 TUNDRA
The occurrence of ice ages in the history of the Earth

Approximate time
Era Geological period of occurrence
Cenozoic Pliocene/Pleistocene 10 million–10,000 years ago

Mesozoic Jurassic 150 million years ago
Paleozoic Carboniferous/Permian 300 million years ago
Ordovician 450 million years ago
Proterozoic (Precambrian) 600 million years ago
750 million years ago
900 million years ago
2,300 million years ago
Note: Here the term ice ages is used to indicate times when ice sheets (and therefore tundra habitats) have been pres-
ent on Earth.
billions of years ago, showing that ice ages have occurred

right from the very early history of the Earth. The Earth is
considered to be about 4.6 billion years old. From around 3.8
billion years ago, the first true rocks were being formed and
the very first living organisms were present on the planet.
Between then and 2.3 billion years ago there was at least one
ice age, and possibly as many as three. It is very difficult to
construct a precise climatic history of the Earth at such an
ancient stage in its development, but it is certain that the
Earth was already occasionally experiencing a sufficiently
cold climate to result in the formation of glaciers in at least
some parts of the world. These early ice ages are not shown in
the graph because their precise timing and duration are so
uncertain.
The timing of ice ages, particularly the more recent ones,
which are better recorded in the geological strata and better
dated, shows a distinct periodicity. The overall pattern sug-
gests a cycle in which cold periods alternate with warmer
ones. The Earth seems to experience ice ages roughly every
150 million years, but as yet, no one has come up with a
fully acceptable explanation of the mechanism that causes
this long-term regularity. Many theories have been put for-
ward to explain why the Earth might occasionally develop
glaciers and ice caps, and these ideas are extremely varied.
The Earth’s crust consists of a series of plates that are con-
stantly on the move, sometimes separating the landmasses
and sometimes leading to collisions. It is possible that the

configuration and spacing of the plates is involved in deter-
mining ice ages. Perhaps there have to be large landmasses
in the right position (which is not necessarily at the poles)
for ice to begin forming. The appropriate location would
have to be cold (therefore close to the poles) but also where
the precipitation is high, permitting the abundant snowfall
to form ice (which means some distance away from the
poles, which are virtually deserts). Ocean currents also play
an important part. These help to distribute heat energy (ulti-
mately derived from the Sun) over the surface of the planet,
and any disruption to the circulation of these currents could
lead to an ice age. Again, the right positioning of continents
could do precisely this by modifying the direction in which
warm waters flow. When mountain ranges arise, they lead to
changes in precipitation patterns and create high landforms
where glaciers can form. Finally, there is always the possibil-
ity that astronomical factors are involved, such as the dis-
tance between the Earth and the Sun, or the angle of the
Earth upon its axis. Further geological research may help to
sort out exactly what causes the Earth to enter an ice age at
certain points in its history.
Glacial history
Although the present conditions on Earth are relatively
warm, there is no reason to believe that the ice age of
Pliocene and Pleistocene times is over. It is entirely likely that
the present warmth is simply the latest interglacial in a con-
tinuing ice age. More is known about this current ice age
than any other, simply because the records have been better
preserved than for any previous cold episode. This ice age has
so far lasted about 10 million years, beginning in the
Miocene, becoming more evident in the Pliocene, and then
really making its mark on the Earth during the last 1.8 mil-
lion years of the Pleistocene.
It may come as a surprise that an ice age continues despite
all the current discussions of global warming. Ice sheets and
tundra are still present, however, and the possibility remains
that they may one day expand again. During the past 10,000
134 TUNDRA
years (a mere blink of the eye in geological terms), conditions

have been warm and ice caps and glaciers have retreated far
from their former positions. There have been many warm
interruptions to the general cold of the last couple of million
years, however, and there is no reason to believe that the
present warm period is anything more that another such
interruption. The cyclic pattern of recent history, in which
cold and warm episodes have alternated, suggests that the
cold will return. To be able to predict the behavior of the
world climate is obviously very important to people, so
understanding the detailed pattern of climate change—and
with it the fate of the tundra biome—is an important area of
scientific research.
Geologists divide the history of the Earth into convenient
sections, trying to use “natural” breaks wherever possible.
These are points where the fossil content of the rocks
changes relatively abruptly, often indicating a sudden change
in the climate or other conditions on Earth. The separation
of the Cretaceous period from the Tertiary period some 65
million years ago is a good example of a natural break
because the dinosaur fossils disappear and other groups of
organisms exhibit considerable change at that point. This
boundary is now believed to have been caused by a cata-
strophic cosmic event, specifically the collision of a massive
meteor, or bolide, from space onto the surface of the Earth.
The kind of abrupt and cataclysmic environmental change
that resulted from this catastrophe, however, is relatively
rare, and the divisions of geological time are usually far less
abrupt and less easy to identify. This is certainly the case with
the progression of the current ice age, involving the forma-
tion of permanent ice caps and the development of glaciers
in various parts of the world. Although ice began to appear
back in the Miocene, the first evidence for extensive glacia-
tion occurs in the late Pliocene, especially after about 2.4 mil-
lion years ago, and cold conditions became even more acute
around 1.8 million years ago, which is generally regarded as
the beginning of the Quaternary period and the Pleistocene
epoch within that period. But there is still much argument
about this, and some geologists feel that the change at 2.4
million years would be a better marker for the beginning of
the Pleistocene. This argument illustrates that Earth’s cooling

over the last few million years has been gradual and that any
subdivision is somewhat artificial and arbitrary.
Naturally, much more is known about the Pleistocene
glaciations than about the older glacial events. Yet it is not so
long ago that the idea of any past glaciation was unheard of
in scientific debate. In the 18th century, when great advances
were being made in many of the physical sciences, geology
was still in its infancy. A Scottish scientist, James Hutton
(1726–97), should perhaps be regarded as the father of geol-
ogy as a science, and he developed a very important concept
called uniformitarianism. This simply means that past events
can be interpreted by studying what is going on in the pres-
ent, because the mechanisms involved are essentially the
same. This may seem obvious to a modern reader, but it was
quite a novel way of thinking in the late 18th century. The
history of the Earth was considered to be the outcome of a
series of abrupt and unrepeatable events, such as the biblical
Creation and the Flood of Noah. The features of geology,
including fossils, landforms, sedimentary rocks, and so on,
were interpreted entirely within the framework of these cata-
strophic events. Hutton proposed that the erosion of moun-
tains and valleys could be accounted for by the gradual
action of streams and rivers and that the timescales involved,
although extremely long, could be calculated by simple
mathematics.
The idea was taken up by the British geologist Charles Lyell
(1797–1875), whose work greatly influenced his close friend
Charles Darwin (1809–82). Like many geologists at the time,
he was concerned with the problem of how some rocks
(erratics) were found far from their source areas, despite
being massive in size and very difficult to move. What mech-
anism available on Earth today could account for such move-
ments? The great Flood of Noah had been used to account for
these in the past, but could water movements have shifted
such large objects? Lyell wondered whether they might have
been transported in icebergs floating on the floodwaters.
It is not surprising that the real advances in the understand-
ing of ice and ice movements came from the mountainous
country of Switzerland. The Swiss geologist Louis Agassiz
136 TUNDRA
(1807–73) grew up in a land of glaciers, U-shaped valleys,

hanging valleys, moraines, and ice-scratched rocks (see
“Effects of glaciation,” pages 40–43). In 1837, he presented a
research paper to the Swiss Society of Natural Sciences in
which he came to the logical, but extremely novel, conclusion
that the glaciers he knew so well had been much more exten-
sive in times past than they were in his day. He coined the
term Eiszeit, the German equivalent of “ice age,” and proposed
that there had been such a cold event in the geologically
recent past.
Others at that time were making similarly radical claims,
but they were often regarded as somewhat unbalanced. Some
German geologists, for example, claimed in the 1830s that
Arctic ice had once extended as far south as Germany, but
their views were ridiculed. Then Agassiz stepped in and con-
fidently stated that ice masses from the far north had not
only entered Germany but also had covered much of north-
western Europe, parts of Asia, and the northern regions of
North America. American geologists accepted the ideas of
Agassiz much more readily than those of the other Euro-
peans, and in 1846 Harvard University invited him to take up
the Chair of Geology. From there he conducted pioneering
work on the glaciation of North America, although argu-
ments about his very controversial ideas continued well into
the 1860s.
The next question to be answered, once the concept of a
past ice age was well established, was whether it had been
one single event or whether there had actually been several
ice ages. Agassiz tended to assume that a global ice expansion
had taken place on just one occasion, but work being con-
ducted in southeast England showed that there were several
different types of boulder clays, or tills, that could be distin-
guished from one another by the alignment of the stones
embedded within them. These different tills must have been
laid down during separate glacial advances, each moving in a
different direction. Workers in Scotland also found fossil evi-
dence that there had been warm intervals between the glacial
advances.
Geological studies in the European Alps also made it evi-
dent that the recent ice age consisted of more than one gla-
cial advance and retreat. In 1909 this work dealing with the
Alps was formally published, proposing that there had been
four main ice advances. The publication named these after
the rivers Gunz, Mindel, Riss, and Würm, all of which are
tributaries of central Europe’s Danube River, and it identified
the rivers as the advance limits of the different glacial stages.
For about 50 years, this interpretation of the climatic fluctua-
tions of the Pleistocene was regarded as firmly established,
and geologists attempted to fit all new evidence into this
framework. But in science, the test of time often proves new
ideas to be wrong or too simple, and such was the case with
this fourfold interpretation of the Pleistocene. With the
development of more sophisticated techniques for the study
of climate change, such as the use of oxygen isotopes in
marine sediments and in ice cores, it became evident that the
true picture is much more complicated. Within the past mil-
lion years there have indeed been at least a dozen major
glacials, interspersed with warm interglacials. But shorter-
term fluctuations have taken place within these stages as the
climate has displayed a high degree of instability. These
shorter intervals of cold and warmth, which are too short or
too insignificant to permit major biological responses, are
termed stadials and interstadials, respectively, rather than
glacials and interglacials—terms that are reserved for major
changes in climate resulting in major biological responses,
such as the extension of polar tundra during the cold inter-
vals and the expansion of forests in the warm ones, including
the present interglacial.
This understanding of the complexity of climatic fluctua-
tions in the Pleistocene has emerged only since about 1970,
and much still remains to be discovered about its precise pat-
tern, the dating of the glacial advances, the causes of the
observed fluctuations, and the biological and ecological con-
sequences of these changes. Research is currently focused on
the periodicity of warm and cold episodes in the hope of clar-
ifying the pattern of change (see sidebar on page 138) and
providing a basis for future forecasting.
Detailed study of the older glacial episodes is difficult
because so much of the evidence has been destroyed by the
more recent ones. Every time the glaciers advance they are
138 TUNDRA
Cycles of climate change

Using a range of physical and chemical techniques, including the rate at which certain
radioactive compounds in the rocks decay, scientists have now worked out the general
timescale of events in the Pleistocene. The “wavelength” of the glacial/interglacial cycles
(involving one full cycle of cold and warm stages) of the past million years has been about
100,000 years, with the warm episodes occupying only a relatively small proportion of
this time span. The change from warm to cold is usually quite gradual, involving a series
of steadily intensifying cold episodes, so glacial advances probably take place in a series of
stages interrupted by minor retreats. The glacial maximum is eventually achieved late in
the cold part of the cycle, followed by relatively rapid warming, with the maximum
warmth of the interglacial being achieved within around 5,000 to 10,000 years, after
which cooling begins again.
This general pattern seems to have applied to the majority of the glacial episodes of the
later Pleistocene, and it is interesting to note that Earth is 10,000 years into the present
interglacial, so according to the established pattern, things should be on the cooling run
by now. Perhaps the planet was moving in this direction before the global warming of the
past 150 years began when human industrial activities started to pump carbon dioxide
into the atmosphere. We may be holding the next glaciation at bay.
likely to carve away many of the tills and other evidence laid
down in former times. In the case of the most recent glacia-
tion, the evidence remains more fully intact. Glacial
advances have taken different directions, however, so that
some previously glaciated regions have been left undamaged.
This means that the evidence of earlier glaciations remains
undisturbed and can be used to reconstruct the patterns of
past glaciation. The study of recent glacial movements (those
of the last 40,000 years) has been greatly assisted by the
development of the radiocarbon method of dating. This
method depends upon the gradual decay of heavy, unstable
atoms of carbon into their normal, lighter form. The decay
occurs at a constant rate, so the content of heavy carbon in
any organic material gives a reasonable estimate of its age.
Using this and other dating methods, it has been possible to
reconstruct the history of the ice, and hence the tundra, in
some detail for these recent times.
The last major glaciation (called the Wisconsin Glacial in

North America) reached its greatest extent only 22,000 years
ago, which is very recent in geological terms. By 14,000 years
ago it was beginning to melt in response to a rapid rise in the
temperature of the planet. The evidence for this warming is
obtained mainly from the sediments laid down in lakes that
formed around the margins of the melting ice. The mud that
settled in these lakes contained portions of dead plants and
animals that were preserved there, and these provide a
detailed record of the vegetation and animal life of the region
at the time. Where once there was ice, tundra appeared.
Abundant pollen from grasses, sedges and other Arctic herbs
and shrubs was preserved in the mud. Scientists can extract
the pollen grains from sediment cores, identify them, and
count them to supply a picture of the vegetation that fol-
lowed the melting ice northward. In places dwarf shrubs and
even fully developed trees, especially birch, became abun-
dant. But around the Atlantic seaboard, especially in the east-
ern Atlantic, on the European mainland, there is strong
evidence for a sudden drop in temperature back to glacial
conditions between 10,800 and 10,000 years ago on the
radiocarbon timescale. Not only does the tree and shrub
pollen vanish from the record, the very nature of the lake
sediment changes. Instead of the organic mud formed by the
productive growth of aquatic plants, gray clay and stone frag-
ments appear, indicating severe climate and the erosion of
bare soils into lake basins. Clearly, conditions became so cold
during this short episode that there was a return to per-
mafrost and unstable soils that slid down slopes over the
frozen subsoil each summer.
There was also a return to the growth of glaciers, but this
cold period was short and should be regarded as a stadial
rather than a true glacial episode. Very rapid warming fol-
lowed; the evidence from fossils of highly mobile animals,
such as flying beetles, which are able to take rapid advantage
of a change in climate, suggests that the temperature rose by
around 9°F or 10°F (5°C) within a matter of a decade or so.
This is a very sudden change in climate, and it illustrates just
how quickly conditions on Earth can alter. But why did con-
ditions change? Exactly what was taking place that could
140 TUNDRA
cause such massive changes in climate? These are important

questions because the answers will help us understand the
mechanisms that underlie climate function.
One widely accepted theory about the cause of the stadial
at the close of the Wisconsin glaciation is that the melting
of the ice over North America actually caused the return of
cold conditions. At the height of the Wisconsin Glacial, an
enormous ice sheet (the Laurentide Ice Sheet) covered much
of what is now Canada and the northern part of the United
States. As this began to melt, the large volume of water pro-
duced flowed mainly southward into the Gulf of Mexico. As
the ice retreated, however, an alternative outlet became
available, namely the St. Lawrence River leading into the
North Atlantic. The sudden flushing of large quantities of
cold freshwater into the North Atlantic would have had a
massive impact on the circulation of the ocean currents. As
shown in chapter 2, warm waters from the Caribbean move
northward in the Atlantic Ocean and carry their warmth
right up into the Arctic Ocean north of Scandinavia (see the
illustration on page 12). The influx of cold North American
freshwater would have interrupted this important flow and
effectively switched off the oceanic circulation system
shown in the illustration on page 141. If the northward
flow of warm water stopped, the Arctic Ocean would freeze
over and cold polar waters could penetrate southward, cool-
ing the Atlantic seaboard. This is probably how the cold sta-
dial was initiated.
This short but severe climatic fluctuation just 10,000 years
ago has some important lessons for today. It demonstrates
that the uninterrupted flow of currents in the oceans is vital
for the maintenance of the climatic balance of the Earth and
that sudden changes in climate can occur if that balance is
upset. It also shows how easily the balance can be disturbed,
and in days when human activities are altering so many
global processes, people need to beware. Humans must avoid
crossing an environmental threshold that could set in
motion considerable and unstoppable changes.
Because of their sensitivity to climatic conditions, glacial
movements are thus effective monitors of changing climate.
Geologists have spent much effort in recording past move-
ments of glaciers, using various types of evidence, in order to

reconstruct the history of glacial movements. Studies of the
past glacial limits in the valleys of the Swiss Alps have pro-
vided a detailed record of climatic changes over the past
1,000 years. They were in retreat in the 10th to the 13th cen-
turies, the so-called Medieval Warm Period, but then
advanced to achieve greater extent in 1385. The glaciers
advanced and retreated on various occasions during the 17th
century and made their most recent advance around 1860.
These glacial advances correspond with many other pieces of
evidence for particularly cold conditions at these times, and
this part of history, falling between the 14th century and the
mid-19th century, has come to be known as the Little Ice
Age. But since 1860 the glaciers in the Swiss Alps, together
with those of Norway and even the high altitude but low lat-
itude glaciers of the African mountains such as Kilimanjaro,
have been in retreat.
The oceanic conveyor

belt that circulates
around the world’s
oceans. Cold, salty
water is dense, so it
moves along at depth,
warm, less salty
currents at surface
while warm, less saline
cold, deeper salty
currents water moves at the
surface of the ocean.
142 TUNDRA
1925 Tarr Inlet 0 3 6 9 miles
1948
1892
Adam’s Inlet
1907 1892 1929
1907 1907
1963 1892
1948
1907
Muir Inlet
1907
1929
1892
1948
1860
1892
Glacier Bay 1857
The northward retreat 1892

1845
of ice in Glacier Bay,

Alaska, since the 18th
century. As the Earth 1892
has become warmer,
following the Little Ice
Age of the 14th to the 1794
19th centuries, glaciers
1760–80
have been retreating
Icy Strait
around the world.
One glacier that has been studied in considerable detail is

the Jotunheimen in southern Norway, which has retreated by
about a mile (1.5 km) in 200 years. Geologists have made
even more detailed and long-term observations at the Glacier
Bay glacier in Alaska. Its northward retreat has been recorded
since the 18th century and the limits of the glacier can be
accurately traced (see illustration above). During the 200
years between 1760 and 1960, this glacier withdrew about 45
miles (72 km). Glaciers and their movements are thus prov-
ing to be one of the most sensitive indicators of the current
global warming.
Causes of glaciation
Although there have been several episodes of glacial activity
in the Earth’s history, during which ice caps have been pres-
ent in the polar regions, these are the exception rather than
the rule. For the bulk of the Earth’s existence there have been
no ice sheets or glaciers, so present-day conditions are by no
means typical for the planet. Why the Earth should have
periodically entered such a state over the past 2 billion years
remains something of a mystery, and observers have put for-
ward various theories to account for the rare instances of
glaciation.
The basis of one such theory is incomplete oceanic circula-
tion. One thing that is evidently needed for polar ice caps to
develop is the presence of a continental landmass at or near the
poles or for the landmasses to be so arranged that the free cir-
culation of waters around the Earth is interrupted. This pre-
vents the redistribution of energy from the Tropics to polar
regions. The relative isolation of the Arctic Ocean by the ring of
continents that currently surround it, for example, may have
played an important part in encouraging the Pleistocene glacia-
tions. Other possible explanations relate to the circulation of
the atmosphere. The elevation of mountain ranges, such as the
Himalayas and the Tibetan Plateau, can also create new atmos-
pheric circulation patterns that may induce major global cli-
matic changes, as can periods of excessive volcanism. It is
interesting to note that the onset of the most recent Ice Age
coincided with the elevation of the Alps and the Himalayas. Yet
another hypothesis takes into account events in space. Inter-
stellar dust and belts of asteroids could be responsible for peri-
ods of poorer penetration of the Sun’s energy to the Earth’s
surface. Clearly ideas abound, but a full explanation of the
occasional episodes of glaciation on Earth is still some way off.
Intensive study of the Pleistocene glaciations, however,
does provide some information about the pattern of glacial
cycles once an ice age has commenced. The last 2 million
years have shown a distinct pattern of glacial and interglacial
cycles. These have a cycle wavelength of about 100,000 years.
As long ago as 1864, Scottish geologists suggested that ice
ages could be caused by astronomical factors, particularly the
way in which the Earth wobbles on its axis in a regular and
predictable way. But it was a Yugoslav physicist, Milutin
Milankovitch (1879–1955), who finally put together a full
explanation of the cyclic pattern of the glacials during the
Pleistocene (see sidebar on page 144).
144 TUNDRA
The Milankovitch theory

Milutin Milankovitch (1879–1955) proposed the first comprehensive explanation of the
cyclic pattern of the glacials observed in the Pleistocene. In essence, Milankovitch claimed
that there are three patterns, all superimposed on one another, that cause the observed
pattern of glacial and interglacial cycles. First, the Earth follows an elliptical orbit around
the Sun, not a circular one. The result is that the Earth is closer to the Sun every 100,000
years. Second, the Earth is not upright on its axis, and the tilt of the Earth varies resulting
in stronger seasonal differences in temperature at certain times in history; this fluctuation
operates on a cycle of about 40,000 years. Third, there is the wobble of the Earth on its
axis, like a spinning top that has hit a bump, which occurs in a cycle of 21,000 years.
When these three cycles are combined to create a predicted model of how climate should
have changed over the past million years, they form a picture that closely resembles the
geological facts.
The expected timing of glacial episodes on the basis of the Milankovitch calculations
corresponds quite well with the observed dates obtained from field surveys. There are
complications, of course, but basically the astronomical theory devised by Milankovitch
forms an excellent basis for explaining glacial patterns. When he first proposed his model
in the 1930s, few geologists were impressed, but as more information has accumulated
the Milankovitch theory has gained much ground. It is now generally accepted as the
major underlying cause of the Pleistocene pattern of glacial and interglacial cycles.
Obviously, many other factors besides astronomical ones

influence the pattern of glacials and interglacials, so the
astronomical theory must be regarded simply as a starting
point in explaining the history of ice on Earth. As already dis-
cussed, ocean circulation patterns can change global climate
patterns as can mountain building episodes that alter the cir-
culation of the atmosphere. When volcanoes erupt, they
eject huge quantities of dust into the atmosphere that can
alter the climate of the whole planet. For instance, in 1982
the volcano El Chichón in the Yucatán Peninsula of Mexico
forced an estimated 16 million tons of dust into the atmos-
phere and prevented much solar energy from reaching the
Earth’s surface. Such dust veils can reduce the Earth’s temper-
ature substantially. Major eruptions of volcanoes in 1783
resulted in a temperature decline of 3°F (1.5°C) over the
Northern Hemisphere. So, while the climate of the Earth is

what primarily controls the extent of ice on the planet and
therefore the distribution of the tundra biome, many other
factors can also affect the climate.
Biological history of tundra

Ice ages, as discussed earlier in this chapter, are relatively rare
and widely spaced events in the history of the planet (see
table on page 131). Consequently the organisms able to sur-
vive in tundra habitats have had an interrupted history.
There is very little information about the types of organisms
that may have inhabited the tundra regions in the early his-
tory of the Earth, in Precambrian times, but presumably they
were limited to primitive bacteria. When the polar ice caps
returned in Carboniferous times, however, land vegetation
was present, and fossils of the cold regions have survived.
At this time, some 300 million years ago, many of the
present-day continents were all joined together in massive
supercontinents. North America and Europe lay together,
forming the supercontinent of Laurasia. They had recently
become separated from another, and larger, landmass to
their south. This southern landmass, called Gondwana, con-
sisted of what are now South America, Africa, Antarctica,
India, and Australia, and it lay over the South Pole. Perhaps
the location of this huge landmass over a polar position
accounts for the development of an ice age. In any event, an
ice sheet formed over what is now Antarctica and spread out
over the rest of Gondwana. Laurasia lay over the equator; its
climate, therefore, was a hot tropical one, and the extensive
swamp forests that formed at this time became the great
deposits of coal buried within North America and Europe.
While the equatorial supercontinent of Laurasia was occu-
pied by peat-forming swamps, the conditions on Gondwana,
to the south, were very different. In the northern part of
Gondwana (what is now northern South America and
Africa) was a dry desert. But farther south, around the ice
sheet, conditions were much cooler and the region bore very
different vegetation. Woody shrubs grew here, particularly a
plant called Glossopteris, which has given its name to an
146 TUNDRA
entire vegetation type, the Glossopteris flora. These plants

belong to an extinct group called seed ferns. Their leaves
looked like ferns and they were long regarded as true ferns
until fossils were found that showed they bore seeds. They
are now believed to be more closely related to the conifers.
The wood of these plants from the permafrost regions of the
Gondwana tundra shows distinct series of rings, indicating a
strong seasonality of growth (unlike the plants of the tropi-
cal coal swamps, which had no discernible growth rings).
Being quite close to the South Pole, the Glossopteris shrubs
must have survived through long polar nights in which little
or no growth was possible and then grown vigorously when
the summer season arrived. What is remarkable is the size of
these woody plants, considering how close to the South Pole
they grew. The Carboniferous/Permian Ice Age of about 300
million to 280 million years ago included at least five cycles
of glacial advance and retreat. The Glossopteris flora repre-
sents the first known tundra vegetation on Earth. As condi-
tions became warmer in Permian times, and the Earth
emerged from that ice age, the Glossopteris flora gradually
disappeared, presumably evolving new forms in response to
the changed set of conditions or giving way to other plant
types that were better equipped to cope with the changes,
such as Dicroidium, another seed fern.
After the Permian, when the first tundra vegetation died
out, Earth’s climate did not enter another ice age for a long
time. There is some evidence for glaciation in what is now
Antarctica in Jurassic times, 150 million years ago, but it was
not extensive. A different story emerges around 32 million
years ago, when the Antarctic Ice Sheet began to form once
again. The climate of the Earth was evidently on a gentle cool-
ing trend and glaciation commenced on high mountains in
Alaska by about 12 million years ago. This was when the tun-
dra plants and animals known today must have begun their
evolution. This tundra, therefore, is the youngest of all the
Earth’s natural biomes. For many millions of years there had
been no equivalent habitat on Earth—and this is true of no
other biome. There have been tropical forests, temperate
forests, open scrub, and deserts of one type or another
throughout the history of the planet, at least since land plants
and animals first evolved. But the history of this recent tun-
dra has been repeatedly interrupted, and each time an ice age
has hit the Earth, the plants and animals have had to begin
over again in their adaptation to the new environment.
In the recent ice age (the last 10 million years), the likely
origins of plants and animals appropriate to tundra condi-
tions are the mountains and the coniferous forests. The high
mountains of the world, such as those of Alaska 12 million
years ago, were becoming colder, subjecting the natural vege-
tation to increasing stress. Forests had occupied even the
high mountains in Miocene times, but conditions must have
made tree growth increasingly difficult, producing a more
open canopy. The increased light penetration would have led
to the evolution of new forms in the ground vegetation.
Plants of the blueberry family (Ericaceae), the saxifrages (Sax-
ifragaceae), the pinks (Caryophyllaceae), the cress family
(Brassicaceae), the grasses (Poaceae), the rushes (Juncaceae),
and the poppies (Papaveraceae) were all important in the
founding of tundra vegetation. Plants of wetlands, especially
the sedges (Cyperaceae), also found conditions in the devel-
oping tundra suitable for their needs. Among the trees,
birches (Betulaceae) and willows (Salicaceae) proved most
able to develop new forms that could cope with cold and
wind exposure. Joining them, of course, were the mosses and
lichens that could survive even under the harshest condi-
tions. Most important among these were the bog mosses
(Sphagnum species) and the “reindeer moss” group of lichens
(Cladonia species).
Many mammals evolved tundra forms, among them some
surprising groups. The bears proved adaptable, as did deer
(such as caribou), the elephants (mammoths and
mastodons), foxes, cats, rodents, and especially the mustelids
(including stoats, weasels, and wolverines). Birds adapted to
fit the conditions, including songbirds (such as buntings),
owls, raptors (such as falcons and eagles), and grouse,
together with a wide range of seabirds and waders. As the
tundra habitats developed and extended, some mammals
and birds developed migratory patterns, allowing them to
exploit the productivity of the tundra without having to
experience the hardship of polar winters.
148 TUNDRA
Alpine tundra on the lower-latitude mountains also devel-

oped as the world grew cooler. Many of the plants and ani-
mals that were adapting to the polar conditions also
succeeded in the mountains, but their geographical isolation
produced some differences. Mountaintops are rather like
islands in the ocean. They can be widely separated from one
another and the intervening landscape can be difficult to
cross. As a result many mountains began to develop their own
plants and animals in isolation from the general evolution of
tundra organisms. Many alpine species, unable to interbreed
with other members of their groups on neighboring moun-
tains because of the difficulties of travel, began to evolve sep-
arately and produce new forms. The East African mountains
(see “Tundra plants,” pages 86–94) are an excellent example
of this process, in which strange and unique forms arose.
Species that have developed in and remain restricted to their
geographical ranges in this way are called endemics. Mountain
tundra is a habitat rich in endemic species, which makes
them important as reserves of the world’s biodiversity.
People in the tundra

The human species, Homo sapiens, first appeared during the
Pleistocene and is likely to have originated in Africa. Genetic
evidence suggests that all modern humans have a common
ancestor. Humans proved very adaptable to the changing cli-
mate; modifying the environment by wearing the skins of
animals and living in caves heated by fire meant that
humans were able to cope with cooler climates, even the
extremes of the tundra. During the last ice age, humans occu-
pied much of Europe and Asia, hunting many of the large
herbivores of the tundra. In what is now Russia, people
preyed heavily on mammoths and even used their bones to
construct dwellings. Cave-dwelling people in the west of
Europe painted pictures of their prey animals on the walls of
their caves, perhaps hoping thereby to obtain some magical
power over them in the chase. They used primitive weapons
of stone, but by hunting in packs and driving game into con-
fined spaces or traps or over cliffs, they were able to achieve
much more than might have been expected of such a rela-
tively small predator. They also domesticated the wolf and

used this animal as an assistant in driving game.
Late in the last glaciation (probably before the maximum
glacial advance 20,000 years ago), humans crossed from Asia
into North America. World sea levels were low at that time,
so the narrow seas that separate Alaska from Russia were then
mainly dry land, offering no serious barrier to the movement
of animals, including humans. From these invading people
the whole of the Americas was populated, possibly in several
waves. A study of the ancient languages of the Native North
American populations suggests that there were three main
groups: one centered in the far north and west, one on the
Pacific coast, and one in the larger southern region of what is
now the United States. These were the areas that were not
covered by ice during the final (Wisconsin) glaciation, and
the Stone Age peoples occupying these tundra landscapes
around the edge of the massive ice sheet probably survived
by hunting the tundra animals.
The impact of hunters upon these animals, especially the
very large ones such as mammoths, proved fatal. All over the
world, within a few thousand years of human settlement,
the great mammals or “megafauna” of the tundra were all
extinct, including mastodon, cave bear, giant elk, and mam-
moth (see sidebar on page 150). The debate about these
extinctions, and the role of humans in causing them, is still
not finally settled, but many pieces of evidence point to a
human agent. The timing of the extinctions is remarkably
consistent, occurring largely between 12,000 and 10,500
years ago. This was when human populations were growing
and people were developing more sophisticated stone
weapons. Climate, however, was also changing rapidly at
this time. Could the loss of these large tundra animals have
been due to the reduced area of tundra habitat? Certainly
this would have constrained their populations, but several
other warm interglacial periods during the past 2 million
years had occurred that had not proved fatal to these
species, so what was different about this one? The answer
seems to be humans. Never before had the large mammals
faced such a formidable predator. For these cumbersome
beasts, which were becoming increasingly restricted in their
150 TUNDRA
The woolly mammoth

The woolly mammoth had the appearance of an elephant with long, shaggy hair. It stood
about 13 feet (4 m) high at the shoulder and its tusks could grow to a length of about 15
feet (4.5 m). These tusks can still be found buried in the soils of the tundra regions, espe-
cially in Siberia, where local people have made an income by selling them to ivory traders.
It was one such seeker of ivory who first discovered an intact body of a woolly mammoth
embedded in permafrost in 1799. It was extracted relatively intact and studied by scientists.
Since then many such bodies have come to light and the possibility has been discussed
that the DNA could be used to reconstruct the species, possibly using the elephant as a
surrogate mother. These “Jurassic Park” plans, however, have not yet proved viable
because DNA is a relatively fragile molecule, and even when preserved in ice it has usually
severely degenerated over the course of many thousands of years.
area of occupation as the tundra retreated, human presence

was the final nail in their coffin.
The Inuit people of the Arctic regions (one of the three
North American language groups) settled the northern
regions of what is now Canada in prehistoric times. They
spread from Alaska, where they arrived after crossing the
Bering land bridge from Russia, and gradually extended their
range eastward, reaching Ellesmere Island to the west of
Greenland about 4,000 years ago. Using ice as their winter
building material and skins and bark in summer, they left lit-
tle evidence of their occupation for archaeologists to study.
But more permanent settlements, such as campsites that used
boulders in protective circles, still survive and supply evi-
dence of a way of life that must have changed little over
many thousands of years. Ellesmere Island lies only 470 miles
(756 km) from the North Pole, so the prehistoric people who
occupied the region must have been extremely hardy to per-
sist in the face of the extremely harsh climate. Like the mod-
ern Inuit peoples, they would have subsisted largely on
animal flesh obtained by hunting. Seals, birds, caribou, and
fish probably formed the major part of their diet.
The Viking, or Norse, people of Scandinavia were great
travelers and colonists, and they spread out across the islands
of the North Atlantic around the 10th century C.E. They set-
tled Iceland and southern Greenland, surviving there for sev-
eral centuries. Dispute still surrounds the claim that they
traveled farther and even reached North America. One of
their traditional tales, or sagas, tells of the land of Vinland,
where the wild grape was discovered. Archaeological evi-
dence indicates that they settled Newfoundland, but penetra-
tion farther south cannot be conclusively proved.
Antarctica essentially has no prehistory. People did not Inuit hunters with their
reach this continent—or, indeed, confirm its existence—until harvest of caribou
the late 15th century. It is the one continent that has a his- carcasses in Alaska.
tory devoid of the impact of humankind until very recent Caribou have long
times. been an important prey
Although archaeologists know something of the people item for the Inuit people.
of the Arctic tundra during prehistoric times, until very (Photo by Michio
recently they knew next to nothing of human activities in Hoshino/Minden
the alpine tundra regions. The high mountains of the world Pictures)
152 TUNDRA
seemed, like Antarctica, to be devoid of prehistoric human

life. In September 1991, however, two German moun-
taineers were climbing in the Italian Tyrol district of the
Alps in Europe. They were descending from a mountain
peak they had climbed close to the Italian/Austrian border
and were crossing an ice field when they saw something
brown sticking out of the ice. It was the head and shoulders
of a dead body emerging from melting ice. Assuming that it
was an unfortunate mountaineer who had met with a
recent accident, they reported it to the police in Austria
where they descended. When the body was recovered and
examined by pathologists and archaeologists, it proved to
be the victim of a very old mountaineering accident, not a
recent one. In fact, the body was about 5,000 years old and
had been preserved in the ice ever since the unfortunate
traveler had perished, probably in a snowstorm while cross-
ing from Austria to Italy. He wore clothing made of chamois
skins, padded with grass to increase the thermal insulation.
His boots were also filled with grass, and he wore a bearskin
hat. He carried a bow and a quiver full of arrows, so he was
either a huntsman or was expecting trouble on his journey.
He also had equipment for lighting a fire and an ice ax
made of copper. Forensic scientists who analyzed the DNA
in samples from his body confirmed that he had originated
from north-central Europe. It seems he was either crossing
into Italy or was on the return journey to his northern
home.
This discovery in the Alps has provided a rare, if not
unique, glimpse into the life of prehistoric humans in the
alpine tundra regions. There is much archaeological evi-
dence for settlements in the Alpine valleys, where ancient
people farmed. They apparently needed to cross the high
mountains on their travels and they may have also used the
high pastures and alpine tundra for grazing animals in sum-
mer. These prehistoric people were familiar with the change-
able and dangerous weather of the mountains and had
developed equipment to cope with these conditions. The
fact that so few bodies have been found suggests that they
coped fairly well in lands that presented so many survival
problems.
Tundra exploration
The tundra regions of the Arctic, then, were inhabited by
people long before the world’s civilizations were fully aware
of the existence and geography of such lands. The northern
lands were largely known through vague myths and stories.
Global exploration, until very recent times, was driven
mainly by the need to expand the opportunities for food pro-
duction to support human populations, or to gain wealth
through trade, so the far north had few attractions. The
Viking people of Scandinavia were the only European culture
to exploit the economic opportunity of colonizing the
remote North Atlantic islands and seaboards. The discovery
(or perhaps more accurately, the rediscovery) of North Amer-
ica by Christopher Columbus (1451–1506) in 1492 came at a
time when global travel and trade by sea were increasing rap-
idly. European colonialism became an incentive for further
exploration that lasted for several centuries and led, among
other things, to the first modern expeditions by Europeans
into the far north. Most such voyages of discovery proved
disastrous, but explorers made enormous efforts to discover a
northwest passage that would link the North Atlantic to the
North Pacific. If found, it would provide a trade route to the
Far East, allowing ships to avoid the lengthy and dangerous
passage around Cape Horn at the southern tip of South
America or eastward around the Cape of Good Hope in South
Africa.
Exploration in the Arctic in search of the Northwest Pas-
sage accelerated in the 18th century, and many lost their
lives in trying to win the prizes European governments
offered to anyone who found it. Most expeditions set off
from the Atlantic and many ended in Hudson Bay, but at the
end of the 18th century attention moved to the North
Pacific. The British naval officer George Vancouver (1757–98)
took his ship, HMS Discovery, into the Icy Strait of Glacier Bay
in Alaska in 1794 and was impressed by the 10-mile-long (16-
km) ice wall where glaciers entered the ocean (see the illus-
tration on page 142). The American naturalist and
conservationist John Muir (1838–1914) visited the region 85
years later and found that the glaciers had retreated 48 miles
(77 km), probably as a consequence of global warming. The
154 TUNDRA
main glacier of Glacier Bay subsequently became known as

the Muir Glacier.
Exploration of the northern regions also proceeded on
land, particularly by companies interested in fur trading. In
1789 the Scot Alexander Mackenzie (1755–1850), who was
employed by the North West Fur Company, set out with
Native American and Canadian colleagues in birch-bark
canoes to seek a river route from the continental areas of
North America into the North Pacific. He found a great river
that led westward from the Great Slave Lake, but it gradually
turned northward and eventually led into the Arctic Ocean
instead of the Pacific. He named it the “River of Disappoint-
ment,” but it later became known as the Mackenzie River.
The expedition may not have discovered a link to the West
Coast, but it did represent a major advance in the explo-
ration of the tundra regions. The discovery also opened up
the region to fur trappers, whale hunters, and gold prospec-
tors. It is now an important region for oil extraction.
The 19th century saw a great rise in public interest in
exploration in general, and the inaccessible tundra regions of
the world rivaled the inaccessible center of Africa in its attrac-
tions for the intrepid explorer. The romantic image of the far
north, with its mountainous icebergs and bleak isolation, was
stimulated to some extent by the publication of Mary Shel-
ley’s novel Frankenstein in 1818, which depicted its now-
famous monster in the icy tundra setting. The British
launched a series of expeditions to try to reach the North
Pole during the early part of the 19th century, but they all
became stranded in sea ice, having to either turn back their
ships or abandon land-based excursions over the ice. The
British were determined to discover a northwest passage to
the Pacific, their interests being more economic and compet-
itive (beating the Russians to the discovery) than scientific.
Perhaps the best known of these remarkably brave explorers
is John Ross (1800–62), whose name now identifies several
geographical and biological features, including Ross’s gull
and Ross’s goose. The first European to spend a winter in the
High Arctic (in 1820) was William Edward Parry (1790–1855),
who wrote of the “deathlike stillness” and the “dreary desola-
tion” of the experience.
In the late 19th century the media entered into the realm
of exploration. James Gordon Bennett, Jr. (1841–1918), edi-
tor of the New York Herald newspaper, was a great supporter of
exploration. His most famous exploit was to support Henry
Morton Stanley (1841–1904) in his search for Dr. David Liv-
ingstone (1813–73) in remotest Africa, but he also sponsored
Arctic exploration. In July 1879 he provided the financial
backing for an attempt to reach the North Pole by George
Washington De Long (1844–81). Believing that the North
Pole was surrounded by open ocean, De Long set out north-
ward from the Bering Strait, but by September his boat
encountered masses of pack ice, and he spent two years in
the Arctic Ocean north of Siberia before the boat was eventu-
ally frozen in and crushed. Three open boats set off from the
wreck. Only one of the boats reached safety; De Long and his
party died before help could reach them.
The successful discovery of the Northwest Passage did not
occur until the voyage of Roald Amundsen (1872–1928), a
Norwegian who later became the first man to reach the South
Pole. His voyage took place between 1903 and 1906 and it
took him north of Baffin Island and Hudson Bay, through
Baffin Bay, and then through the channels separating Victo-
ria Island and Banks Island from the mainland. Eventually
Amundsen arrived at the Mackenzie River delta. To travel
from there to the Bering Strait, he had to wait until the ice
had melted, and then he finally sailed down the West Coast
to San Francisco. By the time he arrived, the news of his suc-
cess had already become a little stale. Moreover, the excite-
ment that might once have surrounded the achievement had
diminished: Although the existence of the route had now
been demonstrated, it clearly could never become commer-
cially important. (Flight has changed this. Perhaps ironically,
jets from Europe to the west coast of North America essen-
tially follow the course of the long-hoped-for Northwest Pas-
sage through the Arctic tundra.)
The North Pole itself eluded human contact until April
1909, when a U.S. naval officer, Robert Edwin Peary
(1856–1920), led an expedition from Fort Conger on Ellesmere
Island in Canada. Using relays of dogsleds and aided by the
Inuit people, the team laid out a route with supply stations.
156 TUNDRA
Peary himself, together with one companion, covered the final

155 miles (250 km). The achievement was marred to some
extent by a claim (that later proved false) by another explorer
that he had reached the North Pole in the previous year. But
Peary and his companion are now regarded as the first people
ever to reach this remote and ice-bound location. The seasonal
timing of such expeditions is critical, since they must take
place after the worst of the winter weather and yet before the
Arctic Ocean pack ice breaks up and becomes dangerous.
Besides the dangers of collapsing or rafting ice, there is the
ever-present risk of wandering polar bears who can prove
aggressive.
People had long been aware of the existence of the Arctic
tundra. The Vikings were familiar with the tundra, and tales
of the bleak north were present in Greek and Roman mythol-
ogy, which suggests that even these early civilizations had
some contact with the far north. The Antarctic, however, was
shrouded in mystery until very recent times. The Greek
philosopher and naturalist Aristotle speculated in the fourth
century B.C.E. that the great northern landmasses must be
“balanced” by a landmass in the deep south. This supposed
continent later became known as Terra Australis Incognita,
the “Unknown Southern Land.” Aristotle called it “Antark-
tikos.” The northern lands lay under the constellation Ark-
tos, “The Bear,” so the southern lands must be the opposite
of this, he reasoned.
When the Portuguese explorer Vasco da Gama (1469–1525)
sailed around the southern tip of Africa and eventually
reached India in 1497, he proved that Africa was not joined
to the Unknown Southern Land. Ferdinand Magellan
(1480–1521), another native of Portugal, sought the southern
tip of South America in 1519. He sailed through the straits
that now bear his name, which separate the mainland of
South America and the island of Tierra del Fuego to its south.
But he was not sure whether this was truly an island or
another great landmass, the Unknown Southern Land. The
English sailor Francis Drake (1540–96) settled this issue when
his ship was blown south in the Pacific by a great storm and
ended up rounding Cape Horn, thus demonstrating that
there was no connection between South America and the
Southern Land. He also provided the first description of pen-

guins, noting that they were easy to catch (being flightless)
and good to eat.
By the mid-17th century, Dutch traders were regularly sail-
ing in the South Pacific, traveling along the west coast of
Australia, and they considered it possible that this was the
Unknown Southern Land of Aristotle. Among these Dutch
explorers, Abel Janszoon Tasman (1603–59) was the first to
circumnavigate Australia and to encounter Tasmania and
New Zealand. But these lands were temperate in latitude and
climate, not the equivalent of the Arctic wastes.
The notion that the Antarctic continent might exist con-
tinued to appeal to explorers, but it did not have the eco-
nomic appeal of the Arctic in terms of the new sea routes it
might possibly offer. The fishing and whaling potential of
the Southern Ocean, however, eventually led to its system-
atic exploration. The British navigator James Cook (1728–79)
spent much of his time exploring the Pacific Ocean, includ-
ing its southern regions, during the latter half of the 18th
century. He was the first to sail south of the Antarctic Circle,
in 1773, but had to turn back on his southbound voyage
because of pack ice. He little realized that he was only 80
miles (130 km) from the legendary continent. Cook suc-
ceeded in circumnavigating the fringing ice packs of Antarc-
tica but was never convinced that the southern polar regions
contained more than floating ice.
It was a Russian, Fabian Gottlieb von Bellingshausen
(1778–1852), who first sighted land in 1820. In unusual con-
ditions of excellent visibility (something denied to Cook), he
spotted high cliffs of rock, thus proving that a southern con-
tinent existed. The first person to set foot on the continent
was John Davis, an American sailor, who landed on the
Antarctic Peninsula (see the map on page 3) just a year later,
in 1821. James Ross, of Arctic tundra fame, was also eager to
add the Antarctic to his polar explorations, and in 1841 he
led an expedition that aimed to reach the South Pole. As he
approached the Pole, however, he found his passage blocked
by an enormous mass of pack ice, now known as the Ross Ice
Shelf, and he was unable even to approach the region of
Antarctica in which the Pole lies. He did, however, see a great
158 TUNDRA
volcano that was active at that time, and he named it Mount

Erebus. Erebus is the name the Greeks gave to the region of
darkness through which the dead must pass, which gives
some indication of the awe in which Ross and his fellow
sailors held these inhospitable regions. The volcano remains
periodically active, but its main claim to fame is a tragic acci-
dent in 1979, when a plane carrying tourists became
enveloped in a blizzard and crashed on its slopes, killing 257
passengers and crew.
The South Pole itself was not reached until 1911, as a
result of a dramatic and tragic race between two teams of
explorers. From Norway came a team led by Roald Amund-
sen, and from Britain, one led by Robert Scott (1868–1912).
The Norwegians set out on October 19, using four sledges
pulled by dogs. Their most difficult section consisted of the
climb up to the polar plateau, over the dangerous ice mass of
the Axel Heiberg Glacier. From there, the going became eas-
ier as Amundsen’s team crossed the relatively flat plateau,
eventually reaching the South Pole on December 14 and
raising the Norwegian flag. The British team reached the
Pole on January 17, having hauled their sledges over the
entire course, without the use of dogs. Scott’s comment on
reaching the South Pole was, “This is a terrible place.” Ill for-
tune plagued the return journey of Scott’s team. Despite acts
of heroism and self-sacrifice, blizzards and frostbite took the
lives of all members before they could reach their base. Once
again, the remarkable success of Roald Amundsen was some-
what overshadowed in the news media by the romantic yet
tragic failure of Scott’s team. In both his Northwest Passage
expedition and in his attainment of the South Pole, Amund-
sen seems to have missed out on the public acclaim that was
his due.
With the Amundsen and Scott expeditions the great
romantic age of Antarctic exploration ended and the period
of commercial and scientific investigation took over. This
was also the beginning of a period of human history in
which the human population has expanded dramatically
and the need for food and raw materials has led people to
exploit even the most unlikely of the world’s wildernesses.
Having discovered the vastness of the tundra in general and
Antarctica in particular, humanity now asked how the tundra

could be used to human advantage.
Conclusions
Ice ages, periods when ice sheets and tundra are present on
Earth, are relatively rare events in the history of the planet
and seem to occur in a regular pattern, roughly every 150
million years. The causes of these long-term climate cycles
are not understood, but the events may relate to the posi-
tions of continents on the surface of the Earth and the way
they influence ocean currents. Earth is currently occupying a
warm interval within an ice age, and it is possible to detect
more detailed patterns of climatic fluctuation, with alternat-
ing peaks of warmth and cold at intervals of approximately
100,000 years. This scale of climate pattern appears to be
related to astronomical factors, especially the varying dis-
tance between the Earth and the Sun.
Tundra plants and animals have also experienced an inter-
rupted history, and the present vegetation and fauna of tun-
dra is relatively recent in origin, perhaps less than 10 million
years old. High mountain organisms may well have been the
first tundra dwellers, expanding in range and abundance as
the cold of the current Ice Age intensified, especially over the
past million years.
Humans lived in tundra habitats of Europe and Asia
throughout the last glacial period, which reached its maxi-
mum extent around 20,000 years ago. They spread from the
eastern tip of Asia into North America as that glacial episode
began to recede, passing over what was then a dry land
bridge between Asia and Alaska. The Inuit people took up
permanent residence in the tundra and adapted their culture
to life in the bitter cold of the far north. Viking exploration
around the 10th century C.E. led to the invasion of European
peoples into the North Atlantic tundra regions, but these set-
tlements proved to be temporary. Renewed exploration
began in the 15th century when commercial traders sought
alternative routes to the Far East, and this led to a fuller
understanding of the geography and biology of the Arctic
tundra.
160 TUNDRA
The Antarctic has never been populated, even in prehis-

tory. The existence of a great southern continent was
hypothesized since classical times, but evidence was lacking
until the intrepid sea voyages of the 18th and early 19th cen-
tury. The first human to reach the South Pole, the Norwegian
Roald Amundsen, achieved this goal only in 1911, less than a
century ago. Human history in the Antarctic, therefore, is
very recent and brief.
CHAPTER 6
USES OF THE TUNDRA

Whenever people have discovered and explored a new region
of the world, their first concern has been its exploitation. The
hope of becoming rich has often motivated those setting up
exploratory expeditions, even though the geographers and
biologists involved in them may have had higher and more
scientific aims. The tundra regions of the world are no excep-
tion, and many of the early expeditions into the Arctic were
concerned with the discovery of new trade routes that would
bring faster travel and greater riches. Once these hopes were
abandoned, the potential of the Arctic wilderness for hunting
and mineral prospecting soon took over. Understanding the
economic value of the tundra helps us to appreciate its his-
tory and also provides an insight into the problems that this
biome will face in the future.
Hunting and trapping

Hunting has been the basis of human life in the tundra since
the human species first invaded this realm. Tundra plant life
offers few opportunities for food gathering, apart from some
berries in the late summer. Agriculture, as the Norse people
invading Iceland and Greenland were to discover, was of little
use in these northern latitudes. But herds of large grazing
mammals, such as mammoths and caribou, provided a reliable
source of food for human cultures prepared for a largely car-
nivorous diet. The Inuit people of Alaska and northern Canada
have traditionally been coastal in their distribution and
marine-based in their culture. The most northerly settlement
of people in North America is on Ellesmere Island, where Inuit
settlers have been present for at least 4,000 years.
Later, when European explorers and pioneers penetrated
north through the boreal forests of Canada, they discovered
161
162 TUNDRA
lands rich in mammals whose skins were extremely valuable

in the fashionable cities of the south. Similarly, the wastes of
Siberia provided a supply of furs for the citizens of Moscow
and St. Petersburg in Russia, so trappers lived hard lives but
made good profits. Trading companies such as the Hudson’s
Bay Company developed more formal routes for the move-
ment of furs from the outback to the consumers in the cities.
For many decades, the wealth of the tundra lands, north of
the forests, lay mainly in their mammals.
In Europe one animal, the caribou or reindeer, formed the
main source of sustenance for the Sami (sometimes called
Lapp) people of northern Scandinavia. They followed the
migratory herds over the tundra landscape, using the animals
as a source of meat and hides. They protected the herds from
other predators, such as wolves, thus ensuring that young
calves survived and increased the size of the herds. Thus
began a kind of symbiotic relationship between the Sami
people and the reindeer, resulting in a partial domestication
of this docile mammal. Herds are now carefully protected
and managed in northern Europe and Asia and remain the
foundation on which Sami culture is based.
In Antarctica, uncontrolled hunting of marine mammals
for their skins became a major industry within a few decades
of Captain Cook’s first penetration of the Antarctic Circle in
1773 (see “Tundra exploration,” pages 153–159). Seals,
including fur seals, were abundant in the sub-Antarctic
islands at the beginning of the 19th century, but many of
these island colonies had been completely exterminated by
1830. When the seal populations had been depleted, hunters
turned to penguins, which provided a source of oil from their
stored fat. Protection of seals and penguins in recent times
has fortunately led to the recovery of many species, includ-
ing fur seals and king penguins.
Whales abounded in the southern oceans, and whaling sta-
tions were set up in the early 20th century on the Antarctic
islands to assist in the harvesting and butchering of these
enormous marine mammals. The major impact on whale pop-
ulations began with the development of sophisticated hunting
methods, powerful harpoons, and robust ships. The major
problem with whale harvesting is the animals’ very slow rate
of reproduction, which means that populations recover from

losses very slowly. Since 1949 the International Whaling Com-
mission has attempted to regulate whale harvesting and since
the 1960s some species have been fully protected. Some whale
species are now increasing their populations, but whales have
slow breeding rates, so it will take a long time for these animals
to recover fully from the impact of hunting.
Native peoples of the Arctic regions have a long history of
hunting seals and cetaceans (whales and dolphins). For
many species, hunting by traditional methods involves a
limited harvest that is sustainable. In the case of the beluga,
for example, a regulated harvest is possible. But for some
scarcer species, such as the bowhead whale, the largest of
the Arctic sea mammals, even a small harvest could damage
the limited population. This means that even the tradi-
tional hunting by native tundra peoples needs to be moni-
tored and controlled. Seal hunting for skins that can be
exported, which has traditionally played an important part
in the economy of the Inuit people, has suffered a major
downturn in recent years. The reason is that the demand for
seal skins around the world has declined as people have
moved from animal skin to synthetic materials for their
clothing. Ecological concerns among the consumers of the
world are having a substantial impact on the economy of
the people of the far north.
Hunting in alpine tundra has also resulted in drastic falls in
the populations of some mammals. Bighorn sheep have suf-
fered in North America, and chamois and ibex have under-
gone considerable declines in the Alps of Europe. All of these
species have increased as a result of protection in recent years.
Hunting is still permitted, but it is controlled and is often
directed at the large males that are prized as trophies. Even
this activity, however, could have an impact on the genetic
constitution of the species because it involves the consistent
elimination of the biggest and perhaps the fittest males.
Mineral reserves
Mineral prospecting in the tundra regions developed along-
side hunting and trapping in the early history of tundra
164 TUNDRA
exploitation. In the early days, the remoteness and the trans-

port difficulties meant that only the most valuable of geolog-
ical resources were worth pursuing, and chief among these
was gold.
On July 17, 1897, a steamship arrived in Seattle harbor
from the extreme northwest of Canada. It carried news of a
considerable strike of gold on the Klondike River, a tributary
of the Yukon River of Alaska. The news became exaggerated
when journalists claimed that the boat contained a ton of
solid gold, and the outcome was a crazed rush into the north-
ern lands of men lured by the prospect of untold riches. The
strike was indeed a rich one, but the Klondike River lay in an
extremely remote region of mountains, and those who set off
faced extreme hardships of travel and survival in the inhos-
pitable tundra. Nevertheless, prospectors arrived not only
from North America but also from as far as Europe and Aus-
tralia, and the settlement of Dawson City soon became a
boomtown to service the gold rush of the Klondike.
The extraction of mineral wealth inevitably involves geo-
logical disturbance and results in the contamination and silt-
ing of streams and rivers. This was not a primary concern in
the late 19th century, but it has become an increasingly
important consideration in the environmentally conscious
days of the 21st century. A modern understanding of the
fragility of the tundra ecosystem has added to these con-
cerns. Although the extraction of gold from rocks is now
more sophisticated and efficient than in the days of the
Klondike gold rush, the efficiency of extraction means that
larger masses of rock can be treated and more waste is pro-
duced. Extraction of enough gold for a pair of wedding bands
can generate a truckload of waste rock. The environmental
damage associated with gold mining, however, has now
moved from the tundra regions to the Tropics, in Brazil and
Africa.
The tundra habitats of high mountains are also subject to
mining activities. The mountains of Utah contain the largest
hole that has ever been created by human mining activities,
Bingham Canyon. More than 2.5 miles (4 km) in diameter
and half a mile (0.8 km) deep, it was created to permit copper
extraction from the rocks. Open-pit and strip-mining activi-
ties of this type create the most visually destructive impacts

on the landscape, but even subsurface mining creates the
problem of waste disposal and results in the accumulation of
heaps of discarded rock. These piles are often slow to be colo-
nized by plants because of the rock’s high metal content. Col-
onization in the cold tundra habitats is even slower than that
associated with lowland spoil heaps. Apart from metals, some
mountain areas are mined for the rock itself, which is used
for building or decorative purposes. Granite, limestone, and
slate are particularly in demand.
The possibility of commercial mineral exploitation in
Antarctica has raised many difficult questions about the
management of this wilderness continent. In 1959 all inter-
ested governments, including the United States and the
Soviet Union, signed the Antarctic Treaty. Its main concern
was that the continent should be used only for peaceful pur-
poses and scientific research, denying any country the right
to establish military bases or to conduct weapons testing on
the continent. But environmental protection required more
than this, and the question arose of whether and how to
exploit the likely mineral resources of Antarctica, a subject
not covered by the treaty. During the cold war the Antarctic
provided an important point of contact for the great world
powers, and the Antarctic Treaty System evolved, which
sought to develop international cooperation on management
strategies for Antarctica. The signatories of the Antarctic
Treaty worked through the 1980s to establish the Conven-
tion on the Regulation of Antarctic Mineral Resource Activi-
ties, which assumed that mining would occur but tried to
keep environmental damage to a minimum. Then, in 1989,
France and Australia took a tougher stand and demanded a
ban on mining activities. The other nations involved eventu-
ally agreed to this policy, but only on the condition that the
ban should have a time limit permitting future reconsidera-
tion of the use of the mineral resources of Antarctica. The date
agreed upon was 2048, a mere 50 years after the agreement
came into force. For the present the Antarctic is safe from
mineral extraction, but the battle for the protection of the
tundra is not completely over. The global demand for mineral
resources continues to grow, so there will undoubtedly be
166 TUNDRA
renewed pressures upon the Antarctic wilderness by the mid-

dle of the 21st century.
In the northern tundra regions, a new resource has been dis-
covered that is creating even greater environmental problems
than the gold of the past, namely oil and gas reserves. Major
consumers of oil, in particular the United States, have been
eager to discover resources within their own national bound-
aries and thereby to avoid the expense and the vulnerability
associated with depending on imports. The tundra regions of
Alaska and Canada contain important reserves of oil that are
now being extracted. Other reserves will undoubtedly be dis-
covered. It is important to bear in mind, however, that these
reserves are limited and will be exhausted relatively quickly.
The North Slope of Alaska, for example, is estimated to contain
only enough oil to supply all of the demands of the United
States for three years. There are major oil fields beneath the
Beaufort Sea, along the coastal region of northeastern Alaska
and northern Canada. But this region of Alaska comprises the
Arctic National Wildlife Refuge, so the goals of providing the
nation’s energy needs and conserving its wildlife heritage are in
conflict. It remains to be seen whether the two goals can coex-
ist. Meanwhile, additional fields are being discovered out
among the islands of the Canadian Arctic, so the debate and
the problem will continue for some time to come. Global
warming and the continued melting of the ice cover in the Arc-
tic Ocean should permit continued exploration of the region
(see “Consequences of climate change,” pages 175–178).
The exploitation of oil reserves in the tundra lays it open to
many harmful environmental impacts. The most obvious of
these is spillage and pollution, both at the point of extraction
and during transportation. Oil is particularly harmful in
marine situations. Having a low density it floats upon water,
and, depending on its viscosity and stickiness, it can produce
either masses of thick, coagulated rafts or a thin film distrib-
uted over a large area. Floating oil is especially harmful to
seabirds, because it can coat their feathers and leave them
unable to fly or to dive, so oil-contaminated birds are in dan-
ger of starvation. In addition, as they preen their feathers the
seabirds ingest toxic chemicals from the oil and become poi-
soned. The oil may also wash onto shores, where it damages
marine life along cliffs and beaches. Cleaning up after oil spills
usually involves the use of detergents that emulsify the oil, dis-
persing it into very small globules that eventually decompose
in the water. But the detergents are often more harmful to
wildlife than the oil itself, and many conservationists believe
that the cleaning operations do more damage than good.
Oil extraction in the tundra also involves the establish-
ment of settlements and the development of roads or other The Alaska oil pipeline
transport systems. Settlements generate waste and, as has between Prudhoe Bay
been described, waste matter decomposition is slow in the and the port of Valdez,
tundra, so waste mountains can develop. Apart from being is owned by Alyeska
unsightly, waste heaps attract pests, from rats and gulls to Company. It crosses the
polar bears. Road development on tundra soils is also diffi- Brooks Range of
cult because of the freezing and thawing that takes place each mountains, North
fall and spring. Hard surfaces in the winter turn into wet- Slope, and heads south
lands in the summer, and roads quickly break up under the toward the Gulf of
strain. Oil pipelines provide an alternative to the use of truck Alaska. (Photo by
transport, especially where long distances are involved. The Yva Momatiuk and
Trans-Alaska Pipeline runs from Prudhoe Bay on the North John Eastcott/
Slope south to the Gulf of Alaska and avoids the need for Minden Pictures)
168 TUNDRA
either fleets of tankers to pass through the Bering Strait or

lines of trucks to traverse the Alaskan wilderness, but the
pipeline nonetheless presents certain problems. Pipelines
fragment the landscape and can create barriers to the move-
ment of large mammals, such as migratory caribou. This frag-
mentation could affect the survival of some herds, isolating
them genetically and perhaps exposing them to new levels of
predation.
Ecotourism and recreation

Television, books, and films have provided a wealth of infor-
mation about the Earth and its wildlife, making the wonders
of the natural world more accessible, even those wilderness
regions remote from human habitation. It is perhaps
inevitable that people increasingly desire to visit such places
and experience the wilderness firsthand. As wealth, leisure
time, and the availability of global transport increase, more
and more people are indulging in an activity that has become
known as ecotourism (see sidebar at right).
In the late 19th century, the prospect of visiting the Arctic
tundra as a tourist was almost as ambitious as space tourism
is today. But in 1892 a German shipping company began
running tourist trips to the Arctic island of Spitsbergen. Inter-
rupted only by two world wars, these and similar tourist voy-
ages continued until 1975, when an airport was opened on
the island. Cruise ships remain the most popular means of
visiting the polar regions of both Russia and North America,
but the services now provided usually include educational
information supplied by expert lecturers and guides. The
North Atlantic provides the best opportunities for approach-
ing the polar regions, concentrating on Greenland, Spitsber-
gen, and Baffin Island in Canada. Voyages in the North
Pacific usually focus on Alaska, particularly Glacier Bay, but
some trips pass north through the Bering Strait. In recent
years, Russian, Canadian, and American icebreakers have
even succeeded in taking paying passengers to the North
Pole.
One great advantage of using cruise ships as a means of
ecotourism is that they can carry in all the supplies needed
What is ecotourism?
Essentially, ecotourism has two distinct features. First, it is a type of tourism in which the
focus of attention and interest is the natural world and its associated human cultures.
Ecotourists seek out wilderness areas of natural beauty in order to appreciate the scenic
wonders, exciting wildlife, and distinctive culture of the resident people. Second, eco-
tourism sets out to be environmentally friendly, seeking to avoid any ecological damage
to the area visited: to look but not touch. In some respects ecotourism has replaced the
game-hunting of former times. The gun has been replaced by the camera, and the hunt-
ing no longer results in the death of an animal.
The desire to avoid any harmful impact, however, is very difficult to fulfill. When
tourists from developed countries visit wilderness areas they usually demand high stan-
dards of hygiene, good food, clean water, and effective waste disposal, all of which are
difficult to achieve in remote areas. The natural wilderness found in the tundra regions of
the world, both polar and alpine, provides an ideal target for the development of eco-
tourism. But providing for the needs of the ecotourists, especially waste disposal, places a
strain on the resources of the regions they have come to enjoy.
for the tourists and remove all the waste generated by the
same means. Waste disposal is a particularly important con-
sideration in such cold climates where natural decomposi-
tion is slow, and tourists who wish to visit such sites can
disturb the natural ecosystems by depositing waste materials.
The ecotourist is also in danger of disturbing the wildlife by
approaching animals too closely, or by trampling sensitive
plants. In the Antarctic, for example, visits to seal and pen-
guin colonies are very popular, and the animals often show
little fear of humans. But this can encourage very close
approaches, which cause stress to seals and penguins with
young. There is also the constant danger that visitors will
inadvertently bring diseases that will endanger wildlife popu-
lations. Heavy trampling on fragile tundra soils and the pos-
sibility of contamination from spills of fuel and toxins add to
the dangers inherent in wilderness tourism. Conservation
authorities often impose strict controls on both the number
and the activities of visiting parties in the Arctic and the
Antarctic.
170 TUNDRA
Alpine habitats have an even longer history of eco-

tourism. Mountaineering, climbing, and hiking became pop-
ular activities in the 19th century among those rich enough
to afford the travel, the equipment, and the local guides.
People who visited mountains were often naturalists who
were eager to collect both the plants and animals they dis-
covered, thus destroying some of the wildlife that had
attracted them to these locations. Others were “sportsmen”
whose main concern was to kill and collect larger specimens
of mammals and birds as trophies. As in the case of polar
ecotourism, the modern emphasis of alpine ecotourism is
upon observation rather than destruction. Despite this, the
greater accessibility of the mountains to a larger proportion
of the population places new kinds of stress upon alpine
tundra habitats. Excessive trampling erodes trails. Waste dis-
posal becomes a particular problem because it is not always
possible for visitors to take all of their waste products away
with them. In areas of the Himalaya Mountains in Asia, for
example, regions of picturesque mountain landscape have
become scarred by deposits of litter and other waste materi-
als along the well-used trails. Ecotourism in the mountains
therefore requires careful planning and investment. Tourist
sites must provide properly equipped camping areas, includ-
ing latrines and waste-disposal systems, or they will become
littered and spoiled. But local people may find great diffi-
culty in providing such facilities in remote and poor coun-
tries like Nepal.
The economics of ecotourism form an important consider-
ation. If the wild and beautiful places of the world are to be
conserved there must be some financial incentive for the
people who live there. In areas like the Himalayas, for exam-
ple, the arrival of visitors can result in economic gain for the
local people, but if this leads to the development of sophisti-
cated hotels to house the visitors, then the money may flow
into the pockets of developers from outside the region. Even
food may be imported because local produce may not be to
the tastes of foreign visitors. There is also the danger that
local communities will be disrupted and cultures changed by
the presence of tourists. In many instances, local cultures
effectively become converted into museum displays, or even
circus shows, for the sake of visitor entertainment. The dan-

gers of ecotourism are therefore considerable.
Ecotourism is one way in which tundra habitats are pro-
viding leisure activities for people; sport is another. Moun-
tain tundra has also become an important location for winter
sports, especially skiing, and this has led to new threats to
this wilderness habitat. The skiing industry now attracts large
numbers of people to locations that were once deserted or
occupied only by small farms and villages. This new influx of
people has led to extensive development of roads and resi-
dential complexes. Even more serious for the tundra habitat
has been the construction of ski runs and ski lifts that greatly
modify the landscape. Most of the activity, of course, takes
place in winter when the tundra is covered with snow, but
the major damage occurs early and late in the season when
the snow cover is thin and bare areas of grass are visible.
These brown patches on ski runs are the sites of major dam-
age to the underlying vegetation and soil, as pressure, fric-
tion, and physical wear degrade them. In summer, the ski run
is usually easy to recognize because of its poorer vegetation
cover and much lower plant diversity, caused by this damage
from the skis. Recreation is important for people and it is
inevitable that the mountain landscape should attract winter
sports. As in the case of ecotourism, however, excessive use
can damage the very resource that provides the pleasure, and
users must bear this in mind. Additionally, reserves for
wildlife conservation need to be established to ensure that
disturbance and damage do not extend into all available
habitats.
The value of all tundra habitats, like that of other wilder-
ness areas, is real but it is difficult to translate into economic
terms. Wildness has a great appeal to the human spirit,
which is why artists, writers, and poets have long extolled
the beauty and the appeal of remote locations free from the
impact of human society. Most people are inspired by pic-
tures or descriptions of wild places and many will go to great
lengths in order to visit such locations and experience silence
and solitude at first hand. But even people who may never
have the opportunity to visit the wilderness may gain pleas-
ure from the simple knowledge that it exists and that there
172 TUNDRA
are still relatively uncontaminated places on the Earth. Polar

and alpine tundra, therefore, are of great value to all human-
ity as a source of inspiration and inner peace.
Tundra as a carbon sink

Another way in which the tundra serves humanity and,
indeed, the general health of the planet, is by acting as a
“sink” for atmospheric carbon. The value of tundra in this
manner may not be immediately apparent, but is nonethe-
less important.
Scientists have been monitoring the level of the gas carbon
dioxide in the atmosphere for many years. They have also
been able to extract trapped bubbles of atmospheric gas from
ice sheets and have analyzed the bubbles to produce a record
of atmospheric composition in the more distant past. These
studies have shown that carbon dioxide has been rising
steadily for about 200 years. This period corresponds to that
of human industrial development powered by fossil fuels,
which contain carbon. People have been burning ancient
reserves of carbon in the form of coal, oil, and gas, buried in
the rocks, to generate the energy used for daily living. The
increase in atmospheric carbon dioxide over those 200 years
may not seem very great, from around 0.028 percent to 0.038
percent by volume of the atmosphere. This is a very small
concentration of this gas, and a relatively small increase, but
most atmospheric scientists believe that its impact on the cli-
mate has been considerable. Global temperature has risen by
about 3°F (1.5°C) during the same 200 years, and it seems
likely that the rise in carbon dioxide has been an important
contributing factor to this increase. Carbon dioxide, together
with several other gases, acts as a kind of thermal blanket
around the Earth, allowing the Sun’s rays to penetrate and
warm the Earth’s surface but preventing the heat generated
from escaping back into space. This is the same principle that
operates in a greenhouse, hence the expression “greenhouse
effect” for this process.
If atmospheric carbon dioxide concentrations continue to
rise, then additional increases in global temperature and con-
siderable changes in the Earth’s climatic patterns are likely,
most of which would be harmful for human populations and

for wildlife conservation. Any ecosystem that takes up more
carbon dioxide from the atmosphere by photosynthesis or by
any other chemical process than it releases to the atmosphere
by respiration (called a “carbon sink”) must be regarded as
valuable and worthy of protection. Growing forests are car-
bon sinks, as are wetlands. The forest is storing up carbon as
new tree biomass, while the wetland creates organic mud and
peat in which carbon is stored. The tundra biome, especially
the tundra habitats of the Arctic, is also currently a sink for
carbon. As discussed in chapter 3, decomposition in tundra is
slow and organic materials can accumulate in soils (see “Tun-
dra food webs and energy flow,” pages 72–76). In wetter sites,
which are frequent in the tundra regions of the Arctic, peat
develops and forms a store of carbon. The tundra, therefore,
is acting as a sink for atmospheric carbon and will continue
to do so while its conditions remain cold and damp. One of
the major concerns about the future of the tundra is whether
this will continue to be the case.
Conclusions
Despite the fact that tundra habitats are generally remote and
difficult to access, people have used them for a variety of pur-
poses. Tundra has been and continues to be a source of eco-
nomic profit in many respects, but it also has certain values
that are not immediately evident.
Tundra supports many large mammals, including caribou,
arctic fox, seals, and polar bears, and the meat and skins of
these animals have formed the basis of certain human cul-
tures since the Ice Age. In more recent times, hunters and
trappers have continued to exploit this resource, which has
been the basis of economically important trade. In most
cases, the hunting and trapping of animals in the tundra has
concentrated on truly wild animals, but in the case of the
European and Asian herds of caribou, or reindeer, a degree of
domestication has taken place. Reindeer herding by the Sami
people is the only type of agriculture practiced in the tundra.
The rocks of the tundra contain many minerals that have
attracted human attention. Metals such as gold have generated
174 TUNDRA
mass human migrations in search of this precious material, and

others, such as copper, have led to massive destruction of
mountain habitats. In recent times, prospectors have discov-
ered oil and gas reserves in the Arctic. Extractive industries
have developed some of these reserves, giving rise to a new
series of environmental problems.
The wild quality of the tundra has become a focus for a
rapidly expanding tourist industry, both in the polar and the
alpine tundra regions of the world. The remote wildness of
this habitat is an important source of human inspiration, but
it is also fragile and the ecotourism industry needs to be care-
fully controlled if damage to the source of such recreational
and educational enjoyment is to be avoided.
The tundra is one of the few habitats that absorbs more
carbon out of the atmosphere than it releases, and this makes
the tundra biome particularly valuable. This is a hidden use
of the tundra that must be appreciated.
The tundra biome thus has many uses and is valuable to
humans for a wide range of reasons. Although one of the
most remote and inaccessible of the Earth’s biomes, it is still
in danger of damage and destruction by human activities, so
its future lies in human hands.
CHAPTER 7
THE FUTURE
OF THE TUNDRA
The tundra is home to many highly adapted plants and ani-
mals, which have developed the attributes needed to survive
in an extreme environment. But despite the hardy nature of
many of its inhabitants, the tundra is a fragile ecosystem. It is
low in biodiversity, so it is in danger of collapsing if any of its
component species become extinct. Its nutrient capital and
its productivity are low, so the ecosystem may be sensitive to
disruption, whether by changing climate or human distur-
bance. Its soils are unstable, easily damaged and easily
eroded. The tundra, therefore, is a biome that is in danger,
and the problems that face it need to be considered carefully.
Consequences of climate change

The Earth is becoming warmer. As discussed in chapter 6, this
is probably due in part to human activity, which is increasing
the amounts of carbon dioxide and other greenhouse gases
in the atmosphere (see “Tundra as a carbon sink,” pages
172–173). But the Earth’s climate has experienced many
shifts in the past, some of them quite rapid, so the present
change in climate may also have a natural component that
represents a continuation of the cyclic changes that have
long been taking place. Whatever the causes of climate
change, its consequences could be very considerable for the
Earth’s biomes and especially for the tundra biome.
It is impossible to predict with accuracy the climate of the
future, but if the present warming trend continues, then the
global mean temperature is likely to become between 2°F and
4°F (1°C to 2°C) warmer in 2050 than it is at present. This
may not seem very much, but it could have considerable geo-
graphical and biological consequences, and these conse-
quences will not be evenly spread over the face of the Earth
175
176 TUNDRA
because the warming process will not be the same in all loca-
tions. Observations of the increased temperature over the
past 50 years show that the high latitudes, which include the
regions of polar tundra, have become warmer much faster
than the equatorial latitudes. If this trend continues into the
future, then the polar latitudes may undergo a greater change
in climate than anywhere else on Earth.
The tundra biome contains the world’s reserves of ice, and
a warmer climate will mean that some, perhaps most, of this
reserve will melt and join the oceans. It is quite possible that
the Greenland ice sheet will have melted by the end of the
21st century, and this would add considerably to the level of
the world’s oceans. The floating pack ice of the Arctic Ocean
is melting and, if the process continues at the present rate,
may disappear completely by about 2070. The Antarctic ice
sheet is more difficult to predict. Current evidence shows
that the surrounding sea ice is breaking up extensively, but in
the long term much will depend on levels of precipitation.
Warmer sea temperature means faster evaporation of water,
and this could bring more precipitation over Antarctica.
More snow would fall over the vast ice sheet, so the ice vol-
ume might not decline as fast as expected on the basis of
future temperature calculations. So there are still some
important unknown factors in the equation. Water expands
when it is heated, so increasing the temperature of the
world’s oceans will cause an expansion that will raise sea lev-
els globally. Taking into account the combined effects of
melting ice and expanding water, observers widely expect the
level of the Earth’s oceans to rise by four to 10 inches (10–25
cm) by 2050. A sea level rise of this magnitude would flood
many coastal areas of Arctic tundra. But, on the other hand,
the loss of ice and snow over the tundra landscape will
expose more elevated land areas where tundra vegetation can
invade.
While the rising sea level erodes the polar tundra from the
Arctic Ocean in the north, vegetation changes will also take
place from the south. At the boundary between the tundra
and the boreal forest (or taiga) in the Low Arctic, warmer
conditions will enable trees to bear fruit and spread their
seeds into new regions. The forest will advance northward
into regions that are now covered by low shrubs and cushion
herbs. There is already evidence of spruce invading beyond
its former limits in Canada and extending the boreal forest
into the tundra zone. The tundra biome, therefore, will
become squeezed between the forest and the sea, and it is
likely to occupy an increasingly narrow zone as climate
warms.
Meanwhile, in the mountains of the world the warmer
temperatures will cause the tree line to extend upward and
the line of permanent snow will retreat. This means that the
belt of tundra will gradually occupy higher altitudes. But this
process can continue only until the permanent snow zone is
completely lost, and snow loss will depend upon the latitudi-
nal position of the mountain (equatorial mountains will lose
their snow first) and the overall altitude of the mountain
(higher mountains will retain their snow longer). In the case
of low mountains in warm, low-latitude climates, increased
warmth will eventually eliminate alpine tundra vegetation.
As the trees spread upward the tundra plants will have
nowhere to retreat to and will eventually become shaded out
by the expanding forest. Studies of mountains in the high
latitudes, especially those in the polar regions, suggest that
high winds may prevent forest expansion even though the
air temperature may rise. These mountains, particularly those
of the Arctic, may become the final refuge for tundra in a
warmer world.
The prospects for the tundra biome in the event of contin-
ued climatic warming may therefore appear rather bleak. But
the tundra has survived as a biome for several million years,
despite episodes in the Earth’s recent past that were much
warmer than the present world. This suggests that the tundra
has a higher degree of inertia, or resistance to change, than
one might expect. Two possible reasons have been put for-
ward to explain this. First, the very low levels of nutrients in
tundra soils may actually help in the survival of this biome.
Chapter 4 proposed that low nutrient reserves might make an
ecosystem fragile; but poor soils could also make the tundra
difficult to invade. Trees generally need more chemical nutri-
ents than smaller plants do, and most trees find it difficult to
establish themselves in poor soils. Even the tough birches,
178 TUNDRA
pines, and spruces that occupy the border regions of the Arc-
tic tundra find it difficult to germinate and survive in very
poor soils, especially when they are being grazed upon by
voracious herbivores such as arctic hares and caribou. So,
nutrient poverty may actually protect the tundra from tree
invasion.
The second feature that the tundra biome has in its favor is
a high level of genetic diversity in its flora and fauna. Earlier
discussion has stressed that the number of species of plants
and animals in the tundra is low, but among the species pres-
ent there are many subspecies and races. The environment
itself is quite diverse, with wet locations and dry ones, salty
and fresh areas, high altitudes and low, exposed and pro-
tected areas, snow-covered and open locations. The species of
plants and animals present, though low in number, have
developed a whole range of genetically adapted forms, called
ecotypes, that are able to cope in each of these different micro-
habitats. This genetic diversity will greatly assist the tundra
as it faces the challenge of climate change. Whatever new
sets of conditions are generated by the changes, it is likely
that many existing species will have the right set of genetic
adaptations to take advantage of the new opportunities. Per-
haps this high genetic diversity is what has enabled the tun-
dra to survive the many occasions when climate change has
threatened this biome in the past.
Ozone holes
Apart from climate change, another aspect of changing
global conditions that has attracted attention in recent years
is the development of gaps in the ozone layer, especially in
polar regions. These may present a further threat to the
health of the tundra biome.
It is easy to become confused about the gas ozone. It is a
very reactive material, having the chemical formula O3 and
possessing great powers of oxidation. It attacks many types of
materials and can even decompose rubber. When people
breathe it in, therefore, it damages the lungs and can prove
fatal to those with weak respiratory systems. Produced from
the chemical reactions of oxides of nitrogen, ozone is one
component of automobile exhaust fumes, and when coupled

with strong sunlight and other products of combustion it can
contribute to the formation of photochemical smog. So,
ozone close to the Earth’s surface, especially in sunny cities,
is a pollutant gas that is a health risk and needs to be
avoided.
But ozone also occurs naturally, high in the stratosphere,
which is a layer of the Earth’s atmosphere lying about six to
30 miles (10 to 50 km) above the ground. Even in the strato-
sphere, ozone is only a trace gas, and if all the ozone present
in the stratosphere were concentrated at ground level under
normal atmospheric pressure, it would form a layer only 0.1
inch (3 mm) thick. But this small amount of ozone in the
stratosphere performs a vital function for all the living things
on the land surface of the Earth; it absorbs the harmful ultra-
violet (UV) radiation from the Sun. In the absence of such
protection the UV radiation would cause genetic damage
among all living things, apart from those living deep in the
oceans, where they are protected by the ozone-screening
effect of water.
Research into the changes in atmospheric ozone began in
1957 when atmospheric scientists set up an international
network of monitoring stations, including one in Antarctica.
By 1985 sufficient data had been accumulated to demon-
strate that the overall quantity of ozone in the stratosphere,
particularly over the Antarctic, had severely declined during
the period of monitoring. Its concentration had halved since
1957. Detailed studies showed that the loss of ozone was sea-
sonal, with the strongest decline occurring in the Antarctic
spring. Two possible causes were investigated: first, the
increasing abundance of stratospheric jet traffic, and second,
the release by humans of increasing quantities of a group of
chemicals called chlorofluorocarbons (CFCs), used in aerosol
sprays, refrigerators, and the production of plastic foam. Lit-
tle evidence could be found to support the jet plane hypoth-
esis, but CFC release became an increasingly likely
explanation for the development of an “ozone hole” over the
Antarctic. CFCs are released at the Earth’s surface and then
diffuse upward through the atmosphere, gradually breaking
down to release chlorine, which then interacts with ozone,
180 TUNDRA
causing its decomposition to oxygen. Due to circulation pat-

terns in the atmosphere ozone destruction is greatest in the
South Pole region, but an ozone hole has also been develop-
ing above the Arctic in recent years. In September 2000 the
ozone hole over the Antarctic extended over a record 11 mil-
lion square miles (29 million km2). This is an area three times
that of the United States and it covered the whole of Antarc-
tica, together with much of the Southern Ocean and even the
southern tip of Chile. The threat to the health of people and
wildlife is so severe that many nations have agreed upon a
policy to reduce the rate of ozone destruction, which has
been set out in the Montreal Protocol (see sidebar below). By
2005 there are some small signs of ozone recovery.
Pollution
Pollution by human waste products and accidental spills
threatens all of the Earth’s ecosystems, but particularly the
sensitive tundra biome. As discussed in chapter 6, the exploita-
tion of the mineral resources of the tundra regions has often
been accompanied by pollution and consequent environmen-
tal damage (see “Mineral reserves,” pages 163–168). Oil spills
at the site of extraction, or during transport by pipeline or
The Vienna Convention and the Montreal Protocol

In 1985, 20 nations agreed that they needed to act swiftly to protect human health and
control the expansion of the polar ozone holes. They signed the Vienna Convention,
which was a statement of intent to take action, though it did not state precisely what
action was needed. In 1987 a further international meeting in Canada led to the Montreal
Protocol, an agreement to restrict the production and use of compounds such as CFCs
that are known to affect the ozone layer. Those signing the protocol have put its provi-
sions into effect, especially the developed nations, which are responsible for the bulk of
CFC production. In total, 155 nations have now agreed to the proposals of the Montreal
Protocol, of which more than 100 are developed nations. The outcome has been a slow-
ing in the rate of ozone destruction, so that the polar tundra, and the humans who live
and work in the tundra, appear to have been rescued from a most unpleasant fate.
ship, are statistically inevitable, given the large quantities

taken out of the rocks beneath the tundra and the long dis-
tances the materials have to be moved for their treatment and
consumption. Contamination of rivers and soils from mining
waste is likewise an unavoidable outcome during the extrac-
tion of metals from the rocks beneath the tundra. Waste prod-
ucts from human settlements are difficult to dispose of, and
they create pollution and health problems. All of these sources
of pollution are local to the tundra, and the best way to pre-
vent them is to control the extent of exploitation of the habi-
tat, possibly even denying humans the right to commence
such exploitation, as in the case of mineral extraction in
Antarctica. But there are other sources of pollution that are dis-
tant from the tundra regions and more difficult to control.
Pesticides are extensively used throughout the world as a
means of increasing the yield of crops or the health and pro-
ductivity of domesticated animals. They are toxins that are
selected to kill some organisms (such as insect pests) without
causing harm to others, including people. But scientists and
agriculturalists have often miscalculated and used pesticides
that had a harmful impact beyond the target organism. The
use of DDT to kill insects in the 1940s and 1950s, for exam-
ple, was very successful in the control of many insect-borne
diseases, including malaria, so it undoubtedly saved many
thousands of human lives, including those of soldiers during
World War II. But the compound proved very durable in the
environment. Its concentration built up in the fat reserves of
birds and mammals until it eventually impaired their breed-
ing and sometimes even caused their death. DDT also spread
around the world, finding its way through marine food webs
to the tundra regions and accumulating in the fatty tissues of
seals and penguins. Fortunately, the harmful effects of this
compound were recognized in time, and most developed
countries have now banned the use of DDT. The persistence
of this chemical in the environment is limited; within 10
years half of any residual DDT will decompose, so the threat
of global contamination is now reduced. The withdrawal of
this compound in many parts of the world should prevent
the destruction of tundra mammals and birds, as well as
those of other biomes.
182 TUNDRA
Most pesticides, including DDT, are artificial compounds,

so their origin as pollutants is never in doubt, even when
they are found in remote areas such as the Antarctic. But peo-
ple produce other materials that also occur naturally, so it is
less easy to be sure of their source. Nitrogen compounds,
including ammonia and oxides of nitrogen, for example,
occur naturally in the atmosphere, but exhaust gases from
traffic, together with fertilizers that farmers spray, have
injected more of these materials into the atmosphere. They
are pollutants, but because nitrogen is an important element
for the growth and development of plants they are also
growth stimulants for vegetation (see “Nutrient cycling in
the tundra,” pages 76–81). Tundra soils are poor in nitrogen,
so the arrival of the element from the air can change the veg-
etation composition; more robust and fast-growing plants
outcompete the slower and weaker species. This can lead to
the extinction of the less competitive species.
Scientists studying the aerial fallout of nitrogen com-
pounds in the Arctic have found that the quantities arriving
are minute, usually between 0.0027 and 0.027 ounces per
square yard (0.1 to 1.0 g/m2) in a year. This may seem very
small, but in fact it is as great as the combined total of all
other sources of nitrogen in the ecosystem, such as the break-
down of soil materials by decomposition and weathering and
the fixation of atmospheric nitrogen by bacteria. Air pollu-
tion, in other words, has roughly doubled the nitrogen sup-
ply to the tundra ecosystem. In the future, this change in
nutrient cycling is likely to encourage more robust vegeta-
tion growth, permitting more shrubs and trees to invade the
open tundra. This fertilization of the soils by pollutants,
combined with the likely effects of climate warming, raises a
strong possibility that the forest will spread into the tundra.
Here lies the greatest threat to the future of the tundra
biome.
Tundra conservation
For the tundra to remain as one of the Earth’s major biomes,
people must conserve it. Conservation is not quite the same
thing as preservation. In preservation, one isolates an object
from injurious influences and protects it to ensure that it

remains unaltered by events. In the natural world this would
be an impossible task because the natural environment itself
is constantly changing. Climate changes partly because of
natural cycles, and unpredictable events such as wind, earth-
quake, fire, and flood can disrupt natural ecosystems.
Humans, as has been seen, extend their influence far beyond
the regions they inhabit, so even the most remote of wilder-
nesses cannot be fully protected from the effects of human
activity. Conservation, therefore, aims to protect but also to
manage the ecosystem in order to modify or control the pres-
sures that come from outside. There was a time when ecolo-
gists believed that the best form of conservation was to leave
an ecosystem alone and to avoid any form of human inter-
vention. It is now recognized that this attitude is based on an
impossible ideal and that people need to actively manage
ecosystems in order to maintain levels of biodiversity on
Earth.
In the polar tundra, protection from the harmful human
impacts is clearly the first priority. The hunting of mammals
needs to be controlled if their populations are to remain sta-
ble. In the case of some animals, such as the great whales,
controlling hunting may mean stopping it entirely, but with
others, such as seals and caribou, there may be a case for lim-
ited harvesting of the herds. Indeed, there are occasions
when culling of populations is a humane reaction to over-
population. For instance, the red deer (elk) population on the
alpine tundra habitats of Scotland has recently expanded,
with numbers increasing fourfold between 1960 and 2000.
The alpine tundra habitat is becoming damaged, and each
winter a proportion of the elk inevitably starve. The imbal-
ance was the result of past human activity, namely eliminat-
ing the wolf from Scotland (the last wolf was shot in 1743).
Now this large deer has no natural predators. There is, how-
ever, a new herbivore in the alpine tundra that people have
introduced, namely the sheep, and herds of sheep contribute
to overgrazing and vegetation destruction. In response,
human management of the Scottish Highland tundra must
reduce either sheep grazing or deer numbers, or both.
Already, hunters take about 50,000 red deer every year in
184 TUNDRA
Scotland, representing approximately 13 percent of the pop-

ulation. Clearly, however, this is not enough to stabilize the
population, and further culling may prove necessary.
The control of Scottish red deer is an example of the need
to manage ecosystems, especially if they have already been
modified by human activity and left unbalanced. Sometimes
it is necessary to try to put right errors of the past, such as the
local extinction of a species. Reintroduction of the lost ani-
mal is always controversial for many reasons. It may bring
problems for human populations. If the wolf were to be rein-
troduced to Scotland, for instance, sheep farmers would
undoubtedly object, just as farmers have done near Yellow-
stone National Park, where wolves have been successfully
reintroduced (see sidebar on opposite page). The musk ox is a
species that has attracted less controversy than the wolf but
has suffered even more at the hands of human hunters. Once
widespread through the Arctic, it became very restricted as a
result of a long history of hunting from prehistoric to mod-
ern times. It was completely lost from Europe, Asia, and
Alaska, but the surviving herds on Greenland have formed
the basis for reintroduction to Russia and to Alaska. In some
sites the introduced musk oxen have found it difficult to
compete with native caribou herds, but generally the species
seems to be holding its own. In the case of the musk ox,
where humans have been responsible for local extinctions, it
seems only reasonable that humans should try to set things
straight by reintroduction. One of the dangers of this strat-
egy, however, is that the stock may be taken from herds with
slightly different genetic makeup. In the process of reintro-
duction, the conservationist may be taking animals and
plants to regions where they are not fully suited for survival.
In alpine tundra habitats, local extinction is a very com-
mon problem because of the isolation of mountain peaks. If
an animal or plant is lost at one site, it may be difficult for
the species to reinvade from surrounding but isolated moun-
tains. Reintroduction of animals lost as a result of human
persecution has therefore been extensively used in mountain
regions. In North America, the mountain goat has been
reestablished in Oregon, Nevada, Utah, Colorado, Wyoming,
and South Dakota from its residual populations in the north-
Bringing back the wolf

Throughout the world, people generally regard wolves as pests. Livestock farmers, in par-
ticular, are invariably eager to eliminate this intelligent, pack-hunting predator. In the
United States (with the exception of Alaska) the gray wolf had been reduced to a few
small populations in the northern parts of the country by the middle of the 20th century;
even in Yellowstone National Park, the last wolf was shot in 1926. But after 70 years with-
out wolves, conservationists became concerned that the whole ecological balance of Yel-
lowstone had been altered because of the loss of wolves. One of the main prey species in
the park had been elk, and in the absence of the top predator, elk populations expanded
and forests were overgrazed. Wolves, it seemed, had an indirect impact on the balance of
forest trees. In 1995 the U.S. Fish and Wildlife Service introduced 15 gray wolves from
Canada into Yellowstone, and they proved so successful that they now number around
300. Elk populations have reduced and new habitats have been created, especially around
lakes where elk grazing was particularly intense. The remains of dead elk, killed by wolves,
provide food for bears, ravens, and eagles. In short, the return of the wolf has increased
the biodiversity of the park.
ern Rocky Mountains and Cascades. The closely related

chamois of the European Alps was similarly severely reduced
by hunting but has recovered much of its former range as a
result of reintroduction.
Conservationists have also helped some birds to spread
back into mountain areas where they had previously been
exterminated. The bearded vulture of Europe and Asia is a
massive bird that feeds on broken bones that it drops onto
rocks from the air, and it has benefited from human trans-
port and care in bringing it back to several mountain ranges
from which it had been lost as a result of human persecution.
Human beings are now so numerous and so influential in
polluting the atmosphere and the oceans that there is no part
of the planet left unaffected. Even the remote areas of tundra,
both polar and alpine, have experienced change as a result of
the arrival of humans on the scene. As discussed in chapter 6,
the tundra is a useful biome for people because of its mineral
wealth, its recreational opportunities, and its role in the over-
all balance of the planet. People, therefore, will need to play
186 TUNDRA
an increasing part in the conservation of tundra, both pro-

tecting it from further harm and putting right some of the
problems this biome has already suffered as a consequence of
human errors. The future of the tundra, as is the case for all
biomes, now lies in human hands.
Conclusions
The tundra biome, both in the polar regions and in the
mountains of the world, faces a number of problems, some of
which threaten its very survival. Continued climatic warm-
ing is almost certain, and this will have a particular impact
on the Earth’s coldest biome. Tundra lies at the world’s
extremities, either close to the poles or near the highest parts
of mountains. There is no place to which tundra can retreat if
warming continues; it may be eliminated from the tops of
mountains (windy Arctic mountains are a possible exception)
and become confined to an increasingly narrow belt of land
south of the Arctic Ocean. The likely rise in global sea level in
the warmer world can only make things worse. Added to this,
it seems probable that the polar regions will experience more
rapid rises in temperature over the next century than most
other places on Earth. Many of the highly adapted cold-toler-
ant species of plants and animals will find themselves under
climatic and spatial stress, eliminated by stronger, more com-
petitive, warmth-loving species. The one hope for tundra bio-
diversity is its great genetic variability. The organisms of the
tundra are not only highly adapted but also highly adapt-
able, and this may prove vital for their survival.
Ozone depletion in the polar stratosphere has posed a new
threat to the tundra because it permits higher levels of ultra-
violet radiation from the Sun to penetrate the atmosphere.
This can cause genetic damage to exposed tissues. The
human causes of ozone thinning appear to have been identi-
fied, and the efforts of many nations to reduce this poten-
tially serious threat to tundra survival may prove successful.
Pollution, both in the form of direct deposition of harmful
compounds and in the less obvious form of atmospheric and
oceanic contamination, is a further source of damage to the
tundra. People must make pollution control a priority to pro-
tect this biome.
Conservation of the tundra, therefore, will involve the

enforcement of protective measures at an international level,
and many of these are already agreed upon and in place. But
conservation demands more than just protection; it requires
proactive management. Human activities have reduced the
ranges of certain species, and these organisms may require
human assistance to reestablish themselves over their former
ranges. Meanwhile, some species may need to be controlled
to avoid excessive use of resources, and some habitats may
need to be modified to ensure the survival of sensitive
species. The tundra is useful to humans, as well as being valu-
able in its own right, and the future of the tundra lies very
much in human hands.
GLOSSARY
ablation the loss of ice at the base of a glacier, where the
warmth of the Earth causes melting
active layer the upper soil layers in Arctic permafrost environ-
ments that melt in the summer and freeze in the winter
aestivation a period of dormancy that certain animals under-
go to avoid the unfavorable conditions of summer drought
(equivalent to hibernation in winter)
albedo an index of the degree of reflectivity of a surface to
light. Snow has a high albedo; dark-colored vegetation has a
low albedo
allochthonous describing material that has originated away
from the site in which it eventually settles, such as leaves car-
ried into a lake; the opposite of autochthonous
allogenic describing forces outside a particular ecosystem that
may cause internal changes; for instance, rising sea level can
influence water tables in freshwater wetlands farther inland,
and is therefore considered an allogenic factor
alpine tundra vegetation dominated by herbs and dwarf
shrubs found above the tree line of high mountains, where
conditions are too cold for tree growth
anaerobic lacking oxygen
anion elements or groups of elements carrying a negative
charge, such as NO3– or HPO3–
annual an organism (usually a plant) that completes its life
cycle in a single year
anoxic lacking oxygen
Arctic-alpine describing an organism that is found in both
Arctic and alpine tundra habitats
Arctic tundra a region with vegetation dominated by herbs
and dwarf shrubs found in the polar regions where conditions
are too cold for tree growth
aspect the direction of the compass that a slope faces
autochthonous describing material that has originated in the
site where it is deposited, such as bog moss peat in a bog; the
opposite of allochthonous
189
190 TUNDRA
autogenic describing forces within an ecosystem that bring

about changes. For instance, the growth of reeds in a marsh
results in increased sediment deposition. See also FACILITATION
biodiversity the full range of living things found in an area,
together with the variety of genetic constitutions within
those species and the range of habitats available at the site
biogeography the scientific study of the spatial distribution of
living animals and plants
biomass the quantity of living material within an ecosystem,
including those parts of living organisms that are part of
them but are strictly nonliving (such as wood, hair, teeth, or
claws) but excluding separate dead materials on the ground or
in the soil (termed litter)
biosphere those parts of the Earth and its atmosphere in
which living things are able to exist
blue-green bacteria (cyanobacteria) microscopic, colonial,
photosynthetic microbes that are able to fix nitrogen; once
wrongly called blue-green algae. They play important ecologi-
cal roles in some wetlands as a consequence of their nitrogen-
fixing ability, such as in rice paddies
bog a wetland ecosystem in which the water supply is entirely
from rainfall (ombrotrophic). Such wetlands are acidic and
poor in nutrient elements. They accumulate a purely organic
peat with very little mineral matter (derived solely from air-
borne dust), so are prized for horticulture
boreal northern, usually referring to the northern temperate
regions of North America and Eurasia, which are typically
vegetated by evergreen coniferous forests and wetlands.
Named after Boreas, the Greek god of the North Wind
calcareous rich in calcium carbonate (lime)
capillaries fine, tubelike air spaces found in the structure of
partially compacted peat or soils
carbon sink an ecosystem that absorbs more carbon from the
atmosphere than it releases in respiration. Some wetland
habitats operate in this way
catchment a region drained by a stream or river system
(equivalent to watershed)
cation an element or group of elements with a positive charge,
such as Na+, NH4+, or Ca++
cation exchange the substitution of one positively charged
ion for another. Certain materials (such as peat and clay) have
the capacity to attract and retain cations and to exchange
them for hydrogen in the process of leaching
GLOSSARY 191
chamaephyte a plant that grows close to the surface of the

ground, below a height of one foot (25 cm), and in this way
escapes the effects of intense wind-blasting in tundra habitats
charcoal incompletely burned pieces of organic material (usu-
ally plant). These are virtually inert and hence become incor-
porated into lake sediments and peat deposits, where they
provide useful indications of former fires. Fine charcoal parti-
cles may cause changes in the drainage properties of soils,
blocking soil capillaries and leading to waterlogging
climate the average set of weather conditions over a long peri-
od in a region
climax the supposed final, equilibrium stage of an ecological
succession. Many would question whether real stability in
nature is ever achieved
community an assemblage of different plant and animal
species, all found living and interacting together. Although
they may give the appearance of stability, communities are
constantly changing as species respond in different ways to
such environmental alterations as climate change
competition an interaction between two individuals of the
same or different species arising from the need of both for a
particular resource that is in short supply. Competition usual-
ly results in harm to one or both competitors
conservation human protection and enhancement of a habi-
tat or a species
cyanobacteria see BLUE-GREEN BACTERIA
day length the period from sunrise to sunset. This may be
nonexistent in a polar winter
deciduous describing a plant that loses all its leaves during an
unfavorable season, which may be particularly cold or partic-
ularly dry
decomposition the process by which organic matter is reduced
in complexity as microbes use its energy content, usually by a
process of oxidation. As living things respire the organic mate-
rials producing carbon dioxide, other elements such as phos-
phorus and nitrogen return to the environment where they
are available to living organisms once more. Decomposition is
therefore an important aspect of the nutrient cycle
detritivore an animal (usually invertebrate) that feeds upon
dead organic matter
diatoms a group of one-celled photosynthetic organisms that
form an important part of the phytoplankton in wetland
habitats
192 TUNDRA
DNA deoxyribonucleic acid, the molecule that contains the

genetic code
drumlin a deposit of till beneath a glacier that is often carved
into a linear shape by the movement of the ice above
ecosystem an ecological unit of study encompassing the liv-
ing organisms together with the nonliving environment
within a particular habitat
ecotone boundary regions where one type of habitat gradually
blends into another
ecotourism tourism to wilderness areas of the world that tries
to avoid damaging the environment in its development
emergent aquatic plants wetland plants that are rooted in
soil that lies underneath but have shoots projecting above the
water surface
energy flow the movement of energy through an ecosystem,
from sunlight energy fixation in photosynthesis to its acquisi-
tion by consumer organisms and its release by respiration
erosion the degradation and removal of materials from one
location to another, often by means of water or wind
erratic a rock that is carried far from its original position by
the movement of a glacier and is eventually deposited when
the glacier melts
esker a ridge of glacial detritus running along the edge of a gla-
cier or beneath the ice
eutrophication an increase of fertility within a habitat, often
resulting from pollution by nitrates or phosphate from
runoff of these materials into water bodies from surrounding
land. Although the term most often describes wetland habi-
tats, it can also be applied to terrestrial ecosystems, such as
the tundra
evaporation the conversion of a liquid to its gaseous phase,
especially the loss of water from terrestrial and aquatic sur-
faces
evapotranspiration a combination of evaporation from land
and water surfaces and the loss of water vapor from plant
leaves (transpiration)
evergreen a leaf or a plant that remains green and potentially
able to photosynthesize throughout the year. Evergreen
leaves do eventually fall, but may last for several seasons
before they do so
facilitation the process by which a plant species alters its
local environment such that other plants can invade. For
example, when a water lily grows in a lake, its leaf stalks slow
GLOSSARY 193
the movement of water, causing suspended sediments to set-

tle. The lake consequently becomes shallower, permitting
other plant species to invade and eventually supplant the
water lily. Facilitation is one of the forces that drives ecologi-
cal succession
firn powdery or granular deposits of snow that accumulate on
the surface of a glacier
fjord a deep, steep-sided valley flooded by the sea
floodplain the low-lying lands alongside a river over which
the river water expands when water flows faster than the river
can carry it away
fluvial outwash the detritus washed out of a glacier as it melts
food web the complex interaction of animal feeding patterns
in an ecosystem
forest tundra the ecotone (border region) of the forest and the
tundra. Trees that survive here are usually dwarfs, stunted by
strong winds, and are referred to as KRUMMHOLZ
fossil ancient remains, usually the buried remnants of a
once-living organism; the term can also be applied to
ancient buried soils or even the organic remains termed fos-
sil fuels
fragility an expression of the ease with which an organism or
a habitat may be damaged. Fragile ecosystems, such as many
wetlands, need careful conservation
frost heaving the mechanism of freezing and thawing of a soil
that forces stones and unfrozen layers of soil to the surface
fundamental niche the potential of an organism to perform
certain functions or to live in certain areas. Such potential is
not always achieved because of competitive interactions with
other organisms. See also REALIZED NICHE
genes the store of hereditary information of living things,
which is made up of DNA and contained within an organ-
ism’s cells
glacier a mass of permanent ice found in cold conditions,
often occupying valleys through which the ice moves slowly
under the influence of gravity
glacial a period in the Earth’s history when conditions are
cold enough for glaciers to be widespread
gley a pale gray soil produced by permanent waterlogging
greenhouse effect the warming of the Earth’s surface due to
the interaction between radiation and the Earth’s atmosphere.
Short-wave solar radiation passes through the atmosphere
unchanged, but the Earth’s surface radiates it as long-wave
194 TUNDRA
radiation (heat). The atmosphere then absorbs the long-wave

radiation because of the presence of greenhouse gases
greenhouse gas an atmospheric gas that absorbs long-wave
radiation and therefore contributes to the warming of the
Earth’s surface by the greenhouse effect. Greenhouse gases
include carbon dioxide, water vapor, methane, chlorofluoro-
carbons (CFCs), ozone, and oxides of nitrogen
groundwater water that soaks through soils and rocks, as
opposed to water derived directly from precipitation and
present on the surface of the soil
habitat the place where an organism lives
habitat structure the architecture of vegetation in a habitat.
The height and branching patterns of plants contribute to the
complexity of vegetation architecture, and this complexity
creates microhabitats for animal life
halophyte a plant that is adapted to life in saline conditions
as a result of its physical form, its physiology, or both
hibernation a period of dormancy that certain animals under-
go to avoid cold winter conditions
High Arctic the northern regions of the Arctic, where the
growing season for plants is less than two and a half months
hydrology the study of the movement of water in its cycles
through ecosystems and around the planet
ice sheet an extensive cover of permanent deep ice. Only two
ice sheets currently occupy the Earth, one covering Antarctica
and the other based on Greenland
ice wedge water that freezes in a tundra soil and expands,
forming a wedge shape that forces its way down into the soil
and may split the landscape into a series of polygons
inertia the property of resistance to disturbance in an ecosys-
tem. A stable ecosystem is difficult to disturb
interception a function of plants in which the plant canopy
catches rainwater and prevents it from reaching the ground
directly
interglacial a prolonged period of Earth’s history in which the
climate is warm; interglacials are preceded and followed by
glacials
interstadial a geologically short period of warmth, preceded
and followed by stadials
invertebrate an animal lacking a backbone
ion a charged element or group of elements. See also ANION and
CATION
GLOSSARY 195
kettle hole a hollow in glacial detritus deposits resulting from

the melting of a block of ice in that position. It may become
filled with water to form a deep, steep-sided lake
krummholz a vegetation dominated by trees that have been
distorted and stunted by strong winds. See also FOREST TUNDRA
lapse rate the rate at which atmospheric temperature drops
with altitude
latitude imaginary lines drawn horizontally around the Earth
that are named according to the angle they make with the
center of the Earth. Thus, the equator is 0°N and S latitude
and the poles are 90°N and 90°S. The polar regions thus have
higher numbers and are referred to as high latitudes
leaching the process by which ions are lost from soils and sedi-
ments as water (particularly acidic water) passes through them
lichen an organism that consists of a combination of an alga
or a cyanobacterium with a fungus. The combination may
have a leafy form or may look like paint on a rock. Lichens are
generally resistant to cold and drought
limestone sedimentary rocks containing a high proportion of
calcium carbonate (lime)
litter the accumulation of dead (mainly plant) organic materi-
al on the surface of a soil
loess windblown dust and sand carried by winds over bare,
glacial terrain
Low Arctic the southern regions of the Arctic, where the
growing season for vegetation is generally between three and
five months
macrophyte large aquatic plants that can be observed without
the use of a microscope
management the deliberate manipulation of an ecosystem by
humans in order to achieve a particular end, such as
increased productivity or nature conservation
megafauna extremely large animals, many of which became
extinct at the end of the last glacial
methane a gas produced by some living organisms as a result
of the incomplete decomposition of organic matter. It is a
greenhouse gas, a gas that increases the heat-retention prop-
erties of the atmosphere
methanogenic bacteria bacteria that produce methane gas as
a result of their metabolism
microbes microscopic organisms such as bacteria, fungi, and
viruses
196 TUNDRA
microclimate the small-scale climate within habitats, such as

beneath forest canopies or in the shade of rocks. The micro-
climate is strongly affected by habitat structure
migration the seasonal movements of animal populations,
such as geese, caribou, or plankton
mire a general term for any peat-forming wetland ecosystem
mire complex a wetland that consists of a series of different
mire types
moraine an unsorted mass of glacial debris deposited at the
end of a melting glacier (terminal moraine) or beneath the ice
mass (hummocky moraine)
niche the role that a species plays in an ecosystem. The con-
cept of niche consists of both where the species lives and how
it makes its living (such as feeding requirements, growth pat-
terns, or reproductive behavior). The niche may be regarded
as FUNDAMENTAL or REALIZED
nidifugous describing newly hatched birds that rapidly leave
their nest
nitrogen fixation the process by which certain organisms are
able to convert nitrogen gas into organic molecules that can
be built into proteins
nunatak the peak of a mountain projecting from a mass of
ice
nutrient cycle the cyclic pattern of element movements
between different parts of the ecosystem, together with the
balance of input to and output from the ecosystem
occult precipitation precipitation that is not registered by a
standard rain gauge because it arrives as mist, condensing on
surfaces, including vegetation canopies. See INTERCEPTION
ombrotrophic fed by rainfall. Bogs are ombrotrophic mires,
receiving their water and nutrient input solely from atmos-
pheric precipitation
organism any living creature, from bacteria to mammals and
plants
ozone hole extreme thinning of the ozone layer over the polar
regions in their respective summers, which allows excessive
ultraviolet radiation to reach the Earth’s surface. Ozone (O3)
is an unstable form of oxygen gas (O2)
paleoecology the study of the ecology of past communities
using a variety of chemical and biological techniques
palsa a wetland type found only within the Arctic Circle.
Elevated peat masses expand as a frozen core develops within
GLOSSARY 197
them. Palsas pass through a cycle of growth and then col-

lapse, forming open pools. Compare PINGO
paludification a process in which an ecosystem becomes
inundated with water
peat organic accumulations in wetlands resulting from the
incomplete decomposition of vegetation litter
periglacial describing the climatic conditions found around
the edges of a glacier
permafrost permanently frozen subsoil. The upper layer
(active layer) thaws during the summer and freezes in winter
pH an index of acidity and alkalinity. Low pH means high
concentrations of hydrogen ions (hence acidity), while a high
pH indicates strong alkalinity. A pH of seven indicates neu-
trality. The pH scale is logarithmic, which means, for
instance, that pH four is 10 times as acidic as pH five
photosynthesis the process by which certain organisms trap
the energy of sunlight using a colored pigment (usually
chlorophyll) and use that energy to take carbon dioxide from
the atmosphere and convert it into organic molecules, initial-
ly sugars
photosynthetic bacteria bacteria possessing pigments enab-
ling them to trap light energy and conduct photosynthesis.
Some types are green and others purple
physiological drought a condition in which water is present
in a habitat but is unavailable to a plant, for instance because
it is frozen
phytoplankton the collection of microscopic, often one-
celled photosynthetic organisms that live in the well-lit sur-
face layers of water bodies. They form the basis of many
aquatic food webs
pingo a structure formed in the tundra soil by water freezing
under pressure, often fed by a spring. As the water turns to
ice, it forces the surface of the ground to rise into an extensive
mound
pioneer a species that is an initial colonist in a developing
habitat
plate tectonics the theory that the crust of the Earth is divid-
ed into plates that move over the surface, occasionally collid-
ing and buckling to form mountain chains or deep rifts
podzol a type of soil, common in the boreal forest (or taiga)
zone and in the ecotone with the tundra. It consists of a
series of layers formed by the leaching of iron, organic matter,
198 TUNDRA
and clay from the upper layers and their deposition lower
down
pollen analysis the identification and counting of fossil
pollen grains and spores stratified in peat deposits and lake
sediments
pollen grains cells containing the male genetic information of
flowering plants and conifers. The outer coat is robust and
survives well in wetland sediments. The distinctive structure
and sculpturing of the coats permit their identification even
in a fossil form
polygon mire patterned wetlands of the Arctic regions in
which raised polygonal sections are separated by water-filled
channels, particularly apparent from the air
polygons patterns of stones caused by frost heaving on level
ground in tundra habitats
population a collection of individual organisms all of the
same species
precipitation aerial deposition of water as rain, dew, snow, or
in an occult form
primary productivity the rate at which new organic matter is
added to an ecosystem, usually as a result of green plant pho-
tosynthesis
radiocarbon dating a technique for establishing the age of a
sample of organic matter, based upon the known decay rate
of the isotope 14C (carbon-14)
raised bog a mire in which peat accumulates to form a central
dome that raises the peat-forming vegetation above the influ-
ence of groundwater flow. The surface of the central dome
thus receives all its water input from precipitation; it is
ombrotrophic
realized niche the actual spatial and functional role of a
species under competition from other species in an ecosys-
tem. Compare FUNDAMENTAL NICHE
reclamation the conversion of a habitat to a condition appro-
priate for such human activity as agriculture or forestry
rehabilitation the conversion of a damaged ecosystem back to
its original condition
relict a species or a population left behind following the frag-
mentation and loss of a previously extensive range
replaceability the ease with which a particular habitat can be
replaced if it were to be lost
representativeness the degree to which a site illustrates the
major features characteristic of its habitat type
GLOSSARY 199
resilience the ability of a stable ecosystem to recover rapidly

from disturbance
respiration the release of energy from organic food materials
by a process of controlled oxidation within the cell. Under
aerobic conditions carbon dioxide is released, while anaerobic
respiration may lead to the formation of ethyl alcohol
rheotrophic describing a wetland that receives its nutrient ele-
ments from both groundwater flow and precipitation. In
rheotrophic mires the groundwater flow is usually responsible
for the bulk of the nutrient input
rhizopods microscopic organisms resembling Amoeba, but
with a protective shell around their one-celled bodies. These
shells are often preserved as fossils within peat deposits
salt marsh coastal intertidal wetlands dominated by herba-
ceous plants
sediment material that is deposited in an ecosystem, such as a
lake or a peat land, and accumulates over the course of time.
Sediments may be organic and/or mineral in their nature
sedimentation the process of sediment accumulation
snow patch an accumulation of snow that is sufficiently deep
to survive well into the summer season and may even last for
several years. Its edges melt each summer
solifluction the movement of soils down slopes in tundra
conditions, which occurs because the surface of the soil melts
while the lower layers remain frozen
spore the dispersal structure of algae, mosses, liverworts, ferns,
and fungi, from which a new individual can grow
stadial a cold period in the history of the Earth that is less
severe or shorter than a glacial episode
stone stripes lines of stones following the contours of a slope
that are produced by frost-heaving in tundra soils
stratification the layering of lake sediments and peats in the
chronological order of their accumulation
stratigraphy the study of layering in sediments and the
description of sediment profiles. Stratigraphy can provide
information on the developmental sequence of a mire over
time
stratosphere the part of the Earth’s atmosphere lying above
the troposphere, from about nine to 30 miles (15 to 50 km)
subalpine the zone immediately below the alpine tundra zone
on mountains
submerged aquatic plants freshwater plants that are rooted
in soil that lies underwater and grow toward but not above
200 TUNDRA
the water surface, although some submerged aquatics bear

flowers that extend above the water surface. Compare EMER-
GENT AQUATIC PLANTS
succession the process of ecosystem development, which is
driven by the immigration of new species, facilitation by
environmental alteration, competitive struggles, and eventu-
ally some degree of equilibration at the climax stage. The
stages of succession often follow a predictable sequence, and
the process usually involves an increase in the biomass of the
ecosystem
tephra the glasslike dust particles emitted from erupting vol-
canoes. Layers of tephra in ice stratigraphy can serve as time
markers, since the dates of eruptions are well known and the
chemistry of tephra often indicates the precise volcanic erup-
tion involved
terrestrial occurring on land
terrestrialization the process of succession whereby aquatic
ecosystems gradually fill in with sediment
till the detritus carried in and on the ice of a glacier, which is
dumped as an unsorted mass when the glacier melts; some-
times called boulder clay
timberline see TREE LINE
topography the general form of a landscape, including hills
and valleys
transpiration the loss of water vapor from the leaves of terres-
trial plants through the stomata, or pores, in the leaf surface
tree line (timberline) the altitude at which general tree
growth ceases on mountains, thus forming the boundary
between open forest and alpine tundra. Isolated trees may
survive beyond the tree line
trophic level the collection of organisms that occupy a partic-
ular stage of the energy flow through an ecosystem, such as
primary producer, herbivore, predator, and decomposer
troposphere the lower layer of the Earth’s atmosphere, up to
about nine miles (15 km)
tundra the open vegetation of cold, Arctic conditions, found
in the polar regions (Arctic tundra) and on high mountains
(alpine tundra). Trees are absent, apart from dwarf species of
willow and birch
ultraviolet radiation short-wave radiation from the Sun that
is largely absorbed by the ozone layer in the stratosphere. It is
harmful to living organisms
GLOSSARY 201
uniformitarianism the theory that geological processes such

as erosion and sedimentation took place in essentially the
same way in the past as they do at present. The present can
thus act as a clue to the past
vertebrate an animal with a backbone
vulnerability the degree to which an ecosystem is threatened
with conversion to alternative uses, such as drainage of a wet-
land for use in agriculture or forestry. Contrast FRAGILITY
watershed the geographical region from which water drains
into a particular stream or wetland (equivalent to catchment).
The term is also used to describe the ridge separating two
catchments—literally the region where water may be shed in
either of two directions
water table the upper boundary of groundwater in the soil
weathering the breakdown of rock into smaller particles in
soils due to the activity of chemical, physical, and biological
processes
wetland a general term covering all shallow aquatic ecosys-
tems (freshwater and marine) together with marshes,
swamps, fens, and bogs
wildlife both the wild animals and wild plants of a habitat
xeromorphic structurally adapted to resist drought
zonation the banding of vegetation along an environmental
gradient, as in the transition around a shallow water body
from submerged and floating aquatic plants, emergent aquat-
ics, then to reed bed, and finally swamp
FURTHER READING
General biogeography
Archibold, O. W. Ecology of World Vegetation. New York: Chapman &
Hall, 1995.
Bradbury, Ian K. The Biosphere. 2d ed. New York: Wiley, 1998.
Brown, J. H., and M. V. Lomolino. Biogeography. 2d ed. Sunderland,
Mass.: Sinauer Associates, 1998.
Cox, C. B., and P. D. Moore. Biogeography: An Ecological and Evo-
lutionary Approach. 7th ed. Oxford: Blackwell Publishing, 2005.
Gaston, K. J., and J. I. Spicer. Biodiversity: An Introduction. 2d ed.
Oxford: Blackwell Publishing, 2004.
Tundra history
Houghton, J. Global Warming: The Complete Briefing. 3d ed.
Cambridge: Cambridge University Press, 2004.
Imbrie, John, and K. P. Imbrie. Ice Ages: Solving the Mystery.
Cambridge, Mass.: Harvard University Press, 1979.
John, Brian S. The Winters of the World: Earth Under the Ice Ages. New
York: Wiley, 1979.
Pielou, E. C. After the Ice Age: The Return of Life to Glaciated North
America. Chicago: University of Chicago Press, 1991.
Roberts, N. The Holocene: An Environmental History. 2d ed. Oxford:
Blackwell Publishing, 1998.
The tundra ecosystem and its inhabitants

Barbour, M. G., and W. D. Billings. North American Terrestrial
Vegetation. 2d ed. Cambridge: Cambridge University Press, 2000.
Crawford, R. M. M. Studies in Plant Survival. Oxford: Blackwell
Scientific Publications, 1989.
Knystautas, A. The Natural History of the USSR. London: Century,
1987.
Sparks, J. Realms of the Russian Bear. London: BBC, 1992.
Polar tundra
Aleksandrova, V. D. Vegetation of the Soviet Polar Deserts. Cambridge:
Cambridge University Press, 1988.
203
204 TUNDRA
Chernov, Y. I. The Living Tundra. Cambridge: Cambridge University

Press, 1985.
McGonigal, D., and L. Woodworth. Antarctica: The Complete Story.
London: Frances Lincoln, 2003.
Ritchie, J. C. Past and Present Vegetation of the Far Northwest of
Canada. Toronto: University of Toronto Press, 1984.
Young, S. B. To the Arctic: An Introduction to the Far Northern World.
New York: Wiley, 1994.
Alpine tundra
Bowman, W. D., and T. R. Seastedt. Structure and Function of an
Alpine Ecosystem: Niwot Ridge, Colorado. Oxford: Oxford Univer-
sity Press, 2001.
Hambrey, M., and J. Alean. Glaciers. Cambridge: Cambridge Uni-
versity Press, 1992.
Larson, D. W., U. Matthes, and P. E. Kelly. Cliff Ecology. Cambridge:
Cambridge University Press, 2000.
Matthews, J. A. The Ecology of Recently Deglaciated Terrain. Cam-
bridge: Cambridge University Press, 1992.
WEB SITES
Antarctican
URL: http://www.antarctican.com
This Tasmanian web site carries current news from Antarctica.
Antarctic Philately
URL: http://www.south-pole.com
This is primarily a philatelic web site, but it contains a wealth
of information on polar exploration.
Arctic Research Consortium Austria

URL: http://www.arctic.at
This is a good site for a wide range of links.
Conservation International
URL: http://www.conservation.org
Particularly concerned with global biological conservation.
Earthwatch Institute
URL: http://www.earthwatch.org
General environmental problems worldwide.
Gateway Antarctica
URL: http://www.anta.canterbury.ac.nz
This site is based in the University of Canterbury in New
Zealand and has much information on wildlife and manage-
ment in Antarctica.
205
206 TUNDRA
Glacier
URL: http://www.glacier.rice.edu
Based at Rice University, in Texas, this is a good educational
site.
The International Union for the Conservation of Nature

(IUCN) Red List
URL: http://www.redlist.org
Many links to other sources of information on particular
species, especially those currently endangered.
National Park Service of the United States

URL: http://www.nps.gov
Information on specific conservation problems facing the
national parks.
Scott Polar Research Institute

URL: http://www.spri.cam.ac.uk
This is the world’s leading database on the Antarctic.
Sierra Club
URL: http://www.sierraclub.org
Covers general conservation issues in the United States and
also issues relating to farming and land use.
United Nations Environmental Program World

Conservation Monitoring Center
URL: http://www.unep-wcmc.org
Good for global statistics on environmental problems.
U.S. Antarctic Program

URL: http://www.polar.org
The site includes information on the activities of U.S. vessels
in the region.
WEB SITES 207
U.S. Fish and Wildlife Service

URL: http://www.fws.gov
A valuable resource for information on wildlife conservation.
U.S. Geological Survey

URL: http://www.usgs.gov
Covers environmental problems affecting landscape conser-
vation.
U.S. National Science Foundation

URL: http://www.nsf.gov
This site includes satellite images.
INDEX
Note: Italic page numbers refer to illustrations.
A conservation of 183–185 altitude

ablation 38 energy flow in 72 and climate 18, 20, 21
active layer, of soil xvi, 48, factors of distribution of and vegetation 26–27,
56 5–6 28
adiabatic cooling 19 geology of 33–35 American bighorn sheep
Africa, mountains in 3, 37 global warming and 177 125, 163
Agassiz, Louis 135–136 grazing animals in 26 American mountain goat
agriculture 96, 161 hunting in 123, 124, 124, 184–185
air pollution 79 125–126, 163, 170 amino acids, antifreeze 93
air pressure 22 landscapes of 22 ammonia 182
Alaska location of 1–3, 4 Amundsen, Roald 155, 158
birds in 103, 104 mammals in 112, Angara Shield 31
climate of 13, 21 121–126 animals 98–128
ecotourism in xix, 168 minerals in 164–165 adaptations to cold 85,
exploration of 153–154 origin of term xvii 98, 102, 112, 113, 115,
glaciers in 37, 38, 39, permafrost in 47 120, 123–124,
142, 142 prehistoric people in 126–128
mammals in 117, 119, 151–152 in energy flow 70–71
124 productivity of 73 evolution of 147, 148
minerals in 164, 166–168 recreation in xix–xx, grazing. See grazing ani-
pingos in 51 170–171 mals
prehistoric people in vegetation in xvii, 1, 18, hibernating 85, 113
149, 150, 151, 161 26–29, 28, 88, 89, migratory. See migration
shrub tundra in 24, 25 90–92, 92 oxygen density and 22
Alaska hare 119 Alps xvii winds and 85, 101
albatrosses 110–111 formation of 34 annual plants 89
albedo 7, 8, 14–15 glaciers in 37 Antarctica. See also South
alder 23, 25 hunting in 163 Pole
Aletsch (glacier) 39 mammals in 120, 122 birds in 103, 108–111,
algae 56 prehistoric people in 110
alpine pasque flower 91 152 climate in 14, 16
alpine tundra recent ice age in exploration of 156–159
birds in 103, 105 136–137 geology of 32–33
climate in xvii, 16, rock falls in 55 hunting in 162
18–22, 29 vegetation in 89 ice melting in 176
209
210 TUNDRA
Antarctica (continued) invertebrates in 99–100, avens, mountain 56, 127

lack of prehistory of 151 101 Axel Heiberg Glacier 158
landscapes of 23 landscapes of 22–23
location of 1, 2, 3, 14 location of 1, 2, 2 B
mammals in 121 mammals in 75,
minerals in 165–166 111–121 bacteria
ozone holes above 179, minerals in 164, blue-green 96, 97
180 166–168 in decomposition 71,
Antarctic Circle 9, 14 nitrogen in 57, 182 76, 79, 94
Antarctic Ice Sheet 32–33, ozone holes above 180 nitrogen fixed by 77, 96
36, 146 permafrost in 46–48, 47 and weathering 56–57
Antarctic jaeger 110 pingos in 50, 51 Baffin Island 22
Antarctic Peninsula 14, 33 prehistoric people in balsam poplar 23
Antarctic Treaty (1959) 165 148–151, 161 bats 111–112, 121
antifreeze compounds productivity of 73 bearberry 27
in invertebrates 99, 102 vegetation in 14, 23–26, bearded vulture 185
in microbes 95 24, 58, 88, 89 beetles 100
in plants 93–94 arctic willow 119 Bellingshausen, Fabian
antlers 116 Aristotle 156, 157 Gottlieb von 157
Apollo 100–101 Asia beluga 163
Appalachian Mountains 26 birds in 104, 105 Bennett, James Gordon, Jr.
Archaean era 32 conservation in 185 155
Arctic Circle 9 glaciers in 41, 136 Bering Strait 155
arctic fox xvi, 75, 118, 120 mammals in 114, 120, bezoar stone 125–126
arctic hare 119–120 124, 125–126 bighorn sheep 125, 163
arctic heather 87 mountains in 3 Bingham Canyon 164–165
arctic lemming 119 prehistoric people in 148 biodiversity xviii, 84–129
arctic poppy 89, 91 tundra vegetation in 24 conservation of xviii–xx,
arctic skua. See parasitic astronomical factors, of 182–186
jaeger glaciation 143, 144 and ecosystem stability
arctic tern 108, 108–109, Atlantic Ocean, currents in 81, 82
109 12, 12, 13 future of 178
Arctic tundra. See also North atmosphere low temperature and
Pole carbon dioxide in 65, 84–86
birds in 74–75, 103–104, 76, 172 microbial 98
105–110 circulation cells of 9–12, primary productivity and
as carbon sink 173 10, 143, 144 73–74
climate in 14, 16, 17 layers of 65–67 biological history of tundra
ecotourism in xix, nitrogen in 65, 76, 77 145–148
168–169 oxygen in 65, 76, 77 biological weathering 56
exploration of 153–156, ozone in 66, 179–180 biomass, as source of nutri-
161–162 pollution of 79 ents 80–81, 82
geology of 31–32 thickness of 7–8 birches 88
hunting in 116, 117, in tundra 64–67 dwarf 24, 87
161–162, 163 atmospheric pressure 11 paper 23
ice melting in 176 aurora australis 67 birds 102–111
ice-wedge polygons in aurora borealis 67 and chemical elements
50, 50–51, 52 avalanches 55 in soil 78, 79
INDEX 211
conservation of 185 mammals in 117, 120, color

evolution of 147 122, 124 of insects 101
insects eaten by 74–75, minerals in 166–168 of mammals 112–113,
100, 103 permafrost in 46, 48 120, 127
migratory xvi, 74–75, prehistoric people in Colorado, average tempera-
103, 104, 105, 150, 161 ture in 16, 18–19
106–109, 107, 108, shrub tundra in 25 common murre 107
147 Canadian Shield 31 common redpoll 104
nidifugous 106 carbon 76 community 69–70
oxygen density and 22 forests as sinks for 173 competition, in community
perching 104–105 tundra as sink for 69–70
plants eaten by 103 172–173 conservation xviii–xx,
predatory 75, 103–104, wetlands as sinks for 182–186
118, 119 173 continents
seabirds 75, 79, 107–111 carbon dioxide collision of 34–35,
seafood eaten by in atmosphere 65, 76, 133
102–103 172 moving away from each
seed-eating 104–105 increase in 172 other 32, 34,
waterbirds 105–107 in photosynthesis 173 132–133
black-capped marmot carbonic acid 64 supercontinents
123–124 Carboniferous period 146 145–146
blackflies 101 caribou 75, 114, 114–117, continuous permafrost 46,
black spruce 23 115, 151, 161, 162, 183 47, 47
blue-green bacteria 96, 97 carnivores. See predators Convict Lake 43
blue hare. See mountain hare cave deposits 44 Cook, James 157
boulder clay. See tills cesium 79 cooling, adiabatic 19
bowhead whale 163 chamaephytes 86–87 copper 164–165
braided ridges 44 chamois 124, 163, 185 cotton grasses 25
brant goose 106 chemical weathering 56–57 cranberry 25, 88
Brooks Range xvii, 22, 27 chironomids 100 mountain 87
brown lemmings 119 chlorofluorocarbons (CFCs) crane flies 76, 100
Bryum argenteum 94 179, 180 Cretaceous period 134
bumblebees 102 Cladonia 116 crevasses 39
buntings, snow 74, 104–105 climate. See also temperature crowberry 25, 27
butterflies 100–101, 102 in alpine tundra xvii, crustaceans 75
16, 18–22, 29 Cryptozoic eon. See
and glaciation 42, 43 Precambrian time
C and ice accumulation 38 cushion-plant tundra 27
calving 39 patterns of 7–13, 9 cyanobacteria. See blue-green
Canada in polar tundra 14–17, bacteria
average temperature in 16, 19–20
15–16, 16 snow and 29
birds in 103, 104 and tundra distribution D
climate of 9, 13 5–6 Darwin, Charles 135
ecotourism in 168 climate change 134, 138, Davis, John 157
exploration in 154, 155, 140–142. See also global DDT xix, 181–182
161–162 warming; ice ages dead ice 43
landscapes in 22 climbing 170 deciduous plants 87–88
212 TUNDRA
decomposition. See also ecotones 24 exploration, of tundra

humus ecotourism xix–xx, 168–172 153–159, 161–162
in energy flow 71, 76 eider ducks 107 extinction xviii, 184–185
factors of 80–81 elk 183, 185
in nutrient cycling emperor penguin 110, 111 F
78–79, 80–81 enchytraeids 98, 99 falcons
in tundra 80–81, 94–95 endemics 148 gyrfalcons 75, 103–104
De Long, George Washington energy flow 70–72. See also peregrine 103, 104
155 food web feces 79
depressions 11 decomposition in 71, fellfields 27
deserts, cold 11, 17, 26. See 76 Fennoscandian Shield 31
also polar tundra between trophic levels ferns, seed 146
detritivores 76, 99 71 firn 28, 35
discontinuous permafrost in tundra 72–76 flies 100, 102
46, 47, 47 environment. See also pollu- flowers 89–91, 90, 91,
dormancy, of plants 72, 85 tion 100–101
Drake, Francis 156 ecotourism and 169, fluvial outwash 40
drought 85–86, 87–88 170, 171 food chain 71
drumlins 43 extraction of mineral food web 71
dry weight, of plants 73 reserves and 164, chemical elements in. See
ducks 105–107 166–168, 180–181 nutrient cycling
eider 107 enzymes 84, 95 in tundra 72–76, 74
long-tailed 107 epiphytism 97 foothill glaciers 40
migration of 75 Erebus, Mount 158 forests. See also timberline
dunlin 105 erratics 42 altitudinal limit to
dwarf birches 24, 87 eskers 43–44 26–27
dwarf shrubs 53, 63, 64, 87, Europe. See also Scandinavia boreal, soil of 58
97 birds in 104, 105, as carbon sinks 173
dwarf shrub tundra 25 109–110 global warming and
dwarf trees 24–25, 26 climate of 13 176–177
dwarf willows 56 conservation in growing season and 5
183–184, 185 history of 147
E ecotourism in 168 ocean currents and
eagles, white-tailed 104 glaciers in 39, 40, 41, 12–13
Earth 135–138 soil needs of xvi,
age of 132 hunting in 116, 117, 177–178
atmosphere of 9–12, 10, 124, 125–126, 162, temperature and 5, 14,
65–66 163 18
climate patterns of 7–13, mammals in 114, 120, forest tundra 23–24
8, 9 124, 125–126 fossils 33, 44, 135, 139
earthquakes 35 mountains in 3, 21 foxes 119
earthworms 76 permafrost in 47 arctic xvi, 75, 118, 120
ecology 69 prehistoric people in frost heaving 48–49, 51
ecosystem 69–83. See also 148, 152 fungi
energy flow; nutrient tundra vegetation in in decomposition 71,
cycling 24 76, 79, 94
concept of 69–72 evergreen leaves 86, 87, 88 in lichens 96, 97
stability of 81–82 exfoliation 56 and weathering 56–57
INDEX 213
fur trading 154, 162 Gondwana 145–146 harp seal 112

future, of tundra 175–187 granite 165 hawks, rough-legged 75, 118
grasses heathers 88
G in alpine tundra 27 arctic 87
Gama, Vasco da 156 cotton 25 herbaceous plants 90–92, 92
gas reserves 166–168 grass-moss tundra 25–26 herbivores 70, 71
geese 105–107 gray wolf 185 hibernation xix, 85, 113,
brant 106 grazing animals 123–124, 127
migration of 75 in alpine tundra 122–126 High Arctic 17
snow 106 domestic 26, 183–184 semidesert in 26
white-fronted 106–107, in food web 75, 80 vegetation in 25–26
107, 127 hunting of 161 high latitudes 3–4, 7, 8
gentians 89 in polar tundra 114–120 hiking 170
spring 90 Great Slave Lake 154 Himalayas xvii, 34, 37, 143,
geography 1–30 greenhouse effect 172 170
geology 31–68 Greenland history of tundra 130–160
glacial period 130, 137, 143, birds in 104, 106–107 hoary marmot 122
144 climate of 13 hoary redpoll 104–105
Glacier Bay 39, 142, 142, ecotourism in 168 horizons, of soil 58
153–154, 168 glaciers in 38 horned lark 74
glaciers 37–39 ice sheets in 176 Hubbard Glacier 38
advances of 40–41, 42, landscapes of 22 humans
43, 137–138 mammals in 117 prehistoric 148–151,
causes of 142–145 pingos in 51 152, 161
effects of 40–43, prehistoric people in in tundra 148–152
130–132 151, 161 hummocky moraine 43
formation of 29 rocks in 32 humus 58, 63–64
history of 133–142 Greenland Ice Sheet 36–37 hunting 161–163, 170
impurities in 35–36 growing season control of 183, 185
location of 37 of annual plants 89 of mammals 116, 117,
movement of ice in 37, and forest growth 5 123, 124, 125–126,
37–39, 39, 140–142 in polar tundra 15–16, 161–163, 183, 184,
region around edge of. 17, 25, 72–73 185
See periglacial region guillemot, pigeon 107 by prehistoric people
retreats of 41, 41–46, 42, Gulf Stream 12, 12 148, 149, 161
62, 62–64, 63, gulls Hutton, James 135
130–132 ivory 107
size of 37–38 Ross’s 107 I
gley soil 60–61, 64 gyrfalcons 75, 103–104 ibex 125–126, 163
global warming 142, 172, ice
175–178 H and albedo 14–15
Glossopteris trees 33, Hadean era 32 dead 43
145–146 hair, of mammals 112–113, global warming and
gneiss 32 115, 126 176
goats, American mountain hare loss of. See ablation
124, 184–185 Alaska 119 reserves of 36, 176
godwits 74 arctic 119–120 and rocks 41, 42
gold 164 mountain 120, 122 in wind 85
214 TUNDRA
ice accumulation 35–43. See invertebrates 98–102 lapse rate 18, 19

also glaciers; ice caps; ice detritivorous 99–100 lark, horned 74
sheets in food web 74 latent heat 19
climate and 38 in mountains 101 latitudes
and “rock flour” 31 oxygen density and 22 and circulation cells of
ice ages 130–133, 131, 132 plants as habitat for xvi, atmosphere 11
most recent 45, 45–46, 87, 98 high 3–4, 7, 8
133–137 reproduction in 101–102 low 4, 7, 8
prehistoric people in soil as habitat for 98–100 and seasonal variation in
148–151, 152 wetlands as habitat for climate 8–9
timing of 132 100 Laurasia 145
icebergs 39, 42 iron, in soil 61 leaching 64
ice caps 35, 37 isotherm 14 lemmings 75
ice cliffs 39 ITCZ. See Intertropical arctic 119
Iceland Convergence Zone brown 119
glaciers in 35–36, 38 ivory gull 107 Norwegian 117–118,
prehistoric people in 121–122
161 J population explosion of
Iceland purslane 89 jaegers 75 81, 117–119
ice sheets 36–37, 176 Antarctic 110 lichens xv, 96–98
ice shelf 33, 36 long-tailed 118 caribou eating 116
ice snouts 39 parasitic 109–110 epiphytism by 97
ice-wedge polygons 50, Jan Mayen Island 22, 32 formation of 96, 97
50–51, 52 Jurassic period 146 in High Arctic semidesert
ice wedges 49–51, 50 26
inertia 81 nitrogen fixed by 63
K
insecticides 181–182 on palsa mound 53, 54
insects 74 kame terraces 44 photosynthesis in 96,
bats eating 121 Kangshung Glacier 63 97–98
birds eating 74–75, 100, Kenya, Mount 27, 28 pigments in 97
103 kettle-hole lakes 43 and weathering 56
colors of 101 Kilimanjaro, Mount 19 limestone 165
detritivorous 99–100 Klondike River 164 lithosphere 34
in mountains 101 knot, red 105 Little Ice Age 141
plants eaten by 100–101 krummholz 26 little penguin 111
as pollinators 89–90, 91, Lobelia plants 90, 92
100–101, 102 L loess soil 44
reproduction in 101–102 Labrador tea 25 longspur, Lapland 74,
size of 101 lakes 104–105
interglacial period 130, 137, formation of 43 long-tailed ducks 107
143, 144 kettle-hole 43 long-tailed jaeger 118
International Whaling with ramparts 51 loons, Pacific 108
Commission 163 landmasses, and temperature Low Arctic 17
interstadial period 137 13 soil of 58
Intertropical Convergence landscapes 22–23 vegetation of 23–25, 24,
Zone (ITCZ) 11 Lapland longspur 74, 58
Inuit people 113, 150, 151, 104–105 low latitudes 4, 7, 8
161, 163 Lapp people. See Sami people Lyell, Charles 135
INDEX 215
M evolution of 147 nitrogen

Mackenzie, Alexander 154 of mammals xvi, in Arctic tundra 57, 182
Mackenzie River 154, 155 111–112, 115–116, in atmosphere 65, 76, 77
Magellan, Ferdinand 156 127, 147 bacteria fixing 77, 96
mammals xvi, 111–128 Milankovitch theory 143, lichens fixing 63
color of 112–113, 120, 144 in pesticides 182
127 minerals, in tundra 163–168 plants absorbing 71
conservation of mining 164–165 in urine 78–79
183–185 Miocene epoch 133, 134, Norse people. See Vikings
evolution of 147 147 North America. See also
hibernating 113, mollusks 56 Alaska; Canada
123–124, 127 Montreal Protocol (1987) birds in 105, 106–107
hunting of 116, 117, 180 climate of 13
123, 124, 125–126, moraine 42, 43 conservation in 184–185
161–163, 183, 184, moraine lakes 43 ecotourism in 168
185 mosquitoes 100 exploration in 153–154,
marine 75, 112, 121, mosses xv, 94 155, 161–162
162–163 in High Arctic semidesert glaciers in 41, 136,
migratory xvi, 111–112, 26 139–140
115–116, 127, 147 reindeer 116 hunting in 116, 117,
oxygen density and 22 moths 100 161–162, 163
predatory 75, 112–114, mountain avens 56, 127 mammals in 114,
120, 125 mountain cranberry 87 114–116, 115, 117,
prehistoric 149–150 mountaineering 170 119–120, 121, 122,
reproduction in 120, mountain goat 124, 124, 125
127 184–185 minerals in 164,
mammoths 148, 149, 150, mountain hare 120, 122 166–168
161 mountains. See also alpine mountains in 3, 21
marine mammals 75, 112, tundra prehistoric people in
121, 162–163 formation of 33–35 149, 150, 151, 161
marmots 122–124 glaciers in 37, 40 tundra vegetation in 23,
black-capped 123–124 insects in 101 24
hoary 122 location of 2–3, 4 North Atlantic Drift 12
yellow-bellied 122 rock weathering in 55 “northern lights” 67
Matterhorn (glacier) 40, 55 mountain sheep 125 North Pacific Drift 12–13
meadow browns 101 Muir, John 153–154 North Pole
meadows 27 Muir Glacier 154 exploration of 154–156
Medieval Warm Period 141 murre lack of vegetation at 1
mesosphere 66 common 107 latitude of 4
metamorphic rocks 32 thick-billed 78 location of 14
microbes 94–98. See also bac- musk ox 117, 184 precipitation at 11
teria; fungi seasonal variation in cli-
Mid-Atlantic Ridge 32, 34 mate of 9
midges 100, 101 N Northwest Passage, search for
migration 85 nematodes xvi, 98–99 153–155, 158
of birds xvi, 74–75, 103, Newfoundland 21 Norwegian lemming
104, 105, 106–109, New Zealand, glaciers in 37 117–118, 121–122
107, 108, 147 nidifugous birds 106 nunataks 36
216 TUNDRA
nutrient cycling 71–72 peat 53, 57 nutrients absorbed by 79

decomposition in 78–79, peat bogs 51–52 and weathering 56
80–81 penguins xvi, xix, 111, 162 plate tectonics 34, 132–133
and ecosystem stability emperor 110, 111 Pleistocene epoch 133,
82 little 111 134–135, 137, 138, 143,
in tundra 76–81, 77, 78 perching birds 104–105 144, 148
peregrine falcon 103, 104 Pliny 123
O periglacial region 46–53 Pliocene epoch 133, 134
permafrost xvi, 46–48 plovers 74
ocean currents
continuous 46, 47, 47 podzol 58–61, 60
and permafrost 47–48
depth of 46 polar bear xvi, xix, 75,
and temperature 12,
discontinuous 46, 47, 47 112–114, 127
12–13, 133, 141, 143,
geographical distribution polar front 11
144
of 46–47, 47 polar tundra. See also
oceans
offshore 46, 47 Antarctica; Arctic tundra;
primary productivity of
and rocks 48–49, 51 North Pole; South Pole
75
Permian period 33, 146 atmosphere of 66–67
sea level of, global warm-
pesticides xix, 181–182 birds in 74–75, 103–104,
ing and 176
phalaropes 105–106 105–110
offshore permafrost 46, 47
Phanerozoic eon 32 climate in 14–17, 16,
oil pipeline 167, 167–168
phosphorus 19–20
oil reserves 166–168
in Arctic tundra 57 conservation of 183
owls, snowy 75, 103, 104,
plants absorbing 71 energy flow in 72
118
and population explo- factors of distribution of
oxen, musk 117, 184
sions 81 5–6
oxygen
photosynthesis 70 geology of 31–33
in atmosphere 65, 76,
in blue-green bacteria 96 growing season in
77
carbon dioxide in 173 15–16, 17, 25, 72–73
in decomposition 80
duration of daylight in ice melting in 176
density of 22
72 landscapes of 22–23
produced in photosyn-
in evergreen plants 88 location of 1, 2, 2, 3, 3–4
thesis 65
in lichens 96, 97–98 mammals xvi, 75,
for respiration 65, 76–77
in mosses 94 111–121
in soil 61
oxygen produced in 65 minerals in 164,
ozone 66, 178–180
sunlight needed for 85 165–168
physical weathering 55–56 ozone holes above 178,
P phytoplankton 75 179, 180
Pacific loon 108 “piedmont” glaciers 40 productivity of 73
Pacific Ocean, currents in pigeon guillemot 107 vegetation in 1, 14, 17,
12, 12–13 pigments 23–26, 24, 58, 85, 88,
palsa 52–53, 54 in blue-green bacteria 96 89, 145–147
Panagrolaimus davidi 99 in lichens 97 pollution xix, 79, 180–182.
paper birch 23 pines 23–24 See also environment
parasite 69 Scots 24 polygons
parasitic jaeger 109–110 pingos 50, 51 ice-wedge 50, 50–51, 52
Parry, William Edward 154 plant roots stone 49, 51
Peary, Robert Edwin in active layer of soil 48 pomarine 118
155–156 growth of 73, 87 poplars, balsam 23
INDEX 217
poppies, arctic 89, 91 recreation 168–172 sandpipers 74

population 69 red deer. See elk Satyridae butterfly family 101
explosions of 81, red knot 105 sawflies 100
117–119 redpolls saxifrages 87, 88
variation of 120 common 104 in High Arctic semidesert
Precambrian time 32, 33, hoary 104–105 26
131–132 reindeer. See caribou purple 89
precipitation. See also ice; reindeer mosses 116 Scandinavia
snow; water resilience 81 birds in 105
atmospheric pressure respiration hunting in 162
and 11 in energy flow 71 landscapes of 22–23
chemical elements in 79 oxygen for 65, 76–77 mammals in 117–118,
global warming and “rock flour” 31, 41 121–122
176, 177 rock ptarmigan 74, 103 prehistoric people of
and surge 38 rocks 150–151, 156
temperature and 6, 6–7 effects of glaciers on Scots pine 24
in tundra 6, 11, 17, 21, 41–42, 42 Scott, Robert 158
29. See also deserts, cold metamorphic 32 seabirds 75, 79, 107–111
predators 70, 71 permafrost and 48–49, 51 seals xvi
birds 75, 103–104, 118, Precambrian 33, 131–132 conservation of 183
119 shield 32 in food web 75, 112
invertebrates 74 in soil. See skeletal soil harp 112
mammals 75, 112–114, weathering of 55–57 hunting of 162, 163
120, 125 Rocky Mountains 21, 121, sea swallow. See arctic tern
population explosion of 122, 124, 125 sedge marsh 52
118 Ronne Ice Shelf 33 seed-eating birds 104–105
preservation 182–183 Ross, James 157–158 seed ferns 146
primary production 70, Ross, John 154 Senecio plants 90, 92, 92
73–74, 75, 76 Ross Ice Shelf 33, 36, 157 shear lines 39
Priscoan era 32 Ross’s gull 107 sheep
profile, of soil 58 rough-legged hawk 75, 118 American bighorn 125,
proline 93 Russia. See also Siberia 163
Proterozoic era 32 birds in 103, 106, 107 mountain 125
ptarmigans 75, 81 climate of 13 reducing number of
rock 74, 103 ecotourism in 168 183–184
willow 74, 103 glaciers in 38 snow 125
purple saxifrage 89 landscapes of 22–23 shield rocks 32
purslane, Iceland 89 mammals in 116, 117, shields 31–32
123–124 shrub tundra 24, 62
Q prehistoric people in 148 in alpine tundra 27
Ruwenzori Mountains 19 dwarf 25
Quaternary period 134
evolution of 145–147
soil of 58
R S tall 24–25
radiocarbon method of dat- salt, as antifreeze compound Siberia
ing 138–139 93 average temperature in
ranker soil. See skeletal soil Sami people xv, 162 15
razorbill 107–108 sanderling 105, 106 birds in 107
218 TUNDRA
Siberia (continued) formation of 55–57, 62, spruce

fur trading in 162 77–78 black 23
landscapes of 23 frozen. See permafrost white 23
mammals in 114, 122 glacial retreats and 62, stadial period 137, 140
permafrost in 47, 48 62–64, 63 stone, bezoar 125–126
shrub tundra in 25 gley 60–61, 64 stone flies 101
snow in 29 as habitat for inverte- stone polygons 49, 51
Sierra Nevada xvii, 18, 20, brates 98–100 stone stripes 49, 51
34–35, 43, 125 humus in 58, 63–64 stratosphere 66
skeletal soil 57–58, 63 loess 44 subduction zone 34
skiing 171 pesticides and 182 sucking bugs 101–102
slate 165 profile of 58, 59 sugars, as antifreeze com-
snails 56 skeletal 57–58, 63 pounds 93–94, 95
snow snow and 29 sulfuric acid 64
and albedo 14–15 stability of 81–82 sunlight. See solar energy
in alpine tundra 27–29 and timberline xvi, supercontinents 145–146
burial by 85 177–178 surge 38
chemical elements in 79 types of 57–61, 59, 60 Surtsey (island) 32
and climate 29 water drainage in Svalbard 22, 78
depth of 29 60–61, 64, 79–80, swans 106
firn 28, 35 85–86 tundra 107
global warming and solar energy Switzerland, glaciers in 39,
176, 177 distribution of 7–8, 8, 40, 135–136
and growing season 5 21
impurities in 35–36 insects obtaining 101, T
melting 25 102 tall shrub tundra 24–25
persistence of 29. See lack of 72, 84–85 tallus scree 55
also ice accumulation reflected. See albedo tamarack 23
as thermal insulation solar-tracking flowers 90, 91, Tasman, Abel Janszoon 157
29, 52, 85 91 temperature. See also climate
snow avalanches 55 solar wind 67 albedo and 14–15
snow bunting 74, 104–105 solifluction 55 in alpine tundra xvii,
snow crystals 35 South America, mountains 16, 18–20
snowfinch 105 in 3, 21 changes in global 131
snow goose 106 south polar skua. See jaegers circulation cells of
snow patches 28–29 South Pole atmosphere and 9–12,
snow sheep 125 exploration of 157–158 10
snowy owls 75, 103, 104, lack of vegetation at 1 daily variation in 19–20
118 latitude of 4 and decomposition
soil 6 location of 14 80–81
active layer of xvi, 48, precipitation at 11 global, future of
52, 56 seasonal variation in cli- 175–176
in alpine tundra 26–27, mate of 9 lapse rate of 18, 19
29 “south wind” 67 ocean currents and 12,
atmosphere of 64 Spitsbergen (island) 168 12–13, 133
of boreal forests 58 spores 95 in polar tundra 15–16,
chemical elements in spring gentian 90 16, 19–20
77–80 springtails 76, 99–100 and precipitation 6, 6–7
INDEX 219
seasonal variation in 5, location of 1–4, 2, 3, 4 fur trading 154, 162

15–16, 16, 18–19 origin of word xv mineral extraction
and tree growth 5, 14, uses of. See uses, of tundra 163–168
18 tundra swan 107
temperature, low tundra vegetation 86–94 V
and animals 85, 98, 102, adaptations to cold xix, valley glaciers 37, 40, 41
112, 113, 115, 120, 84–86, 92–94 Vancouver, George 153
123–124, 126–128 adaptations to drought vegetation. See also forests;
and biodiversity 84–86 86, 87–88 photosynthesis
and tundra distribution 5 alpine xvii, 1, 18, 26–29, birds eating 103
and vegetation xix, 28, 88, 89, 90–92, 92 dry weight of 73
84–86, 92–94 continental landmasses insects eating 100–101
tephra 35–36 and distribution of 13 roots of. See plant roots
tern, arctic 108, 108–109, in dormancy 72, 85 as source of nutrients 80
109 evolution of 145–147, tundra. See tundra vege-
Tertiary period 134 148 tation
Tetracanthella arctica 99–100 global warming and Vienna Convention (1985)
thallus 96 176–177 180
thermosphere 66–67 growing season and 5 Vikings 150–151, 156, 161
thick-billed murre 78 as habitat for inverte- volcanic dust 35–36
Tibet 47, 63, 105, 143 brates xvi, 87, 98 volcanoes 31, 32, 34,
Tierra del Fuego 21 height of xv, 86–87 144–145, 158
tillites 41–42, 131 polar 1, 14, 17, 23–26, vulture, bearded 185
tills 41–43, 131 24, 58, 85, 88, 89,
timberline 145–147 W
in alpine tundra 18, water supply and 6 wading birds 105–107
26–27 winds and xv, 20–21, 24, water. See also ice; precipita-
global warming and 25, 86 tion; snow
176–177 tussock-heath tundra 25 adiabatic cooling and 19
in polar tundra 14 availability of xvi, 6, 86
soil and xvi, 177–178 and decomposition 80
topography and 21
U
drainage of, in soil 60–61,
vegetation of 23–24 ultraviolet radiation 66, 179 64, 79–80, 85–86
winds and 20–21, 24 Umbilicaria aprina 97–98 from glacial melt 43–44
topography 21, 29, 38 uniformitarianism 135 locked in ice sheets 36,
Torngat Mountains 22 United States. See Alaska; 176
Transatlantic Mountains 33 North America melting snow and abun-
trapping 161–163 Unknown Southern Land dance of xvi, 25
trophic levels 71 156–157 waterbirds 105–107
tropopause 66 urea 79 weathering 55–57
troposphere 65–66 uric acid 79 biological 56
tundra urine 78–79 chemical 56–57
definition of xv–xviii uses, of tundra 161–174. See physical 55–56
factors of distribution of also hunting wetlands
4–7 agriculture 96, 161 as carbon sinks 173
future of 175–187 carbon sink 172–173 formation of xvi, 25
importance of xviii–xx ecotourism and recre- as habitat for inverte-
landscapes of 22–23 ation xix–xx, 168–172 brates xvi, 100
220 TUNDRA
whales 75, 162–163 wind chill 85 wolverine 121

white-fronted goose winds 5–6 wolves 75, 149, 184, 185
106–107, 107, 127 and animals 85, 101 wood nymphs 101
white spruce 23 glacial detritus carried by woolly mammoth 150
white-tailed eagle 104 44 worms xvi, 98–99
willow ptarmigan 74, 103 and snow patches 29
willows 87, 88 solar 67
arctic 119 and tundra vegetation Y
dwarf 56 xv, 20–21, 24, 25, 85, yellow-bellied marmot 122
in tall shrub tundra 86 Yellowstone National Park
24–25 Wisconsin Glacial 139–140 184, 185

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BIOMES OF THE EARTH

Copyright © 2006 by Peter D. Moore

Library of Congress Cataloging-in-Publication Data

Text design by David Strelecky

This book is printed on acid-free paper.

From Richard Garratt:

form a complicated network of feeding interactions, and, at

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

What is the tundra?

that find shelter in the crevices. Within these patches, inver-

found. This is called alpine tundra. The word alpine is taken

biological interest simply because it is capable of withstand-

Why is tundra important?

be conserved. If humans need to improve crops, or domesti-

humans place upon this fragile habitat can be harmful to its

Where is the tundra biome found?

found most abundantly in the mountain ranges of Alaska; in

Ronne Ice Shelf

Map of the Antarctic

and the distribution of

beneath the ice. Some of

the ice sheet lies directly

Ross Ice Shelf

level. The ice shelves

bedrock above sea level

What factors determine tundra distribution?

dra biome is evidently different, being found at high lati-

suspended in the fast-moving air, and this can be extremely

precipitation. The 40 grassland and savanna

temperature and low 0

are therefore restricted to the upper left region of the dia-

The Earth’s climate patterns

not upright, on its axis. This angle varies and is currently

low polar front

The circulation cells of

face is converted to heat, and the air in contact with the

the Intertropical Convergence Zone (ITCZ). Hot air is capable

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

moving cold waters from the Arctic. It is deflected southeast-

Climate in the polar tundra

Fresh snow 0.75–0.95

generally have a lower albedo. The heat absorption by the

because they have to complete their reproductive cycles in a

The polar tundra of the Northern Hemisphere is found in

Climate in the alpine tundra

high Ruwenzori Mountains of Uganda or Mount Kilimanjaro

20,000 Mt. Kenya,

Brooks Range, Scotland

may completely eliminate trees and produce a more tundra-

One final factor of the physical environment of alpine

Diversity of tundra landscapes

northward in the form of the islands of Novaya Zemlya. To

Arctic tundra vegetation

lows, together with alder. Among these are other shrubs,

Away from the wetland sites, however, the High Arctic

Alpine tundra vegetation

summits of some mountains. These areas have shallow soils

elephant acacia tree

scape, often in a sheltered hollow where the wind is not too