Biomes of The World-Arctic and Alpine Biomes

Arctic and Alpine Biomes
GREENWOOD GUIDES TO
BIOMES OF THE WORLD
Introduction to Biomes
Susan L. Woodward
Tropical Forest Biomes
Barbara A. Holzman
Temperate Forest Biomes
Bernd H. Kuennecke
Grassland Biomes
Susan L. Woodward
Desert Biomes
Joyce A. Quinn
Joyce A. Quinn
Freshwater Aquatic Biomes
Richard A. Roth
Marine Biomes
Susan L. Woodward
Arctic and
A L P I NE B I O ME S
Joyce A. Quinn
Greenwood Guides to Biomes of the World
Susan L. Woodward, General Editor
GREENWOOD PRESS
Westport, Connecticut London
Library of Congress Cataloging-in-Publication Data
Quinn, Joyce Ann.
Arctic and alpine biomes / Joyce A. Quinn.
p. cm.
Includes bibliographical references and index.
ISBN 9780313338403 (set : alk. paper) ISBN 978
0313340178 (vol. : alk. paper)
1. Tundra ecology. 2. Tundra ecologyArctic regions. 3.
Mountain ecologyAlps Region. 4. Alpine regions. I. Title.
QH84.1.Q56 2008
577.5
0
86dc22 2008027510
British Library Cataloguing in Publication Data is available.
Copyright C 2008 by Joyce A. Quinn
All rights reserved. No portion of this book may be
reproduced, by any process or technique, without the
express written consent of the publisher.
Library of Congress Catalog Card Number: 2008027510
ISBN: 9780313340178 (vol.)
9780313338403 (set)
First published in 2008
Greenwood Press, 88 Post Road West, Westport, CT 06881
An imprint of Greenwood Publishing Group, Inc.
www.greenwood.com
Printed in the United States of America
The paper used in this book complies with the
Permanent Paper Standard issued by the National
Information Standards Organization (Z39.481984).
10 9 8 7 6 5 4 3 2 1
Contents
Preface vii
How to Use This Book ix
The Use of Scientific Names xi
Chapter 1.
Introduction to Arctic and Alpine Biomes 1
Chapter 2.
Arctic and Antarctic Tundra 33
Chapter 3.
Mid-Latitude Alpine Tundra 83
Chapter 4.
Tropical Alpine Biome 157
Glossary 201
Bibliography 209
Index 213
v
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Preface
In the course of traveling throughout the United States and the world, I have been
fortunate to experience arctic and alpine environments firsthand. I have visited
Lapland and the Land of the Midnight Sun, climbed mountains in the Sierra Ne-
vada and the Pyrenees, and hiked over many high passes in the mountains of
North America, Europe, and Asia. Walking through a flower-filled meadow and
hearing the whistle of a marmot from a rock pile is a treat for the senses.
This book can be only an introduction to a vast topic that would take volumes
to thoroughly explore. It covers a small part of the variety in the natural world,
much of which is at risk because of human activities, whether purposeful or inad-
vertent. A little knowledge of a subject arouses curiositya desire to learn more.
And as one learns more, the subject takes on greater importance. Conservation
issues become more meaningful when the public is aware of the interactions
among elements in the natural world. Hopefully, the information in these pages
will instill an appreciation for arctic and alpine environments and their natural
inhabitants. Instead of seeing individual plants in a botanic garden or animals in
captivity, readers will have a feeling for the value of each in its natural place in the
world.
Arctic and alpine biomes are the cold regions beyond the limit of tree growth,
either because of latitude or elevation. The first chapter explains elements, such as
temperature, precipitation, and forms of life, common to the Arctic, Antarctic, and
alpine regions. It also mentions differences, which are elaborated on in subsequent
chapters. Selected geographic regions are described in chapters dealing with Arctic
and Antarctic, mid-latitude alpine environments, and tropical alpine regions.
vii
Textual material is illustrated with numerous maps, diagrams, photographs,
and line drawings. The intended audience is not only advanced middle school and
high school students, but also university undergraduates and anyone else who is
interested in the natural environment of high mountains.
I would like to thank Kevin Downing of Greenwood Press for his insights and
constant support in the completion of this project. Jeff Dixon did a masterful job of
deciphering my sketches to produce meaningful illustrations. Bernd Kuennecke of
Radford Universitys Geography Department prepared the maps of the worldwide
distribution of arctic and alpine regions. Several people generously provided pic-
tures for the book, while others read drafts and offered suggestions. I am deeply
indebted to all individuals who enabled me to finalize this volume. Any errors,
however, are solely my own.
viii
Preface
Howto Use This Book
The book is arranged with a general introduction to arctic and alpine biomes and a
chapter each on the Arctic and Antarctic Tundra Biome, Mid-latitude Alpine Tun-
dra Biome, and the Tropical Alpine Environment Biome. The introduction
describes unifying characteristics, such as the physical environment of the biome
and plant and animal adaptations. Subsequent chapters begin with a general over-
view at a global scale and descriptions of features specific to that biome. Regional
descriptions are organized by the continents on which they appear. Each chapter
and each regional description can more or less stand on its own, but the reader will
find it instructive to investigate the introductory chapter and the introductory sec-
tions in the later chapters. More in-depth coverage of topics perhaps not so thor-
oughly developed in the regional discussions usually appears in the introductions.
The use of Latin or scientific names for species has been kept to a minimum in
the text. However, the scientific name of each plant or animal for which a common
name is given in a chapter appears in an appendix to that chapter. A glossary at the
end of the book gives definitions of selected terms used throughout the volume.
The bibliography lists the works consulted by the author and is arranged by biome
and the regional expressions of that biome.
All biomes overlap to some degree with others, so you may wish to refer to
other books among Greenwood Guides to Biomes of the World. The volume enti-
tled Introduction to Biomes presents simplified descriptions of all the major biomes.
It also discusses the major concepts that inform scientists in their study and under-
standing of biomes and describes and explains, at a global scale, the environmental
factors and processes that serve to differentiate the worlds biomes.
ix
The Use of Scientific Names
Good reasons exist for knowing the scientific or Latin names of organisms, even if
at first they seem strange and cumbersome. Scientific names are agreed on by inter-
national committees and, with few exceptions, are used throughout the world. So
everyone knows exactly which species or group of species everyone else is talking
about. This is not true for common names, which vary from place to place and lan-
guage to language. Another problem with common names is that in many instan-
ces European colonists saw resemblances between new species they encountered in
the Americas or elsewhere and those familiar to them at home. So they gave the
foreign plant or animal the same name as the Old World species. The common
American Robin is a robin because it has a red breast like the English or Euro-
pean Robin and not because the two are closely related. In fact, if one checks the
scientific names, one finds that the American Robin is Turdus migratorius and the
English Robin is Erithacus rubecula. And they have not merely been put into differ-
ent genera (Turdus versus Erithacus) by taxonomists, but into different families. The
American Robin is a thrush (family Turdidae) and the English Robin is an Old
World flycatcher (family Muscicapidae). Sometimes that matters. Comparing the
two birds is really comparing apples to oranges. They are different creatures, a fact
masked by their common names.
Scientific names can be secret treasures when it comes to unraveling the puzzles
of species distributions. The more different two species are in their taxonomic rela-
tionships the farther apart in time they are from a common ancestor. So two species
placed in the same genus are somewhat like two brothers having the same father
they are closely related and of the same generation. Two genera in the same family
xi
might be thought of as two cousinsthey have the same grandfather, but different
fathers. Their common ancestral roots are separated farther by time. The important
thing in the study of biomes is that distance measured by time often means distance
measured by separation in space as well. It is widely held that new species come
about when a population becomes isolated in one way or another from the rest of
its kind and adapts to a different environment. The scientific classification into gen-
era, families, orders, and so forth reflects how long ago a population went its sepa-
rate way in an evolutionary sense and usually points to some past environmental
changes that created barriers to the exchange of genes among all members of a spe-
cies. It hints at the movements of species and both ancient and recent connections
or barriers. So if you find two species in the same genus or two genera in the same
family that occur on different continents today, this tells you that their fathers or
grandfathers not so long ago lived in close contact, either because the continents
were connected by suitable habitat or because some members of the ancestral
group were able to overcome a barrier and settle in a new location. The greater the
degree of taxonomic separation (for example, different families existing in different
geographic areas) the longer the time back to a common ancestor and the longer
ago the physical separation of the species. Evolutionary history and Earth history
are hidden in a name. Thus, taxonomic classification can be important.
Most readers, of course, wont want or need to consider the deep past. So, as
much as possible, Latin names for species do not appear in the text. Only when a
common English language name is not available, as often is true for plants and ani-
mals from other parts of the world, is the scientific name provided. The names of
families and, sometimes, orders appear because they are such strong indicators of
long isolation and separate evolution. Scientific names do appear in chapter appen-
dixes. Anyone looking for more information on a particular type of organism is
cautioned to use the Latin name in your literature or Internet search to ensure that
you are dealing with the correct plant or animal. Anyone comparing the plants and
animals of two different biomes or of two different regional expressions of the same
biome should likewise consult the list of scientific names to be sure a robin in
one place is the same as a robin in another.
xii
The Use of Scientific Names
1
Introduction to Arctic
and Alpine Biomes
Arctic and alpine biomes, also called tundra, are treeless regions characterized by
several factors that limit plant growth, including low temperatures, shallow
nutrient-poor soils, low precipitation, and drying winds. Incomplete plant cover
gives the impression of a desert, but it is not always dry. Low temperatures limit
evaporation, and there may be moisture or ice deeper in soil and rock. Although it
does not refer to the poles but to high latitudes near and above the Arctic and Ant-
arctic Circles, polar tundra may be a better term for treeless areas in high lati-
tudes because it includes the Antarctic as well as the Arctic. Unless otherwise
specified, arctic is used here to refer to both polar regions. Alpine refers to treeless
areas at high elevations in mountains at any latitude.
It is often stated that alpine climate and vegetation on mountain tops is the
same as in the arctic tundra. Mean annual temperature and duration of snow cover
may be similar, but differences outnumber similarities (see Table 1.1). Both arctic
and mid-latitude alpine tundra have mean annual temperatures below freezing.
Because of their mid- to high-latitude positions, both have extreme differences
between summer and winter temperatures. Winters can be very cold with tempera-
tures never rising above freezing, while summers are consistently cool. Summers
average less than 50 F (10 C) even though air temperature during the day may
reach 6070 F (15.521 C). The growing season, roughly the time between the
last substantial frost in spring and the first in fall is 610 weeks, but freezing tem-
peratures are possible even during the brief summer. Soils are shallow and nutrient
poor, with variations in rockiness and drainage. Both types of tundra have plant life
dominated by dwarf shrubs, mosses, lichens, sedges, and perennial forbs and both
1
Table 1.1 Environmental Comparison between Arctic Tundra and Alpine, Both Mid-Latitude and Tropical
CHARACTERISTIC ARCTIC TUNDRA MID-LATITUDE ALPINE TROPICAL ALPINE ANTARCTIC TUNDRA
Treeline Sea level to 330 ft
(100 m)
Average for 4050 lat
N 5,0009,850 ft (1,500
3,000 m) S 3,3006,550
ft (1,0002000 m)
9,85013,000 ft (3,000
4,000 m)
No trees
Permafrost Continuous and
discontinuous
Discontinuous None Rare
Length and daily mean
temperature of grow-
ing season
Short, with low
temperature
Short, with low
temperature
Year-round, with low
temperature
Short, with low
temperature
Diurnal temperature
variation
Little change because
there is no day or night
Extreme depending on lati-
tude and light regime
Extreme Little change because
there is no day or
night
Seasonal temperature
change
Extreme Extreme Little Extreme
Topography Level to rolling Bare rock, cliffs, deep gla-
cial valleys
Bare rock, cliffs, deep
glacial valleys
Level to rolling, some
mountains, valleys,
and nunataks
Soils Shallow and nutrient
poor or boggy
Shallow and nutrient poor Shallow and nutrient
poor
Shallow and nutrient
poor
Soil drainage Impeded by permafrost Depends on topography Depends on topography Well drained
Light regime (day
length)
24 hours of daylight or
darkness depending on
season
Hours of daylight and dark-
ness vary according to
latitude
12 hours of daylight and
darkness all year
24 hours of daylight or
darkness depending
on season
Sunlight intensity and
ultraviolet light
Low, especially when
cloudy
Intense except when cloudy Intense except when
cloudy
Low, especially when
cloudy
Atmospheric content Normal Decreasing O
2
and CO
2
with elevation
Decreasing O
2
and CO
2
with elevation
Normal
2
Pressure Normal Decreasing with elevation Decreasing with
elevation
Normal
Microclimate in sun vs.
shade
Minor Extreme depending on
elevation
Extreme depending on
elevation
Minor
Wind Calm to strong depend-
ing on location of
Arctic Front
Depends on topography
and latitude
Calm Calm to strong depend-
ing on location of
Antarctic Front
Precipitation Low Increasing with elevation Increasing with elevation
to a cloud level, then
decreasing
Low
Snow cover Thin Varies, thin to deep Varies, thin to deep Thin to none
3
have patterns of wet meadows, dry heath or
shrubs, and rocky habitats according to microcli-
mate and substrate characteristics. Alpine vegeta-
tion patterns are on a smaller scale, with less
extensive wet tundra because of less permafrost
(permanently frozen soil) and better drainage.
Vegetation belts in the Arctic and Northern Hemi-
sphere mid-latitude alpine tundra also have simi-
lar floras. Zones of vegetation successively
change from forests to stunted trees (krummholz)
to tall shrubs to low-stature tundra plants, often
with the same genera or species. Alpine vegeta-
tion in the Tropics and in the Southern Hemi-
sphere, however, is quite different. The biggest
similarity among all arctic and alpine tundra is
the lack of trees, which together with dominance
of particular growthforms and generally cool tem-
peratures categorize these environments into the
same biome.
Differences between arctic and alpine tundra
are also numerous. Light regime and receipt of so-
lar radiation are vastly different. Arctic regions
experience up to six months of daylight or dark-
ness, while alpine regions have seasonality in day-
length and sun angle according to latitude.
Seasonal temperature variations in polar regions
are much greater than diurnal variations. The op-
posite is true for alpine areas in the Tropics; temperatures change more from day to
night than they do from winter to summer. Because no standard daylight and night
exist for a 24-hour period in the Arctic, temperature remains the same, which is
especially important during the growing season in the summer with perpetual day-
light. At high elevations, incoming solar radiation is intense and temperatures are
high during the day, while at night, infrared energy is more easily lost to space,
causing temperatures to drop. Climate characteristics of alpine locations in the
Tropics are distinctive because of a lack of seasonality. Tropical alpine environ-
ment temperature regimes are sometimes described as summer every day and
winter every night. Alpine areas have greater variety in local climates related
to slope steepness and aspect. Steep slopes may intercept low-angle solar
radiation at a more direct angle, thereby absorbing more energy than a flat
area would. Depending on latitude and position north or south of the Equa-
tor, north-facing or south-facing slopes may be always sunny, always shady,
or variable according to season. Some, but not all, alpine areas have strong
winds, while wind in the Arctic is usually negligible. Little snow falls in
.................................................
Sun or Shade?
Most of the energy coming from the sun, short
wavelength, freely passes through Earths
atmosphere to the ground where it is
absorbed. The Earth then reradiates that
energy in longer wavelengths, called infrared.
The atmosphere does not allow longer wave-
lengths to easily escape back to space, instead
absorbing the energy and keeping the air
warm. At high elevations where the air is thin-
ner, even more of the shortwave energy is
absorbed at ground level, raising surface tem-
peratures in sunny locations. When you stand
in sunlight on a high mountain, you feel
warmth because your body is absorbing solar
radiation. Because fewer atmospheric particles
are in the thin air, however, more infrared
energy radiates through the air and back to
space, leaving the surface and air cold. Step into
the shade and you feel the cooler air tempera-
ture. Alpine vegetation experiences these tem-
perature extremes daily. Fog or clouds moderate
temperature by blocking both incoming short-
waves and outgoing longwaves.
.................................................
4
Arctic regions, while some alpine areas receive several feet each winter. Per-
mafrost is often lacking in alpine environments, as are large-scale wet habi-
tats caused by poor drainage. Long Arctic nights may put constraints on
animal life, while thin atmosphere at high elevations affects both plants and
animals.
The term tundra is not easily defined, and it becomes even more problematic at
the Equator and in the Southern Hemisphere. The Russian word tundra, meaning
land of no trees, may be derived from either the Finnish tunturi meaning barren
lands or from the Lapp tundar meaning marshy plain. Strictly speaking, the term
should be used to refer to treeless landscapes in the Arctic or Antarctic as opposed
to mountain tops
The term alpine stems from the European Alps and is frequently used to
describe entire mountain areas. In a more restricted sense, it refers to areas on
mountains above the level where trees grow. The term alpine tundra is also used
for treeless areas at high elevation. Many scientists limit the term alpine to refer to
mountains with steep slopes, excluding high-elevation flat plateaus such as Tibet at
16,400 ft (5,000 m). Because alpine areas are found in all parts of the worldpolar,
tropical, oceanic islands, continental areas, wet, and dry regionsclimate varies
widely. Alpine tundra can be subdivided into two major regions: Arctic-alpine tun-
dra associated with mid-latitude mountains in the Northern Hemisphere and tropi-
cal alpine environments on mountains in the Tropics or Subtropics, primarily in
South America and Africa. Smaller alpine areas on high mountains in the Southern
Hemisphere are a minor third subdivision. Because of north-south migration corri-
dors in the Northern Hemisphere, alpine tundra in those regions bears many simi-
larities to the Arctic, including many of the same plant species. In contrast, tropical
mountains and Southern Hemisphere alpine regions have unique floras because of
their isolation.
Location
Arctic landscapes are circumpolar, surrounding the North Pole on both North
American and Eurasian continents and on Arctic islands. Alpine tundra has a
global distribution and is found at all latitudes, but it can be fragmented and con-
fined to isolated mountain peaks. Northern Hemisphere alpine environments
merge with circumpolar tundra where mountains extend to high latitudes, but they
are isolated on mountain ranges as far south as Nepal and southwestern China.
Except for areas used for mountain pasture, remoteness and inaccessibility of al-
pine areas has helped preserve native and natural vegetation. High mountains are
located in many floristic provinces, which is one factor that contributes to plant di-
versity; but diversity is also enhanced by geographic isolation, climatic change,
mountain building, glaciation, and microhabitats. A small-scale map showing tun-
dra may be misleading because delimitations are frequently broad ecotones rather
Introduction to Arctic and Alpine Biomes
5
than abrupt boundaries, and mountain top areas are often too small to be depicted
accurately.
Definition and Boundaries
Terminology used by various scientists to refer to zones of either arctic or alpine
tundra is not consistent. Terms such as hemi-arctic tundra, southern tundra, mid-
alpine, or low-alpine are imprecise. No precise meaning exists for even the simpler
terms of arctic, subarctic, alpine, and subalpine, and not all authors or scientists
use the same definitions. Arctic stems from the Greek word for bear, Arktos, and is
now used to refer to the high-latitude lands under the North Star in the Ursa Major
and Ursa Minor constellations. Alpine comes from the Latin Alpes for snow-
covered or white mountains north of Italy, the Alps. The sub preface, meaning
not quite infers less of the conditions to make it full arctic or full alpine tundra.
Subarctic and subalpine are often slightly equatorward in latitude or just below
tundra in elevation. Antarctic and subantarctic refer to the Southern Hemisphere
equivalent of arctic and subarctic.
In terms of latitude, arctic refers strictly to the area between the Arctic Circle
and the North Pole, but in terms of biomes, it means the treeless zone, which may
extend south of the Arctic Circle. Tundra may or may not be underlain by perma-
frost. Boundaries of permafrost sometimes correlate with treeline, but some perma-
frost may be a factor of past, not present, climates.
Alpine areas are most often defined by treeline because climate data are sparse
in mountains. The use of treeline as a boundary is generally satisfactory in
the Northern Hemisphere but this criterion cannot be applied worldwide. Tall
rosette plants in tropical mountains make definition of trees and treeline difficult.
Because the western slopes of the Andes rise from desert rather than forest, no tree-
line can occur. The 50 F (10 C) summer isotherm is an often-used criterion. It
sometimes coincides with tundra vegetation because trees cannot grow with cooler
summer temperatures. Subarctic and subalpine refer to the krummholz zone,
where tree species are dwarfed by climatic conditions, but these areas also include
sheltered groups of full-size trees in the Arctic.
High arctic and polar desert refer to bare rocky areas supporting only lichens,
with small pockets of bog or flowering plants in favorable or sheltered habitats.
Equivalent in high mountains is the nival or aeolian zone above the limit of vascu-
lar plants. Although the terms are often used interchangeably, nival refers to snow-
fields and exposed rock, while aeolian refers to high-elevation environments to
which wind transports insects and nutrients. The aeolian zone is most extensive in
the Andes and Himalayas because of their higher elevations.
Indicator plants may be used to delimit tundra. Typical arctic-alpine species
such as moss campion, alpine sorrel, spike trisetum, and purple saxifrage, which
can withstand cool summers and winter winds, are good indicators of Northern
Hemisphere tundra (both arctic and alpine) but not of tropical or Southern
Hemisphere alpine zones.
6
Treeline
The term treeline is indistinct, especially in places where human history of forest
destruction blurs the boundary between natural and artificial and results in an
increase in treeless extent, as in Eurasia. Treeline refers to the transition zone
between forest and the arctic or alpine environment where trees, defined as single
upright stems at least 20 ft (3 m) tall, cease to grow. Some of the same tree species
may continue to occur above treeline as dwarfed or prostrate shrubs. The term tim-
berline, referring to sizeable logs useable for timber, is not synonymous with tree-
line, but the two are often used interchangeably. Treeline varies with latitude,
generally occurring at sea level to 330 ft (100 m) in the Arctic or subarctic to 9,850
13,000 ft (3,0004,000 m) in the Tropics. In the mid-latitudes, treeline is higher in
the Northern Hemisphere, 5,0009,850 ft (1,5003,000 m) at 4050 N compared
with 3,3006,550 ft (1,0002,000 m) at 4050 S (see Figure 1.1).
Several factors are responsible for variation in treeline, especially in the North-
ern Hemisphere mid-latitudes. Slope aspect, coastal versus inland location, and
precipitation add climate variables other than those caused by latitude. Treeline is
highest in continental areas because of more accumulated summer heat. Snowline
shows a similar pattern, again with more variation in the Northern Hemisphere.
Figure 1.1 The elevation of treeline generally increases from the Arctic and Antarctic to-
ward the Equator. However, it peaks in the low mid-latitudes and declines slightly in the
Tropics. Treeline is higher in the Northern Hemisphere because of larger landmasses that
retain more heat. The highest limit of vascular plants follows a similar pattern. (Illustration
by Jeff Dixon. Adapted from Price 1981.)
7
Many factors such as temperature, elevation, daylength, and growing season
are interrelated, making it impossible to conclude that one factor is responsible for
treeline. Treeline is controlled by wind, snowdrifts, rocks, and soils, but trees also
affect the local climate by shading the ground, trapping snow, providing wind pro-
tection, intercepting ultraviolet light, and moderating diurnal temperature changes.
Factors significant to treeline in the mid-latitudes may be irrelevant in the Tropics.
Although only mean values are often available, extremes or duration of climate
factors are usually more significant. Past climates may determine the position of
current treeline because trees live for hundreds of years, but they may not repro-
duce under current conditions. It is frequently stated that a mean temperature of
the warmest month of 50 F (10 C) is the limit for tree growth. While this figure
may be accurate for temperate zones, a mean temperature of 43 F (6 C) for the
growing season, a period ranging from 2.512 months, may be a better worldwide
measure. During the Pleistocene, temperate zone mountains were deforested and
the alpine zone extended to the lowlands, providing migration routes for arctic and
alpine plants. In the Tropics, treeline was 3,3005,250 ft (1,0001,600 m) lower.
With one possible exception, no one factor can account for treelines worldwide.
Why trees do not grow above a certain elevation involves possible causes such as
depth and extent of snow cover, which may shorten the growing season too much.
Deep, long-lasting snow may also harbor molds and fungi, which might inhibit
plant growth. Wind subjects plants to stress, abrasion, blowing ice, and desiccation.
Wind can also blow seeds away from suitable germination sites. Biotic activities,
such as animals eating bark, seeds, or seedlings, or insect infestations are other fac-
tors that can be responsible for the development of treeline. Human activities such
as repeated burning or cutting trees for timber or firewood may have devastated
natural treeline in some areas. Variations in solar radiation intensity or cloud cover
are yet other possibilities. A common explanation is damage from repeated freez-
ing; however, treelines form in maritime mountain climates where frost is rare.
Similarly, mechanical breakage by ice and wind is a potential factor, but treelines
occur in regions that have little wind and ice-scouring. A short growing season that
limits reproduction may contribute to treeline in mid-latitude mountains, but the
concept cannot be applied to tropical mountains that have no seasonality.
Trees need time for soft new tissue or shoots to complete growth and physiolog-
ically winter harden, so they become able to withstand low temperatures and desic-
cation. Trees with a high canopy prevent solar energy from penetrating down to
the soil. At high elevations and cooler temperatures, the root zone does not become
warm enough to allow time for winter hardening and growth is limited. Small-
stature plants in the alpine zone actually have higher temperatures beneath their
canopies, which are lower to the ground, than do taller trees. One response is for
individual trees to be more widely spaced at treeline, which allows the root zone to
become warmer.
Temperature, specifically summer temperatures in the mid-latitudes, usually
determines treeline, but aridity such as in the Andean puna or in central Tibet is
8
also a major factor. Treeline is higher than 14,750 ft (4,500 m) in Tibet and almost
16,400 ft (5,000 m) in the South American puna. Snowlines in those regions are
higher than 19,700 ft (6,000 m), also because of aridity.
Treeline species in the Northern Hemisphere are mostly conifers, dominated by
pines, spruce, and fir. Warmer areas of Eurasia and western North America also
have larch, the only deciduous conifer. Birch is more common in cool, moist cli-
mates such as Fennoscandia (the peninsula that includes Norway, Sweden, and
Finland); and alder and beech are prominent in central Europe and the Caucasus.
Rhododendrons occur at treeline in both Europe and the Himalayas. Tropical and
Southern Hemisphere treeline species are different because of their isolation from
each other. The tropical high Andes have Polylepis, whereas a deciduous southern
beech dominates in the southern Andes. New Zealands treeline is dominated by
Podocarpus and southern beech. Treeline in tropical Africa mountains is represented
by ericaceous shrubs of tree heather and Philippia species, although the zonation
pattern is complicated by tree-like plants that occur in the alpine zone. The highest
tree on the dry Drakensburg Plateau in southern Africa is a Protea.
Physical Environment
Role of Glaciation
Continental glaciation. Continental glaciers, or ice sheets, influenced the land-
scape of not only the arctic and antarctic tundra but also much of northern North
America and Eurasia as far south as the middle latitudes in the United States,
Europe, and Russia. The western part of North America and the eastern part of
Asia, however, were too dry for glaciers to advance that far south. Several times
during the Pleistocene, the climate in the Northern Hemisphere cooled sufficiently
that winter snowfall failed to melt completely the
following summer. Over thousands of years, the
snow accumulated and compressed into ice more
than a mile thick. The sheer weight depressed
the center of the ice mass causing the edges to
move away in all directions. Ice must move over
land to be called a glacier, while the same ice
mass moving into the ocean is called an ice shelf
or ice floe as it floats on water. It is important to
remember that in a glacier, ice is always moving
away from the center; it never moves backward,
toward the area of snow accumulation. If the rate
at which snow and ice accumulates exceeds the
rate of loss (ablation by melting, calving, and sub-
limation) along the edges of the ice, the ice front
advances and moves farther away from the center.
.................................................
How Can Ice Move?
Solid ice in a continental glacier moves very
slowly across the landscape similar to how
semisoft butter or bread dough would move
across a table if pressure were applied to the
center of the mass. Ice moves downslope in a
valley glacier in response to gravitational pull.
The manner of ice movement is complex but
includes sliding over a thin layer of water
melted from friction, like a skier on snow, and
by ice sheer, similar to how each card in a deck
of cards will slide over the others when pres-
sure is applied to the side of the deck.
.................................................
9
If the rate of accumulation equals the rate of ablation, the ice front remains station-
ary even though the ice itself is still moving forward. When the rate of accumula-
tion is less than the rate of ablation, the ice front retreats, or melts back, toward the
center of its origin. These movements occur today in the glaciers covering Antarc-
tica and Greenland.
Specific movements of the ice did different things to the underlying landscape
(see Plate I). As the ice advanced, particularly in Canada near its origins in North
America, it scraped and gouged the land, scouring rocks bare of soil. At the edges
of the ice and during times of glacial retreat, the debris of rock, sand, and clay (gla-
cial drift) that was carried in the ice was deposited as the ice melted. Material de-
posited directly by the ice was not sorted; it became a jumble of different sizes
called boulder clay. Hills and rolling landscapes of boulder clay are called mor-
aines, with several different types depending on specific ice movement. Rivers and
streams carrying meltwater away from the glacier sorted sediments according to
size, leaving behind flat sandy or gravelly landscapes generally called outwash
plains. Smaller areas of sorted sediment within and between moraines were some-
times caused by temporary lakes. Moraines composed of unsorted boulder clay
have variations in drainage and often contain lakes and bogs. Lake beds may be
underlain by poorly drained clay deposits, while sandy and gravelly outwash is
usually well-drained habitat.
Except on highlands above the ice, called nunataks, continental ice sheets obli-
terated all living things on the landscape, forcing arctic plants and animals to move
farther south as the proximity of ice cooled the adjacent climate over a time span of
thousands of years. The last continental glaciers melted away in the Arctic less than
10,000 years ago, leaving little geologic time for plants to recolonize some areas.
Alpine glaciation. The major difference between continental and alpine glaciers is
that alpine glaciers do not cover the entire landscape but are confined to valleys (see
Plate II). Snow and ice accumulates in the upper reaches of steep river valleys with a
V-shape profile. As the ice moves downslope, it carves the sides of the valley as well
as the bottom, resulting in a U-shaped profile with steep, sometimes vertical sides.
As glaciers enlarge and erode more of their valleys, the original rounded landscape
of the uplands between the valleys is eaten way by frost action, eventually becoming
steep, rocky, serrated ridges called aretes. A rich terminology exists to describe vari-
ous alpine glacial landforms. The amphitheater-shaped depression at the headwaters
of the valley where the glacier began is a cirque, which may contain a lake called a
tarn. Moraines develop along the valley walls and at the furthest extent of ice
advance, although meltwater from the ice usually destroys and redistributes the de-
bris in the center of the valley. The rounded, deep valley floor may partially fill with
outwash sediments and become quite flat. Uplands and aretes are usually dry and
windy, while valley floors may have wet habitats such as meadows and bogs.
When arctic plants followed continental glacial retreat back north, they also
retreated upward into the mountains and found refuge during warm interglacial
10
periods in the cooler climate at higher elevations.
Isolation in different mountain ranges and different
habitats in the alpine glacial landscape afforded
opportunities for evolution and speciation.
Snow
Snow is possible at any time of the year at high
latitudes and elevations, even in tropical alpine
environments. Because some snow may persist in
any given summer, plants living in snowy areas
may spend all summer covered with snow. Snow
cover, however, is generally protective. Plants
beneath it have less exposure to extreme tempera-
tures, less winter desiccation, and less wind and
blowing ice. For example, one study showed that
a 14 in (35 cm) snow layer in arctic tundra kept
the temperature at the top of soil near freezing
even though winter air temperature fell as low as
27 F (33 C). In Siberia, less than 8 in (20 cm)
of snow can delay the date soil freezes by two
months, even though air temperature may be
40 F (40 C). The deeper the snow is, the
shallower the frozen soil will be. With little or no
snow, soil freezes deeply, subjecting plant roots to
physical and physiological stress. Some of the
coldest air temperatures occur on clear nights just after the passage of a storm.
Even a thin layer of snow helps protect plants. During spring snowmelt, however,
a snowbank will have a negative effect on soil temperature nearby because heat is
used to melt snow rather than warm the soil. Snow accumulations that melt late
also shorten the growing season.
Plant Effects on Microclimate
Plants themselves affect and alter their own microclimate, and cold arctic and al-
pine climates are not always as cold for the plants as official temperature measure-
ments indicate. Plant stature and canopy structure affect air flow and heat
exchange, changing the conditions plants experience. Different parts of plants fre-
quently have different temperatures. Leaves at ground level may be 85 F (30 C),
while the roots of the same plant are in frozen soil. Short plants closer to the
ground usually accumulate more heat than tall plants. Surface temperature near
colonies of haircap moss in Antarctica can be 45 F (25 C) higher than that of the
surrounding air. Cushion plants, prostrate dwarf shrubs, and herbaceous rosettes
(see p. 15 on Growthforms) have leaf temperatures that differ the most from air
.................................................
Snow as Insulation
What do a snow cave and a potholder have in
common? Why does a potholder help you
remove a hot pan from the oven? It is not the
thickness of the material, but the air inside. Air
is a poor conductor of heat, so the air spaces in
the potholder keep the heat in the pan away
from your hand. Water is a good conductor of
heat, so if the potholder is moist, heat will be
transferred in seconds and your hand gets
burned.
Fresh snow has up to 10 times more insulat-
ing capacity than old, compacted snow because
it has a lot of air spaces. With fewer air spaces,
old snow conducts more heat from the ground
and offers less protection. An ice crust also low-
ers thermal protection because fewer air spaces
exist and heat is more efficiently conducted
away from the soil. Because wet snow is a better
conductor of heat, it has less insulating capacity.
A snow cave will keep you warm by not trans-
mitting your body heat to the outside air.
.................................................
11
temperatures. Surface temperatures compared
with air temperatures are more pronounced on
sunny days and almost negligible under cloudy
conditions. Heat accumulates whenever it is
sunny. Cushion plants are particularly efficient at
trapping that heat. Their smooth, closed canopy
inhibits radiation loss. Some plants retain heat by
means of hollow stems which may be 36 F
(20 C) warmer than the outside air. Fine hairs or
fibers hold heat and also shade plants from strong
solar radiation. Rosettes and cushions both hold in
heat and decrease wind. Warmer conditions in the
ground canopy, however, result in a steep temper-
ature gradient between the ground and air, which
promotes water loss via more evaporation and
transpiration. Root temperatures are also warmer
under low-stature vegetation. Deep roots and rhi-
zomes, however, are colder than the shoots.
Tussock grasses, taller shrubs, and krummholz experience temperatures more
similar to those measured at standard weather station height and do not undergo
extreme diurnal temperature changes. The lower, dead leaves of tussock grasses
serve as both insulation and windbreak, keeping the interior growing shoots warm.
Measurements in a fescue tussock in East Africa illustrated that temperature of the
outer leaves had a large diurnal range, while temperatures at the base varied little.
Giant rosettes in tropical alpine environments are warmer than the air because
their large leaves absorb heat. They also usually grow in areas with little wind so
the heat is not easily dispersed.
Darker leaves, possessing red pigment mixed with green, absorb more solar
radiation and have a distinct advantage in absorbing heat under a snow cover.
Some plants have thick and waxy leaves (sclerophyllous), enabling them to better
withstand wind and blowing ice crystals. Dead leaves and branches often serve
as a wind break. Other plants, especially mosses and lichens, are able to absorb
water through their leaves, a distinct advantage if soils are dry or soil water is
frozen.
Soils
In general, soils are poorly formed entisols or inceptisols, because most arctic and
alpine tundra areas have only recently been free of ice with little time for soil devel-
opment. The cold climate inhibits chemical actions that release nutrients from par-
ent material such as rock or sediment. Organic matter is limited because of sparse
or small vegetation. Soil development is often interrupted by solifluction or other
processes related to permafrost. Needle ice is common on moist, fine-textured
.................................................
Weather Measurements
Weather data used in climate records are meas-
ured at a standardized height of 5 ft (1.5 m)
above the ground in a shaded location. A ther-
mometer absorbing solar radiation from direct
sunlight would not accurately measure the
temperature of the air. Ground temperatures
are more extreme than air temperatures
because the surface absorbs solar radiation
during the day and reradiates it at night. The
consistency in measurements reduces the
effects of the surface and ensures that data
from a wide variety of locations worldwide can
be compared with no complications.
.................................................
12
soils. As surface soil freezes overnight, small ice pedestals lift soil particles and
small stones up to 1 in (2.5 cm) above the surface.
However, physical location, parent material, and drainage do cause variation
in tundra soils. Rocky arctic areas scraped bare during the Pleistocene remain so,
with little or no soil development. Gravelly surfaces are common in the coldest and
driest polar deserts. In contrast, sediment overlying permafrost is saturated and
boggy. Alpine areas also have extremes, with exposed rocky ridges having little to
no soil. Sediment collecting deep between boulders, however, may form a rudi-
mentary soil in small pockets. Former lake beds on valley floors where accumu-
lated sediment supports dense meadows often have initial development of soil
horizons. Poorly drained areas form bogs with histosols.
Although not much is known about the nutrient requirements of arctic and al-
pine plants, nitrogen and phosphorus are the most limiting. Bird and animal waste
enriches the soil with nitrogen, stimulating plant growth. Species richness increases
around animal dens or burrows and bird nesting grounds. After population explo-
sions of lemmings in the Arctic, surviving plants grow more luxuriantly. Nitrogen
enrichment is significant because few nitrogen-fixing plants occur above treeline.
Cold temperatures also limit the activity of nitrogen-fixing bacteria in the soil or in
root nodules.
Permafrost, a condition in which the subsoil is permanently frozen with only
the surface layer thawing in summer, is widespread and may be continuous in the
Arctic. Patchy in alpine zones, some areas of permafrost may be relicts of past gla-
ciations. Permafrost may be continuous in mountain areas above the upper level of
plant growth. Permafrost not only prevents deep rooting but also locks up nutrients
in frozen soil. Melting of the abundant ground ice may disrupt the surface, causing
collapse, buckling of vegetation, or soil movement. Destruction of vegetation, road
construction, off-road vehicle use, or any activity that alters the heat at the surface
can cause the underlying ice to melt. See Chapter 2, Arctic and Antarctic Tundra,
for a thorough discussion of permafrost.
Origins of Arctic and Alpine Flora and Fauna
Although regional variety exists in arctic and alpine floras, so do commonalities.
The most important arctic-alpine plant families worldwide are sunflower, blue-
grass, mustard, pink, sedge, rose, and buttercup. Also significant and widespread
are gentian, carrot, mint, primrose, bellflower, and buckwheat families. Shrub fam-
ilies are primarily heather and sunflower. Representatives of other families are
characteristic in parts of the Southern Hemisphere.
Alpine floras are a mixture of plants, including widespread elements, immi-
grants from adjacent regions, and recent evolution. It is estimated that before the
Pleistocene, circumpolar flora included about 1,500 species of vascular plants. Ice
sheet advances caused arctic species to migrate south. With glacial retreat, alpine
13
survivors migrated back north to the Arctic. Complex north and south migrations
occurred several times, intermixing the arctic-alpine flora. As the climate warmed
and cold-adapted species retreated upward in elevation, alpine tundra vegetation
became isolated on separate mountains, allowing the evolution of forms distinct to
each locale. Glaciation also isolated flora in higher-elevation refugia or nunataks,
from which they recolonized when the ice retreated. Low-elevation corridors that
retained cool climates for a limited time after glacial episodes also allowed plants
to spread. For example, edelweiss, originally native to central Asia, extended its
range westward to the Alps. Relicts of Carline thistle remain in the Alps, but the
species is now more extensive in Siberia. Because of the east-west orientation of
Asian mountains, flora is more fragmented, with more regional variation than in
North America. American mountain ranges with a north-south orientation have
more similar floras because the continuous mountain chains provided more con-
nected migration routes. Areas that stayed ice-free, such as the north slope of
Alaska, central and western Alaska, and the western and central Canadian archi-
pelago in North America, served as refugia. Nonetheless, Pleistocene glaciations
and climate changes reduced the circumpolar flora to about 1,000 species.
The ability to cope with environmental changes and extremes demanded evolu-
tionary adaptation. Many arctic-alpine plants evolved from lowland species. The
incidence of polyploidy (more than the normal number of chromosomes) increases
with latitude. In the low-latitude Arctic, 60% of vascular plants are polyploid,
while 70% are so in the high-latitude Arctic. Polyploidy may have provided more
genetic variation to enable plants to withstand increasingly severe conditions.
Some mountain plants are ecotypes, plants with genetic variations adapted to
harsher conditions compared with lowland members of their species. Ecotypes rep-
resent an early stage in evolution because they have not yet evolved into different
species. Deserts and arctic-alpine plants may have some common ancestors. Both
types of plants require physiologies that tolerate
dehydration, either because of drought or freezing
temperatures. Endemics are plentiful.
Plant Adaptations
Arctic and alpine plants grow in stressful environ-
ments, with low air and soil temperature, low nu-
trient availability, extremes of atmospheric and
soil moisture, and a short growing season, all of
which limit both growth and sexual reproduction.
Some plants grow in cold water-saturated soils
with little available oxygen, while others grow in
habitats that change from wet after snowmelt to
dry in mid-summer. Arctic and alpine plants share
.................................................
Chromosome Numbers
Cells are normally diploid; they have two sets
of chromosomes, one each from the male and
female parent. Some organisms, however, are
polyploid. They have more (3, 4, 5, or even 6)
copies of each chromosome. Triploid means
there are three sets, tetraploid means there are
four sets, and so on. Polyploidy is common in
plants and can occur by natural means or can
be induced artificially in a laboratory. Because
the plants have more chromosomes, more
combinations are possible, and they may have
greater adaptive ability.
.................................................
14
many adaptations, but some may be more impor-
tant than others because of variations in local con-
ditions. Arctic soil and air temperatures tend to be
lower than those in alpine habitats because of per-
mafrost. Alpine soils are generally better drained
because of slopes and less permafrost, so drought
stress is more common in alpine environments.
However, drought stress is important in polar
deserts (extremely cold tundra), where it is also
dry. Both arctic and alpine regions can be windy
environments, especially on exposed ridges, which
usually are dominated by plants of low stature such
as cushions and rosettes. Light regimes vary, and
plants such as alpine sorrel and spike trisetum may
have different ecotypes adapted to continuous
summer light in the Arctic or 1518 hours in mid-
latitude alpine situations. Some alpine plants more
efficiently use carbon dioxide and have higher rates
of photosynthesis because less carbon dioxide is
available in the thin air at higher elevations.
Growthforms
Chamaephytes, plants that hold their regenerating buds just above soil level, are
the most common growthform in arctic and alpine biomes. Hemicryptophytes,
which hold buds at the surface, are also common, but they are less important in al-
pine tundra than in arctic environments. The two most common growthforms in al-
pine tundra are short or prostrate woody shrubs and herbaceous perennials of
several types. Less common or regional growthforms include giant rosettes on trop-
ical mountains, geophytes on mid-latitude mountains with pronounced seasonal-
ity, stem and leaf succulents, and annuals or biannuals, which become rarer at
higher elevations. Although most arctic plants use the C
3
pathway in photosynthe-
sis, they can usually photosynthesize at lower temperatures than their temperate
counterparts. Succulents, which are capable of the Crassulacean Acid Metabolism
(CAM) method of photosynthesis, live in dry niches at all latitudes, including sub-
arctic. Bryophytes and lichens, which are nonflowering and often desiccation toler-
ant, are common and often extend to higher elevations. Because alpine tundra is
defined as above treeline, few or no phanerophytes (plants such as trees that hold
buds high above the ground) occur. Some growthform characteristics may be due
to poor nutrition as well as to low temperatures.
Prostrate shrubs are well adapted to arctic-alpine conditions. The fact that arc-
tic willow is found north of 83 N, almost to the limit of plant growth in the Arctic,
indicates its hardiness and resistance to cold conditions. A major advantage of the
small shrub growthform is its permanence because new tissue is not needed every
.................................................
Upper Limit of Plant Growth
The highest vascular plant in the world, Saus-
surea gnaphalodes, is found on scree at 21,000
ft (6,400 m) on the north side of Mount Everest.
Widespread in the Himalayas and other moun-
tains, it is an herbaceous perennial rosette
cushion in the sunflower family with thick
white hairs. Other plants holding records for
life at high elevations are a sandwort (Arenaria
musciformis) at 20,413 ft (6,222 m) and greater
stitchwort (Stellaria holostea) at 20,132 ft (6,136 m),
both in the Himalayas. Lichens and mosses
grow to even higher elevations than vascular
plants because they can tolerate complete des-
iccation. They are found at 19,350 ft (5,900 m)
on Mount Kilimanjaro, 22,000 ft (6,700 m) in
the Andes, and 24,250 ft (7,400 m) in the Hima-
layas. More than 50 species of lichens grow
above 13,100 ft (4,000 m) in the European Alps.
.................................................
15
year. The plant can wait out bad years before
resuming growth and reproduction. This advant-
age is even more significant if the shrub is ever-
green. Ground-hugging or prostrate plants are
covered and protected by winter snow. They are
also able to absorb more heat from the ground in
summer.
Plants may be evergreen or deciduous. Some,
like arctic bell heather or moss plant, withdraw
chlorophyll from their leaves at the end of the
growing season, leaving winter foliage red. Others
remain green. Evergreen plants do not replace
their leaves every year and need fewer nutrients;
they grow on acidic, low-nutrient soils. Decidu-
ous willows and birches grow more quickly but
also need better soil. Several shrubs, especially in
New Zealand, transfer carbohydrates and
nutrients from leaves to the woody parts of the
plants for winter, returning them to the growing
buds in spring. Plants that maintain a few leaves
above ground begin photosynthesis before snow
fully melts because radiation can penetrate snow
up to 20 in (50 cm).
The most typical tundra growthform, how-
ever, is the herbaceous perennial, which depends
on long roots and rhizomes. Four major types
graminoid, cushion forb, mat forb, and leafy
forbdominate along with a few small ferns and
bulbs (see Figures 1.2 and 1.3). Most have a deep
root system that serves both to store carbohy-
drates and extract deeper soil moisture. Surface
roots absorb heat from the soil layer that warms
quickest, while large tap roots store energy and
nutrients. Thick taproots may stabilize plants
under frost-heaving or needle-ice conditions, but,
thick roots may break when frozen.
Graminoid plants include both grasses and
sedges, usually growing in clumps or tussocks.
Grasses have hollow round stems, while sedges
are usually triangular in shape and have a noticea-
ble difference to the touch.
Low-growing with a streamlined shape, cush-
ion plants such as the circumpolar moss campion,
.................................................
Lichens
Lichens are organisms composed of a symbi-
otic association between fungi and algae that
takes several forms. Crustose lichens are crust-
like, growing flat against the rock or soil.
Foliose lichens are leaflike, and fruticose lichens
are shrubby or hair-like. In addition to green,
they can be yellow, orange, or brown, with
many color variations.
.................................................
.................................................
Cryptogams and Phanerogams
The term cryptogam, literally meaning hidden
seed, refers to various algae, fungi, mosses,
lichens, ferns, and other lower plant or plant-
like organisms that lack flowers and reproduce
by means of spores too small to be seen. Blue-
green bacteria, green algae, slime molds, fungi,
lichens, molds, and yeasts are in a category
called thallophytes. Liverworts and mosses are
bryophytes, structurally simple green, seedless
plants. Many cryptogams, especially algae, are
tiny and must be observed with a hand lens or
microscope. Most others are small, 0.82 in (25 cm)
tall or less than 4 in (10 cm) long. They form
flattened mats, spongy carpets, tufts, or turfs
according to humidity and sunlight. Most cryp-
togams are nonvascular plants. Although cate-
gorized as cryptogams because they do not
produce seeds, club mosses, horse-tails, and
ferns are vascular plants called pteridophytes.
Cryptogams form a significant part of arctic-
alpine floras.
In contrast, the term phanerogam refers to
plants with visible seeds, like flowering plants
such as angiosperms and gymnosperms. Vas-
cular plants such as phanerogams have differ-
entiated tissue designed to transport water
and nutrients from roots to leaves.
.................................................
16
are common pioneering plants in exposed, windy areas. Formed of many short
branches that fit tightly together, they are not soft like the name cushion implies
but present a firm rounded surface. The plant may be sculpted by wind and ice
because any twig projecting above the general level is quickly cut back. The tightly
packed branches expose a lot of leaf area at the cushion top, but beneath the can-
opy, it is warm and protected from the wind. Cushion plants improve their micro-
habitat by accumulating plant debris and nutrients beneath their canopy. Moss
campion grows in size by adding more branches, but growth rates are slow. Only
0.5 in (1.3 cm) in diameter after the first five years, the plant may need 25 years to
reach 7 in (17.8 cm). Initial growth is slow because energy is directed toward devel-
opment of a taproot system. A 10 in (25.4 cm) diameter plant can have a 45 ft
(1.21.5 m) taproot, good for both anchoring the plant and obtaining subsurface
water. Cushion plants in South America and New Zealand, however, grow much
larger.
Figure 1.2 Four major growthforms dominate the arctic and alpine tundra: (a) grami-
noid, (b) cushion, (c) mat, and (d) leafy forb. (Illustration by Jeff Dixon. Adapted from
Zwinger and Willard 1972.)
17
Mat plants, such as dwarf clover and alpine sandwort, are a looser tangle of
prostrate branches that root over a large area. Ground-level branches and short
upright stems are not so tightly packed as in cushions and are not dependent on
one long taproot. The plant grows larger as prostrate branches root where they
touch the ground. Because they have a short structure of woody twigs, cushions
and mats are often called tiny shrubs.
Rosettes, such as tundra dandelion, are a major type of leafy forb. They grow
flat to the ground, sometimes closer to the ground than cushions or mats, in the
warmest microenvironments. Overlapping leaves, which form a circular pattern on
a very short stem, present maximum exposure of leaves to light. Smaller rosettes
are offset from the mother plant. Some plants, especially rosettes such as snowball
Figure 1.3 Four major growthforms of arctic and alpine tundra include (clockwise from top left)
sedge, moss campion, alpine sandwort, and big-rooted spring beauty. (Photos by author.)
18
saxifrage, big-rooted springbeauty, and yellow stonecrop, have succulent leaves or
a thick taproot. The taproot of big-rooted springbeauty can be 6 ft (1.8 m) long or
more, while the succulent leaves of stonecrop form marble-size balls. Waxy leaf
surfaces contribute to moisture retention.
Plant hairs both offer protection from intense radiation and drying wind and
also help to absorb or conserve heat (see Figure 1.4). Dark hairs such as found
on draba, fleabane, and arctic poppies are less reflective and absorb more solar
radiation. In contrast, light-colored, silky, wooly, or transparent hairs such as on
pasqueflower, some lousewort, cottongrass, and some willow catkins allow solar
radiation to pass through where it is then absorbed by the darker surface of the
plant. Longwave energy reradiated from the plant cannot escape and the energy
is trapped in a mini-greenhouse of hairs. The fluffy tops of arctic cottongrass tus-
socks retain heat that delays soil freezing. Freezing of the surrounding soil pushes
the mound of cottongrass upward where it occupies a position to get more of the
suns rays in spring. It emerges from snow before adjacent, slightly lower-lying
plants do, increasing its growing season by 410 days. Some of the hairiest plants
Figure 1.4 Like pasqueflower, many arctic and alpine plants have hairs to both protect
from excessive sun and retain heat. (Photo by author.)
19
are the earliest to flower because they trap more energy or heat. Hairiness takes
many different forms.
Coping with Climatic Stress
Both arctic and alpine plants are subjected to extremes of low temperature. Plants
are more susceptible when they are not frost hardened. During winter, the plants
physiology alters, usually through dormancy, so cold can be endured. A decrease
in the photoperiod (hours of daylight) and lower temperatures at the end of
summer also harden plants. Soil drought may be another factor in winterhardening
for alpine plants. From spring to fall, and every night in the Tropics, growing tissue
is more susceptible to freezing. Aboveground tissue, especially new growth, may
die, but the plant is usually not killed. If new growth has not yet begun, or if the
plant is still protected under snow, it will not be affected.
Tolerance limits of alpine plants vary with geographic location and plant parts.
The stem and root are usually more tolerant of low temperatures. Low-stature
plants (cushions, dwarf shrubs, rosettes, and sedges) have similar tolerance limits
regardless of family or genera relationships. Mid-latitude alpine plants in the tem-
perate zone such as the Alps can tolerate 14 to 25 F (10 to 4 C) without
damage. Earlier flowering plants, which are more likely to be exposed to colder
early summer temperatures, are most cold hardy. Tropical alpine plants in the
Andes, Africa, and Hawaii can tolerate temperatures even lower, 2 to 16 F
(9 to 19 C). A more stable temperature during the growing season is responsi-
ble for the higher tolerance limits in the mid-latitudes. Low temperatures in the
Tropics are less predictable, which explains the lower tolerance limits.
Plants have several ways of coping with low temperatures. The first is to avoid
exposure. In nontropical mountains with seasons, phenology (the date of growth
activity such as flowering and setting seed) is important. Plants are dormant in win-
ter in spite of periodic warm spells and potentially high radiation and cannot be
damaged by extreme cold. With warming temperatures and longer daylight hours
in spring, plants become active and are most sensitive to cold in summer. A
decreasing photoperiod in late summer signals the plant to finish its growth cycle
before fall. As nutrients in the leaves are reabsorbed by the roots or stems when the
leafy parts die back, the plants hardiness to cold is increased. Plants can avoid ex-
posure to cold by their morphology, particularly plant size and position of regener-
ating buds. Tall plants are above the snow cover and exposed, while short ones are
protected from air temperatures. Most tundra plants carry their regenerating buds
at or close to ground level. Graminoids and perennial forbs, both of which are
abundant in arctic and alpine environments, carry regenerating buds below ground
level, pulled back into the soil as roots contract. Cushion plants store heat under
their low canopy. Tropical rosettes curl their leaves over the center growing point,
and dead leaves insulate the stalk. Liquid inside the rosettes keeps the plants warm.
Microhabitat preference (open versus sheltered, snowbeds versus bare) is a simple
way to avoid exposure.
20
Many plants have the ability to avoid freezing even when temperatures are low.
By accumulating sugars in tissues, the freezing point is lowered, not by much, how-
ever, and this adjustment is least effective. A more efficient method is supercooling.
In a physiological process, the leaves and stem cool to below-freezing tempera-
tures, but tissues do not freeze and suffer damage. Common in areas where low
temperatures regularly drop to 10.5 F (12 C), such as encountered in the South
American paramo, this method is used by the giant rosette Espeletia and some
shrubs. If temperatures drop below the limit of supercooling, however, tissue
freezes quickly and damage occurs. Giant rosettes in tropical alpine areas in Africa
experience even lower temperatures but do not supercool, nor do higher-elevation
Andean plants.
Some plants are able to tolerate freezing. In all plants, 2530% of the plant is
space between cells. Water in intercellular spaces (that is, not inside the cells)
freezes, and in the process, releases heat to the cells and takes water from the cells.
Water or ice between the cells makes the leaf appear darker in color. When thawed,
the cells reabsorb the water within one to four hours and the normal green color
returns. Some plants can begin growth in spring, then cease growth and wait out
temperatures below freezing at no harm to the plant until conditions improve.
Plants adjust to the cool temperatures of the short growing season by preform-
ing flower buds at the end of the summer. Growth of buds and flowers usually
takes more than one season. Catkins of some prostrate willow species develop over
four growing seasons compared with two seasons for shrubby species in better envi-
ronments and one year for lowland species. Protected by the plant and snow cover
over the winter, buds quickly elongate and flower early the following season. Some
plants bloom just two weeks after beginning to grow in spring. Solar radiation can
penetrate up to 2.0 in (50 cm) of snow, an important factor for buttercup, which
begins growth before the snow melts. However, the upper layer of soil must be
thawed for growth to take place. Preformed flower buds are especially common in
New Zealand alpine tundra. Spring growth, beginning with elongation of pre-
formed leaves and flowers, depends on the supply of nutrients from storage organs
as well as increasing receipt of solar energy.
Roots, rhizomes, corms, and bulbs store energy
and nutrients. The root-to-shoot ratio increases
with both latitude and elevation. Root organs
become larger as the aboveground part of the
plant becomes smaller, resulting in more stored
energy. Old leaves serve a similar purpose in
small shrubs, recycling the nutrients, but even
prostrate shrubs are absent from the harshest
climates.
A rare adaptation in arctic-alpine plants is to
repair or replace tissue damaged or killed by
freezing. Preformed leaf buds and flower buds lost
.................................................
Heat Resistance
Because surface temperatures can be extreme,
tolerance of some alpine plants to high tem-
peratures is similar to plants found in tropical
deserts. Lethal temperatures of leaf surfaces
(that is, temperatures at which plants die) in
the Central Alps is 112 F (44 C), the same as
the average in Mauretania in the Sahara. In the
Hindu Kush mountains, the lethal limit is 102 F
(39 C), and in central Norway, 106 F (41 C).
.................................................
21
to frost will not be replaced because the growing season is too short. For the same
reason, flowers or fruit frozen later in the summer cannot be replaced. Fat roots,
however, do store nutrients and to some extent can replace frozen or eaten leaves.
Graminoids and rosettes can activate belowground growing points if damage is not
extensive.
Alpine plants are also subject to heat stress. Heat-trapping structures can be det-
rimental during peak solar radiation, causing ground and plant temperatures to
soar. Establishment of seedlings where high surface temperatures prevail is difficult
at best and is another indication of why vegetative propagation is more common.
Patches of bare soil are frequent in alpine regions, and seedlings are commonly
found in disturbed sites, such as animal burrow entrances and tire tracks. Any ac-
tivity that destroys vegetation opens the ground to accumulation of surface heat.
However, such bare spots can be permanently damaged by erosion before they can
be reclaimed by vegetation.
Surface soils, particularly in alpine environments, can become quite dry, but
because deeper moisture is often available, plants have few adaptations related to
tissue desiccation. Plants showing drought stress usually grow on exposed sites or
in shallow soil. Drought stress in the high-latitude Arctic is common in some
grasses, rosette species, and cushion plants in bare, well-drained soils.
To gain the most energy from solar radiation, some plants orient their leaves
vertically to catch the low sun at a more efficient angle. This vertical orientation
also allows light to be reflected to and absorbed by other leaves instead of back to
space. The parabolic form of several flowers is beneficial for both pollination and
development of seeds. The white and yellow flowers of avens, poppy, and butter-
cup have good reflectivity and concentrate energy toward the flowers center. Some
flower heads maintain an orientation toward the sun like a lowland sunflower.
Many alpine plants not considered evergreen retain some green color in winter.
Examples include some leaves of cinquefoil and matgrass. The bark of young
shoots of dwarf shrubs, especially the green stems of blueberry, can conduct photo-
synthesis. Mosses and lichens undertake photosynthesis in winter, and all alpine
plants are capable of photosynthesis processes at temperatures near freezing. While
much solar radiation is reflected from snow, some wavelengths penetrate, depend-
ing on snows thickness. If snow is shallow, some, but not all, alpine plants can
carry on photosynthesis.
Lichens and some mosses can withstand colder or shorter growing seasons, but
they need water. Lichens have advantages in the low stature of their lifeform, close
to warm daytime surface or rock temperatures, and those warmer temperatures
may melt snow needed for water. Their optimum photosynthesis temperature is
about 41 F (5 C), a temperature easily reached at the surface. Low night or winter
temperatures do not harm lichens, and they can remain dormant while frozen.
They are also long lived and can survive unfavorable conditions in a dormant state.
In the Alps, map lichen lives up to 1,300 years, and in western Greenland, up to
4,500 years. Lichens grow beyond the limits of vascular plants but eventually are
22
limited by the lack of water from snowmelt due to low temperatures and
sublimation.
Reproduction
About 90% of the vegetation is perennial, including grasses, sedges, flowering
plants, mosses, lichens, and prostrate shrubs. Age estimates based on growth rate
of leaves and rhizomes suggest that many arctic species are long lived, delaying the
need to reproduce either sexually or vegetatively. Stems of birch and willow may
be 200400 years old, and dwarf birches rising only a foot high may be hundreds
of years old. Other plants may have equally long life spans: northern woodrush,
90130 years; avens, 80120 years; and cottongrass, 120190 years.
While the most common method of reproduction is vegetative rather than by
seed, most arctic-alpine plants reproduce both ways. Flowering and seed produc-
tion, however, may not be successful every year. Most plants are insect pollinated,
but wind and birds are also factors. Long-tubed flowers are usually pollinated by
bumblebees, which are attracted to colorful flowers. No long-tongued bees such as
bumblebees are native to New Zealand alpine tundra, and unlike the mid-latitudes
and the Arctic, which both have colorful flower displays, the mostly small, flat,
white or yellow flowers in New Zealand are pollinated by small insects, flies, and
short-tongued bees.
Some plants are biennial, taking two years to flower, but few are annuals
because of the short growing season. In Peary Land in Greenland, only 1 of 96 vas-
cular plants, the Iceland purslane, is annual. In the alpine environment of the
Rocky Mountains in Wyoming, only 3 of 121 plants are annual. In general, annu-
als comprise only 12% of the flora in alpine or arctic areas and plants are typically
small. Drier alpine regions have slightly more annuals, for example, 6 of 108 spe-
cies in the Sierra Nevada in California. Some species, such as rock primrose, that
are normally annuals have perennial ecotypes in arctic or alpine tundra.
Arctic annuals have a low root-to-shoot ratio with no need to store energy
because the plant only lives one season. Their growth is indeterminate, meaning
that they will continue to grow flowers and produce seed until killed by either cold
or drought. They use water sparingly, regulating their stomata as the soil dries, and
have a high rate of photosynthesis that allows them to complete seed ripening. No
annuals are endemic to the Arctic.
Germination of most plants takes place soon after snowmelt when soils have
warmed and water is sufficient. Seed dormancy requirements for chilling, light, or
seed coat scarification are present in only 2040% of arctic-alpine species.
An additional safeguard for most plants is that not all seeds germinate every
year, a survival strategy that ensures that a disastrous summer season will not
destroy the species. Seeds may remain viable for many years. Arctic lupine seeds
that were found frozen and preserved for 10,000 years germinated within 48 hours
after being thawed and moistened. Seedlings are rare in the Arctic, but Bigelows
sedge and cottongrass are an exception and can be seen in abundance on bare soils
23
in northern Alaska. Seeds, distributed by wind or animals, are usually small with
no burrs or fleshy fruits.
Clonal growth is supplemental to sexual reproduction, and few plants rely solely
on vegetative means. Most alpine plants in western North America reproduce by
seed, but vegetative reproduction is more common in the Arctic because of the diffi-
culty of setting seed there. Alpine sorrel has two ecotypes, one reproducing by seed
in alpine tundra, the other by rhizomes in the Arctic. Plants which reproduce sexu-
ally at lower elevations may rely on vegetative means in alpine tundra. Vegetative
reproduction is also common where wet areas curtail the growing season. Percent-
age of species with some form of cloning varies from 5080% of total flora in both
arctic and alpine environments, but it may be less important in alpine tundra.
Cloning takes several forms. Many tussock-forming grasses, sedges, and rushes
form dense clusters. As new shoots grow at the edges, old ones in the center die.
Over time, the tussock becomes a ring of living plants surrounding dead centers or
a sinuous pattern of lines (see Figure 1.5). As the most important clonal strategy in
the world, it is used by carex and kobresia sedges, rushes, hairgrass, fescue, mat-
grass, bluegrass, and needlegrass. Some graminoids and forbs grow new plants
from stolons or rhizomes that extend either below or above ground away from the
mother plant. The invasive nature of mint plants and Bermuda grass in a home gar-
den illustrates the efficiency of reproduction by rhizomes.
Young plants can become completely separated from the parent by rock or soil
movement. Graminoid examples include carex sedges, woodrush, bentgrass, and
fescue. Examples of forbs that clone this way include primroses, avens, snow lotus,
ragworts, dock, bellflower, and candytuft. Some mat-forming forbs such as pussy-
toes, Celmisia, and strawflower form herbaceous clusters of rosettes. As plant
Figure 1.5 Because the center dies as the edges expand with growth, vegetative repro-
duction of many types of grass tussocks produces rings or a sinuous pattern of living
plants. (Illustration by Jeff Dixon. Photo courtesy of Susan L. Woodward.)
24
clusters grow larger, they disintegrate into separate plants. Other examples are gen-
tians, plantains, and speedwell.
Dwarf shrubs, which are small, close to the ground, and buried in litter, may
grow adventitious roots from the woody stems, providing the soil is moist. Typical
plants include Coprosma, avens, azalea, and willows. In a similar manner, woody
stems of taller prostrate shrubs (220 in, 550 cm, high) may also become buried in
litter or soil. They first grow shoots from buried buds, with adventitious roots
developing much later. Common plants are crowberry, rhododendron, blueberry,
Hebe, St. Johns wort, willow, and Styphelia.
Viviparous plants form vegetative propagules on inflorescences. A subspecies
of alpine bluegrass produces plantlets, and viviparous bistort produces bulbils
(seed-like shoots) instead of flowers. Some plants that do not ordinarily reproduce
vegetatively, such as glacier buttercup and alpine sorrel, sometimes grow roots
from flower stalks that become accidentally buried.
Many lichens reproduce by means of fragmentation. Broken pieces subse-
quently root to create clones of the parent. As it dries, staghorn lichen contracts
into a ball form that is blown around the tundra by the wind, breaking off pieces as
it rolls. With moisture, it unfurls and continues growth.
Animal Life
Arctic and alpine environments have distinct assemblages of animals. With the
exception of high-latitude mountains that merge with arctic tundra in northern
Alaska, northern Urals, and northeastern Siberia, no mammals and few birds are
common to both regions. Many animals in the Arctic have a circumpolar distribu-
tion, meaning they are found in all or most regions surrounding the North Pole. Al-
pine animals, however, which derive from adjacent lowlands rather than from the
Arctic, have patchy distributions because the alpine areas of high mountains are
like islands in a sea of unsuitable habitat. Isolated areas of alpine environment
are more difficult for animals to colonize than the continuous expanse of the arctic
tundra.
Animal life of arctic and alpine regions can be categorized as permanent resi-
dents, seasonal breeding visitors, or seasonal but nonbreeding visitors. Patterns of
distribution can be continuous, disjunct, or disperse. Distribution areas of terres-
trial arctic birds and mammals are usually continuous, especially on a given conti-
nent, although each species will occupy only a particular habitat within that large
region. Lapland Long-spur, for example, is found in both North America and Eur-
asia but not over the oceans. In contrast, marine birds and mammals, such as Arc-
tic Skua, Black Guillemot, and ringed seal, are found on both the oceans and
continents, with seals being limited to coastal areas. A continuum exists between
the terms disjunct and disperse, with disjunct referring to broad distribution areas
that are widely separated, such as caribou in Canada and Alaska and reindeer in
25
Fennoscandia. Disperse distributions refer to widely scattered areas, such as mar-
mots or pikas limited to mountain tops in mid-latitudes.
Alpine regions of the high plateaus of Central Asia have the largest variety of ani-
mals. Because this is the richest tundra, it is believed to be the origin of animal life
now found in the Arctic. During the Pleistocene, while most of North America and
Europe was covered in ice, northeastern Siberia was not because the climate was too
dry to accumulate enough snow. Animals thrived in this cold, but ice-free zone. As
the Bering Strait land bridge emerged due to dropping sea level during the Pleisto-
cene, several cold-adapted animals moved across to Alaska, where they could then
spread throughout North America. Along with extinct animals, such as wooly mam-
moth and sabre-tooth cats, North America gained several modern species, including
moose, lemming, hare, muskox, fox, and caribou from Eurasia.
Adaptations to Cold
Morphological or physiological adaptations. Tundra animals must be adapted to a
short season of plant growth and to a cold winter too long for hibernation. The
most obvious adaptation in these cold environments is the need to regulate body
temperature by maintaining a balance between heat loss and heat production. Most
animals have a small surface-to-volume ratio, which minimizes heat loss, as do
short appendages such as ears and limbs. Arctic fox, for example, can tolerate tem-
peratures of 58 F (50 C). Many animals, such as muskox and brown bear,
have a thick layer of fat beneath the skin that serves as insulation and stored
energy. Mammals often have a double coat of fur, one of which is a dense and fine
undercoat for insulation. The amount of heat lost depends on the steepness of the
temperature gradient between the animals warm
body and the cold air. Warm skin will radiate and
conduct heat quickly, but an insulating layer of
fur or feathers will lower the temperature gradi-
ent, allowing the animal to retain more heat. Fur
and feathers along with air trapped in the pelt are
poor conductors of heat, so with no wind to dis-
turb the insulating layer, the animal stays warm.
Because an efficient winter coat would trap too
much heat in summer, many birds and mammals
undergo a seasonal change. Mammals like
muskox have longer or thicker fur in winter. Bird
feathers do not change in terms of thickness of
plumage, but the structure of the feathers traps
more heat in winter. Muscular contractions that
fluff up feathers increase the thickness of insula-
tion, but thick fur on mammals cannot be so eas-
ily fluffed.
.................................................
Ptarmigan Camouflage
Seasonal color changes help camouflage both
birds and animals. Along with the weasel
(called ermine in winter), arctic fox, and arctic
hare, the ptarmigam changes to white in win-
ter (see Plate III). The three species of ptarmi-
gan have three seasonal plumages, changing
not only color but also thermal properties from
winter, to spring, to autumn. The loss of pig-
ment in the white feathers leaves air spaces
that trap heat. The birds legs are feathered for
warmth, and feathers around their feet in win-
ter act as snowshoes. Because the female sits
on the nest first, she is also the first to change
to speckled camouflage plumage in spring.
.................................................
26
Animals may control the amount of metabolic heat their bodies produce by
increased activity such as running, digging, or shivering. Ptarmigan feathers, how-
ever, provide such good insulation that the birds have no need to shiver. Compar-
ing the same thickness, feathers are better than fur for insulation, so mammals
must be larger to produce enough metabolic activity to keep warm. Arctic fox and
arctic hare are the smallest animals to survive with just their basal metabolism, but
they are 10 times the weight of ptarmigan. Some birds go into torpor by reducing
their body temperature at night, which in turn decreases their need for energy by
2550% of normal.
Many body extremities have either no insulation or are covered with thin hair.
Examples include nostrils, toe pads of arctic foxes, soles of brown bear and polar
bear feet, hooves of caribou and bighorn sheep, and feet and legs of most birds. To
avoid losing heat through these areas, a physiological adaptation controls the volume
and temperature of blood flow. Constriction of blood vessels limits flow and allows
these extremities to become cooler (see Figure 1.6). An exchange of heat also takes
place from arterial blood to venous blood. Blood in arteries moving toward extrem-
ities transfers heat to veins transporting blood to the body core. Therefore, heat is
retained in the body and cooled blood is sent to the extremities. Gulls feet, for exam-
ple, can be close to freezing with no harm, thereby lowering the temperature gradient
between the animal and the icy surface and limiting heat loss.
During the warmer summer or periods of increased metabolic activity such as
running, animals may need to expel extra heat. Blood vessels then dilate and the inter-
nal heat exchange process is bypassed so excess heat can be taken to and lost through
the extremities. Another major means is by panting, used by canines and even by
ptarmigan. Evaporative cooling through sweat or saliva on the fur or feathers is not
used because the moisture would interfere with the insulating qualities of the pelt.
Figure 1.6 A system of heat exchange between arteries and veins helps animals in arc-
tic or alpine environments maintain body temperatures. (Illustration by Jeff Dixon.)
27
Behavioral adaptations. Some animals depend on behavior to keep warm. Small
animals have neither a thick fur coat nor a high enough metabolism and must
avoid exposure to cold conditions. In true hibernation, body temperature decreases
until it approximates that of the shelter, and with little to no temperature gradient,
the animal loses little to no heat or energy. Several marmots and ground squirrels
in alpine tundra hibernate, but only one arctic species does, the arctic ground squir-
rel in eastern Siberia and western North America. Arctic environments, especially
with permafrost, offer few places to find a shelter that will not freeze, and the arctic
ground squirrel is restricted to well-drained sandy soils well covered by snow. The
burrow entrance is below the level of the chamber, so heat is trapped as it rises from
sleeping bodies. Because the soil is well drained, burrows and sleeping chambers
are dry with no snow or ice to steal heat for either melting or evaporation. As a
result, temperatures do not drop below 10 F (12 C) regardless of outside condi-
tions. Although not true hibernators, bears reduce their metabolism as they den up
for winter. Some mammals, such as chipmunks and hamsters, are torpid but wake
up periodically to eat. Others, such as deer mice and pikas, are active all winter
and cannot become torpid.
Many rodents, such as lemmings, avoid winter cold by maintaining active life
in tunnels beneath the snow where they continue to feed on roots, stems, and buds.
Animals that do not hibernate must have food to carry them through the winter.
Pikas, deer mice, wood rats, Old World hamsters, and narrow-skulled voles store
food, not fat, for the winter. They live in insulated nests on or above the ground or
sometimes in shallow burrows. Piles of grass left to dry on rocks by pikas before
storage are common sights in North American mountains. Lemmings do not store
food but continue normal foraging behavior beneath an insulating layer of snow
where temperatures are not much lower than freezing and nests are lined with grass
and old fur.
Arctic and alpine animals have similar adaptations to cold although they may
not be taxonomically related. Cold steppe and desert animals in the mid-latitudes
have many of the same adaptations to cold winters, and it would have been rela-
tively easy for them to adapt to alpine conditions.
Role of Snow Cover
Snow cover can be either a benefit or detriment depending on the depth and the
animal. Many small animals need an insulating snow cover in winter, with each
species (such as voles, lemmings, deer mice, and pikas) finding it in a preferred hab-
itat (such as depressions, krummholz, or patterned ground). Snow-free areas are
uninhabited. Distributions and behavior of large animals are also related to snow
cover. Snow in the Arctic is reworked by wind into mobile drifts or a crusty cover.
While drifting snow may uncover vegetation, herbivores most often must dig
through snow to forage. If snow is too crusty, caribou migrate to softer snow at the
edge of the forest, but in areas of soft snow, animals do not need to migrate unless
it is too deep for easy movement. Others herbivores such as mountain goats in
28
mid-latitude alpine environments browse on subalpine trees and krummholz
branches that project above the snow cover. Mountain sheep either paw through
the snow or graze on ridges that are blown free of snow. Ptarmigan, one of the few
winter bird residents, can burrow into soft snow or migrate to taller willows for
food or shelter. Most predatory birds migrate, but Snowy Owls hunt where snow is
thin or at rodent air holes through the snow. Mammal predators such as wolves
migrate with the prey, but arctic fox and least weasel remain on the tundra to hunt
the occasional rodent.
Reproduction
Because most mammals, unlike birds, in both the arctic and alpine environments
are nonmigratory, or only travel a short distance like caribou to the forest edges,
the short growing season plays only an indirect role in reproduction. Reproduction
is more closely controlled by quantity and quality of food resources. Large animals
with long gestation periods breed only once a year, and time of birth is correlated
with the season with maximum forage. Small mammals such as rodents with short
gestation periods have more than one litter during the short summer breeding sea-
son. Both the number and size of litters are related to quality of diet, which is also
correlated with length of growing season. Some small animals, such as brown lem-
mings in arctic tundra, which breed under snow all winter, are a major exception.
Such breeding does not take place every year but the reason why is as yet
unknown. The montane vole in alpine areas breeds beneath snow in the early
spring as well as in summer, but probably does not do so all winter.
Birds
Because of the vast expanse of permafrost and boggy ground in the Arctic, water
birds and shore birds (ducks, geese, swans, plovers, and sandpipers) are numerous.
No such habitat exists in alpine environments, either mid-latitude or tropical.
In both arctic and alpine environments, many migrant breeding birds make the
most of the short summer season by arriving on the arctic tundra or alpine habitat
already paired. In other species, such as the Pectoral Sandpiper in the Arctic, court-
ship time is shortened. In the alpine zone, Rosy Finches and White-tailed Ptarmi-
gan pair after arrival, but Water Pipits may already have mates. Unlike birds in
climates with a longer summer season, pairs attempt to nest only once, but clutches
are usually larger than birds of more temperate climates. For most passerines in
both arctic and alpine environments, the female incubates the eggs but both male
and female feed the young. For other birds, males, females, or both may share
duties of caring for eggs and chicks. Most birds undergo molting after breeding sea-
son is over, avoiding a double expenditure of energy.
Invertebrates
Adaptations of insects to alpine environments have parallels in the Arctic and Ant-
arctic. Both morphological and behavioral adaptations enable them, like plants, to
29
survive cold winters but grow and reproduce in short, cool summers. Insects have a
variety of ways of surviving long cold winters that last eight to nine months in the
Arctic and northern mountains. Some live in or adjacent to unfrozen lakes and riv-
ers where the temperature is around freezing and snow cover protects them from
cold. Many rely on supercooling, meaning that the temperature at which they
freeze is decreased. An insects freezing point of 17.5 F (8 C) in the summer
may adjust as temperatures drop in the fall, reaching 30 F (35 C) in mid-winter.
The process reverses as temperatures warm in the spring. Other insects are tolerant
of freezing, can survive frozen ice in their tissues, and are able to survive tempera-
tures 23 to 30 F (5 to 35 C), depending on the species, in a frozen state.
When the upper soil layer freezes to an ice cover in winter, insects enveloped in the
ice become oxygen deprived but can survive several weeks in an anaerobic state.
This mechanism is used by insects living in ponds or lakes that freeze solid.
Morphological adaptations include small size, making it easier for those insects
to find enough food and shelter. The Colias species of butterflies are largest in the
subtropics and smallest in the Rocky Mountains, Himalayas, and Arctic. Pierine
butterflies in the Andes are also small with smaller wings than lowland species.
They hide in vegetation on the Altiplano to escape low temperatures and windy
conditions. The percentage of species having small wings, or none, increases with
elevation. About 60% of insects above 13,200 ft (4,000 m) in the Himalayas have
small wings, and several are unable to fly at all, compared with lowland counter-
parts with normal wings. Mount Kilimanjaro in tropical Africa has a similar insect
component. Small wings reduce the risk of being carried off by wind. More dark
colored insects, which absorb more heat energy, occur at higher elevations. Red-
brown is better than black in raising temperature because it more efficiently absorbs
infrared radiation, while also protecting from excessive ultraviolet radiation. Al-
pine butterflies typically have darker bodies and wings than lowland relatives,
because they need to absorb more heat energy in order to fly. Similar patterns of
temperature regulation are found in alpine blowflies in the Eastern Pamir Moun-
tains and grasshoppers in the North American Rockies.
Predatory insects such as carabid beetles and some spiders that are insensitive
to cold are nocturnal, preying at night on insects immobilized by low temperatures.
In dry environments, insects avoid desiccation by sheltering in crevices and vegeta-
tion. Some grasshoppers in the Andes and Rockies, however, are equivalent to de-
sert insects in resistance to aridity and desiccation. Similar to how plants may need
several seasons to develop flowers and buds, some insects take two to four years to
complete their life cycles because summer is short. Overwintering can take place at
different stages in development. Conversely, some insects, perhaps because of
more productive habitat, complete their life cycle in just one year. Several types
overwinter as adults, ensuring survival of the species.
Small animals like spiders can be found above the limit of plant growth, living
on wind-blown debris in the aeolian zone. Bacteria can also survive if wind-blown
organic matter is available.
30
Amphibians and reptiles are rare in both arctic and alpine environments. In
North America, the wood frog is the only amphibian even on the fringes of the Arc-
tic, while only a few toads and the sagebrush lizard can be found in mid-latitude
mountains. The Old World has few to none of these species in the Arctic, but more
occur in Palearctic alpine environments.
Human Impact
Whether arctic or alpine, tundra vegetation is easily damaged, and because of its
very slow growth, it is also slow to recover. Arctic areas are threatened by oil and
mineral exploration. Resulting development disrupts animal migration routes, and
oil and chemical spills can be devastating. Permafrost presents problems for con-
struction. Eurasian tundra is further degraded by overgrazing by domestic reindeer.
Much of Antarcticas ice-free land is used for research stations with buildings, air-
fields, garbage, trampling, and noise. In some countries, alpine regions are sub-
jected to heavy grazing, and woody shrubs at the edges of the alpine zone are often
burned in an attempt to create more pasture. In some areas, it is hard to determine
what the natural vegetation is. The rising popularity of ecotourism may be benefi-
cial in preservation of both arctic and alpine ecosystems, but it requires careful
management unless increased numbers of visitors disturb the natural landscape the
travelers have come to see.
Further Readings
Benders-Hyde, Elizabeth M. n.d. http://www.blueplanetbiomes.org.
Encyclopedia of Earth. n.d. http://www.eoearth.org.
Palomar College. n.d. Waynes Word. http://waynesword.palomar.edu/index.htm.
Weather Base. n.d. http://www.weatherbase.com.
World Wildlife Fund. n.d. http://www.worldwildlife.org/wildworld/profiles/terrestrial_na
.html.
31
2
Arctic and Antarctic Tundra
Tundra refers to several types of vegetation of different low-stature growthforms,
including tall shrubs (616 ft, 25 m), dwarf-shrub heath (28 in, 520 cm), and
graminoids and cryptogams (both 420 in, 1050 cm). Plants, including vascular
plants, lichens, and mosses, may cover 80100% of the ground. The term polar
desert (or semidesert) is used to describe extremely barren tundra. Vegetation
includes dwarf shrubs, sedges, perennial forbs, mosses, and lichens. Tundra
environments are located where climate conditions are too severe for growth of
trees.
Polar Comparisons
Although one might expect both northern and southern polar regions to be similar
in terms of climate and environmental conditions, several major differences exist.
The Arctic is an ocean almost surrounded by large landmasses, while the Antarctic
is a high, ice-covered continent isolated from other continents by cold water which
is covered by ice in winter. Sea ice in the Arctic is reduced by one-half in the
summer, but by only one-seventh in the Antarctic. Northern Hemisphere tundra
lies generally north of 60 N. While the North Polar ice cap thaws in summer and
the land warms, allowing plant growth to a high latitude, the ice cap in Antarctica
depresses summer temperatures, creating tundra conditions at lower latitudes, to
50 S in some areas. Temperatures on land fringing the Arctic rise well above freez-
ing in the summer, supporting more than 400 kinds of flowering plants. Most
of the ice-free zone of Antarctica is too dry and cold for vegetation, and life is
33
limited to small patches of ice-free ground. Only two species of flowering plants
are known; vegetation is primarily cryptogams. The Arctic has a variety of mam-
mal and bird life. The Antarctic has several sea birds, but no land mammals. Per-
mafrost is characteristic of the Arctic but is rare in the Antarctic because there is
little moisture. The lack of trees in the Antarctic renders the concept of treeline
irrelevant.
Permafrost
Permafrost is ground that is permanently frozen. Widespread in the Arctic, perma-
frost covers half of both Canada and Russia. Beneath the boreal forest south of the
tundra, permafrost is usually discontinuous or sporadic, depending on local condi-
tions and insulating effects of vegetation. Poleward of treeline, permafrost is gener-
ally continuous, underlying the entire landscape with the exception of large bodies
of water. The thickness of permafrost, which can extend down more than 2,000 ft
(600 m), depends on several factors, including mean annual temperature, type
of soil and rock, proximity to ocean, and topography. Permafrost can be wetin
which case abundant water forms ice bodiesor dryin which case water in
gravel, soil, or solid rock is limited. In either case, because of the cold climate, too
little heat is available during the summer to completely thaw the soil.
Summer heat only thaws the surface part of the soil, below which the ground
remains permanently frozen. The thawed surface, called the active layer, is often
waterlogged because the permafrost below impedes drainage. Depth of the active
layer varies but is generally 824 in (2060 cm), except along rivers where it can be
6 ft (2 m) or more. On even slight slopes, the active layer may slowly flow, only
0.52 in (15 cm) per year, in a process called solifluction. Permafrost and solifluc-
tion exert both a negative and positive influence on tundra plants and animals.
While continual disruption of surface soil creates difficulties for small seedlings,
differences in topography or texture provide a variety of microhabitats.
Permafrost creates specific features on the landscape collectively called pat-
terned ground, which can be circles, polygons, nets, hummocks, steps, or stripes
(see Figure 2.1). Circles, nets, and polygons form on level ground. On steeper
slopes, the patterns may be elongated until they become stripes. Although the par-
ticles making up the patterned ground may be sorted or unsorted, resulting in either
a border of stones or no border at all, most patterns are caused by differential freez-
ing of the active layer. Because they hold more water, fine-textured soils contract
and expand with temperature changes more than coarse-textured soils do. As the
soil freezes, any intermixed larger particles are pushed either upward or outward
from a center. This process is most important in sorted patterns. Desiccation crack-
ing, similar to how mud shrinks as it dries and cracks into polygons, is more signifi-
cant in creating unsorted patterns. On a smaller scale, needle ice, which disrupts
34
soil surfaces, occurs on moist surface soils when
night temperatures drop below freezing. Several
explanations for formation of patterns have been
proposed, and different processes may produce
similar results.
Usually small (15 in [35 cm] in diameter),
circles may be outlined by coarse stones or by
plants at the more stable edges. The centers where
frost-heaving disturbs the soil are usually bare.
Polygons can be various sizes. Desiccation crack-
ing in summer creates polygons only 8 in (20 cm)
in diameter, but polygons larger than 3.3 ft (1 m)
are common and usually have ice wedges at their
borders. Fine-textured soil between polygons may
shrink and crack from either dryness or cold. The
cracks collect water that when frozen enlarges the
crack, exaggerating the polygon shape. Depend-
ing on the processes involved, either the edges or
centers can be lower, and sometimes a lower cen-
ter contains a pond in the summer. Frost-heaving
may push rocks to the surface in the middle of the
polygon. Larger rocks slowly move downslope
away from higher centers and collect at the edges,
creating a sorted polygon. Large polygons may be
up to 100 ft (30 m) in diameter, with rims 3.3 ft
(1 m) high. Different soil textures and water avail-
ability support different types of vegetation.
Deeper snow accumulates in the troughs between
polygons, providing more insulation and shelter
for lemmings. Interconnected circles or polygons
form a net or mesh pattern on the landscape.
Solifluction may create steps in the landscape
where the active layer overrides more stable soil.
Vegetation often concentrates at the base of the
riser, which may be 15 in (40 cm) high, where
snow accumulates and provides protection in
winter. The tread, which is more exposed to wind,
may be bare or may support cushion plants.
Other major features on tundra landscapes
where water is abundant are hills called pingos. A
horizontal subsurface layer of water freezes into a lens of ice. As it draws more
water from the surrounding soil, the lens expands and pushes the overlying soil
upward, forming a hill with an ice core. Although most are smaller, pingos may be
.................................................
Alaska Pipeline
Permafrost in Alaska posed serious problems
for construction of the pipeline from the north-
ern coast, where oil is extracted, to the south-
ern ports from which it is shipped. The pipeline
was elevated in areas of ice-rich permafrost,
because a buried pipeline of warm oil would
thaw frozen ground. Heat conducted down
support columns of an elevated system, how-
ever, would also melt the permafrost, dislodge
the columns, and disrupt the pipeline. The
problem was solved by putting ammonia
inside the columns. Evaporation and condensa-
tion processes involve an exchange of heat
energy. Like water, liquid ammonia takes heat
from its surroundings to evaporate, and the
environment becomes cooler. When the am-
monia gas condenses back into liquid, it
releases that heat back into the environment.
Inside the support column tube, liquid ammo-
nia trickles down to well below ground level,
where it absorbs heat from the ground and
evaporates. The ammonia gas then rises inside
the tube to above ground level, where the gas
is chilled by cold arctic air surrounding the col-
umn. As the gas condenses back into liquid
ammonia, it releases the heat taken from the
ground into the atmosphere. The liquid then
trickles back down to rewarm and reevaporate.
In this never-ending cycle, the ammonia takes
excess heat that would thaw the permafrost
layer and transfers it to the air above. Ammonia
has a very low freezing point and remains liq-
uid in spite of temperatures that maintain per-
mafrost. This closed system has successfully
kept the permafrost frozen and the columns
and pipeline in place.
.................................................
35
as much as 150 ft (45 m) high and 1,800 ft (550 m) in diameter at the base. Tops are
relatively warm and well drained and are home to such animals as arctic ground
squirrels and arctic foxes. A breach in the surface soil may let heat penetrate and
melt the ice lens, resulting in a small hill with a pond at its center. More common
in discontinuous permafrost and smaller than a pingo, a palsa is a low hill in peat
or boggy areas that has an ice core.
Climate
Temperatures in the Arctic and Antarctic are primarily controlled by the light re-
gime. Both arctic and antarctic tundra regions experience 24 hours of daylight dur-
ing some part of their respective summers, but the number of days with that
extreme varies with latitude, ranging from one day at the Arctic and Antarctic
Circles (66
1
=
2
N and S) to six months at the poles (90 N and S). At 70 N and S,
the sun is above the horizon continuously for two-and-a-half months and at 75 N
and S for four months. The same figures apply to winter darkness, that is, days
with no sunlight at all. Because of the angle of the suns path, however, long hours
of twilight in winter when the sun is only a few degrees below the horizon are
characteristic. With no incoming solar radiation during the dark winter, tempera-
tures drop. Even the continuous daylight of summer provides little warmth.
Figure 2.1 Frost action in permafrost contributes to the development of patterned
ground, from polygons to stripes, which determines vegetation patterns. (Illustration by
Jeff Dixon. Adapted from Sharp 1938. Photo courtesy of the U.S. Fish and Wildlife Service.)
36
Temperature contrasts between summer and win-
ter may be more related to degree of continental-
ity than they are to latitude: Higher latitude areas
in coastal locations, especially those affected by
warm currents, are warmer in winter than lower-
latitude regions farther inland. Mean temperatures
of the warmest summer month vary considerably,
but generally average between 40 F (4.5 C) and
55 F (13 C). Even though temperatures on a
hot summer day occasionally may reach 70 F
(21 C), the average for the short summer season
is below 50 F (10 C).
For much of the year, the region experiences
a negative energy balance (see sidebar Arctic
Energy Budget). The low angle of the sun in the
sky imparts little energy to the arctic landscape.
The high albedo of ice and snow means that
much incoming solar radiation is reflected and
sent back to space. Infrared energy continues to
radiate from both land and sea surfaces all year,
even during winter darkness. The result is a con-
tinual energy loss most of the year. The energy
balance does not become positive until after
snow melts, usually in June, even though incom-
ing solar radiation increases from the time of the
spring equinox. This makes the summer growing
season even shorter because about one-half of
the summers solar radiation comes before snow
melts and is lost either by reflection off snow cover
or is used as energy to melt snow rather than warm
the air andplants. After snowmelt, albedo decreases
abruptly and photosynthesis takes place continu-
ously during the 24 hours of sunlight.
Cold winter temperatures in arctic tundra rarely have a direct effect on plant
and animal life because most organisms are adapted to extremes of cold, dormant
during the winter, or protected under an insulating cover of snow. Some animals
migrate to escape extreme cold or lack of food. Because the tundra environment is
on the fringes of the continents, winter temperatures are not as extreme as they are
in lower-latitude boreal forest climates where many animals and birds live and
hibernate, including some that migrate from the tundra. Because boreal forest trees
lack a protective snow cover and are exposed to the weather, they must be more
cold tolerant than tundra plants. The most significant factors affecting life in the
Arctic are cool summer temperatures and the short time period during which
.................................................
Arctic Energy Budget
Temperature of Earths atmosphere is deter-
mined by the energy budget, basically a com-
parison of how much energy enters Earths
atmospheric system and how much energy
leaves. In spring and summer when the sun
is high in the sky and days become long,
more energy reaches Earth than the amount
that exits as infrared radiation. The surplus of
energy causes temperatures to rise. In fall and
winter when the sun drops lower in the sky
and days become shorter, less energy comes
in. Infrared energy is always being radiated
back to space, however, and when the out-
going amount exceeds what comes in, there is
a deficit of energy and temperatures begin to
drop. Reflection, evaporation, and melting of
snow complicate this pattern. Incoming solar
radiation striking a white snowy surface will
reflect back into space and not enter Earths
energy system at all. This phenomenon is
called albedo. A sunburn on the bottom of
your chin is from solar radiation reflected off
snow or water. Incoming radiation may also be
used to melt snow or evaporate water, and
thus cannot be used to warm the air. That is
why it is cooler around a swimming pool in the
summer.
.................................................
37
temperatures remain above freezing. The short summer and lack of heat for active
growth also affect chemical activity and food sources, indirectly limiting life
as well.
Arctic precipitation is generally low because cold air cannot hold much water
vapor. Coastal regions along warm currents, such as Iceland, Norway, and southern
Alaska, are wetter, with up to 30 in (750 mm) or more precipitation a year, but the
cold airmasses more common to the Arctic produce 10 in (250 mm) or less. Avail-
ability of moisture for plants, however, also depends on substrate and wind condi-
tions. Wind often redistributes sparse snow cover, leaving some regions quite dry.
Sublimation of frozen water into dry air also robs soil of needed moisture by elimi-
nating snowmelt. Regions underlain by extensive permafrost, however, are boggy
because water that cannot percolate into the ground remains in the surface soil.
Circumpolar Climate Variation
Temperature. Although the climate is cold, temperature and precipitation vary
according to latitude, airmasses, wind direction, coastal or continental location,
mountains or valleys, and protected inlets or exposed coasts (see Figure 2.2).
Because the sun angle is low, even small topographic irregularities cause differen-
ces in microclimate, such as sunny or shady slopes. The following transect
describes changing climate conditions eastward around the Arctic region. Iceland
and the northeast Atlantic all the way north to Svalbard and east to Murmansk,
Russia, are influenced by the warm North Atlantic Drift current and are mild for
the latitude. Eastward to the Ural Mountains in Russia, continental influence
increases. Winters are colder, and the summer frost-free period is shorter. East of
the Ural Mountains in northwestern Siberia, winters become even colder. Temper-
atures do not rise above freezing until June and the frost-free period is only 3040
days. Northeastern Siberia between the Taymry Peninsula and the Kolyma River
has the most extreme climate. Cold air drains from adjacent mountains, settling
into valleys and causing severe temperature inversions with cold air underlying
warm air. Summers are also cool.
East of the Kolyma River in northeastern Siberia and into Alaska and the Yukon,
the landscape is characterized by many mountains and intervening valleys. The
mountains are high enough to effectively separate cold Arctic airmasses in the north
from warmer Pacific airmasses to the south. Northern coasts have an unmodified
arctic climate, inner valleys have a continental influence, and Pacific coastal areas
are moderate. In spite of many lakes in the area, the region between the Mackenzie
River and Hudson Bay in the Northwest Territories and western Nunavut is conti-
nental with severe winters. Almost the entire Canadian archipelago, from Banks
Island on the west to Ellesmere Island on the east, is surrounded by thick sea ice
from December to April. The area is persistently cold but is not extreme. The eastern
part of the Canadian Arctic, particularly the Ungava Peninsula in Quebec and Baffin
Island, has more maritime influence because the sea there is not frozen. Cyclonic
38
Figure 2.2 (a) Pangnirtung, Nunavut, Canada, is typical of the Low Arctic, whereas
(b) moderate temperatures and more precipitation at Vard o, Norway, are characteristic of
maritime Arctic. (Illustration by Jeff Dixon.)
storms coming from the south bring warmer airmasses and temperatures near or
above freezing even in winter, especially in the Ungava plateau.
Precipitation. Precipitation varies according to continental or coastal position and
location with respect to wind direction and mountains, but is generally low. Most
precipitation falls as snow, even in summer. Precipitation in Iceland, the northeast
Atlantic, and Russia west of the Urals is high for the Arctic, 1530 in (380750
mm), and decreases toward the east. In northwestern Siberia east of the Ural Moun-
tains, which are a precipitation barrier, annual precipitation drops to about 10 in
(250 mm). The cold air continues to be dry eastward to the Kolyma River. Although
winter air is extremely dry, ice fog over cities is frequent in December and January.
Moisture generated by urban areas quickly freezes into the cold air. Precipitation in
northeastern Siberia and adjacent Alaska varies because of topography. The north
coast, influenced by Arctic airmasses, is fairly dry. Interior mountains have upslope
precipitation, and south-facing mountains have extensive snowfall. Anchorage,
Alaska, gets 5 ft (1.5 m) of winter snow, the equivalent of 6 in (150 mm) of rain.
Because of the dominance of high pressure and absence of cyclonic storms, lit-
tle precipitation falls in northwestern Canada and the Canadian islands during the
winter months, although frequently blowing snow reduces visibility. Cyclonic
storms do bring up to 2 in (50 mm) of rain in the short summer, and increased
moisture from melting snow and ice causes frequent fog. Higher rainfall in the east-
ern Canadian Arctic is caused by mountainous topography and paths of cyclonic
storms from the Great Lakes or Atlantic coast. Precipitation primarily falls in
summer and is variable, depending on storm tracks.
Strength and occurrence of wind depends on topography and frequency of cy-
clonic storms. It is not exceptionally windy in the Arctic, but wind is more noticea-
ble because low vegetation does not block it. Infrequently a warm winter wind will
melt snow, which then recrystallizes into a hard crust, preventing animals like
muskoxen and lemmings from accessing food. Except for steep slopes, wind-
exposed ridges, and among the tallest shrubs along rivers, most of the Arctic is
snow-covered from late August to May or June, or even early July.
Tundra Soils
Soil development is slow and limited because of little plant cover, a short growing
season, permafrost, and the short time land has been free of glacial ice. Soils are
thin, young, and nutrient poor. The cold and short growing season inhibits both
chemical weathering of mineral material and biologic activity. Sparse vegetation
contributes little organic matter and nutrients. Clear soil profiles rarely develop
because frost-heaving and solifluction continually overturn soil layers. Soil classi-
fication systems for the Arctic are varied but generally mirror patterns of vegeta-
tion. Minimally developed spodosols (podzols) exist on well-drained soils with a
deep active layer dominated by dwarf-shrub heath and dwarf birch. Inceptisols
(Arctic brown soils) occur on uplands and dry ridges where cushions and heath
shrubs grow. Other inceptisols (tundra soils), where the dominant process is
40
gleization, are common where drainage is impeded. They support cottongrass,
tussock grasses, dwarf-shrub heath, and some sedges. Histosols (bog and half-bog
soils), which also develop through gleization, occur along shores of lakes and
ponds where drainage is better and peat accumulates from sedges and mosses. On
drier and better-drained areas, a soil profile begins to develop, with an iron and
clay layer at the surface. In polar deserts, calcium carbonate and magnesium fre-
quently accumulate on undersides of rocks as the surface dries in summer, espe-
cially in areas of sedimentary rocks and recently uplifted marine sediments. In
contrast to most tundra soils, which are acidic, these polar desert soils are basic,
mainly because they lack nitrogen and phosphorus.
Tundra Vegetation
Arctic tundra plant life is dominated by dwarf shrubs, mosses, lichens, sedges, and
perennial forbs. The region was still covered by ice only 8,00015,000 years ago.
The circumpolar Arctic is divided into two major zones according to latitude, cli-
mate, and vegetation. Covering the largest area in lower latitudes, the Low Arctic
has the most variety of growthforms. The active layer is deep on rolling upland,
allowing good drainage and root penetration. Vegetation cover is 80100%, with
less on rocky continental shield areas. The general appearance of the landscape is
that of a grassland with cottongrass tussocks and birch and willow shrubs up to 5 ft
(1.5 m) tall (see Figure 2.3). Subshrubs, mostly heaths, are shorter, less than 8 in
Figure 2.3 The tundra near Churchill, Manitoba, in Canada is grassy with stunted
trees and low shrubs. (Photo by author.)
41
(20 cm) high. Fruticose lichens such as reindeer moss and snow lichen grow in dry
sites and mosses such as haircap grow where it is wet.
While Low Arctic vegetation is commonly called tundra, the higher latitudes of
the High Arctic are often described as polar semidesert and polar desert. The High
Arctic is mostly bare rock, angular from frost action; plants are herbaceous. Most
of the area is polar semidesert, covered by lichens and mosses. Flowering rosettes,
cushions, and mat-forming forbs, mostly saxifrages, are only found in protected
sites and are no more than 3 in (8 cm) high. In true polar deserts, even cryptogams
are sparse, and total cover is 03%.
On a large scale, the circumpolar tundra zone has similar plant communities;
however, regional and local differences occur.
Tundra Animals
Large Mammals
Caribou and reindeer. North American caribou and European reindeer, the same
species, are the most common large mammal (see Figure 2.4). They spend winters
at the forest edge and summers on the tundra. Bulls average 350 lb (160 kg);
females are smaller. Unlike other deer, both sexes have antlers, and except for preg-
nant females, both genders shed them in fall. Pregnant females keep their antlers
until late spring when they calve, using them as deterrent to keep others away from
foraging areas scraped free of snow. Calving grounds are on the tundra, which is of-
ten still snow-covered in May when births occur. Calves are precocious and can
run soon after being born. After a week of nursing their mothers rich milk, they
can forage on their own.
Large, concave hoofs give support in deep snow or marshy land, and a thick
coat with hollow hairs provides insulation. The caribou winter diet consists of
starchy lichens rich in carbohydrates, which gives them energy to survive the cold.
During spring and summer, they eat high-protein shoots of sedges, grasses, dwarf
willows, and birches. Mouth muscles sort living from dead plant tissue; indigestible
parts are spit out. To avoid overgrazing one area, the animals continually migrate
in large herds of thousands, trampling the landscape along regular routes up to 500
miles (800 km) long. Wild herds in the tens of thousands still exist in America,
where their spectacular migrations create conflict between conservation and devel-
opment. Most European reindeer were domesticated 3,000 years ago, although a
few wild animals still remain in Fennoscandia and Russia.
Muskoxen. A fixture in the high north for thousands of years, muskoxen once
shared the tundra with mastodons and mammoths (see Plate IV). They are big ani-
mals, up to 900 lb (408 kg) and 7 ft (2 m) long, but with short legs. Both sexes have
horns. Glands between the bulls eyes produce the musky odor for which they are
42
named. Long brown or black hair almost touch-
ing the ground combined with a soft, wooly
undercoat provide superb insulation. Lying down,
the animal does not even lose enough heat to melt
the snow and ice beneath it. Muskoxen live on the
tundra all year, eating a variety of grasses and
shrubs. In summer, herds consist of about 10 ani-
mals; they increase to 1520 in winter. One male
dominates each herd, breeding with the females,
which each bear one calf in spring. The calf
matures fast, being able to graze within a week,
but continues to take its mothers milk for a year.
The animals natural method of defense, backing
into a circle with horns extended outward, suc-
cessfully protects calves in the center from arctic
wolves, but made them easy prey for rifle-wield-
ing humans. Populations once on the verge of
extinction have been brought back through con-
servation efforts. Unlike caribou, they do not
undertake long migrations, but spend winter on
higher ground where winds blow vegetation free
of snow.
Brown bears. Brown bears, grizzly, Kodiak, and
Kamchatka bears are all the same species regard-
less of geographic location, although scientists
recognize several subspecies. A mature male can
weigh 900 lb (400 kg) and stand 9 ft (3 m) tall on
its hind feet, but females are smaller. Found in
both forest and tundra habitats throughout the
Northern Hemisphere, they may be brown,
cream-colored, or black. Brown bears are distin-
guished from their close relative the black bear by their dished face and a promi-
nent hump on their shoulders. Long, curved claws and muscles in the hump are
adaptations to digging out roots, insects in decaying wood, or small mammal bur-
rows. Best known for gorging on migrating salmon in Alaska, brown bears are
not picky eaters. Although classified as carnivores, they are omnivorous, eating
plants, fruit, insects, birds, rodents, moose, caribou, and carrion, a varied taste
that lures them to human garbage dumps in populated areas. Sedges, grasses, and
roots are tasty treats on the tundra. Because of their size and food requirements,
they are solitary except for mothers and cubs. Although not true hibernators, they
lower their body temperature and metabolism in winter and go into torpor.
.................................................
Caribou Parasites
Caribou have few natural enemies. Golden
Eagles and bears prey on calves, while packs of
wolves can overpower adults. When tempera-
tures rise to 50 F (10 C), swarms of summer
insects in clouds of millions plague the animals
and interfere with feeding. Experiments have
shown that mosquitoes can extract up to 4 oz
(115 g) of blood from one animal in a single
day. They try to escape by twitching, running,
occupying windy locations, clustering together,
or immersing themselves in snow or water. The
two worst insects plaguing caribou are warble
flies and nose bot flies. Warble flies lay eggs on
the caribous leg hairs. After burrowing into the
skin, larvae travel beneath the surface to the
animals back, where they cut air holes through
the skin to the surface. They grow to about an
inch long beneath the animals hide. When de-
velopment finishes, the larvae pop through the
skin and pupate on the tundra into adult flies.
Female nose bot flies lay their eggs in the cari-
bous nostrils, from which hatched larvae move
to the throat, to take advantage of the warm,
moist environment found there throughout
the winter. In March, the larvae grow quickly
and are eventually coughed out by the caribou
to pupate into adults on the tundra.
.................................................
43
Polar bears. As tall as 10 ft (3 m) and weighing up to 1,700 lb (770 kg), adult male
polar bears are the largest members of the bear family. Females are smaller. Popula-
tion estimates are between 22,000 and 27,000. Their only enemies are humans and
now quite possibly climate change. Polar bears are
at the top of the arctic food chain, spending winter
hunting seals on pack ice by waiting at breathing
holes or stalking them as they rest. They prefer
ringed seals, but also eat bearded seals, and if hun-
gry, will eat reindeer, birds, and beached whales.
A bear can eat 100 lbs (45 kg) of seal blubber at a
single sitting, and if hunting is good, leave the rest
of the meat for scavengers. Preference for fat has an
advantage in that its digestion releases metabolic
water, important in an environment where water is
either salty or frozen.
Polar bears are well adapted to frigid arctic
conditions. An insulating layer of blubber up to
4.5 in (12 cm) thick helps maintain normal body
Figure 2.4 Reindeer, shown here in Lapland in northern Norway, and caribou in
North America are the same species. (Photo by author.)
.................................................
Polar Bear Colors
Although usually appearing to be some shade
of white, polar bear fur has no pigment and
takes on the hue of its surroundings, slightly
yellow-orange from the low-horizon sun or blu-
ish in light filtered through clouds or fog. In
late summer or fall, bears can appear gray
because oils from their prey have stained the
coat. Bears in zoos may have a green tint
because of algae growing in the fur. Polar bear
skin is actually black, and if fur is damaged,
black patches may show through.
.................................................
44
temperature of 98.6 F (37 C), the same as a
humans. Under extreme cold or windy condi-
tions, animals curl up in a snowbank for addi-
tional protection, covering their uninsulated
muzzle with a furry paw. Their bulk, with small
ears and tail, helps to conserve heat. Two different
layers of fur provide two layers of insulation. The
undercoat is dense fur, and longer guard hairs are
hollow. With all this insulation, bears can become
too warm, especially when running, and shed
excess fur for summer. Feet are covered with
small bumps, and sometimes hair, giving them
traction on icy surfaces. Black claws are sharply
curved for both clinging to ice and catching seals.
Because of harsh conditions, reproduction is
slow. An adult female may have only five litters in
her 18-year life span. Although they can create a
den on drifting pack ice, pregnant females usually
dig snow dens on coastal land in fall. They give
birth during the winter, not eating, drinking, or def-
ecating until spring when the new family leaves the
den. Normally, the litter is two cubs. Like all bear
cubs, they are small, 12 in (30 cm) and 1 lb (0.5 kg),
blind, helpless, and totally dependent on mothers
milk, which has 31% fat content. By April, they can
weigh 50 lb (22 kg). Cubs remain with their mother
for two-and-a-half years while they learn to hunt
and survive. Neither males nor females hibernate.
Their circumpolar distribution is limited to
ocean areas frozen into sea ice most of the year
and adjacent land in North America and Eurasia.
Because they hunt seals, they prefer shallow water
near the shore or the edge of the ice pack that does not freeze too deeply. Although
classified as marine mammals because of preferred habitat on sea ice, most live on
the tundra in summer while others remain on ice floes all year. Polar bears need
sea ice on which to hunt, and without it they have no access to their prey, seals. As
such, they are vulnerable to climatic warming.
Small Mammals
Small rodents are plentiful. Many remain active in winter but need an insulating
layer of snow. Lemmings and voles survive under the snow, where they are hunted
by weasels. The same areas that have an insulating snow cover, however, are most
subject to spring flooding. Lemmings that fail to move their nests to higher ground
.................................................
Polar Bear Jail
Churchill, Manitoba, is near the site where
waters of Hudson Bay freeze first because of
the fresh water outlet from the Churchill River.
Bears migrate from other parts of Canada to
get on the ice first near Churchill. They have
fasted all summer, living off accumulated fat,
and by early to mid-November, they are hungry
and ready for seals. Although dozens of bears
may congregate near Churchill, the population
can disappear overnight when ice conditions
are right. Bears initially gathered at the town
dump where garbage was available, but they
began to cause trouble. A Polar Bear Alert pro-
gram now uses noise primarily to scare bears
away from town. If a bear continues to be a
problem, especially if a mother is teaching her
cubs bad habits, the animal is humanely
trapped and taken to polar bear jail, a hangar-
like facility away from town. When the Bay
freezes, they are released. If the facility is full,
helicopters airlift the bears in mesh cargo nets
and deposit them in remote locations, with the
hope that they will not return. Adults are
sedated to keep them calm, and their eyes are
coated with petroleum jelly for protection from
cold, dry air during the airlift. Small cubs too
delicate for the net are transported inside the
helicopter.
.................................................
45
in spring risk drowning. The tundra vole, which can swim, lives in the wettest
areas. Brown lemmings occupy better-drained polygon ridges, and collared lem-
mings live on well-drained rocky uplands. Narrow-skulled voles are found on the
best-drained sandy soils along streams. Several voles live in shrubby heath, willow,
or alder thickets. No animals especially suited for digging, such as gophers, inhabit
the arctic tundra because permafrost, extremely cold winters, and shallow snow
cover leave no unfrozen layer through which to dig. Arctic hares in Low Arctic
tundra spend the winter eating twigs that extend above the snow cover.
Lemming cycles. Food chains in the tundra are short and simple. Lemming popu-
lation cycles that peak and crash every three to six years are a good example of the
delicate balance in tundra ecosystems. Small mouse-like animals, lemmings are
only 5 in (13 cm) from nose to short tail. Although they have a reputation of com-
mitting mass suicide, they are merely migrating in an attempt to escape overcrowd-
ing. The animals do not hibernate, but remain active beneath snow all winter
eating green shoots of sedges and grasses. They also mate all year, and within two
years, a female can produce up to 14 litters of eight pups each. However, due to
predation and drowning, most lemmings do not live that long. Nonetheless,
because pups double their birth weight in just four days and females can become
pregnant while still nursing, a colony can potentially increase its population 100
times in a single winter. Lemmings must eat twice their weight in food every day,
which has consequences for vegetation and underlying soil. Digestion is inefficient,
and 70% of the intake is deposited as nitrogen-rich waste, which nourishes tundra
vegetation. By cutting dead vegetation to gain access to new, green shoots, lem-
mings aid the decay process. As a result, lemming foraging areas can become lush,
but if too much plant cover is removed, thawing of permafrost leads to denudation.
Lemming population increases lead to more predators. Populations of arctic
fox, weasels, Snowy Owls, and Pomerine Jaegers increase. Wolves, which usually
attack larger prey, will also eat lemmings because they are so abundant. Owls catch
lemmings on the surface, while the slim bodies of weasels allow them to pursue
lemmings in their tunnels. Eventually the lemmings must migrate to find food or
die. Crowding, insufficient food, or less-nutritious food inhibit breeding, and the
lemming population crashes. A corresponding decrease in predator populations
follows. During the bust period, vegetation recovers and permafrost is reestab-
lished. Lemmings in Fennoscandia migrate in the thousands, while elsewhere they
move in smaller groups.
Birds
Many birds, including waterfowl, jaegers, and ptarmigan, are found in the Arctic,
and some, such as Snow Geese and ptarmigan, influence the vegetation by their
grazing activities. Millions of geese, swans, ducks, gulls, and waders live in the tun-
dra in summer. Waterfowl are the first to arrive in spring, followed by shorebirds
and land birds. Birds that nest in the tundra include several species of plover,
46
sandpiper, geese, and ducks. By necessity, birds must nest on the ground, vulnera-
ble to scavengers and predatory mammals. Several birds of prey such as falcons,
hawks, and Snowy Owls share the same nesting areas with prey species. Food
includes insects, fish, and aquatic vegetation. Mosquitoes, craneflies, springtails,
and stoneflies are plentiful because the numerous ponds and bogs are good breed-
ing sites. Many tundra birds change their diets according to season and availability.
They feed on both seeds and insects in spring, concentrate on insects in summer,
and change to seeds in late summer. Food intake may be determined by habitat,
or habitat may be selected according to food preferences. Willow Ptarmigans eat
willow twigs and birch buds and leaves, while the Rock Ptarmigans diet is alder
catkins and buds. Geese eat grass, sometimes the whole plant, leaving the land bare
except for mosses and lichens. With less grass, cyanobacteria, an important nitro-
gen-fixer, grows on the surface. Because bird excrement is rich in nitrogen, surviv-
ing or colonizing vegetation grows well on nesting grounds.
Among the very few passerines, Lapland Longspur, Snow Bunting, wagtails,
and pipits breed on North American and Eurasia tundra. Many birds fly long dis-
tances. Undertaking an exceptional 21,000 mi (33,800 km) migration each year,
the Arctic Tern breeds in arctic tundra but winters in subantarctic seas. Peregrine
Falcon, a raptor, also summers in the tundra. Arctic birds have various ways of car-
ing for young. Either males or females may sit on the nest and feed the chicks while
the other flies away. Some produce two clutches, one each for the male and female
to tend, in the hopes that some might survive. Leaving the breeding grounds before
the end of the summer season may be a way to reduce demands on scarce food
resources.
Although many birds nest on the tundra, few stay all year. Gyrfalcon, the
worlds largest falcon, eats arctic hare and ptarmigan, while ravens survive on car-
rion, droppings, and human garbage. Snowy Owls, which nest and lay eggs before
snow melts in spring, prefer lemmings and other small animals. Ptarmigan congre-
gate on bare land or areas where the snow is thin because they cannot scratch
through deep snow for food.
In spite of continuous light or darkness during arctic summers or winters, both
birds and mammals maintain a daily activity cycle that corresponds to a 24-hour
period. Daily activities in small mammals are controlled by an innate circadian
rhythm rather than cues from changing light levels.
Animal Effects on Vegetation
Activities of mammals and birds can alter plant communities and extensively dam-
age tundra vegetation. Concentrated grazing by reindeer has the biggest impact on
reindeer lichen. Because lichens grow slowly, 0.2 in (6 mm) per year, where winter
grazing is intense, grazing can totally destroy the carpet in three to four years.
It may require five years of other growth, mostly grasses, before lichens begin to
regenerate, and 1520 years for total restoration. However, most wild reindeer
herds are self-regulating. They graze lichens only under extreme cold and
47
maximum snow cover, quickly moving to summer pastures when conditions
allow. Grazing by domesticated herds does much more damage because of their
concentration in limited areas.
Arctic fox has a local but strong influence on vegetation. The animals deep
entry passage into its burrow disturbs permafrost, resulting in a deeper active layer.
Aeration and accumulation of excrement enrich the soil, which then supports a
dense cover of several grasses and forbs. Bean Geese and White-fronted Geese pre-
fer cottongrasses and other sedges, only eating the succulent parts next to the root.
The accumulation of organic matter in stems and bird droppings then influences
vegetation growth through the addition of nutrients. Geese have large feeding areas
and usually do not do much damage. If concentrated, however, geese can devastate
an area, leaving it barren, trampled, and covered in droppings. Birds of prey use
the same spot for years, and effects of altered vegetation beneath roosts are quite
visible in different and taller growths of forbs.
Lemming population fluctuations are large and significant. When lemming
populations peak every three to four years, they overgraze and reduce the vegeta-
tive cover. They normally do not eat the less-nutritious mosses, but still damage
them in winter under the snow cover by gnawing through the moss layer to reach
the succulent parts of grasses and forbs. Since lemmings can destroy local areas
while leaving frozen parts of others intact, they contribute to the hummocky surfa-
ces so common in arctic tundra. Over time, the untouched turf becomes thicker
with shoots, forming a hummock. The less nutritious upper stems that lemmings
cut and discard become dried plant litter, an important source of organic matter in
the soil.
Human Influences
Air pollution, including radioactive fallout, accumulates in the Arctic due to wind
patterns and lack of precipitation to clean the air. Lichens absorb moisture and
nutrients directly from the air, and because of slow growth, accumulate toxic mate-
rials over many years. Radioactive particles introduced into the food chain when
lichen is eaten by reindeer or caribou can be transferred to human populations eat-
ing the meat. Mining, exploration, roads, and other development disrupt the fragile
balance of tundra ecosystems. In spite of that fragility, alteration and destruction
due to human activities is surprisingly limited due to sparse populations. Intro-
duced plants rarely last more than one or two seasons because they are not adapted
to the climate or soils. Vehicle traffic compresses and eventually destroys the mat
of vegetation, and the main dominants of moss, lichens, dwarf shrubs, and shrubs
are usually replaced by grasses and forbs. In such altered areas, the number of plant
species is only one-half to one-third that of natural tundra. Alteration of the perma-
frost by buildings, roads, or change in vegetation cover can be significant. As the
insulating layer is destroyed, more ice melts, resulting in a phenomenon called
48
thermokarst, in which the surface soil collapses into voids vacated by ice cores.
Problems are only intense near the few populated areas in the Arctic.
Global warming is visibly affecting tundra environments. Ponds in the tundra
on Ellesmere Island have been drying up over the last 25 years, and salt content in
the water has increased due to accelerated evaporation. Sea ice and permafrost in
several arctic or antarctic environments are melting at unprecedented rates. Coastal
regions left unprotected by disappearing sea ice are now exposed to erosion, and
seawater encroachments on formerly fresh water lakes threaten wildlife habitat.
Temperature changes in the oceans will affect land. Snow appears to be melting
earlier in the summer in some places, and permafrost temperatures are rising.
Regional Expressions of Arctic Tundra
North America
Tundra occupies about 20% of the North American continent, from 55 N along
Hudson Bay to Alert on Ellesmere Island at 82 03
0
N. Primarily in Canada, it
stretches from 61 W in Newfoundland to 168 W in Alaska (see Figure 2.5). In
this large territory, climate, topography, ice cover, soils, ecosystems, and diversity
of plants and animals all display much variation. The most expansive region is
Low Arctic, the gently sloped uplands that have good drainage and deeper soils to
permafrost. High Arctic tundra is found on the northern Canadian islands, includ-
ing Ellesmere, Baffin, and Queen Elizabeth.
Climate. In winter, cold temperatures over the continent create a high pressure,
which keeps moist airmasses away. With summer warming, the high pressure
Figure 2.5 Arctic tundra in North America and Greenland. (Map by Bernd Kuennecke.)
49
weakens, and the Arctic Frontthe southern limit of cold, dry airroughly coin-
cides with treeline. Cyclonic storms and moist airmasses can then enter the Arctic
and bring summer rain. This seasonal cycle of pressure and storms results in dry
winters and wetter summers. The northwestern Canadian islands, however, are
dominated by cold, dry arctic air all year and are surrounded by sea ice. The south-
eastern Canadian Arctic has a warmer maritime climate with more cyclonic
storms. The Canadian mainland is continental, with a greater seasonal temperature
range and much warmer summers than experienced farther north.
The Brooks Range falls on the border between arctic and boreal forest biomes,
vegetatively defining north and south Alaska. Cyclonic storms are frequent on the
south coast, where the Bering Sea receives high precipitation, but infrequent on the
Arctic coast because mountains block the storms. North Alaska experiences cold
temperatures with deep temperature inversions. Ice fog is a hazard in industrial
areas when temperatures drop below 5 F (15 C). The west coast facing the
Bering Sea is less severe.
In eastern Canada, precipitation is limited except for maritime Baffin Island,
but because most precipitation falls as light rain or mist every few days in summer,
soils rarely dry out except in the High Arctic. Soil moisture is usually high, espe-
cially where permafrost impedes drainage.
Differences in biotic communities between Low Arctic and High Arctic can be
attributed to differences in severity of climate (see Table 2.1). Radiation, mean an-
nual temperature, summer and winter temperatures, length of growing season cor-
responding to time of snowmelt, and precipitation all decrease from south to north.
Even with a longer period of summer daylight at higher latitudes, the lower sun
angle and high reflectivity off snowy surfaces diminishes radiation gain farther
north, resulting in lower temperatures. Mean annual temperatures in the Low Arc-
tic, both in maritime locations and the more continental interiors, are higher than
those in the High Arctic. Both summer and winter temperatures are slightly
warmer in the Low Arctic. Low Arctic receives more precipitation, and snow cover
is generally deeper. Because snow melts sooner, the growing season is one to two
months longer in the Low Arctic. The date of snowmelt is significant in
Table 2.1 North American Arctic Tundra Climate Summary
CLIMATE CHARACTERISTIC LOW ARCTIC HIGH ARCTIC
Mean Annual Temperature 20 F (6.5 C) 10 F (12 C)
January Mean Temperature 10 F (24 C) 22 F (30 C)
July Mean Temperature 50 F (10 C) 40 F (4.5 C)
Annual Precipitation 18 in (450 mm) 6 in (150 mm)
Depth of Snow Cover 14 in (35 cm) 6 in (15 cm)
Growing Season 34 months 1.52.5 months
Date of Snow Melt mid-June late July or August
Depth of Active Layer 880 in (20200 cm) 860 in (20150 cm)
50
determining the growing season. Average depth of the active layer, however, is
similar in both subdivisions.
Vegetation varies according to temperature, aridity, and snow depth, usually
becoming shorter and more sparse with increasing latitude (see Figure 2.6).
Plant communities of the Low Arctic. North of the closed boreal forest, an open
lichen woodland of widely spaced trees marks the beginning of treeline, here a
broad zone more than 100 mi (160 km) wide. Fruticose lichens grow 6 in (15 cm)
thick on treeless ground. Still farther north, the woodland thins to small, scattered
groves in sheltered sites where snow accumulates. Black spruce dominates these
treeline groves in most of Canada but is replaced by white spruce in the Yukon and
Alaska. Low Arctic tundra vegetation fills the nonforested areas between groves of
trees. Although fires are necessary to maintain the open woodland of the treeline
zone, they kill shrubs, mosses, and lichens. Shrubs resprout within 210 years, but
it takes 60200 years for the lichen community to regenerate.
With the exception of Barrow on the Arctic coast, the Low Arctic includes all
of Alaska, mainland Canada, southern Baffin Island, and southern Greenland.
The term tundra refers to Low Arctic vegetation with a complete cover of mainly
woody species, sedges, and grasses, and excludes the cushion-plant vegetation that
is more typical of High Arctic and polar deserts. The landscape is grassy with low
or dwarf shrubs. Shrubs are significant except in wetlands, where they are replaced
by sedges, bryophytes, and grasses.
Figure 2.6 Vegetation decreases in height from treeline to the polar deserts in the far
North. (Illustration by Jeff Dixon.)
51
Scientists recognize several major vegetation types in which shrub height corre-
lates with snow depth. Tall shrub tundra consisting of alders, birch, and willows
about 10 ft (3 m) high, with a dense understory of forbs and grasses, is most com-
mon along river bottoms, near lakes, and other protected areas where warmer soil
contains more nutrients and the active layer is deeper. Species vary slightly accord-
ing to geographic location and subspecies are recognized. The understory can be
nodding hairgrass and matgrass. This vegetation type is important habitat for
moose in northwest Canada and Alaska. Arctic hare and ptarmigan also feed on
the willows.
Low shrub tundra with various combinations of dwarf birch, low willows,
heath, forbs, and graminoids is found primarily on rolling uplands in Alaska and
northwestern Canada beyond the forest tundra zone where snowcover is not as
deep as in protected areas. An open canopy of shrubs averaging 20 in (50 cm) high
is dominated by bog birch and blue willow. Ground cover is made up of clumps of
carex sedge and cottongrass tussocks, and tiny heath subshrubs only 6 in (15 cm)
high (see Plate V). Arctic bell heather is found where snow is late to melt. Mosses
and fruticose lichens complete the ground cover. In northeastern Canada, limited
snow cover and abrasive winds support different species and a more limited plant
assemblage.
In areas with minor snow cover, dwarf-shrub heath tundra may or may not
include cottongrass tussocks. Small heath shrubs grow on medium-drained soils.
Species vary with geographic location and include mountain azalea and Lapland
rosebay. Evergreen leaves on several species are diagnostic of this community.
Graminoids include red fescue, highland rush, and arctic bluegrass. Large areas of
western and northern Alaska, especially the Mackenzie River delta, are covered
with a community variant that includes cottongrass. Cottongrass is conspicuous
because of its white tufts, but heaths still dominate. A well-developed cryptogam
layer of mosses, including several Sphagnum species, is present. Especially when
they flower in early summer, cottongrass communities are important grazing
ground for caribou, providing good nutrition for lactating females.
Regions with low relief, poor drainage, and a thin active layer over permafrost,
such as the coastal plain and outer Mackenzie Delta support graminoid-moss tun-
dra (mires) with plants 12 in (30 cm) tall. Because of shallow permafrost, this
region is characterized by patterned ground, polygons, nets, and stripes. These wet-
land meadows are dominated by water sedge, few-flowered sedge, and cottongrass,
but they also include marsh cinquefoil. Different soil textures support different
plants. The fine-grained soil in the centers of the polygons or in cracks between
them are too saturated for much plant life other than some mosses. Lichens, dwarf
shrubs, and forbs grow on the coarser, raised, and better-drained rims.
Wind-swept slopes and ridges with little to no snow cover are a cushion plant
cryptogam polar semidesert, with low cushions or mats. This community, similar
to the High Arctic and limited in the west, is more common in the east where more
bare rock and gravel occur. Mountain avens plants, less than 2 in (5 cm) high but
52
more than 3 ft (1 m) in diameter, can account for 8090% of the vascular plant
cover, especially on calcareous soils. Associated low-stature perennial forbs are
northern anemone, viviparous bistort, and moss campion. Lichens are common.
Plant communities of the High Arctic. High Arctic refers to the high-latitude Cana-
dian archipelago and the northern tip of Greenland. Because of more arid condi-
tions and less protective snow, plant cover is sparse. Several vegetation types occur,
two of which are small areas of tundra vegetation similar to those communities
found in the Low Arctic but with few to no tall shrubs. Cushion plantcryptogam
and cryptogam-herb semidesert cover most large areas of the High Arctic. Herb
barrens constitute small areas of true polar deserts.
Graminoid-moss tundra, found wherever drainage is impeded, such as river ter-
races and coastal lowlands, forms major grazing grounds for muskoxen. Adjacent
lakes and ponds provide breeding sites for waterfowl and shorebirds. Sedges are
the most important plants, but mountain foxtail and Fishers dupontia grasses are
also common. Saxifrage cushions, cottongrass, and lousewort are widespread, but
mountain avens and prostrate arctic willows are restricted to hummocks with bet-
ter drainage. Mosses and cyanobacteria are abundant, but lichens are rare. Mires
in the southern and eastern High Arctic are dominated by sedges, but in the north-
west Canadian islands, they are dominated by dupontia.
Dwarf-shrub heath tundra has fewer heath species than in the Low Arctic.
Heath tundra needs snowbed sites that melt by early July, situated near rocks that
absorb and provide summer heat, situations rarely found in the High Arctic. Arctic
bell heather is most characteristic, but other species include short-leaf sedge, arctic
willow, purple saxifrage, and mountain avens cushions. Mosses are plentiful.
Many fruticose lichens, including several Cetraria species, arctic finger, and white-
worm are members of this community. Heath tundra in northern and central Baffin
Island has a richer flora than elsewhere but is still dominated by arctic bell heather.
Polar semideserts are not barren like true polar deserts, but have some plant
cover, a richer flora, and more soil development. They cover about one-half of the
southern Canadian islands and one-quarter of the northern islands. Cushion plant
cryptogam vegetation is similar to that found in limited sites in the Low Arctic.
Predominantly growing on warm gravelly surfaces with a deep active layer, it
becomes a minor component where rocky surfaces, fine soils, cold climate, or late
snowmelt occurs. Vegetation is dominated by mats or rounded cushions of arctic
mountain avens, with associated cushion plants of several saxifrages, sandworts,
and starworts. Vascular plants, however, are less important than cryptogams.
Lichens and mosses are numerous and varied. Semideserts provide the main habi-
tat for Pearys caribou, collared lemming, ptarmigan, and numerous passerine
birds.
Cryptogam-herb semidesert is the general vegetation type on low rolling
uplands and beaches with finer soils rather than gravel or rocks. Vascular plants
account for only 520% of the total; cryptogams make up 5080%. Vegetation is
53
dominated by bryophytes and both crustose and foliose lichens, in contrast with
the dominance of fruticose forms in the Low Arctic. Yet, the general appearance is
that of a grassland because graminoids of mountain foxtail and northern woodrush
are visually dominant. Upland sedges are totally absent. Forbs include rosettes or
cushions of mouse-ear, draba, sandwort, rooted poppy, and saxifrage species.
Bryophytes are abundant, including several mosses. A black cryptogamic crusta
mixture of black crustose lichens and cyanobacteria capable of fixing nitrogenis
common on the soil surface. In the High Arctic, seedlings of vascular plants do best
in deep mosses and desiccation cracks where conditions are more moist and there
is less needle ice. More seedlings die in summer due to soil drought than due to
winter cold.
Found largely in the Queen Elizabeth Islands, polar deserts in the High Arctic
are dominated by herb barrens that form a mosaic with semidesert vegetation.
About 98% of the area is bare soil and rocks, justifying the term desert. Herb bar-
rens are determined by the lack of soil development, low soil-nutrient content, nee-
dle ice, dry surface, and a short growing season of just one to one-and-a-half
months. The few vascular plants are found in relatively protected areas of desicca-
tion cracks or near stones in patterned ground. Vascular plants may include draba
and rooted poppy rosettes and red sandwort and purple saxifrage cushions. Crypto-
gams cover less than 1% of the ground, a major contrast with semideserts where
cryptogams dominate. The barrens were probably still covered by ice during the
Little Ice Age 130430 years ago. Not even lichens grow yet on the rocks.
Animals. North American mammals are those typical of circumpolar tundra. Car-
ibou is the major large herbivore in the Low Arctic, joined by muskoxen at higher
latitudes. Muskox graze sedge-moss meadows, and in winter, paw through as
much as 20 in (50 cm) of snow to feed. Small herbivores in the Low Arctic include
arctic hare, both brown lemming and collared lemming, and arctic ground squirrel.
Brown lemmings graze on graminoids and mosses in wet areas, while collared lem-
mings eat forbs and dwarf shrubs in drier areas. The only mouse-like species in the
High Arctic is the collared lemming, most populous in raised-center polygons with
deep snow in the troughs. Because of harsher conditions, this species has fewer lit-
ters per year than its relatives in the Low Arctic. Their primary foods are mountain
avens, herbaceous willow, and purple saxifrage. Arctic hares are more common in
the Low Arctic, where they feed on willows.
Predators in the Low Arctic include brown bears that venture into the tundra to
feed primarily on roots and grasses. Smaller animals that prey on rodents such as
lemmings include the least weasel and short-tailed weasel, both of which can ma-
neuver into narrow burrows. Because lemming populations are lower and nesting
birds with eggs are less common, fewer carnivores inhabit the High Arctic, in terms
of both species and total numbers. Least weasels are not found in the High Arctic,
but short-tailed weasels are. Arctic fox is common to both environments, as is an
occasional timber (or arctic) wolf. By hunting in packs, wolves are capable of
54
taking larger animals such as caribou. Foxes search out small mammals and follow
polar bears onto sea ice to scavenge leftover seal kills. Avian predators outnumber
mammalian predators. Low Arctic predatory birds numerous in high lemming
years include Snowy Owls, jaegers, Glaucous Gulls, and Short-eared Owls. These
birds winter elsewhere, the Snowy Owls being the first to arrive in early spring, the
rest returning to the tundra by the time snow melts. All but the Short-eared Owl
are found in the High Arctic. The only predators staying in the tundra all winter
are the least weasel, short-tailed weasel, and arctic fox. Polar bears only use the
tundra during summer.
Ptarmigan are birds common to both environments. Willow Ptarmigan is more
common in the Low Arctic where willow shrubs are a significant food source while
Rock Ptarmigan is the major species in the High Arctic. Snow Geese breed in
the High Arctic but winter along the western coast of the United States (see Figure
2.7). Other birds found in the Low Arctic are also typical of circumpolar tundra.
Greenland
Covering more than 23 degrees of latitude from 59 46
0
N to 83 04
0
N, the north-
ern most land on Earth, Greenland has considerable climate variation. A distinct
north-south gradient in temperature and precipitation develops, especially in win-
ter. Almost 80% of Greenland is ice cap, actually islands buried in ice. Areas clos-
est to the ice cap are more continental in climate, while coastal regions are varied
due to a cold current on the east and south coasts and a warm current off the south-
ern third of the west coast. Summers are colder on the east coast.
Figure 2.7 Snow Goose with nest in the High Arctic. (Courtesy of the U.S. Fish and Wild-
life Service.)
55
This large continent in the path of westerly winds splits storm tracks. Radiation
cooling from the ice cap causes a persistent high pressure in the north, while the
southern part experiences cyclonic storms that move north along the west coast
before crossing over the ice cap. Directly in the path of cyclonic storms generated
between polar and maritime airmasses, the southwest coast is wet, with 30 in
(760 mm) or more of precipitation annually. Coastal climate is also influenced by
whether or not land is adjacent to sea ice. All coastal stations have moderate tem-
peratures compared with the rest of the Arctic. Nord (81 36
0
N) is the coldest with
a mean annual temperature of 2.5 F (16.4 C). Stations on the south coast have
mean annual temperatures around freezing. Inner fjords are warmer than the
coasts in summer, due both to warming effects of downslope (f ohn) winds and less
oceanic influence. Although these strong winds originate on the ice cap, they
increase in temperature as they descend and bring warm, dry conditions, raising
temperatures as much as 35 F (20 C) in a few minutes. Such increases in winter
can melt snow, and expose vegetation where snow cover is light. Where accumula-
tions are deeper, only the surface melts, and then it subsequently refreezes forming
an ice crust. The interior gets less precipitation, has a shorter season of snow cover,
and a more continental temperature regime, that is, higher temperatures in summer
and lower in winter. Warmer summers and a longer growing season are important
for plants there.
Northern Greenland is polar desert, with a low mean annual temperature,
strong winds, low precipitation, and high evaporation.
Greenlands topography and history has affected its flora and fauna. Almost all
of the current ice-free land was glaciated during the Pleistocene and was exposed
only in the last 10,000 years. The ice cap is a barrier to most organisms, limiting
colonization and distribution areas. The ice-free edge of the continent is dissected
by fjords and glaciers into peninsulas and islands, further limiting migration.
Because of this relative isolation, fewer species of both plants and animals are
found in Greenland than in other parts of the Arctic. Caribou, arctic hare, and arc-
tic fox are the only animals found throughout ice-free Greenland. Arctic wolf,
short-tailed weasel, collared lemming, and muskox are present only in High Arctic
Greenland.
Most mammals originated in Eurasia and came across the Bering Strait to
North America, and then migrated to Greenland. Three migration routes are
reflected in plant and animal distributions. The route from North America via
Ellesmere Island over Naras Strait allowed access primarily for High Arctic spe-
cies. Davis Strait from Baffin Island and Labrador brought in Low Arctic and bo-
real species from North America. Other low Arctic and boreal species migrated
from Europe via North Atlantic Islands. Wintering grounds of migrating birds indi-
cate their probable origins. Wheatears winter in tropical Africa, and most likely
colonized Greenland from the Old World. Snow Buntings fly from northeastern
Greenland to the southern Russian steppes and to the Great Lakes, indicating that
they probably colonized from both the east and west. Greenland Redpolls winter in
56
both North America and Europe, also indicating a double colonization source.
Lapland Buntings, which winter in North America, probably also colonized from
there.
Any species that migrated to Greenland from Europe must have been maritime
or quite adaptable. Several species that are primarily continental in Eurasia have
become maritime in Greenland, indicating a westward migration. Elements of
Greenlands flora that have come from the west, from continental North America,
have remained continental in character. The south and southeast coasts of Green-
land have a majority of European elements in the flora, both because of proximity
and similarity of climate with northwest Europe. The western flora is more similar
to that of the North American continent.
Tundra vegetation in Greenland varies according to topography and climate.
Most of the ice-free area is mountainous, with rolling uplands and deep glacial
troughs, creating local climates and habitats. Greenland has no extensive land-
scapes comparable to those found in the rest of the Arctic. Geology also varies, and
substrate may be many types of rocks or sediment. Soils are podzolics, arctic
brown, and polar desert, with low organic matter. Because of aridity, some soils
are salty, and salt crusts may be caused by excessive summer evaporation. Wet
soils supporting sedges, grasses, and mosses are rare and only found in broad, flat
valleys in west-central and eastern Greenland.
Inland heads of fjords, which are more continental and have July mean temper-
atures over 50 F (10 C), support low shrub tundra of downy birch and Greenland
ash. Some plants only occur around the hot springs common in basalt areas of
west-central and eastern Greenland, well north of their normal climate limits. A
willow scrub and a variety of herbaceous plants are restricted to this habitat. Hot
spring habitats may have served as refugia for tundra plants during colder climate
periods.
Most plants and animals have narrow environmental requirements. For exam-
ple, wading birds need both snow-free nesting grounds and abundant insects for
food. The only area with good conditions for both is north of Scoresby Sund. Far-
ther north, little food is available, and farther south, both too much snow and cool
summers limit insect numbers. Therefore, most waders in Greenland occur in the
High Arctic rather than Low Arctic or subarctic. Ptarmigan in eastern Greenland
migrate according to availability of food. They spend the summer breeding season
in the interior near the fjords, where they feed on viviparous bistort, especially the
protein-rich bulbils. Throughout the short summer, they follow the bistort bloom
into the mountains. In winter, they move to open plateaus and plains on the outer
coast where vegetation is blown free of snow and the major food is mountain
avens, arctic willow, purple saxifrage, and bistort bulbils, according to snow cover.
Caribou in western Greenland move from coastal wintering grounds to inland
summer calving grounds just as high-quality graminoids emerge.
Three areas of Greenland have been well studied, and two others provide im-
portant habitat. The southern tip of Greenland, considered subarctic, is shrubby.
57
Birch about 5 ft (1.5 m) tall can be found in sheltered snow-covered places. How-
ever, no seedlings are produced and plants propagate vegetatively. The underlayer
consists of dwarf shrubs, grasses, or forbs. Most of this area and habitat was
exploited or cleared for agriculture and pasture as early as 1,000 years ago.
On the southwest coast at about 66 N, the area centered on Sndre Strmfjord
is Low Arctic with a pronounced continental-maritime gradient. The western part
has high mountains, up to 4,500 ft (1,400 m), with alpine glaciers, putting the east-
ern low hills and U-shaped valleys in rainshadow. A steep gradient of precipitation,
temperature, and duration of snow according to topography is reflected in vegeta-
tion, from grassy meadows and dwarf shrub on the coast to grasslands with salt
pans and lakes in the drier interior. Inner fjords that are warmer in summer support
low shrub communities dominated by willow and birch, with an open canopy up
to 2 ft (0.5 m) high. This general region is the most important territory for caribou
in Greenland. Their grazing has changed blue willow scrub into bluegrass com-
munities. The caribou population is unstable due to overgrazing and the lack of
their natural predators, wolves.
Jameson Land in eastern Greenland north of Scoresby Sund at about 70 N is a
transition from Low to High Arctic. The gentle plateau dipping west is a large low-
land area with steep, moist slopes and inland valleys with a warmer and drier conti-
nental climate. Up to 1,300 ft (400 m), it is dominated by dry dwarf-shrub heath
with a 2075% cover of vascular plants. Dominant species include arctic bell
heather, bog bilberry, and mountain azalea (see Plate VI). Moist sites also have
mosses. Snowbed vegetation within 6 mi (10 km) of the coast supports Bigelows
sedge, bistort, and arctic willow. The drier east has open dwarf-shrub heath. Even
drier sites in the east are fellfield with alpine bearberry, mountain avens, arctic wil-
low, and moss campion. Jameson Land is one of the most important areas for
muskox in the world. Winter foraging grounds have little snow cover, and summer
sites are thick forb communities where deep winter snows impart summer mois-
ture. This region also provides important July molting areas for Pink-footed Geese
and Barnacle Geese. The birds are separated by habitatPink-footed Geese along
the coast and rivers with an open view and Barnacle Geese on smaller rivers and
lakes surrounded by hillsand feeding zones. They both eat sedges but in different
parts of the meadows.
Disko is an island off the west-central coast, about 70 N. It is a high (up to
5,900 ft [1,800 m]) basalt plateau dissected by U-shaped valleys and steep slopes.
The west has a High Arctic maritime environment, cool and foggy, with solifluc-
tion stripes and frost boils on the surface. Vegetation is an open, mossy heath
dominated by arctic willow and arctic bell heather along with herbaceous plants.
South-facing slopes have dense forb vegetation and blue willow scrub, especially
around hot springs where water temperature can reach 50 F (10 C). Species more
normally found farther south grow well in these habitats. Peary Land at the north-
ern tip of the continent at 83 N also has a High Arctic environment. Precipitation
near the coast, which always falls as snow, is less than 8 in (200 mm), decreasing to
58
less than 1 in (25 mm) in the interior. What little
snow falls is wind blown, exposing much bare
land. The drier interior has mountain avens,
heath, and graminoids. Snowdrift sites, significant
because they provide the only water in summer,
support sparse communities with sedges, cotton-
grass, arctic bell heather, and mosses. Cover in
both is less than 5%. On the Arctic coast, which is
more humid with frequent fog, mosses and lichens
grow in patterned ground cracks.
Eurasia
Fennoscandia. Although little permafrost occurs
in Fennoscandia (the peninsula that includes
Norway, Sweden, and Finland), mean annual
temperature below freezing and summer months
below 50 F (10 C) qualify the area as a tundra
environment. Much of the mountain backbone of
the peninsula is high enough for alpine tundra,
and arctic and alpine zones merge in the north
(see Figure 2.8). Fennoscandia is positioned in
storm tracks between warm Atlantic air and cold
polar air. Maritime conditions prevail on the
coast, while the lee side of the mountains in the
interior is drier and more continental. For exam-
ple, Hardangervidda, the high-elevation plateau
in southern Norway, has 60 in (1,500 mm) of pre-
cipitation on the west and 20 in (500 mm) on the
east. The coast receives autumn precipitation
caused by cyclonic storms, while summer convec-
tional showers bring moisture to the interior. The
lowest precipitation occurs in the northeast, with
only 12 in (300 mm) received annually at Kauto-
keino, Norway. Except in coastal locations, winter
temperatures may drop below 40 F (40 C).
The length of the growing season is 120 days or
less, and on the Hardangervidda, it is a mere
74 days. Most areas have more than 200 days with
snow cover, but depth and coverage, from inches
to several feet, varies with wind. The time it takes
for snow to melt affects the length of the growing
season, and some snow may remain all summer.
.................................................
Iceland
At 65 N, Iceland is on the southern boundary
of the Arctic. While the island is basically a lava
plateau, plant distributions are complicated by
fjords, isolated mountain peaks, lava flows, gla-
ciation, wind-blown sand deposits, and settle-
ment since 900 AD. For centuries the highlands
have been used for summer grazing of freely
roaming sheep. A small landmass affected by a
warm current, Iceland has mild temperatures
for the latitude. July averages 45 F (7 C), and
January means drop to only 19 F (7 C). An-
nual precipitation is high for the Arctic, 2550
in (6001,200 mm), and snow is deep, averag-
ing 40 in (100 cm). Because of glaciation and
volcanic activity, soils are gently rolling till-
covered moraines, glaciofluvial deposits, or lava
flows. Plant communities are similar to those
found on mainland arctic environments, but
differences exist. Basalt limits acidity, so little or
no sphagnum moss and few heaths grow on
recent lava flows. Lichens are poorly repre-
sented in heath vegetation, partly due to graz-
ing by reindeer. Widespread on the island,
Arctic fox is the only native land mammal. The
fox has two color phases; one becomes white in
winter while the other remains bluish white all
year. Mink, introduced for fur farming in the
1930s, have escaped and are currently found in
the vegetated highlands but not on the barren
plateau. Reindeer have been introduced, and an
estimated 4,000 live in the highlands. There are
no native small mammals. A field mouse was
introduced and its feral distribution today is not
known. The bird fauna is rich and varied.
Approximately 80,000 Pink-footed Geese breed
in large colonies on the plateau, and 10,000
Whooper Swans breed along lakes and ponds.
.................................................
59
Well-defined vegetation belts characterize
Fennoscandia tundra. The subalpine includes the
upper or northern limit of boreal forest, a treeline
with downy birch, which is also typical of arctic
tundra in Russia, Iceland, and Greenland. Tree-
line varies from 4,000 ft (1,200 m) in southern
Norway (61 N) to sea level in the north. The
crooked stems of birches are less than 30 ft (10 m)
high, and the zone is the equivalent to krummholz
in North American alpine areas. In more conti-
nental areas, the treeline and krummholz tree is
Norway spruce. Bog birch, common juniper, and
several heath species such as Scotch heather and
blueberries, make up the shrub layer. An herb
layer consists of nodding hairgrass, matgrass, and
carex sedges. Mosses are found in wetter areas,
replaced by lichens where it is drier.
The low alpine is similar to the subalpine but
without trees. With lower summer temperatures,
shrub size is much smaller, 10 ft (3 m) high down
to 8 in (20 cm). Thickets have tall forbs such as
monkshood, buttercup, globeflower, and grami-
noids. The ground layer, unless covered by litter
or shaded, consists of mosses and liverworts. The
most common heath association in the low alpine
is dominated by European blueberry with a
ground layer of bryophytes. Because less organic
matter accumulates at higher elevations, the
low-alpine belt is the upper limit of mires. Mire
vegetation includes bog rosemary, bog birch,
cottongrass, small cranberry, and sphagnum
moss. Graminoids dominate moderate snowbeds,
but extremely long-lasting snowbeds may have
mosses, the colorful mountain lichen, moss plant,
and buttercups. More continental areas have a
fruticose lichen ground cover of Iceland moss and
Cladonia lichens.
Many communities have an upper limit
between the low- and mid-alpine belts (see Figure
2.9). The mid-alpine is dominated by meadows of
highland rush, but Bigelows sedge, sheep fescue,
and other graminoids also occur, with minor rep-
resentation of heaths. This belt is absent in
.................................................
Coal Mining in the Arctic
Halfway between Norway and the North Pole,
Svalbard was an international whaling base in
the seventeenth and eighteenth centuries. At
74 81
0
N, you would expect the island group
to be bitter cold and uninhabited, but because
it is in the path of the warm North Atlantic
Drift, temperatures are relatively mild. The
warmest summer month averages 40 F (4.5
C), while winters average 7 F (14 C). Plants
are typical of circumpolar Arctic. The islands
have a wealth of natural resources, especially
coal but also wildlife. Although it has been part
of Norway since 1920, several nations have
operated mines, but only Norwegian and Rus-
sian companies still extract coal. More than half
of the population (total population 1,701 in
2005) is employed in coal mining and related
service industries. About 60% of the land is
covered by glaciers and snowfields, and ice
floes often block entrances to ports. There are
no muskoxen, hares, wolves, owls, falcons, or
ravens. The single rodent, a mouse that occu-
pies bird cliffs, was possibly introduced. There
are also few invertebrates, notably no biting
flies that harass reindeer. The single mamma-
lian predator is arctic fox, which preys on birds
and eats carrion. Two endemic subspecies,
Svalbard reindeer and Svalbard Ptarmigan, are
of conservation significance. The reindeer prob-
ably colonized from northeastern Greenland
and the Canadian archipelago and has been iso-
lated for 40,000 years. Even though few mam-
mals live on Svalbard, visitors can see some of
the largest concentrations of birds in the North
Atlantic. Migrating birds that nest on the tundra
include Pink-footed Goose, Barnacle Goose, and
Brant Goose. Polar bears can be seen along the
coast in summer. About 65% of the islands is
environmentally protected in nature reserves,
national parks, and bird sanctuaries.
.................................................
60
Finland. An abrupt transition from low-alpine to mid-alpine exists, noted by the
absence of several plant communities, but a gradual transition separates mid-alpine
and high-alpine belts in which many of the same species are present. Plants grow-
ing in both mid-alpine and high-alpine belts include moss campion and saxifrage
cushions and spiked trisetum and carex graminoids, along with alpine sorrel and
several buttercups. Dwarf willows are the only shrubs in the high-alpine zone
where the number of plant species decreases with elevation. Plant cover, domi-
nated by lichens and mosses, is incomplete. Solifluction, polygons, and stripes
Figure 2.8 Except for the mountains in Fennoscandia, Arctic tundra in Eurasia is lim-
ited to a narrow coastal zone. (Map by Bernd Kuennecke.)
Figure 2.9 Vegetation on high mountains in Norway, here at Gudvangen, merges with
arctic tundra in northern Fennoscandia. (Photo by author.)
61
make the soil rocky and unstable but still able to support a wide variety of vascular
plants.
A more circumpolar flora exists on peaks, while more strictly Eurasian species
appear at lower elevations. Few plants are endemic to the region. Plants originating
in the western Arctic probably survived the Pleistocene by taking refuge on the
northern and southern mountains that extended above the ice. The central moun-
tains were too low and covered by ice. Some species are found in only one high-
alpine area, indicating the location of their refugia.
Animal life in Fennoscandia is more varied than in most of the Arctic, reflect-
ing the connections to alpine environments. However, not all species are present in
both the arctic and alpine regions, and mammals are few. Wild reindeer have been
extinct in Sweden since 1860 and in Finland since 1900. Today, they are only
found in small Hardangervidda areas in Norway. However, domesticated herds
are many and widespread. Small mammals include Norway lemmings, famous for
their massive migrations, shrews, and voles. Digging animals are more common
than in the rest of the Arctic due to the lack of permafrost. The root vole is found in
moist areas near streams or mires in the low- and mid-alpine zones. Other voles
occupy other habitats. Mountain hare are found all the way to the high-alpine
zone, and like ptarmigan, will burrow into snowbanks for warmth.
Several native mammalian predators occur, primarily in alpine environments,
but most large species are endangered. Wolverines were hunted almost to extinc-
tion but are now protected, with populations primarily on the mountainous frontier
between Sweden and Finland. Wolves are nearly extinct, and only a few brown
bears still live in Swedens forests. Lynx is the most common predator in the subal-
pine and low-alpine zones. Lynx, brown bear, and wolf are mainly forest animals,
occasionally venturing into the tundra. Smaller predatory mammals more com-
monly found in both arctic and alpine tundra are ermine and other weasels, red
fox, and arctic fox, although few arctic fox remain. All predatory mammal popula-
tions fluctuate with prey populations.
Nesting birds are plentiful, their local presence determined by vegetation and
habitat. In wet areas, waterfowl and shorebirds dominate. Whooper Swans and
Bean Geese are found in wet areas of birch forests, while Mallards prefer smaller
lakes surrounded by willows. Willow Ptarmigan are found in willow thickets,
while Rock Ptarmigan prefer rocky slopes. Waders such as Golden Plover and
Red-necked Phalarope congregate around bogs and adjacent heathlands. Passer-
ines such as Snow Bunting, Meadow Pipit, and wheatears frequent several
habitats.
Because many predatory birds only reproduce when prey is plentiful, their
numbers correlate with populations of small mammals. Rough-legged Buzzards
and Short-eared Owls catch rodents, while Golden Eagle and Gyrfalcon both prey
on ptarmigan. Lemmings are prey to Snowy Owl and Long-tailed Skua in the Arc-
tic. Ravens are opportunistic, catching live prey and eating carrion. Many preda-
tory birds are now protected by law.
62
European common frog is the only amphibian found above treeline, but it is
rare even in the low-alpine zone. Reptiles are scarce. The common viviparous liz-
ard lives in the low alpine, but grass snake is limited to the subalpine zone.
Russia. Latitudinal belts of tundra are limited to a relatively narrow northern strip
on the Eurasian continent north of the Arctic Circle. Terminology is confusing
because all authors do not use the same terms to define Russian Arctic zones.
Three categories of tundra (southern, typical, and arctic), as well as polar desert to
the north and forest tundra to the south, are described here. All zones are not repre-
sented equally across the continent.
Temperature in all zones is strongly influenced by degree of continentality (see
Table 2.2). Mean annual temperatures are similar, as are January means, which
are considerably warmer than the more continental boreal forest farther south.
Tundra zones thus reflect differences in July mean temperatures, which are suc-
ceedingly cooler from south to north. July in the forest tundra averages 54 F
(12 C), above the 50 F (10 C) limit for tundra, which explains the presence of
trees. Both the western and eastern parts of the Russian Arctic are slightly warmer
due to warmer oceans nearby, particularly the North Atlantic Drift and Barents
Sea on the west. Summer begins when mean air temperatures rise above freezing,
which occurs in early June in southern tundra but is delayed to early July in polar
desert. Temperatures drop below freezing in September. The frost-free season is
short, ranging from two months or less in polar desert to three-and-a-half months
in southern and forest tundra. The western Russian Arctic has a longer frost-free
season, four-and-a-half to five months from mid-May until mid-October, due to the
influence of the ice-free North Atlantic Ocean.
Table 2.2 Russian Arctic Tundra Climate Summary
CLIMATE
CHARACTERISTIC
SOUTHERN
TUNDRA TYPICAL TUNDRA ARCTIC TUNDRA POLAR DESERT
Mean Annual
Temperature
10 F (12 C) 10 F (12 C) 10 F (12 C) 10 F (12 C)
January Mean
Temperature
15 F (26 C) 15 F (26 C) 15 F (26 C) 15 F (26 C)
July Mean
Temperature
48 F (9 C) 48 F (9 C) 39 F (4 C) 34 F (1 C)
Annual
Precipitation
13 in (330 mm) 13 in (330 mm) 9 in (230 mm) 7 in (175 mm)
Depth of Snow
Cover
16 in (40 cm) 16 in (40 cm) 9 in (23 cm) 9 in (23 cm)
Depth of Active
Layer
33 in (85 cm) 30 in (75 cm) 21 in (55 cm) 16 in (40 cm)
63
Precipitation varies with latitude, longitude, topography, and distance from the
ocean. In general, polar deserts are driest and tundra zones are wetter. Precipita-
tion is highest on the mountains in the central Taymyr peninsula and on the
Barents Sea Coast, where up to 20 in (500 mm) may be received annually. About
one-third falls as rain in July and August. Relative humidity is high, typically 8090%,
because cold air quickly reaches saturation.
Snow cover in tundra zones lasts 200280 days, remaining the longest amount
of time in colder Siberia and shortest amount of time in the warmer west. Because
of generally light winter precipitation, snow cover is not deep, but it is variable
depending on geographic location. Less snow falls in the north and polar desert,
and more on the flatter southern tundra. European Russia has deeper snow (up to
24 in [60 cm]) because it is both warmer and wetter. Frequent strong winds redis-
tribute the snow, blowing ridgetops free and piling snow in swales. Snowmelt
begins in early May in the west, but not until the end of June on arctic islands.
Some sheltered areas retain snow all summer. Summer snowfalls are light, and the
temporary cooling is too brief to harm most plants.
The typical Siberian arctic landscape is a gently rolling plain with many
streams, ponds, and lakes. All but the westernmost area is underlain by continuous
permafrost, and various types of patterned ground are characteristic. Polygonal
mires are a pattern of polygons up to 50 ft (15 m) in diameter with rims about 6.5 ft
(2 m) wide and 3 ft (1 m) high. Deep cracks in the rims fill with water that freezes
in winter, perpetuating the polygon, while depressed centers contain ponds or bogs
in summer. The landscape is large scale. One mire may consist of 50100 polygons.
Frost boils are also common. Frost-heaving pushes bare soil to the surface, like a
bubble in boiling water. The bare top may be 25 in (60 cm) in diameter, surrounded
by vegetated soil 4 in (10 cm) higher. Boils are separated by narrow, shallow
troughs. Whole landscapes can consist of frost boils. Groups of small pingos, up to
30 ft (10 m) wide and 1.5 ft (.5 m) high, are relics of past glaciation and create a
hummocky landscape with continuous vegetation cover. Larger pingos are 600 ft
(180 m) in diameter and 230 ft (70 m) high.
The depth of the active layer varies but generally decreases with latitude. The
deepest active layer, 50 in (130 cm), is on south-facing slopes in southern and typi-
cal tundras. Peat bogs have the shallowest active layers, 20 in (50 cm) in the south,
decreasing to only 8 in (20 cm) in the north.
Southern tundra is the ecotone between tundra and boreal forest, the northern
limits of treeline, and is widest on the west and east where climates are slightly
warmer. Tree growth is sparse, with krummholz and prostrate trees along rivers.
Treeline species vary with geographic area but include downy birch, willow-like
Chosenia arbutifolia, several species of larch, Siberian spruce, and Mongolian pop-
lar. This zone is characterized by shrub communities with a three-tier stratification.
Taller shrubs, 2 ft (0.6 m), are several species of birch and willow, as well as alder
and mountain alder. Shrub height depends on depth of winter snow, and plants are
taller where snow provides more protection. Many dwarf shrubs, 6 in (15 cm) tall,
64
such as arctic bell heather, mountain crowberry, and bog bilberry, grow under-
neath as a second layer. Mountain avens cushions, several carex sedges, and cot-
tongrass are also common. A low ground cover, 4 in (10 cm), consists of a rich
variety of fruticose and foliose lichens, along with mosses and some liverworts.
In addition to the more typical shrub community, other plant communities
occur in other situations. Large, flat areas may have palsa bogs with a grassy cover
of cottongrasses and sedges and water-loving mosses such as Calliergon and Drepa-
nocladus. Peat hills are formed by Sphagnum species. Warmer south-facing slopes
have meadows with a rich mix of grasses, legumes, and other forbs, both arctic and
boreal species. Forest enclaves on the European arctic islands or along rivers have
typical treeline species according to geographic area. Northeastern Russia has tus-
sock communities of carex sedges and cottongrass.
Southern tundra has a diverse assemblage of animals. Birds are abundant, espe-
cially water or shore birds such as White-fronted Goose, Brant Goose, Stone
Plover, and sandpipers. Surface-feeding, or puddle ducks, are also common. Many
wading birds and passerines are insectivores, but Willow Ptarmigan feeds on wil-
low stems and buds.
Typical mammals are voles and lemmings. Voles are widely distributed in
southern tundra, especially in European Russia, where their presence increases cer-
tain forbs and grasses. Predators are numerous compared with the relatively few
prey species. Arctic fox, Snowy Owl, Pomatorhine Skua, and Rough-legged Buz-
zard all prey on lemmings, and their populations are generally tied to the lemming
cycle. Skuas only breed when lemming populations are high. Foxes depend on a
variety of food, but increase in population when lemmings peak. Snowy Owls also
breed more in high-lemming years, and rarely nest in low-population years. The
relationship between lemmings and weasel populations is not known.
Proximity to several biogeographic provinces contributes to the rich variety of
species in the southern tundra, and many nonarctic elements are present in both
the flora and fauna, among them several boreal forest mosses and many birds and
insects, especially in the forest enclaves. Plants more typical of northern tundra
zones are absent or restricted to snowbeds. Circumpolar species predominate, but
more geographic variation exists here than in northern tundra zones. Compared
with European Russia, Siberia is more strictly arctic.
Typical tundra is also called moss-lichen tundra because of the prevalence of
mosses and lichens and its low stature, less than 8 in (20 cm) high (see Figure 2.10).
This zone, which lacks trees altogether, has its widest extent in the west; it is frag-
mented in the east. Closed shrub thickets are limited to river valleys or protected
areas. Vegetation is low willow shrubs with semiprostrate birches and willows.
Mosses are prominent and characteristic, making a 510 in (1325 cm) thick layer
on the soil. The liverwort Ptilidium ciliare is a common codominant in the moss
cover, where many species combine to form a mosaic. The moss turf can either be a
continuous layer or be interrupted by frost boils with bare spots. Because the moss
layer is thick, vascular plants must have long rhizomes to reach through to the soil.
65
Carex sedges are common, as are some dwarf shrubs, mainly Lapland bell heather,
polar willow, and alpine avens cushions. The same fruticose and foliose lichens are
found here as in southern tundra. Some dwarf heath shrubs grow in protected loca-
tions, but they rarely flower in this colder zone. A greater circumpolar component in
the flora, similar in both species and appearance all across Arctic Russia, exists in
this belt, in contrast to more geographic variation in southern tundra. In river val-
leys, there may be low willow thickets or low moss carpets of water-loving species,
with a few flowering plants of alpine sorrel, buttercups, and saxifrage cushions.
Drier fellfields have mountain avens cushions but no mosses, and warmer south-
facing fellfield slopes support meadows with a variety of grasses and forbs.
Birds typical of southern tundra, such as White-fronted Goose and Stone
Plover, are absent from typical tundra. Without shrubs, no birds or insects depend-
ent on shrubs can survive. Small mammals are limited to lemmings. The moss is
home to a rich variety of invertebrates, including springtails, mites, spiders, and
rove beetles, but the sparse undergrowth supports few insects.
Arctic tundra is found on the northwestern mainland coast, and on Arctic
islands only in the east. The absence of shrubs and even most dwarf shrubs simpli-
fies the vertical structure of the community. The moss turf is only 12 in (35 cm)
thick, and vegetative parts of flowering plants exist within the moss, not extending
above it except for an occasional flower stem to 3 in (10 cm). No stratification
exists, just one mixed layer of mosses and forbs. Sedges such as the cottongrasses
Figure 2.10 Typical tundra in Russia is characterized by a thick layer of lichens,
mosses, and liverworts. (Photo by author.)
66
common to typical tundra are replaced by grasses and forbs. Growthforms include
tufts, cushions, and dense mats. The tiny dwarf shrub, polar willow, grows totally
protected within the moss layer, with its leaves lying on top. Mountain avens are
found only on fellfields. Fishers dupontia grass gains dominance as carex sedge
numbers decline. The liverwort disappears and is replaced by an abundance of Dre-
panocladus uncinatus moss. Except for sphagnum, other dominant mosses remain the
same as in typical tundra. Fruticose lichens are replaced by crustose types. Ground
may be 50% bare, and no communities have a closed vegetation cover. Plants find
shelter in cracks caused by permafrost action, and polygon patterns are outlined by
vegetation. Depressions may be bare if snow fails to melt early enough to allow plant
growth. Few additional habitats occur in arctic tundra to support different vegeta-
tion. South-facing slopes have a flora much like the general community.
The most northern arctic environment, polar desert, has limited extent, occur-
ring primarily on the north half of Novaya Zemyla and the western arctic islands,
and on Cape Chelyuskun in Taymyr. This extreme environment is characterized
by low summer temperature, little precipitation, a shallow active layer, and a short
growing period. Biotic diversity is low, with the number of vascular plant species
in local floras being about 50, one-half of that found in arctic tundra and one-quar-
ter of that in typical tundra. Less diversity is seen in mosses and lichens. Up to 95%
of the ground is bare; only the cracks bordering polygons are vegetated. Crypto-
gams dominate. Fruticose lichens include Cetraria species, snow lichen, and white-
worm lichen; Parmelia is an important foliose lichen. The moss layer is only 2 in
(5 cm) thick and lacks stratification. Flowering plants are sparse, found either within
the moss layer or lying flat on its surface. A uniform species composition prevails
throughout the polar desert, the vegetation only changing in density depending on
habitat. Only two or three land birds nest in the region. No earthworms, spiders,
beetles, sawflies, or craneflies inhabit this tundra belt.
Reindeer are the most numerous large herbivore on the Eurasian tundra. In
spite of sometimes heavy grazing, wild herds have little influence on tundra vegeta-
tion, and most winter pasture for both wild and domesticated reindeer is in or at
the edge of the northern boreal forest. Reindeer change their diet during migration.
On Taymyr and breeding grounds in the north, they feed on grasses and forbs; in
the southern tundra, they eat sedges, including cottongrass, and willows. In boreal
forest wintering grounds, they feed on lichens. Isolated herds on Novosibirskiye,
Novaya Zemyla, and Sibiryakova islands subsist all winter on polar willow shoots
and buds, because no shrub, lichen, or meadow communities occur there. The pop-
ulation uses multiple feeding grounds, so it rarely damages tundra vegetation.
Trampling only occurs in concentrated trail areas like land bridges between lakes
or river crossings.
Antarctica
Antarctica is a large continent with two geologic zones (see Figure 2.11). East Ant-
arctica is a stable continental shield composed of metamorphic and sedimentary
67
rocks, while West Antarctica is a mountain belt that includes the Antarctic Penin-
sula. Scotia Arc, a chain of peaks forming several islands off the northwestern part
of the Peninsula, has a long history of volcanism. It extends north to the South
Shetland Islands and northeast to the South Orkney Islands, South Sandwich
Islands, and South Georgia, almost to South America. Less than 5% of Antarctica
is ice-free. Most exposed land occurs along the coast, on islands, and on narrow
coastal strips of the Antarctic Peninsula, where the ocean moderates temperatures,
but dry valleys and nunataks (mountains above the ice) can also be found.
The northern boundary of the Antarctic tundra environment is best defined as
the 50 F (10 C) February isotherm, which occurs at the latitude where cold ocean
currents of the Southern Ocean meet warmer waters from the north, initiating a
change in both plankton and types of sea birds. It is also the position of the southern
Polar Front, where polar air meets mid-latitude air, roughly 55 to 60 S. These lati-
tudes, excluding the tip of South America, make up the subantarctic zone and a
transition from Southern Hemisphere cool-temperate climates to Antarctic condi-
tions. The subantarctic and transition zones also include the Kerguelen Islands at
50 S in the Indian Ocean and South Georgia at 54 S in the South Atlantic Ocean.
The Antarctic, subdivided into northern and southern zones (generally south of 60
S), includes the South Sandwich Islands at 58 S and Bouvetya at 55 S in the
South Atlantic Ocean. As in the Arctic, mean annual temperature is below freezing.
Figure 2.11 Tundra in Antarctica is limited to small ice-free areas. (Map by Bernd
Kuennecke.)
68
Diversity of terrestrial life is low compared with similar latitudes in the Arctic
due to the small extent of ice-free area, severe climate, climate history, and geo-
graphic isolation. Organisms that disappeared during more extensive glacial cover
have been unable to recolonize the continent. The milder subantarctic zone has
higher plant diversity, with 24 graminoids, 32 forbs, 250 mosses, 150 liverworts,
and more than 300 lichens. The northern Antarctic zone has only one graminoid,
one forb, 75 mosses, 25 liverworts, and 150 lichens. Only two species of flowering
plants, Antarctic pearlwort and Antarctic hairgrass, grow on the continent, as far
south as 68 12
0
S. Sunny north-facing slopes are the only areas that support turf-
forming mosses and vascular plants. Diversity in the southern zone is reduced to
30 mosses, one liverwort, and 125 lichens. No graminoids or forbs are found there.
Heat, wind, moisture, substrate, slope, nitrogen, and disturbance by birds and
mammals determine which plants actually grow at a particular site.
In the northern or coastal zonewhich includes the northwestern Antarctic
Peninsula, adjacent islands, some islands in the Scotia Arc, and Bouvetyaonly
one summer month averages above freezing. The southern boundary of the zone is
where the average temperature of the warmest summer month (January) fails to
rise above freezing. The coastal zone is fairly wet, especially in the north; it receives
at least 20 in (500 mm) of precipitation and regularly experiences relative humidity
over 80%. Although soils are shallow, frost action, solifluction, and accumulations
of peat are common. The west coast of the Antarctic Peninsula, where it is rela-
tively moist, supports 100 kinds of lichens, while the drier east coast has many
fewer. Two main communities exist in this predominantly maritime zone. A grass-
moss combination that may also include lichens is dominated by the only two flow-
ering plants in Antarctica, both limited to this zone. Sheltered north-facing slopes
lower than 350 ft (100 m) elevation and some flat areas protected by winter snow
support grass-moss or grass-lichen-moss communities. The several subdivisions of
lichen-moss association are based on different species and different percentages of
lichens versus mosses. Lichen crusts and fruticose lichens coexist with moss cush-
ions. Moss communities include a haircap moss and Chorisodontium aciphyllum turf
with peat accumulations up to 3.3 ft (1 m) thick, or thin moss carpets of Drepanocla-
dus, Brachythecium, and Acrocladium. Wet areas may have a moss carpet of Calliergi-
dium, Calliergon, and Drepanocladus. Fruticose lichens include Usnea species, while
foliose types include Xanthoria elegans and Umbilicaria. Granite moss and two crus-
tose lichens (Buellia frigida and Rhizocarpon flavum) may be found on rock surfaces.
The southern or continental zone is found in the southern and southeastern
parts of the Peninsula, on adjacent islands, and on other limited coastal areas. Sea
ice fringes most of the coast in winter, and some may not melt during summer. The
warmest month here never averages above freezing and may be as low as 5 F
(15 C). Interior locations are colder, with January (summer) means of 10 F
(12 C). This zone is drier, with only 6 in (150 mm) of precipitation, almost all
falling as snow. Although cold, the soil is too dry for patterned ground to form and
also lacks an organic layer. The brief growing season, strong winds, wind-blown
69
snow, long-lasting snow cover, and high salt con-
tent in the soil prohibit much plant growth.
North-facing slopes have more vegetation because
they receive more solar radiation, water from
snowmelt or glaciers, and nutrients from bird or
marine mammal droppings. Conditions are too
severe for most organisms, but cyanobacteria,
algae, lichens, and a few mosses survive. Liver-
worts are rare, and there are no flowering plants.
Although most of the surface is bare, crustose
lichens are found. Many lichens are endemic.
Others are either cosmopolitan or also found in
the Arctic. Under the best conditions, lichens
grow only 0.4 in (1 cm) in 100 years. Cryptoendo-
lithic lifeforms are microorganisms and crustose
lichens living within pores of rocks.
No land mammals, birds, reptiles, or amphib-
ians inhabit Antarctica, and no land mammals or
birds breed south of South Georgia in the suban-
tarctic. Terrestrial animals are limited to inverte-
brates such as rotifers, nematodes, water bears,
mites, and springtails. As many as 50 seabird spe-
cies, numbering in the millions, breed on the con-
tinent or islands in the summer, as do seals.
Pelagic or far-ranging birds feed at sea and include
albatrosses, fulmars, petrels, and shearwaters.
Among them are five species of albatross, 10 dif-
ferent petrels, and both the Arctic and Antarctic
Terns. Coastal species remain closer to shore and
include skuas, cormorants, terns, and sheathbills.
The skuas feed on land, stealing eggs from breed-
ing sea-bird colonies. Because of the scarcity of
snow-free ground, breeding takes place in large
concentrations. Huge populations of marine organisms such as zooplankton, ceph-
alopods, and fish support fur seals and southern elephant seals, which rest and
breed on islands, especially Scotia Arc. In the summer, most coastal ice-free areas
are intensely used by birds and seals, which transfer nutrients from sea to land in
their waste.
Of the 17 species of penguin in the world, only four breed in Antarctica: Adelie,
Emperor, Chinstrap, and Gentoo. Adelie and Emperor Penguins are most com-
mon. The others are found in the subantarctic islands, in southwest Africa, and
even at the Equator in the Galapagos Islands. All penguins are unable to fly. Their
shortened wings instead are used like paddles to propel them through water at
.................................................
Emperor Penguins
At more than 3 ft (1 m) tall and 88 lb (40 kg),
Emperor Penguins are the largest in the world
(see Plate VII). They eat fish, squid, and crusta-
ceans captured during deep dives, reaching
depths of 700 ft (200 m) or more lasting up to
20 minutes. Emperors breed on sea ice at the
edge of the continent during winter and have
no nests. Instead, they keep their eggs and
young warm by perching them on their feet
and insulating them with a special layer of skin
called a brood pouch. After laying a single egg
in May, the female returns to the sea, leaving
the male to incubate the egg in total darkness,
keeping it off the ice. When after two months
(the incubation period) she returns, the male
has lost up to one-third of his body weight and
now must walk up to 60 mi (100 km) to the
open sea to feed. After he returns, the pair
alternate regurgitating meals for the chick and
going back to the sea to feed. This takes place
until January when the chicks are mature
enough to walk to the ocean and begin inde-
pendent lives. Ice conditions are important to
successful breeding. If distance from the nest-
ing site to the ocean is too far, the penguins
may not have enough energy resources to feed
the chick. If the sea ice breaks up too early in
spring, the chicks may not be ready to survive
in the open ocean.
.................................................
70
speeds up to 25 mph (40 km/h). Their streamlined bodies have a thick layer of
blubber as insulation and heavy bones to help keep them submerged in ocean
water. Feathers are dense, forming a waterproof cover. Their diet is predominantly
krill, taken from shallow water. They can leap several feet out of the water to get
onto land or raised ice ledges. Although their waddling seems awkward, they can
walk several miles over rocks, ice, or snow to ancestral nesting grounds called rook-
eries. If the surface is snow-covered, they save energy by sledding downhill. Some
penguins are territorial, and their evenly spaced nests are a loose pile of small
stones on snow-free rocky headlands. The normal clutch is two eggs, but only one
chick may survive. Both males and females take turns incubating eggs and caring
for chicks. When they are three to four weeks old, the chicks join nursery group
huddles while both parents go to sea to feed. Parents feed their chicks by regurgitat-
ing into the beaks of their hungry offspring. While healthy adults have no land
predators, several birds prey on eggs and young chicks.
Research stations and bases constructed in the last 100 years have destroyed
much of the tundra, especially on the Antarctic Peninsula and adjacent islands.
The delicate ecosystem is easily disrupted by pollution, research, tourism, and
potentiallymining.
Further Readings
Antarctic Connection. n.d. http://www.antarcticconnection.com.
Arctic Portal. n.d. http://arcticportal.org/en/caff.
Dargaud, Guillaume. n.d. http://www.gdargaud.net/Antarctica/Penguins.html.
Environment Canada. n.d. http://www.mb.ec.gc.ca/nature/ecosystems/da00s04.en.html.
Ley, Willy. 1962. The Poles, Life Nature Library. New York: Time-Life.
National Oceanic and Atmospheric Administration. n.d. http://www.arctic.noaa.gov.
Polar Bears International. n.d. http://www.polarbearsinternational.org/about-us.
Smithsonian National Museum of Natural History. n.d. http://www.mnh.si.edu/arctic/
html/wildlife.html.
71
Appendix
Biota of the Arctic and Antarctic Tundra Biome
(arranged geographically)
North American Arctic Tundra
Some Characteristic Plants of Low Arctic Tundra
Treeline
White spruce Picea glauca
Black spruce Picea mariana
Tall shrubs (510 ft, 1.53 m)
Willow Salix alaxensis
Mountain alder Alnus crispa
Downy birch Betula pubescens
Low shrubs (20 in, 50 cm)
Bog birch Betula nana
Blue willow Salix glauca
Tea-leaved willow Salix planifolia
Heath family dwarf shrubs (48 in, 1020 cm) Ericaceae
Alpine bearberry Arctostaphylos alpina
Black crowberry Empetrum nigrum
Marsh Labrador tea Ledum palustre
Bog bilberry or Alpine blueberry Vaccinium uliginosum
Arctic bell heather Cassiope tetragona
Mountain azalea Loiseleuria procumbens
Graminoids
Nodding hairgrass Deschampsia flexuosa
Matgrass Nardus stricta
72
Red fescue grass Festuca rubra
Arctic bluegrass Poa arctica
Water sedge Carex aquatilis
Sedge Carex lugens
Few-flowered sedge Carex rariflora
Bigelows sedge Carex bigelowii
Common cottongrass sedge Eriophorum angustifolium
Tussock cottongrass sedge Eriophorum vaginatum
Highland rush Juncus trifidus
Forbs
Cushions or mats
Arctic mountain avens Dryas integrifolia
Mountain avens Dryas octopetala
Moss campion Silene acaulis
Leafy or rosettes
Viviparous bistort Polygonum viviparum
Marsh cinquefoil Potentilla palustris
Northern anemone Anemone parviflora
Cryptogams
Cetraria (Fruticose lichen) Cetraria spp.
Reindeer moss (Fruticose lichen) Cladonia rangiferina
Whiteworm lichen (Fruticose) Thamnolia vermicularis
Aulacomnium (moss) Aulacomnium turgidum
Haircap moss Polytrichum juniperinum
Sphagnum moss Sphagnum spp.
Some Characteristic Animals of Low Arctic Tundra
Herbivores
Caribou Rangifer tarandus
Arctic hare Lepus arcticus
Brown lemming Lemmus sibericus
Collared lemming Dicrostonyx groenlandicus
Arctic ground squirrel Spermophilus parryii
Carnivores
Brown or Grizzly bear Ursus arctos
Timber or Arctic wolf Canus lupus
Least weasel Mustela nivalis
Short-tailed weasel or Ermine Mustela erminea
Arctic fox Alopex lagopus
73
Birds
Willow Ptarmigan Lagopus lagopus
Snowy Owl Nystea scandiaca
Pomerine Jaeger Stercorarius pomarinus
Glaucous Gull Larus hyperboreus
Short-eared Owl Asio flammeus
Sandpiper Calidris spp.
Lapland Larkspur Calcarius lapponicus
Snow Bunting Plectrophenax nivalis
Canada Goose Branta canadensis
Some Characteristic Plants of High Arctic Tundra
Arctic willow Salix arctica
Bog bilberry or Arctic blueberry Vaccinium uliginosum
Graminoids
Mountain foxtail grass Alopecuris alpinus
Fishers dupontia grass Dupontia fisheri
Rock sedge Carex rupestris
Sedge Carex stans
Short-leaf sedge Carex misandra
White cottongrass sedge Eriophorum scheuchzeri
Three-flowered rush Juncus albescens
Northern woodrush Luzula confusa
Forbs
Cushions or mats
Arctic mountain avens Dryas integrifolia
Red sandwort Minuartia rubella
Tufted saxifrage Saxifraga caespitosa
Nodding saxifrage Saxifraga cernua
Purple saxifrage Saxifraga oppositifolia
Starwort Stellaria spp.
Leafy or rosettes
Flat-top draba Draba corymbosa
Lousewort Pedicularis spp.
Dwarf willow Salix herbaceae
Rooted poppy Papaver radicatum
Mouse-ear Cerastium alpinum
Cryptogams
Arctic finger lichen (Fruticose) Dactylina arctica
74
Snow lichen (Fruticose) Stereocaulon alpinum
White worm lichen (Fruticose) Thamnolia vermicularis
Dermatocarpon (Crustose lichen) Dermatocarpon hepaticum
Lecanora (Crustose lichen) Lecanora epibyron
Ditrichum (moss) Ditrichum flexicaule
Hylocomium (moss) Hylocomium splendens
Tomenthpynum (moss) Tomenthpynum nitens
Some Characteristic Animals of High Arctic Tundra
Herbivores
Pearys caribou Rangifer tarandus pearyi
Muskox Ovibus moschatus
Collared lemming Dicrostonyx groenlandicus
Carnivores
Polar bear Ursus maritimus
Timber or Arctic wolf Canis lupus
Birds
Rock Ptarmigan Lagopus mutus
Pomerine Jaeger Stercorarius pomarinus
Glaucous Gull Larus hyperboreus
Snow Goose Chen caerulescens
Some Characteristic Plants of Greenland Arctic Tundra
Greenland ash Sorbus groenlandica
Bog bilberry Vaccinium uliginosum
Mountain azalea Loiseleuria procumbens
Alpine bearberry Arctostaphylos alpina
Graminoids
75
Forbs
Cushions or mats
Leafy or rosettes
Some Characteristic Animals of Greenland Arctic Tundra
Herbivores
Caribou Rangifer tarandus
Arctic hare Lepus arcticus
Collared lemming Dicrostonyx torquatus
Carnivores
Timber or Arctic wolf Canis lupus
Birds
Wheatear Oenanthe oenanthe
Greenland Redpoll Carduelis flammea rostrata
Lapland Bunting Calcarius lapponicus
Arctic Redpoll Carduelis flammea hornemanni
Pink-footed Goose Anser brachyrhynchus
Barnacle Goose Branta leucopsis
Eurasian Arctic Tundra
Some Characteristic Plants of Fennoscandia Arctic/Alpine Tundra
Treeline
Norway spruce Picea abies
Low shrubs (8 in to 10 ft, 20 cm to 3 m)
Common juniper Juniperus communis
European blueberry Vaccinium myrtillus
True or Scotch heather Calluna vulgaris
76
Bog rosemary Andromeda polifolia
Small cranberry Vaccinium oxycoccus
Moss plant Cassiope hypnoides
Graminoids
Sheep fescue grass Festuca ovina
Spiked trisetum grass Trisetum spicatum
Tussock cottongrass sedge Eriophorum vaginatum
Forbs
Cushions and mats
Nodding saxifrage Saxifraga cernua
Leafy forbs
Monkshood Aconitum septentrionale
Globeflower Trollius europaeus
Dwarf willow Salix herbacea
Buttercup Ranunculus platanifolius
Alpine sorrel Oxyria digyna
Cryptogams
Mountain lichen (Fruticose) Solorina corceae
Iceland moss (Fruticose lichen) Cetraria islandica
Tree reindeer lichen (Fruticose) Cladonia mitis
Some Characteristic Animals of Fennoscandia Arctic/Alpine Tundra
Herbivores
Reindeer Rangifer tarandus
Norway lemming Lemmus lemmus
Tundra or Root vole Microtus oeconomus
Mountain hare Lepus timidus
Carnivores
Common shrew Sorex araneus
Lynx Felis lynx
(Continued)
77
Least weasel Mustela nivalis
Red fox Vulpes vulpes
Birds
Whooper Swan Cygnus cygnus
Bean Goose Anser fabalis
Mallard Anas platyrhynchos
Golden Plover Pluvialis apricaria
Red-necked Phalarope Phalaropus lobatus
Meadow Pipit Anthus pratensis
Wheatear Oenanthe oenanthe
Long-tailed Skua Stercorarius longicaudus
Snowy Owl Nyctea scandiaca
Short-eared Owl Asio flammeus
Rough-legged Buzzard Buteo lagopus
Golden Eagle Aquila chrysaetos
Gyrfalcon Falco rusticolus
Raven Corvus corax
Reptiles and amphibians
European common frog Rana temporaria
Common viviparous lizard Lacerta vivipara
Grass snake Natrix natrix
Some Characteristic Plants of Russian Arctic Tundra
Treeline
Downy birch Betula tortuosa
Chosenia Chosenia arbutifolia
Siberian larch Larix siberica
Siberian spruce Picea obovata
Mongolian poplar Populus suaveolens
Tall shrubs (2 ft, 0.6 m)
Resin birch Betula glandulosa
Felt-leaf willow Salix alaxensis
Alder Alnaster fruticosus
Mountain alder Alnus crispa
Dwarf shrubs (6 in, 15 cm)
78
Mountain crowberry Empetrum hermaphroditum
Bog bilberry Vaccinium uliginosum ssp.
microphyllum
Polar willow Salix polaris
Graminoids
Fishers dupontia grass Dupontia fisheri
Siberian spike-seed grass Trisetum sibiricum
Tufted hairgrass Deschampsia caespitosa
Carex sedge Carex lugens
Tufted cottongrass sedge Eriophorum vaginatum
Forbs
Cushions or mats
Hawkweed-leaved saxifrage Saxifraga hieracifolia
Arctic saxifrage Saxafraga hirculus
Polar poppy Papaver polare
Leafy or rosettes
Snow buttercup Ranunculus nivalis
Sulphur-colored buttercup Ranunculus sulphureus
Long-stalked starwort Stellaria longipes
Cryptogams
Iceland moss (Fruticose lichen) Cetraria islandica
Gray reindeer lichen (Fruticose) Cladina rangiferina
Arctic finger lichen (Fruticose) Dactylina arctica
Snow lichen (Fruticose) Stereocaulon alpinum
Whiteworm lichen (Fruticose) Thamnolia vermicularis
Northern reindeer lichen (Fruticose) Cladina stellaris
Green dog lichen (Foliose) Peltigera aphthosa
Parmelia (Foliose lichen) Parmelia omphalodes
Biatora (Crustose lichen) Biatora
Lecidea (Crustose lichen) Lecidea
Ochrolechia (Crustose lichen) Ochrolechia
Aulacomnium (moss) Aulacomnium turgidum
Hylocomium (moss) Hylocomium alaskanum
Tomenthypnum (moss) Tomenthypnum nitens
Giant calliergon moss Calliergon giganteum
(Continued)
79
Drepanocladus (moss) Drepanocladus uncinatus
Tritomaria (liverwort) Tritomaria quinquedentata
Ptilidium (liverwort) Ptilidium ciliare
Some Characteristic Animals of Russia Arctic Tundra
Herbivores
Reindeer Rangifer tarandus
Amur lemming Lemmus amurensis
Brown lemming Lemmus sibericus
Tundra or Root vole Microtis oeconomus
Carnivores
Brown or Kamchatka bear Ursus arctos
Birds
Stone Plover Limosa lapponica
Rough-legged Buzzard Buteo lagopus
Arctic Skua Stercorarius parasiticus
White-fronted Goose Anser albifrons
Bean Goose Anser fabalis
Brent Goose Branta bernicola
Curlew Sandpiper Calidris ferruginea
Antarctic Tundra
Some Characteristic Plants of Antarctic Tundra
Graminoids
Antarctic hairgrass Deschampsia antarctica
Forbs (cushion plant)
Antarctic pearlwort Colobanthus quitensis
Cryptogams
Usnea (Fruticose lichen) Usnea aurantiacoatra
Usnea (Fruticose lichen) Usnea antarctica
80
Xanthoria (Foliose lichen) Xanthoria elegans
Umbilicaria (Foliose lichen) Umbilicaria decussata
Umbilicaria (Foliose lichen) Umbilicaria aprina
Buellia (Crustose lichen) Buellia frigida
Rhizocarpon (Crustose lichen) Rhizocarpon flavum
Haircap moss Polytrichum alpestre
Chorisodontium (moss) Chorisodontium aciphyllum
Drepanocladus (moss) Drepanocladus
Brachythecium (moss) Brachythecium
Acrocladium (moss) Acrocladium
Granite moss Andreaea
Some Characteristic Animals of Antarctic Tundra
Sea mammals
Fur seal Arctocephalus gazella
Southern elephant seal Mirunga leonina
Birds
Adelie Penguin Pygoscelis adelias
Emperor Penguin Aptenodytes forsteri
Gentoo Penguin Pygosceles papua
Chinstrap Penguin Pygosceles antarctica
Antarctic Petrel Thalassoica antarctica
Wilsons Storm Petrel Oceanites oceanicus
Antarctic Skua Catharacta antarctica
Wandering Albatross Diomedea exulans
Royal Albatross Diomedea epomophora
Snow Petrel Pagodroma nivea
Arctic Tern Sterna paradisae
Antarctic Tern Sterna vittata
Southern Fulmar Fulmarus glacialoides
81
3
Mid-Latitude Alpine Tundra
The term mid-latitude refers to the areas between the Tropic of Cancer and the Arc-
tic Circle and the Tropic of Capricorn and the Antarctic Circlethe regions
between 23
1
=
2
and 66
1
=
2
in both hemispheres. Mid-latitudes in general are char-
acterized by seasonality of temperature due to a marked change in sun angle and
length of day from summer to winter. Mid-latitude alpine tundra is the life zone on
mountains above treeline. Alpine tundra of most Eurasian and North American
mountains is similar to that of the Arctic but with a richer flora. Limited in extent
in the Southern Hemisphere, mid-latitude alpine life zones have growthforms and
environments similar to those in the north, although the plant species present are
quite different. Small areas of alpine environment on mountain summits are diffi-
cult to illustrate on a world map.
Although it is often stated that altitude mimics latitude, many important differ-
ences exist between arctic and alpine environments. Both regions have low mean
annual temperature and short growing seasons. Both experience a seasonal change
in sun anglehigher in the summer and lower in the winterwhich contributes to
seasonality of temperature. The light regime differs, however. Alpine regions never
experience 24 hours of day or night, and duration of daylight or darkness varies
according to latitude. Although permafrost occurs on high mountains, arctic land-
forms do not occur because few large, flat areas on which they could develop exist.
Disturbances like avalanches and rock slides are common in steep mid-latitude
mountains but are rare to nonexistent on level expanses in the Arctic.
83
Mid-Latitude Alpine Climates
All alpine areas have climate characteristics in common, many of which are inter-
related. Most important are atmospheric pressure, temperature, and moisture con-
tent of the air. The density of air decreases as altitude increases. At 8,550 ft (2,600 m),
air pressure is 26% lower than at sea level, and at 19,000 ft (5,800 m), which is close
to the upper limit of vascular plant life in the Himalayas and the Alps, it is 50%
lower. Less dense air means less carbon dioxide for plants to use in photosynthesis.
Temperature decreases with elevation at an average rate of 3.5 F for every 1,000 ft
of elevation (6.5 C per 1,000 m), known as the average lapse rate. Actual lapse
rates, however, vary depending on inland or coastal location, latitude, and
weather. In spite of sometimes high relative humidity readings, alpine air is usually
very dry. Little water vapor can be held in the thin air, and low temperatures inhibit
evaporation. Temperatures at ground level, however, are higher, which increases
both evaporation and relative humidity.
Few long-term climate records exist for mid-latitude areas above treeline. In
general, soil and air temperatures are low, and the growing season is 410 weeks
long. Niwot Ridge west of Boulder, Colorado, is typical of alpine climates in conti-
nental areas (see Figure 3.1). At standard weather station height of 5 ft (1.5 m), the
mean annual air temperature at 12,300 ft (3,750 m) is 25 F (3.7 C) and mean
July temperature is 46 F (8 C). Temperatures in the Sierra Nevada are slightly
warmer because of a dry summer, while alpine zones in maritime regions such as
Mount Rainier in the Cascade Range have both more precipitation and moderate
temperatures. Temperatures closer to ground level and extremes during the short
growing season are more important for plant life than average temperatures. Plants
8 in (20 cm) high experience July temperatures ranging from 70 F (21 C) during
the day down to 39 F (4 C) at night. Soil temperatures to a depth of 8 in (20 cm)
are similar to the air above. Even without considering extremes, ground-level cli-
mates have a temperature range of 30 F (16 C) from day to night. Although low
temperatures are common during the mid-latitude growing season, freezing tem-
peratures are rare.
Macro-, meso-, and microclimates. It is hard to generalize a mountain climate, and
conventional climate statistics and descriptions are misleading. Alpine climates
can be described on three scales. Macroclimate refers to the general climate of the
geographic location, such as maritime for the North American Cascades and Euro-
pean Alps, continental for the Rocky Mountains, or semidesert for the Great Basin
of North America and Tibet. Mesoclimate takes into account variation with eleva-
tion, slope steepness, aspect, wind, and water, all of which are interrelated. For
example, a north-facing slope in the Northern Hemisphere will be shadier and
more humid, allowing forest, and consequently treeline, to grow at a higher eleva-
tion than on the hotter and drier south-facing slope.
84
Figure 3.1 (a) Niwot Ridge in the Southern Rocky Mountains of Colorado is typical of
mid-continental alpine tundra. (b) The higher precipitation and mild winter tempera-
tures of Paradise Inn on Mount Rainier in Washington illustrate a maritime influence.
(Illustration by Jeff Dixon.)
Rugged topography creates a mosaic of microclimates (climate in a small
area), which may change every few feet. Because meso- and microenvironments
may be different from macroclimate, elevation is not always the best criterion for
plant growth limits. Microclimate on a slope might override climate conditions of
elevation, so that a plant may be found higher than what is considered normal.
Both rocks and vegetation clumps affect wind and snow patterns. Snow often
accumulates in the lee of the obstruction, altering the microhabitat there. An
east-facing niche in the lee of a boulder on a windy west slope, for example, may
accumulate enough snow to protect a plant from temperature extremes. The high-
est vascular plant in the world, snow lotus (Saussurea gnaphalodes), can grow on
scree at 21,000 ft (6,400 m) in the Himalayas because the microclimate there is
similar to conditions at 13,000 ft (4,000 m) where it normally grows. The three
most important microclimatic factors are solar radiation, slope steepness, and
slope aspect or exposure (direction the slope faces). Others include wind speed,
air temperature above vegetation, and soil characteristics of structure, tempera-
ture, and moisture. At any scale, a change in one of these factors may affect
many others. For example, different amounts of incoming solar radiation will
cause differences in soil temperature, air temperature, snow cover duration, and
soil moisture, all of which affect vegetation. Microclimate diversity permits high
biological diversity.
Radiation and temperature. Because of the thin
atmosphere at high elevations, alpine areas are
often subjected to intense solar radiation. Sunny
and shady microclimates experience very differ-
ent temperatures. The shady area measures
actual air temperature in the thin atmosphere,
while the sunny area is directly warmed by the
suns rays. Diurnal temperature changes can be
extreme because of the thin atmosphere. Intense
solar radiation reaches the surface and is
absorbed by the ground, which becomes warm.
At night, clear skies allow much infrared radia-
tion to escape back through the atmosphere and
return to space, making nights cold. Increased
cloudiness limits the passage of both solar and
infrared radiation, narrowing the temperature
difference between both day to night and micro-
climates. A snow cover will reflect much of the
solar radiation during the day and also insulate
the ground, preventing terrestrial radiation from
escaping at night. Both processes reduce temper-
ature and diurnal temperature ranges.
.................................................
Solar Collectors
Energy from the sun is most efficiently
absorbed when the surface is at a 90-degree
angle to the suns rays. Solar collectors never lie
flat on the ground, but instead are inclined at
an angle to intercept the suns rays as directly
as possible. Because sun angle varies both with
latitude and season of the year, inclination of
the collectors must be aligned according to lat-
itude and be able to move seasonally as the
sun moves higher or lower in the sky. For even
more efficiency, they might also adjust
throughout the day to follow the suns path
through the sky from sunrise to sunset. Moun-
tain slopes in the alpine zone cannot adjust to
variation in the incoming solar rays, but plants
have adapted to the different habitats afforded
by different slopes and aspects.
.................................................
86
Slope angle and aspect. The steepness of a slope
affects the angle at which sunlight strikes the sur-
face. Even at a high latitude, a very steep slope
might receive close to a 90-degree sun angle,
increasing the amount of solar energy it absorbs.
The direction of exposure adds another dimen-
sion to temperature variation. Slope aspect affects
the time of day or year sunlight can reach the sur-
face. In mid-latitudes of the Northern Hemi-
sphere, the south-facing slope is sunny during all
daylight hours in both summer and winter. The
north-facing slope will only get sunlight on
summer mornings and evenings, and in winter
will receive none. The opposite is true for the
Southern Hemisphere: the north-facing slope is
sunny while the south-facing slope is in shadow.
An extreme example of temperature differences
according to slope aspect was measured in the
Austrian Alps. Surface temperature on the south-
west slope in summer was 176 F (80 C), while the northeast slope was only 73 F
(23 C). Sunny slopes experience more temperature change from day to night, and
snow melts sooner on sunny slopes.
Wind. In mid-latitudes, strong winds are usually associated with a position under
the jet stream or in the paths of cyclonic storms. Some mountains are windy while
others are not. Windiness is more related to topography than to elevation, being
strongest on exposed ridges. Winds are weaker at ground level where low-stature
plants grow. Wind redistributes snow, affecting the moisture gain or loss for an
area. Strong winds, especially in dry air, encourage sublimation, preventing water
gain from snowmelt, and also increase evapotranspiration from plants, both lower-
ing their internal temperatures and causing them to dry out. Wind affects heat
exchange by mixing layers of air, cooling the surface during day and warming it
at night.
In rugged topography, upslope and downslope winds alternate between day
and night. Intense warming during the day creates a low pressure, causing air to
rise. The resulting upslope winds frequently give rise to thunderstorms in the after-
noon. After sunset, the alpine air rapidly cools, creating a high pressure that causes
air to sink into the valleys below. Downslope winds at night, called cold-air drain-
age, can cause temperature inversions by displacing warm air, making valleys
cooler than the alpine environments at higher elevations. Such inversions may be a
nightly occurrence, literally turning the cooler-with-elevation norm upside down,
and can contribute to the development of microclimates.
.................................................
Surface Temperatures
Because the ground more efficiently absorbs
and loses heat than the air does, difference in
surface temperatures from day to night can be
extreme. That is a major reason why sand on
the beach is hot during the day. The ground
absorbs solar radiation all day, concentrating
the heat at the surface. It is then readily avail-
able to be radiated back to space at night,
causing the surface to be cold. Horizontal sur-
faces lose more radiation at night than vertical
surfaces lose. Frost may form on the sloping
windshield of your vehicle but not on the verti-
cal side windows because more radiation is lost
from the more-horizontal surface, making it
colder.
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87
Precipitation. Precipitation in mid-latitude mountains generally increases with
elevation all the way up to the summits. The potential for evapotranspiration
decreases with elevation, primarily because of cooler temperatures and shorter
growing seasons. Low temperatures and low evaporation indicate a wet climate
with plenty of moisture. Therefore, it would appear that most alpine areas should
have an abundance of moisture, at least on an annual basis. However, that rela-
tionship is misleading because all precipitation may not become available for
plant use. As much as 80% of that which falls in alpine areas ends up as runoff
from snowmelt or percolates through soil and rocks, eventually to reemerge in
rivers and streams. Therefore, alpine zones are often considered arid, a concept
reinforced by widely dispersed vegetation. Furthermore, the higher temperatures
experienced by small alpine plants close to the ground result in higher evapo-
transpiration during the growing season, often resulting in a water shortage by
summers end. Low annual precipitation totals recorded in some mid-latitude al-
pine habitats may be artifacts of measurement errors caused by few weather sta-
tions, strong winds, inaccurate determination of the water equivalence of
snowfall, or evaporation.
Snow protects plants from both wind and cold, and different plant communities
grow on windy snowfree ridges than those found in snow-filled gullies. Lack of
snow exposes plants to daily extremes of full solar radiation and nighttime infrared
loss. Measurements of microclimate temperatures in the Alps during winter show
that plants can be exposed to diurnal temperatures as high as 86 F (30 C) and as
low as 14 F (10 C). Wind and accompanying frigid temperatures often sculpt or
prune any parts of plants that project above snow level as evidenced by krummholz
at treeline. However, plants also have an effect on wind, decreasing its speed and
causing snow drifts to accumulate on their lee sides.
Soils
Because of low soil temperatures and uneven distribution of soil moisture, soil de-
velopment is slow. Microorganism activity is especially inhibited. Most alpine soils
are regularly disturbed by freezing, erosion, and downslope slippage and contain a
mix of large rocks and fine particles. Large rocks derive from onsite weathering,
glacial deposits, and landslides. Sources of fine material include weathering of
parent rock in place and from water, avalanche, and wind deposition. Dust, blown
up from adjacent lowlands or blown in from nearby glacial sediments, can add
nutrients not present in local rocks. Soils typically vary over short distancessand,
peat bogs, shallow carbonate soils, and deep profiles in alpine grasslandsand the
variation contributes to microhabitat development. Wind-blown material may be a
major component of grassland soils where limited water flow fails to bring in sedi-
ment. Cushion plants, abundant where it is windy, trap sediment and nutrients,
enriching both plant and soil. Fine material accumulates in spaces between large
rocks, where plants subsequently germinate and root. If soil, however, is buried too
88
deep below the rocky surface, as commonly hap-
pens on scree slopes, it fails to become warm
enough to support most plants. Except in mead-
ows and bogs, organic matter is extremely limited
in alpine soils because plant cover is sparse.
Frost action, either diurnal or seasonal, dis-
rupts soil formation, making it hard for plants to
colonize. Needle ice occurs almost nightly during
the growing season, uprooting small plants and
churning up seed beds. At a much larger scale on
a seasonal basis, frost-heaving can create pat-
terned ground and solifluction. Most large-scale
patterned ground in the alpine zone, however,
was formed during the Pleistocene and is no lon-
ger active.
At the beginning of the growing season, most
soils are saturated from snowmelt, and areas
where snow was deep or runoff accumulates may
be saturated all summer. Later in summer, after
the snow has melted, soils tend to dry out. How-
ever, plants are rarely subjected to drought stress.
While the top inch (2.5 cm) of soil may dry out
even in the wettest mountains, deeper soils retain
water that can be tapped by deep-rooted plants.
The Pamirs in the interior of Asia are one of the
driest alpine regions. Yet, even with less than 12
in (300 mm) of annual precipitation, little snow
cover, surface desiccation, and low relative hu-
midity, vegetation in the alpine semidesert at 14,000 ft (4,250 m) is not drought-
stressed. Measurements of leaf moisture loss and transpiration rates indicate that
plants are getting sufficient soil moisture.
Three major categories of soils occur in alpine areas. Poorly developed rocky or
stony soils (or entisols) are called lithosols when formed on bare rock of slopes and
ridges and regosols when formed on unconsolidated rock like talus or scree. Parent
material is the only difference, and both types are shallow, dry, and azonal, mean-
ing no defined horizons or layers are apparent. Plant communities on entisols are
usually dominated by lichens, mosses, and cushion plants. Alpine turf and
meadow soils (or inceptisols) are up to 30 in (75 cm) deep and exhibit distinct hori-
zons. The dark A horizon contains finer soil particles and has considerable organic
content, while the B horizon is lighter in color because it contains less humus. Both
turf and meadow soils support herbs and grasses with their mesh of intertwined
roots. Bog soils (or histosols) are found in depressions or where seepage saturates
ground to depths of 3.3 ft (1 m). Mossy bogs often overlie and insulate permafrost,
.................................................
Succulents in Alpine Tundra
Succulent plants are found in alpine regions of
all climate zones. Both in the crassula family,
mountain houseleek grows at 10,500 ft (3,200
m) in the Alps, and yellow stonecrop occurs at
12,250 ft (3,725 m) in the Rocky Mountains.
Also a member of the crassula family, a type of
hen and chicks, Echeveria columbiana is found
at 13,500 ft (4,100 m) in Venezuela. Cacti also
growhigh in the Andes. Several species of Teph-
rocactus, such as paper-spine cactus, which has
long, flat, flexible spines, occur above 14,000 ft
(4,250 m). Several alpine cacti, such as Opuntia
lagopus at 14,750 ft (4,500 m) in Peru, and many
others called old man cactus throughout the
high Andes, have protective coverings of long
white spines or hairs. C
3
plants dominate in al-
pine environments, and plants dependent on
C
4
or Crassulacean Acid Metabolism (CAM)
methods are rare. Treeline is generally the
upper limit of most C
4
plants, and succulent
CAM plants are restricted to hot or dry microha-
bitats. Most alpine succulents deal with surface
drought by storing water.
.................................................
89
which contributes to the poor drainage conditions. In the saturated soils, a lack of
oxygen inhibits decay of plant matter and makes bog soils acidic.
Plant Adaptations
Alpine growthforms are predominantly perennial herbs, graminoids, rosettes,
cushions, and low shrubs. Graminoids (grasses and sedges) are more typical of
moist habitats, while rosettes and cushions occur on windy ridges or areas that lack
protective snow cover. Cushions are prominent where low soil temperatures inhibit
elongation of stems. Small, compact leaves help maintain higher temperatures in
the plants. In an experiment growing the same plants at different elevations, indi-
viduals in the alpine zone were shorter and produced fewer branched flower stems
as well as fewer and smaller leaves than individuals of the same species grown at
lower elevations. Prostrate woody shrubs, either evergreen or deciduous are com-
mon, as are lichens and bryophytes. The alpine flora of any given mountain region
commonly has 200300 kinds of higher plants. Most have temperate origins and
are mainly members of the rose, pink, buckwheat, mustard, and saxifrage families.
Most perennials have more biomass in roots than in leaves and shoots, because
roots are essential to anchor the plant, absorb water and nutrients, and store the
carbohydrates necessary for rapid growth in spring. Annuals make up less than 6%
of the total alpine flora of most mid-latitude mountains; they need moist soil all
summer and bare sites with little competition, an unlikely combination in the al-
pine environment.
Most alpine plants can begin growing at temperatures just above freezing, while
lowland plants usually require temperatures of 4055 F (413 C). Early summer
growth draws on energy supplies stored in the root or tuber, which are then replaced
during the summer. Many plants have a red color due to anthocyanin pigments in
stems and leaves. Anthocyanins convert light to heat, which is especially important
in the spring and an important factor in making plants cold hardy. The red color
shows particularly at the beginning and end of the growing season; it is masked by
green chlorophyll in summer. Hairs on leaves or buds offer protection from strong
solar radiation and also conserve heat. Many plants have a natural antifreeze that
keeps the tissue from freezing when air temperature is below freezing.
Under deep snowpacks that limit the growing season, some plants preform
flower and leaf buds at the end of the previous summer season. Plants may have
microhabitat preferences around a snowbed, related to the average date of melting.
In the Rocky Mountains, for example, alpine avens, creeping sibbaldia, and Rocky
Mountain sage grow at the outer edges of a snowbed, while snow buttercup, which
tolerates a shorter season, grows toward the center. Zones where snow melts faster
not only have longer growing seasons but also more nutrient-rich soils. Humus
content is higher because plants grow for more summer days. Snowbeds are also
good sources of soluble nutrients such as potassium and calcium, which
90
accumulate over the winter and are released as the snow melts. However, the sur-
face beneath extremely late-lying snowbeds may be bare or support only mosses
and lichens.
Because most alpine plants are not constrained by water shortages, desiccation
is rare and only found in specific habitats. Plants have various means by which to
obtain moisture. Cushions, abundant in both humid microclimates and exposed
ridges, are commonly deep-rooted, and also trap water in their compact canopies.
Many lichens absorb their own weight in water, almost directly into the thallus
cells, a much quicker process than via roots and stems. Mosses also absorb water
like a sponge.
At the upper limit of vascular plant growth, small rosettes and cushions are
common growthforms. Wind-swept ridges support plants like saxifrages similar to
those found in the Arctic, but because more rocky crevices are available to trap
snow and provide more water, soft-tissued plants like buttercup also thrive. In wet
protected pockets called snow crannies, buttercups can grow at higher elevations
than even cushion plants. Glacier buttercup at 14,000 ft (4,275 m) is the highest
vascular plant in the Alps, and Grahams buttercup in New Zealand grows in the
permanent snow zone at 9,500 ft (2,900 m). In high-elevation snow crannies, snow-
melt provides moisture in otherwise dry sites while the surrounding rocks absorb
and radiate heat.
Reproduction
Alpine plants have three ways of maintaining their position within the community,
two by reproduction and one by being long-lived. Most plants reproduce sexually
even though conditions may not be favorable for completion of the cycle every
year. Vegetative reproduction is also important for most plants. Plants are slow
growing and may be 1015 years old before flowering.
The three patterns of flowering timesearly summer, mid-summer, and late
summeroverlap in mid-summer because of the shortness of the season. Early-
flowering plants do so at or shortly after the snow melts, mid-season plants flower
at the peak of vegetative growth, and flowers of late-season plants appear late in
the growing period after all leaves have been produced. All early- and mid-season
plants exhibit flower preformation. Inflorescences begin development in the previ-
ous season; how much is preformed depends on how soon in the summer season
the plant flowers. Glacier buttercup flowers early and may have two to three gener-
ations of preformed flowers blooming in a single season. In most mid-latitude
mountains, peak flowering is mid to late July. Late-season plants usually do not
have preformed flowers. Viviparous bistort, whose preformed flowers finish
blooming late in the summer, is an exception. It requires three seasons to form the
bud and does not flower until the fourth.
Temperature and photoperiod (hours of daylight) are two factors that exert the
most control over the time of flowering. Grasses such as alpine bluegrass must
undergo cold winter temperatures. Many plants need a long day in addition to one
91
or two cold winters. Very early bloomers, though,
such as primrose, buttercups, purple saxifrage,
and most carex sedges and woodrushes, are more
dependent on time of snowmelt than photo-
period. Late-season flowers such as alpine blue-
grass and alpine timothy respond to decreasing
photoperiod in late summer. Alpine flower displays
are known for their mosaics of colors and patterns.
Two different sexual reproduction strategies
are common and vary with respect to the number
of seeds produced and the type of pollination
undergone. Under favorable conditions, seeds
are abundant. Early-flowering species produce
fewer seeds but have a greater chance that their seeds will mature and ripen. Late-
flowering plants produce more seeds to increase the chance that some might sur-
vive to maturity. Early-season plants are more likely to be cross-pollinated, while
more late-season plants depend on self-pollination. The number of pollinators
does not decrease with elevation, although the spectrum changes. Butterflies and
beetles are replaced by bumblebees and flies. Bumblebees, however, are immobi-
lized at temperatures below 50 F (10 C). Above 13,200 ft (4,000 m) in the Hima-
layas, bee-pollinated flowers cannot grow, and plants depend on other insects. In
the Alps, 29 different insect families were observed visiting edelweiss. Humming-
birds are important pollinators in some warm-summer mid-latitude mountains
like the Sierra Nevada and southern Rockies, but none occur in polar mountains.
Wind can be important in dispersing pollen from one plant to another.
Germination and seedling establishment are difficult on surface soils prone to
desiccation and needle-ice disturbance. To survive, the seed must become firmly
anchored in deeper soil. Some seeds survive in crevices or in shelter of existing
plants, especially cushion canopies. Germination can be extremely rapid, within
one week after snowmelt. Few plants have seeds that require a dormant period.
Delayed germination until the following summer is environmentally imposed by
low soil moisture, ensuring that the plant will not germinate too late in the short
growing season. Seeds of plants growing in wet meadows, such as glacier lily,
snowball saxifrage, tufted hairgrass, and American bistort, do have a required dor-
mancy period. Even though there is enough soil moisture, these seeds require scari-
fication and do not germinate until the next season. However, seeds of a variety of
tea-leaved willow in the Rocky Mountains both ripen and germinate in July. In
contrast, short-fruited willow seeds ripen in August when the habitat has dried, pre-
venting germination until the next season. Other plants produce immature seeds
that require winter dormancy, not fully ripening until the following season.
High temperatures are required for successful germination. The lower threshold
for most plants is 40 F (5 C), but alpine sorrel requires at least 60 F (15 C). A di-
urnal cycle between cold and warm is usually necessary. Soil warming from solar
.................................................
Seeds
Alpine plants produce a lot of seeds. Average
output from one buttercup plant is 500, and
one alpine willowherb individual will produce
60,000 small plumed seeds. In Alaska, 1,155 via-
ble seeds were recovered in one square meter
of soil in alpine tundra. Such high seed produc-
tion usually is a hedge against high potential
mortality.
.................................................
92
radiation is important, but if surface soil becomes too hot or dry, seedlings die.
Because roots must reach a water source before the surface soil dries, growth dur-
ing the first year of most seedlings is mainly root development.
Because flowering, setting seed, germination, and survival of seedlings are diffi-
cult, most plants also reproduce by vegetative means. Such clonal development
takes many forms, including rhizomes, stolons, off-setting new rosettes, growing
adventitious roots from buried stems, and growth of bulbils or plantlets. The three
most dominant alpine families (sedge, grass, and sunflower) produce obligatory
clones, meaning that the new plants do not remain connected to the original plant
but become totally separate individuals.
Plants live longer at higher elevations, up to 3050 years for tall perennial forbs
according to age estimates based on slow growth rates or DNA. Yellow stonecrop
lives for five years, viviparous bistort for 26 years, and tussocks of kobresia for 200
250 years. Woody dwarf shrubs are also long lasting, 109 years for mountain cran-
berry. Acantholimon diapensoides cushions in the Pamirs are estimated to be 400 years
old. Other slow-growing tap-rooted species live for 100300 years. Some clones of
carex sedges, cottongrass, alpine azalea, or willow can be thousands of years old. In
NewZealand, Raoulia mats are hundreds of years old. The longevity of alpine plants
makes the ecosystems especially susceptible to environmental damage because
plants cannot quickly be replaced.
Animal Adaptations
Animals must contend with reduced oxygen and sometimes strong winds. The
reduction in oxygen content of the air is usually not a factor in limiting distribution
of animals in the alpine. Yaks are permanent residents up to 19,500 ft (6,000 m) in
the Himalayas. The guanaco of the Andes has more hemoglobin in its blood to
help bind more oxygen, and many animals have hemoglobin with an increased af-
finity for oxygen. Llamas more efficiently extract more oxygen from thinner air.
Although more red blood cells to carry more oxygen is not the norm for most al-
pine birds or mammals, it is a common acclimatizing response in lowland species,
like humans, when traveling to higher elevations. Several high-elevation animals
have higher heart and respiratory rates to circulate oxygen more quickly. Humans
who live at high elevations develop larger lungs and therefore more gas-exchanging
alveoli.
Unlike the Arctic, alpine tundra supports several burrowing or digging animals.
Small animals usually require a deep insulating cover of snow over their winter
homes. Meadow voles and water voles in alpine environments are restricted to lee
slopes and depressions where snow accumulates. Red-backed voles and heather
voles find protection under krummholz, while deer mice, pika, and mountain voles
prefer rocky areas. Although low-lying areas are important for snow accumulation
and insulation, those same areas are subject to flooding during snowmelt. Water
93
voles live in especially wet meadows and can even swim, but their burrows must be
elevated above flooding. Pocket gophers live in a narrow habitat zone between too
wet with flooding and too dry with insufficient snow cover. Soil from tunnels they
dig beneath the snow is left on the surface, exposed as long, sinuous piles of dirt
when the snow melts.
Ungulates in mid-latitude alpine environments tend to congregate in small
groups. Many are sure-footed and can maintain their balance on sheer rocky out-
crops and boulders. Chamois and ibex in Europe, thar in the Himalayas, llama in
the Andes, and mountain goat and bighorn sheep in North America, escape danger
by jumping and climbing steep cliffs. Large front hooves, long toes, or hooves with
gripping action are adaptations to their rocky habitat. Rocky areas, common in the
alpine, are preferred nesting sites for many small mammals because they provide
protection from powerful digging predators such as brown bear and badger. Mar-
mots, pikas, and golden-mantled ground squirrels select large rocks to burrow
beneath.
Breeding in alpine birds usually coincides with the peak of insect populations
during the short summer. Flies, stoneflies, beetles, moths, butterflies, and the para-
sitic sawflies and bees are abundant. Arthropods are generally small, enabling them
to hide easily in loose rock. Their dark color absorbs more sunlight, which is im-
portant in the cool air. Many have small wings or are flightless, a response to strong
winds. Others can withstand ice in their tissues or undergo supercooling so their
body tissue does not freeze.
Treeline
Treeline generally occurs at lower elevation with increasing latitude, from more
than 12,000 ft (3,600 m) in the Himalayas down to sea level in the Arctic, but more
factors are involved than just latitude (see Table 3.1). Treeline is not a strictly
defined line but a zone because microhabitats caused by late snowmelt, cold air
drainage, sunny ridges, avalanches, cold or wet soil, and rocky slopes all affect
growth of trees. Treeline is higher in both continental areas and larger mountain
masses such as the Rocky Mountains and Himalayas because they retain more
heat, making summers warmer. A combination of winter snow, summer drought,
and more bare rock in the Sierra Nevada depresses treeline. Treeline is also lower
in maritime areas, caused either by cooler temperatures due to ocean influence or
by heavier snowfall and shorter growing seasons. The Pyrenees, for example, with
both a smaller area and the influence of moist maritime airmasses, have a lower
treeline than the Rocky Mountains at similar latitudes. For the same reasons, per-
haps exacerbated by drought, treeline in the Southern Hemisphere is usually lower
than that of similar latitudes in the Northern Hemisphere.
The woody plants at treeline are predominantly evergreen because of the
growth advantage the evergreen habit affords during short cool summers. Pine is
94
Table 3.1 Mid-Latitude Treeline According to Latitude
MOUNTAIN RANGE LATITUDE
TREELINE
MAIN SPECIES FEET METERS
Northern Fennoscandia,
Norway
71 N sea level sea level White birch
Scotch pine
Norway spruce
Denali, Alaska, United States 64 N 2,500 750 White spruce
Black spruce
Central Fennoscandia,
Norway
61 N 3,600 1,100 Scotch pine
Norway spruce
Grampians, Scotland 57 N 1,600 500 Scotch pine
Northern Rocky Mountains,
Alberta, Canada
50 N 7,200 2,200 Engelmann spruce
Subalpine fir
Alpine larch
Western hemlock
Mt. Olympus, Washington,
United States
47 N 4,500 1,400 Mountain hemlock
Subalpine fir
Alaska cedar
Central Alps, Switzerland 46 N 6,500 2,000 European larch
Stone pine
Prostate pine
Norway spruce
Mountain pine
Carpathians, Romania 46 N 5,600 1,700 Prostrate pine
Norway spruce
Cascade Range, United States 45 N 5,000 1,500 Whitebark pine
Subalpine fir
Engelmann spruce
Mountain hemlock
Alpine larch
Mt. Washington, New Hamp-
shire, United States
44 N 5,000 1,500 Balsam fir
Black spruce
Paper birch
Central Rocky Mountains,
United States
Subalpine fir
Whitebark pine
Limber pine
Southern Alps, New Zealand 44 S 3,300 1,000 Southern beech
Pyrenees, Spain and France 42 N 7,500 2,300 White fir
European beech
Tien Shan, Kyrgyzstan 42 N 9,500 2,900 Central Asian spruce
(Continued)
the dominant family, with species of pine, spruce, and fir most prevalent, but hem-
lock and juniper are also present. Larch (also called tamarack), the only deciduous
conifer, has species in both western North America and in Eurasia. Hemlock forms
treeline in the Cascade and Coastal ranges of North America. Broadleaf deciduous
species are also common. Treeline in Fennoscandia is dominated by birch. Broad-
leaf trees are usually found where competition from conifers is absent, such as on
disturbed areas in avalanche tracks. Their flexible stems and ability to regenerate
from roots to form clones makes them ideally suited to the annual mass movement
of snow. Alder is found on moist slopes where avalanches occur in both the Alps
and British Columbia. Beech is dominant in central and southern Europe and the
Caucasus. Rhododendrons and junipers form treeline in Nepal. Rhododendrons
are trees in the Himalayas, not the shrubs familiar to Americans and Europeans.
Because of the physical and climatic continuity of North America and Eurasia,
especially at the height of the Pleistocene, many plants in Northern Hemisphere
mountains are similar throughout the mid-latitudes. Treeline is no exception, and
different species of common genera may occur in different regions. A relationship
between Clarks Nutcracker and whitebark pine in the Cascades, Sierra Nevada, and
the Rockies is matched by similar relationships in other areas, such as Spotted Nut-
cracker and Swiss stone pine in the southern European mountains, from the Alps to
the Caucasus, and Japanese Nutcracker and Siberian dwarf pine in eastern Asia. By
feeding on and burying the pine seeds, the birds unwittingly disperse the trees.
Conspicuous in Northern Hemisphere mid-latitude mountains is a belt of
stunted, crooked trees called krummholz (see Figure 3.2). Engelmann spruce, a
common treeline species, is a good example. When tall individuals grow densely
packed in the forest, the lower limbs die due to shade. However, isolated trees
Table 3.1 (Continued)
MOUNTAIN RANGE LATITUDE
TREELINE
MAIN SPECIES FEET METERS
Southern Rocky Mountains,
United States
Subalpine fir
Andean Steppe, Argentina 40 S 5,400 1,650 Southern beech
Sierra Nevada, California,
United States
38 N 10,800 3,300 Whitebark pine
Mountain hemlock
Sierra lodgepole pine
Foxtail pine
Himalaya Mountains and
Tibet
30 N 12,000 3,600 Himalayan fir
Himalayan pine
Western Himalayan
spruce
Drooping juniper
Lesotho Plateau, South Africa 29 S 7,500 2,300 Protea
96
maintain healthy lower branches that may then form adventitious roots where they
touch the ground. Over time, a small grove of cloned trees develops near treeline.
In slightly harsher conditions of cold and wind, the growthform changes. A strong
prevailing wind will either break branches on the windward side of the tree or bend
them toward the lee side, forming flag trees. With even more extreme condi-
tions, the plant becomes a shorter tangle of stunted limbs. Any shoot that grows
above the protective winter snow cover is naturally pruned by cold, dry winds.
Weight of heavy snow may also break exposed branches. The prostrate krummholz
vegetatively reproduces by rooting downwind where it is protected by the snow it
accumulates, while the windward side dies. A few trees, such as dwarf mountain
pine in the European Alps, Siberian dwarf pine in eastern Asia and Japan, and
some junipers, are genetically controlled forms that grow prostrate or dwarfed even
under nonalpine conditions.
Treelines in Southern Hemisphere mid-latitudes are more floristically diverse
than those of the Northern Hemisphere due to isolation and lack of convenient
migration routes. Most trees and shrubs are broadleaf evergreens. The most promi-
nent is the Southern beech, distributed in South America as well as Australia, New
Zealand, and as far east as New Guinea, even though it is not adapted to long-
distance dispersal. Deciduous species of southern beech occur in the drier moun-
tains of Patagonia and New Zealand. In southern Africa, treeline in the Drakens-
burg Mountains is a grassland and scrubland because of a dry climate and frequent
fires. The highest tree is a Protea.
Figure 3.2 Under severe windy and icy conditions, trees become stunted, forming
krummholz. (Photo by author.)
97
Habitats and Plant Communities
Alpine plants are usually limited to a specific environment, but plant communities
change from one to another over short distances, perhaps just a few feet, because of
the small-scale mosaic pattern of habitats (see Figure 3.3). A boulder field, also
called felsenmeerliterally a sea of rocksis a large accumulation of angular
rocks shattered by frost action. Rocks completely cover the surface, which may be
either fairly level or on a gentle slope. Boulder field communities are largely limited
to crustose and foliose lichens on the jumble of large rocks. Because of the consist-
ent growth rate of some lichenssuch as map lichen, which requires 1,000 years to
reach a diameter of 0.4 in (1 cm)boulder fields can be dated according to when
they were last covered by glacial ice. A few vascular plants may grow in pockets
between rocks where soil collects, snow accumulates, and the rocks both radiate
heat and provide wind protection. Pikas and marmots take shelter in boulder fields,
especially if close to a meadow for food.
Establishment of plant life on unstable talus and scree rock accumulations at
the base of cliffs and steep slopes is difficult. On talus slopes, the smallest rocks are
closest to the top of the cliff, while the largest are carried by gravity to the base. The
sharp, angular rocks of talus, a product of frost action, are as large as 6 in (15 cm)
in diameter. Scree refers to steep slopes of smaller rocks, down to gravel size.
Although apparently dry, damp soil may be found just 1 in (2.5 cm) beneath the
scree surface. Mountain avens, a circumpolar plant occupying the same habitat in
both arctic and alpine environments, is the best pioneer plant to stabilize scree.
With thick evergreen leaves covered by shaggy hairs on the underside and an elabo-
rate root system intertwined in scree, it survives downslope movement on windy
slopes where other plants would be ripped apart. Avens has nitrogen-fixing nodules
on its roots and enriches the soil with nitrogen. As branches accumulate dirt and
debris, humus piles develop beneath its leaves. Other plants on scree include
Easter-daisies, alpine milkvetch, buckwheat, and phlox. Roots are either a mesh of
shallow rhizomes or big taproots which provide security on the slope.
Figure 3.3 Plant growthforms and species change across environmental gradients. (Illus-
tration by Jeff Dixon.)
98
With 3550% rock, fellfields are flat areas usually remaining bare of snow due
to wind. Soil development between the rocks is poor because wind carries away
fine particles, leaving only shallow coarse gravels. Because of rapid drainage,
summers with little snowmelt, and wind exposure, fellfields are dry environments
where plants are exposed to extremes. Major growthforms are ground-hugging
cushions, mats, succulents, and rosettes, which may also be hairy or have waxy
cuticles. Typical plants, which bloom early in spring before any available moisture
is gone, include moss campion, alpine phlox, and sandwort. Few mammals live in
fellfields because forage is scarce.
Tussock-forming grasses, sedges, and forbs about 8 in (20 cm) tall grow in
meadows and bloom in summer because soils are moist. Lichens and mosses cover
the ground between higher plants. Sedges are dominant but both sedges and grasses
are present and both extend their roots 1 ft (0.3 m) or more into the deep soils, the
best on the alpine tundra. Because leafy forbs are abundant, meadows are some-
times referred to as herb-fields.
Snowbeds develop where snow accumulates in the lee of wind obstructions,
such as plants or rocks, or in nivation hollows that develop where snow accumu-
lates on a slight slope (see Figure 3.4). The weight of the snow pushes saturated soil
downslope into a low ridge, creating a shallow bowl or depression that then accu-
mulates more snow and intensifies the process. The pattern of late-melting snow
causes concentric rings of plants to develop, reflecting the time of snowmelt and
length of the growing season. Plants in winter are protected from intense cold and
temperature changes by deep snow. Temperatures in snowbeds are warmer com-
pared with exposed fellfields. Snow also provides protection from drying winds.
Dust and debris particles that collect on the snow surface add to soil development
when the snow melts. The pink tinge of old snow is caused by green algae encased
in a reddish coating that survives at melting snow temperatures of 32 F (0 C).
The snow surface attracts flies, spiders, mites, and springtails where they feed on
Figure 3.4 Snowbed communities typically have rings of different plants, each grow-
ing according to date of snowmelt and length of growing season. Snow in the center
may not melt all summer. (Illustration by Jeff Dixon. Adapted from Zwinger and Willard
1972.)
99
pollen and spores. Insects in turn attract birds. However, snowbeds also have detri-
mental effects, such as sheer weight of the snow, lower summer soil temperatures,
too much water, solifluction, and a shortened growing season. Snowbed indicator
plants are widespread. Snow liverwort, dimpled lichen, and flake lichen all serve to
bind particles and stabilize soil. Scandinavians use the presence of creeping sibbaldia,
a snowbed plant, as an indicator of soggy soils that would make a poor roadbed.
Animal-disturbed communities develop most commonly in meadows and
snowbeds where soil is both deep and well drained enough for digging. In North
America, pocket gophers are most often the culprits. The gophers bare dirt
mounds are subject to needle ice, which causes further disturbance. The sites often
revert to fellfield vegetation with cushion plants. Water and wind erosion may fur-
ther degrade the soil, but soil disturbance can be beneficial because it increases po-
rosity, allowing rain and snowmelt to soak in. Meadow voles take over abandoned
gopher holes and tunnels. They eat grasses and sedges in wet meadows and
undergo three- to four-year population peaks. Active in winter beneath the snow,
they can destroy cushion plants if their numbers are high enough.
Behind solifluction terraces on permafrost, around lake edges, or beneath snow-
banks are wet marsh communities that remain green all summer because of the
abundant water supply. These areas, with mosses, lichens, sedges, and low wil-
lows, most closely resemble arctic tundra. They are high in organic matter and
peat, which insulates and preserves the permafrost beneath. Soil is thick, black,
and mucky on top with many roots. Below the surface layer, the soil is colored by
blue-gray clay or red-yellow iron stains. With humic acids and no oxygen, the soil
is acidic. A common plant is marsh marigold.
Heath communities are widespread in the northern Rockies in both the United
States and Canada and are also found in the Olympics, Cascades, Sierra Nevada,
and Europe (see Figure 3.5). Because they prefer cool cloudy weather, they are rare
in the southern Rockies where it is too dry and sunny. All heath plants are mem-
bers of Old World genera and grow in acidic and well-drained but moist soils. Pro-
tective winter snow melts early in summer. Blueberries, bilberries, bell heather,
Labrador tea, alpine azaleas, and rhododendron are common. Plants, many of
which are evergreen, are low growing with small leathery (sclerophyllous) leaves.
Their thick cover and short stature prevents establishment of most plants other than
Iceland lichen.
Pleistocene Migrations and Similarities of Floras
Of the approximately 1,000 arctic tundra species, 500 of them are also found in
Northern Hemisphere mid-latitude alpine. A few genera (bluegrass, draba, sand-
wort, cinquefoil) are also in the tropical alpine, and spiked trisetum grass can be
found all the way to Antarctica. Mount Washington in New Hampshire has 75 al-
pine species, almost all of which are arctic. In the Sierra Nevada with more than
100
600 species, only 20% are arctic; most others are related to desert lowlands, the
Rocky Mountains, or the Cascades. The degree of relationship between the Rocky
Mountain flora and that of the Arctic depends on location, with areas farther north
having more arctic elements. In the Wrangel Mountains in Alaska, 70% of the
plants are arctic, while in the Sangre de Cristo Range in the southern Rockies, only
32% are. Eurasian mountains also show a relationship with arctic plants according
to latitude. In Fennoscandia, 63% of species are circumpolar. The Swiss Alps and
the Altai Range have 35% and 40% arctic species, respectively.
Central Asian Mountains (Altai, Himalayas, and Karakoram) have large and
distinctive floras because they were not completely covered by glaciers, and thus
have had a longer time for speciation to occur. The Himalayas are believed to have
Figure 3.5 Heath communities, such as in

Oztal, Austria, in the European Alps, are
characteristic of cloudy conditions and moderate temperatures. (Photo by author.)
101
been a Pleistocene refuge, and many alpine plants may have originated there. Of
the more than 1,000 alpine species in the Himalayas, only an estimated 2530%
are also found in the Arctic.
Arctic to Rockies to Andes
The Pleistocene era probably contributed more to the current distribution of arctic
and alpine plants in North America than any other factor. Many species in both
biomes are identical, and many more are closely related members of the same gen-
era. Many plants originated in the Arctic and migrated to the Rocky Mountains,
while others originated in mid-latitudes and followed the retreat of cold climates
northward at the close of the Pleistocene. These migrations were facilitated by the
north-south orientation of mountain ranges and their continuation into South
America. Even though the mountain chains are not continuous, peaks were close
enough together to allow the passage of plants.
With the onset of the Pleistocene, temperatures in the Rocky Mountains and
other North American ranges decreased and glaciers formed on many peaks and
ranges. The arctic tundra vegetation spread southward along the axes of the ranges,
taking advantage of the cooler climate. Even though no glaciation occurred south
of the San Juan Mountains in southwestern Colorado, herbaceous tundra plants
were able to find suitable habitats all along the peaks in the Rocky and Sierra-Cas-
cade mountain chains all the way to the Andes and even Tierra del Fuego at the
southern tip of South America. At least 19 families of seed plants in the Colorado
alpine are also represented in the Andean paramo above 11,500 ft (3,500 m), and a
minimum of 20 genera are shared by both regions. However, at least 22 additional
families in the paramo are not represented in the Rockies, and few to none of the
same species occur in both the Rockies and the paramo. The greater variety of
plant families in the paramo and their absence in the mid-latitudes seem to indicate
that the general trend of migration was to the south, with little northward move-
ment. A partial explanation may be that in the late Pleistocene, parts of the Andes
were still being built by volcanic activity and suitable habitats were destroyed,
blocking northward migration. (See Chapter 4 for discussion of Andean paramo.)
Mid-latitude plants were also adapting to an alpine climate. Depending on the
time period, these adaptations may have taken place as the general land surface was
uplifted from lower, subalpine, or even desert elevations, or in response to the cool,
periglacial climate. With each glacial retreat, plantsboth the original arctic species
and the newly derived mid-latitude speciesfollowed the cooler climates north,
sometimes migrating from peak to peak. Arctic species most likely advanced and
retreated with each change in climate. Those species originating in the Rockies or in
the Sierra-Cascade ranges may have adapted to the alpine climate at any time during
the Pleistocene, some possibly in the last stages of the era, giving them less time and
hence opportunity to migrate north and become established in the Arctic. This may
partially explain why more arctic species grow in the mid-latitudes than vice versa.
102
North-South Migrations in Europe
The major difference affecting migration in Europe versus the Americas is the trend
of most of the mountain ranges. Unlike the north-south corridors offered by the Rocky
Mountains, the east-west ranges of Europe served as barriers to both northward and
southward migration. The flora of the European Alps is more closely related to the
mountains in central Asia than it is to the flora of Fennoscandia. Similarities in genera
and species do exist, however, between the Alps and the Rocky Mountains. One pos-
sible explanation is migrations from North America over the Bering Strait land bridge
and subsequently west to the Alpsa very long journey. Some species, such as moun-
tain avens, alpine sorrel, moss campion, alpine bluegrass, and nodding saxifrage, are
common to both the arctic and to the alpine areas of Europe.
During the Pleistocene, almost the entire British Isles were covered with ice;
tundra climate and vegetation developed at the edges of the ice sheet. As the ice
retreated and climate warmed, temperate vegetation from Europe invaded, push-
ing the alpine flora to the highlands. Ample evidence survives in bogs to show that
alpine vegetation was formerly much more widespread. Remains of alpine plants
are found buried deep beneath peat that has accumulated since the Pleistocene.
Adaptations of Lowland Species
Plants from some types of lowland environments may have rather easily adapted to al-
pine conditions as mountain ranges rose. Some nonmountain plants are early spring
bloomers that die back during the hottest part of summer. Hot steppes or deserts are
more likely to have species preadapted to life at high elevations because many of their
plants bloom in spring. The fact that Indian paintbrush is found in both southwestern
deserts and in the alpine tundra of the Rocky Mountains is a good example of the simi-
larity between desert and high-elevation floras and environments. It might be expected,
too, that boreal forest plants, adapted to cold temperatures, would also become mem-
bers of the alpine flora, but they live in the shade of the forest and are not exposed to
intense light. In many Asian mountains, the krummholz zone is poor in flora and
many bald peaks occur because forest plants failed to adapt to the alpine zone.
In contrast to slow uplift, a relatively rapid uplift would eliminate the local non-
mountain flora because of the lack of evolutionary time, thereby leaving the alpine
tundra habitat open to colonization by any appropriate immigrating alpine species,
or in their absence, leaving it species poor. In any case, the size and composition of
the alpine flora depend on several factors other than the rate of uplift, including age
of the mountain, proximity of nearest mountains, richness of flora on adjacent
mountains, prevailing winds, and bird migrations.
Regional Expressions of Mid-Latitude Alpine Tundra
North America
Although not exceptionally high, the northern Appalachians in eastern North
America have alpine areas (see Figure 3.6). Mountains in Labrador and on eastern
103
Ellesmere and Axel Heiberg Islands form the eastern fringe of the Canadian Shield.
These mountains can be considered both arctic and alpine. Most North American
mountains high enough for alpine environments, however, are in the western part
of the continent, where two main cordilleras are dominant. Although not always
directly on the coast but still with maritime influence, a coastal region extends from
the Alaska Range south to the Cascades and Sierra Nevada in California. Many
peaks are more than 14,000 ft (4,300 m). Denali (formerly called Mount McKinley)
in southwestern Alaska, at 20,298 ft (6,187 m), is the highest peak in North Amer-
ica. The Cascades are still-active volcanoes up to 14,000 ft (4,300 m) high that sit
atop a lava plateau at 5,000 ft (1,500 m). The Sierra Nevada is a recently uplifted
fault block mountain reaching elevations greater than 12,000 ft (3,600 m). Except
for the Olympic Range in Washington, most mountains directly on the coast in the
United States are lower, and few rise above treeline.
In the interior, an almost continuous stretch of mountains extends from the
Brooks Range in central Alaska south through the Rocky Mountains in western
Canada and United States to northern New Mexico. The Rockies are geologically
complex, with many peaks above 14,000 ft (4,300 m). The Great Basin, with its se-
ries of north-south-trending fault block mountains, lies between the western ranges
and the Rocky Mountains. Some summits rise more than 13,000 ft (4,000 m), and
even though relatively isolated, most of their flora resembles that of the Rocky
Mountains. The volcanic San Francisco Peaks complex in northern Arizona is also
an isolated mountain range between the Sierra Nevada and the Rockies.
Figure 3.6 Alpine tundra in North America is predominantly in the higher mountains
in the West. (Map by Bernd Kuennecke.)
104
All mountain ranges with alpine environments have been glaciated. Coastal
mountains in Alaska still have extensive valley glaciers, as do the Olympic Moun-
tains and several peaks in the Cascades, such as Mount Rainier and Mount Shasta.
The drier Sierra Nevada and interior ranges have only small glacial remnants.
Alpine tundra in North America is geographically widespread, each region or
range having a unique environment. They have much in common but also many
differences. All have low air and soil temperatures, high winds, intense solar radia-
tion, short growing season, and sometimes heavy snow. Geology and substrate
vary and include all kinds of rocks. Northern mountain alpine regions are similar
to arctic tundra where the ecosystems merge. Coastal mountains have frequent
clouds and fog, and consequently less solar radiation. Continental mountains have
fewer clouds and more solar radiation, but also intense summer thunderstorms.
Permafrost is most common in northern mountains. Widespread circumpolar or
arctic-alpine distributions are common, as are areas rich in endemics evolved from
surrounding lowlands. Limitations imposed by the alpine environment restrict the
number of species. The Colorado alpine environment supports about 300 species
of plants, the White Mountains in California have about 150, and east-coast moun-
tains have only about 100.
Because of prevailing westerly winds, the west coast and western sides of
mountains receive more precipitation. Most precipitation in the Sierra Nevada and
Cascades falls in the winter, with limited summer thunderstorms. The Cascades
receive heavy winter snow, while summers in the Sierra Nevada are especially dry.
All the interior mountain ranges, including those in the Great Basin, are suscepti-
ble to intense thunderstorms in the summer because of continental climate and sur-
face heating in surrounding valleys. On the east slope of the Rockies, especially in
Colorado, early fall snow is very wet and heavy because of moist air circulating
upslope from the Gulf of Mexico. While the same source of moisture may cause an
occasional light summer snowfall in the southern Rockies, it will more often gener-
ate thunderstorms. Summer snowfalls can provide much needed water to alpine
vegetation. In contrast, winter snow is dry, icy, and associated with strong winds.
The White Mountains in New Hampshire are not only in the path of cyclonic
storms from the west but also get maritime influence from the Atlantic Ocean.
Consequently, they are wet.
South of 60 N, permafrost is discontinuous and influenced by slope, aspect,
and drainage. Permafrost plays only a small direct role in alpine vegetation because
most plant roots are shallow and not restricted by the active layer, but soil disrup-
tion from frost action has indirect effects. Frost-heaving can affect soils in Colorado
to a depth of 12 in (30 cm).
Origins and plant communities. North American alpine regions have complex flo-
ras with a mixture of arctic-alpine, circumpolar, and lowland elements, with many
endemic genera and species. Each site has a complicated geologic history. During
the Cretaceous, North America continued to move northward into cooler latitudes,
105
eliminating frost-sensitive species. Circumpolar species migrated across the Bering
Strait land bridge both to and from Asia. Mountain building and glaciation isolated
areas, and Pleistocene glaciation pushed arctic species south. During postglacial
warming, arctic species and adapted lowland species were forced to retreat upward
to the alpine. Northern parts of both the Sierra Nevada-Cascade and the Rocky
Mountains are dominated by circumpolar and arctic-alpine elements, which
become less important farther south. Lowlands like the Columbia River Gorge,
currently barriers to alpine plants, were colder and passable during the Pleistocene.
Mountains in the Great Basin are isolated peaks surrounded by desert shrubland.
As such, they have fewer species, both because of isolation and small summits with
limited alpine habitat. Rocky Mountain species extend west across the Great Ba-
sin, perhaps because the Rockies are older and served as a biotic source area for a
long time period. Species from the Sierra Nevada, which is younger than both the
Rockies and Great Basin mountains, are not found far east into the Great Basin.
Herbaceous vegetation is mostly perennial and includes cushion plants, rosettes,
grasses, and sedges. Annuals are a minor component of arctic and alpine flora but
are more commonly found in alpine situations. Some 55 annual species occur in al-
pine North America, 47 in the dry summer Sierra Nevada in California, 10 in the
Rocky Mountains, and only two in New England. At least 10 annuals have both
an arctic and alpine distribution. Based on exposure, snow cover, and moisture, a
gradient from steep rocky slopes to the valley bottom forms a typical vegetation
profile (see Figure 3.3). Woody cushion plants such as moss campion and purple
saxifrage occupy the steep rocky fellfields exposed to wind and generally are snow
free. Drier meadows have a mountain avens turf, while moist areas farther down
the slope support hairgrass meadows. Where water accumulates in bogs sedges
dominate, with willow thickets at streamside. Late-melting snowbanks may be bar-
ren except for lichens and moss-covered rocks because the growing season is too
brief for vascular plants.
North American alpine animals. Although many plant species are shared by both
arctic and alpine areas, few mammals or birds are common to both. Alpine regions
have more diversity of mammals. In the Cascade Range in Washington and Ore-
gon, 47 mammals use the alpine environment, and 32 occupy Niwot Ridge in the
Colorado Rockies. Smaller alpine areas have fewer. Animals can be divided into
three groups: permanent residents, seasonal residents, and occasional visitors.
Rodents are plentiful, and although population cycles similar to those of lem-
mings in the Arctic do not occur, numbers do vary from year to year. Because small
resident mammals need protective cover, habitats with rock piles, krummholz, and
willow thickets have the highest species diversity. Burrowing animals other than
pocket gopher are not common due to rocky soils, but they are important for soil
enrichment. Their digging mixes and aerates the soil, adds nutrients from drop-
pings, buries plants and humus, and distributes seeds. Depending on geographic
location, some small resident mammals have narrowly defined habitats, while
106
others are wide-ranging. Pikas, deer mice, and voles remain active beneath snow in
winter. Mice and voles share communal nests to increase warmth. In contrast, mar-
mots hibernate to escape the winter cold.
Erroneously called cony, which refers to a hoofed animal in the Himalayas,
pikas are in the rabbit family but have small ears in response to the cold (see Figure
3.7). Widespread in North American alpine tundra, their territory can be recog-
nized by the red-orange color of jewel lichens that grow on rocks where the animals
perch. Pikas are solitary during the winter but mate in spring and raise the litter as
a pair. They are restricted to rock piles or rock polygons for protection from preda-
tors but need access to meadows for food. The furry soles of their feet provide trac-
tion as they scurry across rocky terrain. Pikas remain active beneath the snow in
winter, eating hay and seeds they harvested in summer. They cut meadow plants
and place them on rocks in the sun to dry, thereby preventing mold from spoiling
their food stores. Although grasses and sedges dominate, hay piles also include sev-
eral species of forbs. Opportunistic feeders, pikas will take anything edible. Because
their food has little nutrition, they must eat every hour. They also reingest fecal pel-
lets to gain the most nutrients. They conserve moisture by expelling almost crystal-
line uric acids, which can be seen as a white coating on the rocks.
Marmots and woodchucks, or groundhogs, are found only in the Northern
Hemisphere. They probably originated in the Himalayas, but migrated to North
America where they now live in many environments, including grasslands and al-
pine tundra. Marmots grow up to 24 in (60 cm) long and weigh 15 lb (6.8 kg). All
species spend summers storing fat for winter. They have thickly furred bodies,
short ears and legs, and a bushy tail. Strong claws enable them to dig burrows in
boulder fields close to meadows for food. Marmots will stay in their burrows when
biting insects are especially active on windless days. They are not fussy eaters but
prefer forbs over grasses and sedges. Rocks provide lookout stations for protection
against predators, where marmot whistles alert the colony to potential danger.
Marmots are social. Some live in groups of
510 animals usually consisting of one breeding
pair and its offspring, or one male with two to
three females. Females breed at two to three
years, bearing two to six pups depending on the
species. Marmots live about five years, and pups
remain together as a family group for two to three
years before venturing out on their own. New
males moving into a social group often kill any
pups fathered by former resident males.
Marmots have both large and small enemies.
They provide summer food for foxes, coyotes,
wolves, bears, and eagles, and they are also
hunted by humans. Mongolians and Native
Americans use the meat and fur.
.................................................
Groundhog Day
Marmots and groundhogs are true hibernators,
lowering their body temperatures to slightly
above freezing. The hibernation period varies
with species and location, but may be one-half
to two-thirds of the year. Even though in hiber-
nation, marmots and groundhogs do wake up
occasionally. Punxsutawney Phil, a groundhog
in Pennsylvania, is famous for predicting the
duration of winter based on whether or not he
sees his shadow when he comes out of his bur-
row on February 2, Groundhog Day.
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107
Kobresia, hairgrass, and other alpine sedges and grasses provide nutritious for-
age for elk, deer, and mountain sheep. Some alpine areas are currently used for
grazing of domestic stockcattle and sheep. If overgrazed by domestic animals,
the cropped tussocks cannot trap the snow, which is their main source of moisture
on dry, windy ridges and slopes, and subsequently die.
Few large carnivores are found in alpine tundra. Brown (or grizzly) bears are
now limited in distribution. Mountain lions and black bears are occasional visitors
in the summer but migrate to lower elevations in winter. Coyotes venture onto the
tundra in the summer to hunt rodents in wet meadows. Short-tailed weasels hunt
in boulder fields, slipping their slender bodies into rodent burrows. Because prey is
always available, weasels are active all year. Shrews feed on insects and other small
invertebrates.
White-tailed Ptarmigan is one of the few birds to occur in both arctic and alpine
regions. Only the male remains on alpine tundra all winter; the females winter in
willow thickets at treeline. Triggered by longer daylight hours in early May, males
occupy territories in which they are joined by the females. The birds usually pair
for life even though they separate after breeding and the female raises the chicks
alone. By necessity, the nest is a depression on the ground, usually hidden among
plants. Chicks and eggs are preyed on by nutcrackers, falcons, and weasels, and
also succumb to harsh weather conditions. Ptarmigan females often feign a broken
wing to lead an intruder away from the nest. Although ptarmigan is the only per-
manent resident, several breeding birds that winter elsewhere are common to most
of the North American mountains. The Water Pipit, found throughout the alpine
zone in western United States, winters in South America, but from late June to Au-
gust, it resides on the alpine tundra, eating a variety of insects. While some birds
fly long distances to avoid winter cold, others, such as Rosy Finches, just move into
lower elevations. Golden Eagle and Red-tailed Hawk are raptors frequently seen in
the west (see Figure 3.8). Only the Red-tailed Hawk is common on the east coast.
Other birds often seen in krummholz at the edge of western alpine zones are Stel-
lars Jays and Clarks Nutcrackers.
Figure 3.7 Both (a) pikas and (b) marmots are common rodents in rocky areas close to
meadows. (Courtesy of Shutterstock. Copyright: Serg Zastavkin and Sascha Burkard.)
108
As in the Arctic, cold temperatures restrict amphibians and reptiles, both of
which are rare in alpine tundra. Insects are plentiful and include major pollinators,
such as bumblebees and flies. Most prevalent are black flies, mosquitoes, craneflies,
and many types of tiny flies. Butterflies are sometimes carried by wind from lower
elevations. Grasshoppers, ladybugs, beetles, leaf hoppers, and spiders also live at
these elevations.
Coastal ranges. Treeline species in the coastal ranges, dominated by mountain
hemlock, subalpine fir, and Alaska cedar, are typical of the distinct maritime influ-
ence on the west coast. The alpine zone consists of various combinations of low
shrubs and perennials. The northern alpine zones have more affinity to arctic tun-
dra, while areas farther south are more similar to nearby alpine tundras. Windy,
snowfree ridges facing south or west are sites for fellfields or boulder fields where
rosettes, cushions, and prostrate shrubs with a ground cover of foliose lichens domi-
nate. Moister sites on moderate slopes with intermediate snow depths support
meadows of sedges and fescue grasses with small birch or willow shrubs. Alpine
Figure 3.8 Redtail Hawk is a frequently seen predator in alpine environments. (Cour-
tesy of Shutterstock. Copyright: Jemini Joseph.)
109
bittercress and creeping sibbaldia dominate snow-
beds and moist depressions, but an alpine turf of
mountain hare sedge with many flowering forbs,
especially chiming bells, may also develop. Drain-
age sites that get nutrients and water from snow-
melt upslope support sphagnum hummocks with
black crowberry and arctic willow. Fireweed,
tufted saxifrage, and many more species grow on
stabilized rocky slopes, but unstable scree slopes
are sparsely populated. Watered areas with good
drainage and protection from wind are covered by
low carpets of mountain heather.
Rodents, including deer mice, meadow voles,
and pikas, are the most common animals in the
coastal ranges. The hoary marmot is widespread
in Alaska and Canada, but except for the Olym-
pic marmot, which is endemic to the Olympic
Range, no marmots live in the coastal ranges in
the United States. The Mazama pocket gopher is
restricted to the coastal ranges and the Cascades. The mountain goat has been
introduced to the Olympic Mountains. With no natural enemies to restrict popula-
tions, its sharp hooves cause increased soil erosion and alteration of plant com-
munities. Dalls sheep is native to the Canadian and Alaskan mountains. Short-
tailed weasel and dusky shrew are the most prevalent small carnivores. Brown
bears are limited to the coastal ranges in Canada and Alaska.
Birds, such as White-tailed Ptarmigan, Horned Lark, Water Pipet, and Stellars
Jay are similar to those found in most North American alpine tundra. Golden
Eagle and Red-tailed Hawk are commonly seen raptors.
Cascade Range. The Cascades extend from southern British Columbia to northern
California. The western side of the range is maritime and wet, with rainfall equiva-
lents of 100 in (250 mm) a year, while the lee side is drier and more continental,
receiving 20 in (50 mm) or less of precipitation. Several large valley glaciers extend
to elevations below the alpine zone, especially in the North Cascades and on
Mount Rainier and Mount Shasta. Treeline varies with age of the volcanic peaks,
but is about 5,000 ft (1,500 m). Trees are those typical of coastal mountains, but
include whitebark pine and Engelmann spruce on the drier eastern side. Alpine
communities are poorer than those in the Rockies due to substrate conditions.
Soils, undeveloped because of recent volcanic activity, provide few nutrients, have
little humus, do not retain water, and can be abrasive to plant shoots and roots.
Much of the alpine zone on Mount Hood is covered with loose pumice. Both fell-
field and snowbed plants are sparse and small because volcanic soils are unable to
retain water.
.................................................
Olympic Mountains
Although not exceptionally highthe summit
of Mount Olympus is only 7,965 ft (2,428 m)
the Olympic Mountains are the wettest in the
United States with long and active valley gla-
ciers. Directly in the path of westerly winds
from the warm North Pacific Current, the
mountains have a mean annual precipitation of
200 in (5,000 mm). The lower slopes are
densely covered in temperate rainforest. The
northeastern slopes, however, are in a rain-
shadow and receive only 20 in (500 mm) annu-
ally. Because of abundant moisture, alpine
vegetation is dense and rich. Tall glacier lilies,
grasses, paintbrushes, and louseworts grow in
profusion in wet subalpine meadows.
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110
Good volcanic soils support black crowberry,
which helps stabilize steep slopes. Other typical
plants include cliff Indian paintbrush and Lem-
mons rockcress. Tolmie saxifrage and partridge
foot, a tiny shrub, often grow together on steep
scree slopes, helping to stabilize the loose volcanic
soil. Windy fellfield communities have few spe-
cies, mostly grasses, sedges, and rushes, with no
annuals, although prostrate pussypaws, Lyall
lupines, and alpine asters do occur. Bistort can be
found in more protected microhabitats. Newberry
knotweed is characteristic of open pumice areas
in the Cascades where it has no competition.
The Cascades in northern Washington, espe-
cially Mount Baker and Glacier Peak, are geologi-
cally complex and heavily glaciated, with a varied
environment due to different kinds of rocks. A
strong floristic affinity with the Arctic and mountains to the north and east exists,
with few endemics. Deep snowbeds have sedge mats with hairgrass and tiny par-
tridge foot shrubs. Herb communities on moist southern exposures are dominated
by lupine. Dwarf-shrub heath communities are dominated by kinnikinnik, western
moss heather, or mountain heather. Associated species vary with location and con-
ditions. Kobresia sedges form a dense cover in dry meadow communities. Boulder
fields are characterized by patterned ground and high floristic diversity.
Rodents in the Cascades have habitat preferences. Mazama pocket gophers are
common in grasslands, herb-fields, and wet meadows. Rock piles adjacent to
meadows, providing both protection and nearby foraging areas, are home to pikas
and yellow-bellied marmots. Water voles occupy willow thickets, and heather
voles take cover in krummholz. Dusky shrews are restricted to wet meadows,
whereas deer mice are found in rock piles, herb-fields, meadows, willow thickets,
and krummholz. Short-tailed weasels prey on rodents. Pacific winds often bring
storms that cover everything with ice for days or weeks, making foraging difficult.
Mountain goats have been introduced. Ptarmigan is present, and summer bird pop-
ulations are the same as those in the Rocky Mountains, including Rock Wren and
Clarks Nutcracker.
Sierra Nevada. A tilted fault-block mountain primarily composed of granite, the
Sierra Nevada range extends 400 mi (650 km) from northern to southern California.
It contains the highest peak in the contiguous United States, Mount Whitney at
14,496 ft (4,418 m). Precipitation falls in winter, with potential for high snowfall,
although annual totals are variable. The sandy soil is dry, with near-desert condi-
tions throughout most of the summer. Summer drought is responsible for the ab-
sence of many arctic and alpine plants more common in the Rockies. Timberline
.................................................
Mount Rainier
Dormant but still warm at its summit crater,
Mount Rainiers volcanic cone has been exten-
sively eroded by radiating valleys filled with al-
pine glaciers. Mount Rainier receives record
snowfalls, and glaciers cover a significant part of
the mountain. At Paradise Inn (elevation 5,550 ft
or 1,692 m), slightly above treeline, the average
annual precipitation is 116 in (2,945 mm), with
annual snowfall of 50 ft (15 m). In 197172 the
station received the world record of snowfall in
a single season, 90 feet (27.5 m). Glaciers, how-
ever, have been melting over the past several
years.
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111
species reflect the difference between the wetter western slope and the drier eastern
slope. Mountain hemlock is a major timberline tree on the west. On the eastern
slope, foxtail pine resembles bristlecone pine, a treeline species in the Great Basin,
in its age and gnarled appearance, and whitebark pine forms flagged krummholz.
The Sierra has the richest alpine flora in North America, but less than 20% of
the species are of arctic origin, compared with 50% in the Central Rockies. Another
17% are endemic and related to genera at lower elevations in California or the
Great Basin. A few large genera, such as milkvetch, lupine, cinquefoil, penstemon,
and buckwheat, account for many of the species and subspecies. High plateaus not
covered by ice during the Pleistocene (nunataks) have the richest flora because they
served as refugia. Almost all major Rocky Mountain alpine communities are pres-
ent on the unglaciated plateaus.
The Sierra flora has a relatively large component of annual species, predomi-
nantly on dry, sandy sites such as south-facing slopes. Several plant communities
are based mostly on moisture content but to some extent on substrate (see Figure
3.9). Wet marble areas support a dense meadow of short-hair reedgrass and other
grasses and sedges, with some prostrate shrubs of dwarf bilberry. Slightly drier
upland sites with more exposed soil have a saxifrage-pussytoes cushion plant com-
munity. Short-grass sedge and Nutalls sandwort grow with timberline sagebrush
shrubs in dry meadows. Widely spaced rosette and cushion plants dominated by
oval-leaved buckwheat and endemic clubmoss ivesia grow in alpine gravels. Plant
Figure 3.9 Granite fellfields in the Sierra Nevada are too dry to support many plants.
(Photo by author.)
112
cover on bare granite is sparse, primarily phlox, Davidsons penstemon, and ivesia.
Very arid sites are bare except for pussypaws and dwarf knotweed. Carson Pass,
where andesite lava flows replace granite, is the northern limit for many alpine spe-
cies. The andesite supports few plants except for alpine sorrel where snow melts
late. Snowfree andesite sites have plants more closely related to cold deserts at the
base of the Sierra. Common plants are bottlebrush squirreltail, Great Basin violet,
and prickly gilia.
In addition to widespread rodents, such as pika, deer mice, and heather vole,
the Sierra is home to the endemic alpine chipmunk and Sierra pocket gopher.
Yellow-bellied marmots take up residence in rock piles that provide protection
from predators. They feed in nearby meadows. Mountain bighorn sheep are occa-
sionally seen. Bird life is similar to that found in other North American mountains,
including the White-tailed Ptarmigan (which was introduced).
Formerly widespread, brown bears were exterminated from the Sierra. Moun-
tain lions and black bears can occasionally be seen, but the most common carni-
vores are small, such as short-tailed weasel and dusky shrew.
Great Basin. Alpine environments are limited to the tallest mountain ranges in the
Great Basin, such as the White Mountains, Ruby, and Snake Ranges. Summit
areas are small, and the number of plant communities is low. Because the moun-
tain tops are geographically isolated both from one another and from larger ranges,
alpine communities have evolved separately as lowland species adapted to chang-
ing climate conditions. Many plants are endemic. The western region, in the rain-
shadow of the Sierra Nevada, is dry. The east has heavier winter snow and more
summer thunderstorms. With the exception of a few late-lying snowbeds or peren-
nial streams, most habitats are dry. Great Basin ranges have not been well studied,
but the following generalizations can be made. Mountains east of Elko, Nevada,
are more closely related botanically to the Rockies. Alpine floras on the White
Mountains and other western ranges are distinct, even though they share several
plants with the Sierra Nevada. Treeline is frequently hard to define because of arid-
ity, and the lower limit of the alpine zone is sometimes defined by the absence of
woody big sagebrush rather than trees. Wetter areas have a krummholz of white-
bark pine, Engelmann spruce, limber pine, or subalpine fir, while rocky or dry open
areas have a sparse cover of various grasses. Snow accumulation areas support forb
and graminoid plants like mountain hare sedge.
The Ruby Mountains have more precipitation, greater diversity, and greater
similarity to the Rockies. Meadow sites are dominated by arctic willow, elk sedge,
marsh marigold, and American bistort. Cushions of alpine avens and moss cam-
pion account for most of the cover in dry fellfield or rocky ridges. Moist soil around
ponds supports Rocky Mountain sedge.
The White Mountains, named for the color of the rock, not snow, are cold and
dry with sparse alpine vegetation. Patterned ground, a relic of the Pleistocene, is
dramatically apparent with little vegetation to hide it. Although the highest ranges
113
in the Great Basin show effects of alpine glaciation, the White Mountains were too
dry for snow to accumulate. Without glaciation, the alpine landscape remains roll-
ing uplands. Of the 200 species in the alpine flora, 62% are also found in the Sierra
Nevada, but only 28% also occur in the Rockies. Plant distribution is closely
aligned with rock type. Except for stands of bristlecone pine, dolomite (a type of
limestone) substrate is dry and barren with little plant cover except mats of Cov-
illes phlox. Granite fellfields have more cover and are dominated by Mono clover.
Bottlebrush squirreltail, of cold-desert origin, grows on both substrates, but is more
abundant on granite. Three speciesphoenicaulis, Masons Jacobs ladder, and
rambling fleabaneare restricted to elevations above 13,000 ft (4,000 m). Many
plants are hairy in response to harsh winter and summer conditions. Because the
White Mountains receive so little rain and snowfall, no snowbed or marsh com-
munities are found.
Animal life in alpine tundra of Great Basin mountains is limited because of
smaller habitat areas. Pikas, deer mice, and yellow-bellied marmots are common.
Grasslands and herb-fields also support northern pocket gopher, an indication of
affinity with the Rocky Mountains. Isolated areas have rare groups of mountain
bighorn sheep. The short-tailed weasel is the common predator of rodents. The va-
riety in bird species is also less than both the Sierra-Cascades and the Rockies, but
......................................................................................................
Bristlecone Pine
As its scientific name Pinus longaeva implies, bristlecone pine is long-lived (see Plate VIII). The old-
est living tree, with 4,844 rings, was cut down on Wheeler Peak in the Snake Range in 1964. Pine
Alpha in the White Mountains is 4,300 years old. Interpretive sites exist on the White Mountains
and in Great Basin National Park on Wheeler Peak, but bristlecones grow on many high mountains
in the Great Basin, usually at elevations of 9,00011,500 ft (2,7503,500 m) where summer temper-
atures barely reach 50 F (10 C), placing it at the edge of the alpine zone. Unlike alpine plants that
are of low stature, bristlecones stand exposed to harsh weather conditionsbitter cold, strong
winds, blowing ice and snow, and intense radiation. Bristlecones can be large trees or krummholz.
Young trees are upright and straight, but old trees subjected to centuries of environmental abuse
are broken and gnarled, though still massive. Limestone or dolomite is the preferred substrate.
Needles, in bundles of five, can remain on the tree actively performing photosynthesis for 40
years; most other pines replace needles after only a few years. Because of its age and sensitivity to
drought conditions that alter the width of growth rings, bristlecones are used in dendrochronol-
ogy, the science of dating by means of comparing tree rings. Speculation and theories abound as
to why bristlecones are long-lived, but no consensus has been reached. Even old plants, 3,000 or
4,000 years old, continue to flower and set seed. Clarks Nutcrackers feed on bristlecone seeds
when other food is unavailable and may be responsible for helping to perpetuate the species by
caching seeds deep enough that they are protected from the weather, allowing them to germi-
nate and produce seedlings.
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114
Rock Wren, Horned Lark, Clarks Nutcracker,
Golden Eagle, and Red-tailed Hawk are seen.
Rocky Mountains. The most studied alpine
regions in North America are in the Rocky Moun-
tains, particularly Niwot Ridge and Trail Ridge in
the Front Range of Colorado. Because the cordil-
lera was an old and persistent migration route, the
Rockies have a rich alpine flora that is similar all
the way south to their terminus in northern New
Mexico. Alpine sorrel, an arctic species, is one of
the most common plants. Typical in alpine envi-
ronments, a mosaic of communities develops in
response to a variety of microhabitats influenced
by wind, water, snow, and rocks.
The Southern Rocky Mountains include many
ranges in Colorado and New Mexico with a vari-
ety of landscape and climate. Even though the
mountains have been carved by alpine glaciers, not
all of the preexisting uplands were dissected. Large
areas such as Trail Ridge in Rocky Mountain
National Park remain as undulating expanses
above treeline. The San Juan Mountains in south-
western Colorado, directly in the path of west
winds and cyclonic storms, get both heavy snows in winter and thunderstorms in
summer. The Sangre de Cristo range in New Mexico is isolated from the main cor-
dillera but has similar plant communities. Arctic plants are usually limited to wet
marshes in the Southern Rockies and are smaller than the same plants found in the
Arctic. Mountain avens and kobresia, common in the northern ranges and in the
.................................................
Glacial Water
Boulder, Colorado, located on the semi-arid
Great Plains at the base of the Colorado Rocky
Mountains, is unique among American cities in
that it gets almost half of its water supply from
a glacier. Early in the twentieth century, the city
purchased Silver Lake, where meltwater accu-
mulated before flowing into Boulder Creek and
through town. In the 1920s, the Burlington
Railroad promoted visits to the Arapaho Glacier
as a tourist excursion, drawing hundreds of
people from the Chicago area to play on the
snow and ice. Alarmed at the potential despoil-
ment of their water supply, the City of Boulder
purchased 3,869 acres of land from Roosevelt
National Forest, including the Arapaho Glacier
and its entire watershed, for $1.25 per acre. To
maintain purity, the watershed is now closed to
the public, and the surrounding alpine environ-
ment remains unspoiled. Permits may be
obtained for scientific studies, however.
.................................................
..................................................................................................................
INSTAAR
The Institute of Arctic and Alpine Research (INSTAAR) is an interdisciplinary institute in the Graduate School at
the University of Colorado. Students in many fields, including anthropology, ecology, geography, geology,
atmospheric, and oceanic sciences, study and conduct research on how the Earths surface is affected by natu-
ral and human-related processes. Programs focus on ecosystems, geophysics, and past global change. Faculty
have special expertise in polar and alpine research on such topics as arctic and antarctic hydrology, atmos-
pheric dynamics, alpine ecology and climatology, invasive species, arctic climate change and variability, geo-
chronology, and paleo-ecology. Understanding environmental processes is a prerequisite to dealing with
world problems, such as maintenance of water quality and consequences of long-term environmental altera-
tions. Field-based courses and research opportunities are available at the Mountain Research Station, 20 miles
west of Boulder, and the Institute maintains a meteorological station on Niwot Ridge at 12,280 ft (3,743 m).
..................................................................................................................
115
Arctic, are almost absent. The Central Rocky Mountains, extending from Colorado
to Montana, also have a variety of landforms and rock types, and communities vary
accordingly. Calcareous soils in Glacier National Park, for example, support com-
munities similar to dry sites elsewhere because limestone does not retain water.
Principal treeline species in the Rockies in the United States are Engelmann
spruce and subalpine fir. In Montana, limber pine grows on drier sites, while white-
bark pine occupies more moist sites. In the Canadian ranges, alpine larch and west-
ern hemlock grow with an understory of heath shrubs in wetter climates similar to
the Cascades. Treeline in Alaska and the Yukon resembles the transition from bo-
real forest to arctic tundra, where white spruce, black spruce, and tamarack
dominate.
Talus and scree slopes are stabilized by subsurface rhizome networks of several
forbs, including milk vetch, drabas, and ragworts. Plants in boulder fields include
many saxifrages and alpine sorrel. Because they depend on sparse snowmelt in
largely snowfree areas, plants in fellfields bloom early. Rosettes of alpine primrose,
cushions or mats of alpine phlox, alpine candytuft, moss campion, dwarf clover, and
many others dot the ground with colorful flowers (see Plate IX). Although sedge
meadows dominated by kobresia tussocks are limited to areas blown free of snow
during winter, exposing the plants to weather extremes, they contain more species
than any other alpine community. Other plants include one-flowered harebell, arctic
gentian, and several more tiny plants. About one-fifth of the plants are lichens, the
most abundant being Iceland lichen, which is widespread in windy, snowfree areas
in the Arctic and throughout the alpine in North America. One of the most common
plants in snowbeds is the snow buttercup, blooming at the edge of the snow as it
melts. Prostrate, creeping arctic willow also grows near snowbed margins. Tussocks
of tufted hairgrass, found in alpine regions in many parts of the world, form a
meadow-like community in the snowbed. Bistort and creeping sibbaldia are also
common. The center of the snowbed remains saturated all summer and supports
clumps of Drummonds rush. All snowbed plants grow in precise locations accord-
ing to the time of melt, which determines the length of the growing season.
Plants that flourish in soils disturbed by pocket gophers tunneling beneath the
snow cover include sky pilot, alpine avens, and alpine sage. Many of these pioneering
plants are large and showy, attracting bees and flies for pollination. Alpine marsh
environments with saturated soils and standing water are green spots in frequently
dry tundra. Marshes often occur where water flows out of a tarn. Although Rocky
Mountain sedge dominates, other sedges are also found. Succulent sedums such as
rose crown are common. Succulence is an adaptation to conserve moisture and is ben-
eficial because plants have difficulty obtaining water from cold acidic soils. Marsh
marigold and cottongrass are common in wet areas. Several rushes grow in the grav-
elly streams, and several types of willow grow in thickets along stream channels.
The Northern Rocky Mountains in Alberta are primarily sedimentary or meta-
morphosed sedimentary rocks. Vegetation varies widely according to geographic
location, frost action, microhabitat, and substrate. In contrast to the southern and
116
central Rockies, the dominant growthforms are dwarf shrubs, some evergreen such
as bell heather, and others deciduous species such as willows. Moist areas support
grasses and sedges, while rosettes and cushion plants occur on drier ridges. Rocky
and heath tundra, the driest sites, are dominated by mats of mountain avens, arctic
willow and arctic bell heather, with rosettes of alpine locoweed. Several mat-
forming plants more typical of arctic climates include black crowberry and moun-
tain heathers. Elk sedge and mountain hare sedge tussocks form meadows.
Meadow and snowbed plants have the shortest growing season but need both win-
ter protection and summer water.
Arctic and alpine zones in Alaska and the Yukon merge, and the alpine environ-
ment and flora resembles that of the Arctic. The east-west trending Brooks Range in
Alaska at 69 N is a climatic barrier preventing northern airmasses from penetrating
farther south, separating arctic tundra from boreal forest in interior Alaska. A spruce
treeline exists on the south slope, but there is no forest on the north. A mosaic of veg-
etation exists according to slope steepness, aspect, soil depth, and texture. The high-
est exposed elevations have a reindeer lichen turf with moss. Rocky summit areas,
boulder fields and fellfields, are limited to primarily northern woodrush and dotted
saxifrage. The most common community on drier ridges is composed of sparse cush-
ions and mats, 90% of which are mountain avens. Deeper soils downslope have
downy birch, willow, and heath shrubs. Gentle slopes with fine soils, impeded drain-
age, and thick organic horizons have sedge meadows or cottongrass tussocks similar
to that found in Low Arctic tundra. Dwarf shrubs of marsh Labrador tea and bog bil-
berry also occur, and mosses account for 35% of the cover.
Several associations occupy snow accumulation sites. The center of the
snowbed with the shortest growing season has forbs of Bigelows sedge and bistort.
Areas with a longer growing season have arctic bell heather surrounded by a
lichen-heath community. Where snow melts first, a mountain avens and willow
fellfield develops.
Mule deer and pronghorn may be occasional visitors, grazing wet meadows,
willow, and krummholz. Bison and elk formerly occurred at high elevations in sev-
eral mountain areas, but most have been exterminated. However, elk have been
reintroduced in some places. Bull elk occasionally remain on the tundra all winter,
grazing in open meadows.
Mountain goats are native only to the mountains in southeastern Alaska,
Canada, and western Montana (see Figure 3.10). Mountain sheep are plentiful in
remote areas of the Rockies. The all-white Dalls sheep, distinguished by its thin
horns, ranges from Alaska to northern British Columbia, while mountain bighorn
sheep is found in the Rockies in the United States and occasionally in the Great
Basin. Most bighorn sheep in Colorado live on the eastern slope where snowfree
rocky crags provide winter forage. Rams gather a harem of ewes. Lambs, born in
May and July, are capable of jumping and climbing cliffs when only three days old.
Rodents are widespread. Dusky shrew, deer mouse, heather vole, and meadow
vole are found in a variety of habitats. The most common small mammal in moist
117
meadows is the meadow vole, in willow thickets it is the dusky shrew and water
vole, while krummholz is home to yellow-bellied marmot and heather vole. Pikas
live in rock piles. The collared pika occurs in the Yukon and Alaska, and the Amer-
ican pika occupies similar habitats in Alberta, British Columbia, and the United
States. The two most common marmots both live in the Rocky Mountains. The
hoary marmot, native from Alaska to northern Idaho, is replaced by the yellow-
bellied marmot in most of the Rockies in the United States. The Alaska marmot is
limited to northern Alaska.
Brown bears are still common in the remote Northern Rockies and as far south
as Glacier and Yellowstone National Parks in Montana and Wyoming. Short-
tailed weasels hunt rodents, and dusky shrews eat small invertebrates.
Bird life, including White-tailed Ptarmigan, Water Pipet, Rosy Finch, Horned
Lark, and Golden Eagle, is typical of most mountain areas in western North
America.
Although most amphibians and reptiles are restricted by cold temperatures,
some boreal forest toads reach high elevations in the southwestern Rocky Moun-
tains, as do a few salamanders in Colorado. Along with common pollinating
insects, such as bumblebees and flies, four species of ants inhabit the Colorado
alpine.
Mount Washington and other high peaks of eastern North America. The Appala-
chian Mountains in eastern North America are older than western ranges and have
Figure 3.10 Mountain goats are native to the northern Rocky Mountains. (Photo by
author.)
118
rounded topography and lower summits. Plants are more closely related to the Arc-
tic than to the Rocky Mountains. Arctic flora that had been pushed south in
advance of the Pleistocene ice sheets took refuge upslope as the climate warmed
and glaciers retreated. Alpine environments now exist only on isolated summits
that were covered by continental ice during the Pleistocene, including Mount
Katahdin (Maine), the White Mountains (New Hampshire), Green Mountains
(Vermont), Adirondacks (New York), and the higher peaks in Canada, such as the
Notre Dame Mountains on the Gaspe Peninsula and the highlands of southern
Labrador and eastern Quebec. Although elevations are not high compared with
mountains in western North America, cold, snowy, or foggy conditions depress
treeline to lower elevations. As in other alpine areas, the mean temperature of the
warmest month is below 50 F (10 C) and the summer growing season lasts four
to five months. Although dry microclimates exist, the overall climate is moist, with
snow or rain all year and many foggy days. Mount Washington, in the White
Mountains, averages 90 in (2,280 mm) of precipitation a year and frequently expe-
riences foggy or icy conditions.
Patterned ground that developed during the Pleistocene and is no longer active
can be seen on soils formed from crystalline rocks of schist and gneiss. Because of
extreme variation in weather conditions, krummholz or the treeline zone has a
wide elevational range, 4,8005,300 ft (1,4501,600 m), and stunted trees can often
be found in the shelter of large boulders. Several plant communities, typified on
Mount Washington, are present, reflecting the interrelated environmental factors
of fog, snow accumulation, and wind. Because of persistent foggy conditions, tree-
line species are distinct from those of both the western mountains and the Arctic.
Balsam fir dominates, but black spruce, found only in bogs in the Arctic, and
groves of paper birch are prominent. Flagged trees indicate strong prevailing wind
patterns. Even though windy, summits with heavy fog and deep snows are moist
all summer and support sedge meadows dominated by Bigelows sedge, haircap
moss, and Iceland lichen. Even porous sandy soils remain moist because of near
constant foggy conditions. Melting free by early July, snowbank communities
under heavy fog are the most species rich with many heath and willow shrubs,
along with nodding hairgrass.
Several different shrubby heath communities exist in areas with less moisture,
either due to fewer foggy days or less snow to provide meltwater; indeed, they are
the dominant vegetation type in the alpine zone of the White Mountains (see
Figure 3.11). Bog bilberry is the most common shrub in eastern alpine regions,
joined by mountain cranberry and sweet blueberry.
Other low plants such as Labrador tea shrubs, Bigelows sedge, highland rush,
and Greenland sandwort and Lapland diapensia cushions vary with locale. Club-
mosses grow protected beneath the shrubs. Windswept areas without snow have a
sparse cover of diapensia cushions, with highland rush and some low-growing bog
bilberry. Sunny streamsides protected from wind have marshy vegetation, shrubs
of bearberry willow, mountain alder, and three-toothed cinquefoil, along with
119
sedges and viviparous bistort. Bogs in similar locations are limited in extent and
dominated by Bigelows sedge. Because foggy conditions filter solar radiation and
moisture is available all year, these mountain areas have few hairy or succulent
plants. The dominant heath and lichen vegetation on Mount Washington closely
resembles that of the Arctic. About 70% of the vascular plants on Mount Washing-
ton are of arctic origin. Fruticose lichens, also typical of the Arctic, are more com-
mon in the cool, wet environment than crustose forms more prominent in drier
areas.
Animal life on high mountains in eastern North America is limited. Typical
species include deer mice, meadow voles, and groundhogs, solitary animals that
are also called woodchucks. Related to marmots, groundhogs grow to be 20 in
(50 cm) long and weigh less than 6 lb (2.7 kg). Like marmots, they prefer rocky
habitats adjacent to meadows. Short-tailed weasels are common, as is the masked
shrew. Frequently seen birds include Slate-colored Juncos and White-throated
Sparrows. Red-tailed Hawk is the most common raptor.
Europe
The east-west orientation of most Eurasian mountain ranges allowed few migra-
tion corridors to develop between mountain tops and the Arctic (see Figure 3.12).
As a consequence, Eurasian alpine communities contain fewer arctic plants than
Figure 3.11 Low heath shrubs protected among rocks are typical of Mount Washing-
ton and other eastern North America mountains. (Photo by author.)
120
similar habitats in North America. Most plants and animals, however, are close rel-
atives of North American forms. Even with few arctic species, dominant growth-
forms are the same as those in arctic tundra.
European Alps. The European Alps extend only a short distance north to south,
4448 N, but cover a 750 mi (1,200 km) arc from France in the west to Austria in
the east. The widest point is only 150 mi (240 km). Alpine conditions exist above
roughly 6,500 ft (2,000 m). The highest peaks are Mont Blanc in the west (15,771 ft,
4,807 m), Piz Bernina in the middle (13,284 ft, 4,049 m), and Grossglockner in the
east (12,457 ft, 3,797 m); all have alpine glaciers. Like the Himalayas, Andes, and
Rockies, the Alps are young mountains, dating from the Tertiary. High plateaus,
which form step-like levels at 5,900 ft (1,800 m), 8,200 ft (2,500 m), and 9,850 ft
(3,000 m), are old erosion surfaces produced during geologic pauses in mountain
uplift. During the Pleistocene, when ice almost completely covered the Alps, valley
glaciers sharpened the landscape and deposited extensive stony moraines in val-
leys. Calcareous bedrock and soils are characteristic of the outer chains, while
......................................................................................................
Extreme Weather
Although the summit of Mount Washington is only 6,288 ft (1,917 m), it is famous for weather that
can change from warm and sunny to freezing fog and gale-force wind within minutes. More than
100 people have died on its summit or slopes, many during the summer months. The extremes
result from a unique set of circumstances including a north-south orientation and a position
beneath the jet stream where three storm tracks converge. Wind speed in summer averages 25
mph (40 km/hr), increasing to 45 mph (72 km/hr) in winter. However, 100 mph (160 km/hr) winds
can be expected about every three days in winter, and the highest wind ever recorded on Earths
surface, 231 mph (372 km/hr) occurred on Mount Washington in April 1934. Buildings and large
instruments are chained down to prevent their loss in high winds. Even without wind, tempera-
tures are consistently cold. Mean annual temperature is only 26 F (3 C), with a winter extreme
of 47 F (44 C). Wind chill may lower the perceived temperature to 120 F (84 C). The
highest temperature ever reached in summer is only 72 F (22 C). Snowfall is high, averaging 255
in (650 cm) per year, even though it is often redistributed by strong winds. The maximum ever
recorded for one season was 566 in (1,438 cm). The summit is foggy more than 300 days of the
year. If the cold air is clean and lacks freezing nuclei (particles for ice to cling to), it will be super-
cooled, meaning that water droplets are unable to change state into solid even though air tem-
perature is well below freezing. When supercooled fog comes into contact with buildings or
instruments, the water droplets instantly freeze, coating the objects with a thick coat of rime ice
(see Plate X). Weather records were taken 18701892, but the summit was unoccupied until 1932.
An independently operated Mount Washington Observatory now houses scientists and volunteers
who take weather measurements and do research year-round on such diverse topics as ice physics
and cosmic rays.
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121
granitic materials dominate in the inner mountains. All soils are young, reflecting
characteristics of the substrate, and closely correlate with vegetation. The Alps
have been heavily influenced by human activities for thousands of years, so it is
difficult to say what makes up the natural vegetation.
The European Alps, as well as mountains such as the Pyrenees, Carpathians,
Caucasus, and northern Balkan Peninsula in Europe, rise between two biomes: the
Temperate Broadleaf Deciduous Forest to the north and the semi-arid Mediterra-
nean Biome to the south. No other mountains in the world occupy similar posi-
tions. Half of the mountain flora is derived from the southern and central Europe
lowlands, while the other half is dispersed from the north and east. No endemic
plant families and only one endemic genus (Berardia) are found in the Alps. How-
ever, bellflower, draba, lousewort, primrose, buttercup, saxifrage, and violet genera
contain many endemic species. Some endemics are distributed throughout the
Alps; others are concentrated in refugia. About one-half of the plants are restricted
in distribution, while the other half migrated from the Arctic or from central Asian
highlands since the Pleistocene.
Mountain climate varies from the outer regions to the inner valleys and also
from east to west, but generally has enough rain in warm periods for plant
Figure 3.12 Alpine tundra in Eurasia. (Map by Bernd Kuennecke.)
122
growth and is cold enough in the winter for plant dormancy. Both north and
south slopes are wetrain in summer and snow in winterwith annual precipi-
tation up to 100 in (2,500 mm). Parts of the central mountains have continental
climate characteristics with temperature extremes. Plant communities occur at
different elevation levels according to location, at the fringes or interior of the
mountains.
Treeline species in the outer regions are Norway spruce or prostrate pine. The
inner and central Alps have mixed forests of European larch and stone pine or for-
ests of mountain pine. Natural vegetation on steep slopes usually has not been dis-
turbed by human activities, but due to agricultural clearing for hundreds of years, it
is impossible to tell where natural treeline is in most of the Alps. Forests on steep
slopes are cut through by avalanches and rock slides. Huge treeless rock outcrops,
especially in the limestone areas such as the Dolomites of northern Italy also com-
plicate the issue of treeline (see Plate XI).
The alpine area has a lower zone with dwarf-shrub heaths, an upper zone with
dense, steppe-like grasslands or meadows, and a subnival zone. Some of the dwarf-
shrub heath in the lower zone may be undergrowth of forests that have been cut,
but there is no way to tell the difference between those remnants and alpine vegeta-
tion. Both are thickets with dense moss or lichen layers beneath. Dominant shrubs
may be rhododendrons, blueberries, or creeping azalea, depending on microhabi-
tat. Snowbed communities do not occur in the lower zone because snow melts too
quickly, but wetter areas support matgrass in their centers with dwarf shrubs such
as rhododendrons at the edges. Frost action prevents a closed vegetation cover on
exposed ridges.
The beginning of the upper zone is where alpine grasslands form dense carpets
and replace dwarf shrubs. The grasslands, however, are primarily sedges because
of dry habitat. Different species of carex grow according to the acidity of substrate.
Associated with them are rosettes with rhizomes, woody creeping mats, and tiny
dwarf shrubs. Fruticose lichens grow between sedge plants. On windswept ridges
without a protective snow cover, kobresia sedge replaces carex. Ridges may experi-
ence temperatures down to 58 F (50 C), which creeping azalea, purple saxi-
frage, and moss campion can withstand. In contrast, microclimate conditions in
summer can be close to 100 F (37.8 C). Houseleek grows where the surface is hot
and dry, but the succulent plant suffers no drought stress. On steep, sunny slopes,
fescue grasses replace sedges.
Under prostrate green alder, which grows mostly in avalanche paths, the soil is
rich in nitrogen, so forbs grow larger than normal. On disturbed areas such as
scree, which are susceptible to movement, major growthforms are creeping mats
and gravel stabilizers because loose rock surfaces provide only holes or crevices for
rooting. Spherical cushions of rock jasmine grow here, needing little room for root
growth. Lichens and mosses often occur in large numbers. Edelweiss can grow on
both scree slopes and in meadows. Norwegian hairycap moss occurs in the wettest
areas of snowbeds with the shortest snowfree period, while creeping willows grow
123
at the edges. Snowbed plants need protection from frost in winter, and many plants
cannot survive if the temperature drops too low. Sensitive plants have affinities to
the warm temperate or Mediterranean zones. Frost-tolerant plants usually have an
arctic or northern high mountains origin.
The subnival zone, a transition to the region of permanent snow, is character-
ized by patches of grassland and scattered cushion plants. Within this zone, plants
grow in sheltered, sunny niches. In

Otztal, Austria, the permanent snow line is
10,170 ft (3,100 m), but several plants grow as high as 11,480 ft (3,500 m). For-
merly thought to be the highest vascular plant in the Alps, glacier buttercup at
12,073 ft (3,680 m) was later discovered to be topped by two-flower saxifrage grow-
ing at 14,600 ft (4,450 m). Mosses, lichens, bacteria, and algae grow even higher.
Red deer and chamois are common herbivores; they may overpopulate some
areas and damage subalpine forest. In contrast, mountain ibex is rare and only
found in the Alps, although related species occur in the Pyrenees and Caucasus
Mountains. Ibex, formerly widespread, had been reduced to a small herd in the
Gran Paradiso area in northwestern Italy, but now has been successfully reintro-
duced to other parts of the Alps (see Figure 3.13). Marmots are the most conspicu-
ous rodent. Both snow vole and alpine shrew are endemic to the Alps. The snow
voles nearest relative occurs in Eastern Siberia. The presence of endemics illus-
trates the relative isolation of European ranges, while affinities with eastern Asian
species reflect east-west migration routes. Most of the larger carnivores, such as
Figure 3.13 Ibex, seen here near Gran Paradiso in the Italian Alps, are native to many
mountains of Europe and Asia. (Courtesy of Shutterstock. Copyright: ArturKo.)
124
brown bear, lynx, Bearded Vulture, and Golden Eagle, have been hunted to near
or actual extinction. Golden Eagles are now protected. Small predators include the
insectivorous alpine shrew and the wildcat.
The European Alps had already been extensively used prior to Roman con-
quest, with many permanent settlements built close to treeline. Summer grazing
took place on alpine meadows, and hay was harvested on slopes too steep for graz-
ing. Conversion of forest to pasture on steep slopes increased the frequency and
area of avalanches. Approximately one-half of current avalanches can be attributed
to human activities. European countries now use alpine areas in different ways.
Some continue traditional agriculture, while others have abandoned the land.
Tourism and recreation, especially the creation of ski runs, which destroys vegeta-
tion and soil, have had profound effects.
Central Asia
Except for the Himalayas and adjacent ranges, the high mountain systems of Asia,
are generally located in the southern part of the former United Soviet Socialist
Republic. The Carpathians, Altai, Sayanskiy Range, and Tarbagatay Ranges rise
......................................................................................................
Cairngorm Mountains
The highest peaks in the Grampian Mountains, including the Cairngorms, in central Scotland are
Ben Nevis at a mere 4,406 ft (1,343 m) and Ben Macdui at 4,295 ft (1,309 m). Despite the relatively
low elevation, however, remnants of tundra vegetation, relics from the Pleistocene, grow on many
slopes and summits in the range. Great Britain was almost completely covered by glaciers during
the Pleistocene, and ancestors of mountain plants survived conditions at the edges of the ice. Due
to a warm ocean current offshore, todays climate is moderate for the mid-latitudes and is similar
to maritime tundra on arctic coasts. Many of the alpine plants are also found in the Arctic. Birds
include several species, such as ptarmigan, that are also found on arctic tundra.
......................................................................................................
......................................................................................................
The Pyrenees
Separating the Iberian Peninsula from the rest of Europe, the Pyrenees Mountains are a barrier to cy-
clonic storms coming from the northwest. The French side of the mountains receives the brunt of
the storms and is cool and snowy, while the Spanish side in the rainshadow is warm and dry. The dif-
ference in climate is reflected in the vegetation. White fir and European beech forests cloak the
northern slopes, contrasted with shrubs and grasses on southern slopes. Both sides, however, have
deep glaciated valleys and active glaciers. Pico de Aneto, 11,168 ft (3,404 m), is the highest point.
Much of the flora and fauna are similar to the Alps, but many endemic species, such as Pyrenean sax-
ifrage, iris, and blue thistle, occur. The Pyrenean chamois, or isard, is slightly smaller than chamois in
the Alps. Marmots have been successfully reintroduced, and genetically similar Slovenian brown
bears were introduced to supplement the dwindling population of Pyrenean brown bears.
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125
above temperate forest, while the Tien Shan is
surrounded by deserts. The Tien Shan and Carpa-
thians have been most studied, the former being
more continental while the latter has more mari-
time influence.
Tien Shan Mountains. The Tien Shan, at 42 N,
are northeast-southwest trending ranges that form
the border between Kyrgyzstan and the Tarim Ba-
sin in China. Average elevation of the alpine zone
is 13,450 ft (4,100 m), and some peaks rise more
than 16,400 ft (5,000 m). Part of the range is a pla-
teau at 12,500 ft (3,800 m). Because all the high
mountains were glaciated and piedmont glaciers
covered the plateau, moraines and outwash are
common. Continuous permafrost exists above
10,800 ft (3,300 m). About 70% of the total spe-
cies in the Tien Shan alpine zone originated in the
Pamir-Tien Shan region or other mountains of
southern and central Asia. Arctic flowering plants
are only 1020% of the total and are more com-
mon in moist communities. Few plants have
affinities to mountain floras of southern Europe.
In this harsh environment, due not only to high elevation, but also relatively low
latitude and surrounding aridity, solar radiation is intense. Temperatures and length
of growing season are comparable to parts of arctic tundra. Air and soil temperatures
drop below freezing even on most summer nights. The lower limit of the alpine zone
is 9,500 ft (2,900 m), where mean July temperatures are 50 F (10 C) and mean Jan-
uary temperatures are 4 F (20 C). Growing season ranges from six months at
the lower limit to only one-and-a-half months at the upper limit (13,100 ft, 4,000 m).
Winds are strong in summer due to surface heating. Nights, however, are calm due
to radiational cooling and subsiding air. Winters are also less windy because of dom-
inant high pressure caused by cold air. The mountains are dry, with as little as 6 in
(150 mm) of precipitation per year. Even in the wetter summer season, precipitation
may fall as hail or snow. Winter snow cover, which lasts two to six months, is thin
and unevenly distributed. Below 11,500 ft (3,500 m), communities are xerophytic,
but higher elevations are more humid. In the subnival zone, mean monthly tempera-
tures rarely rise above freezing, and the zone of permanent snow and ice begins at
14,100 ft (4,300 m).
Because the mountains rise from desert, there is no subalpine forest. Only a
sparse treeline of central Asian spruce exists on the wetter northern slope. Several al-
pine communities, with overlapping limits, are based on elevation, temperature, and
precipitation. Soil moisture is most significant. The two lowest alpine zones, best
.................................................
Carpathian Mountains
The maritime influence on the Carpathians,
from Slovakia east through southwestern
Ukraine and south to Romania, is illustrated by
its high annual precipitation, 55 in (1,400 mm),
and mild winter and summer temperatures.
Because the mountains are surrounded by forest
rather than desert, about one-half of the alpine
plants are derived from boreal or deciduous for-
est elements. The highest peak, Mount Goverla,
is only 6,762 ft (2,061 m), and variation in tree-
line elevation is due to human activities. Natural
vegetation is dominated by pines and rhodo-
dendron shrubs, with tall herbs that thrive with
plenty of water. Heath and mosses are also typi-
cal. Natural shrubby communities are replaced
by secondary growth meadows where human
influence has been high. When overgrazed,
other grasses are replaced by matgrass, the final
stage in human-induced change.
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126
developed in the eastern region, are desert and semidesert with about 8 in (200 mm)
annual precipitation. Both have salty soils low in humus and basically vary in terms
of the amount of grass cover. In the desert zone, at about 10,000 ft (3,000 m), the
combination of aridity and poor soils supports only a thin cover of sagebrush and
salt-tolerant shrubs with a few sea lavender herbs and low shrubby cushion plants.
Slightly wetter semideserts are still dominated by sagebrush, but also have needle-
grass and a fewother flowering plants.
Higher on the slopes are dry steppes and cold steppes. Soils tend to be saline.
Dry steppes, with 9 in (230 mm) of precipitation, are dominated by tussocks of fes-
cue, spike fescue, and false needlegrass. Few nongrasses occur, but lichens are
common, especially crustose and fruticose. Cold steppes, both slightly wetter and
cooler, have a distinctly different community dominated by aster-like snow lotus,
with locoweed at higher elevations or fescue, reedgrass, and false needlegrass at
lower elevations. Other flowering plants are sparsely represented.
Higher elevations are more moist due to more precipitation and cooler temper-
atures. Steppe meadows receive 11 in (280 mm) and have a four-month growing
season. Snowmelt provides sufficient moisture content in spring, but soils become
dry in summer. Communities are dominated by kobresia sedge or Tien Shan fes-
cue. Because of summer drought, few forbs appear, although asters, gentians, dan-
delions, and some others can be found. Slightly higher and wetter, moist meadows
with deep soils (10 in, 25 cm) are widespread, particularly on shady slopes and
river terraces (see Figure 3.14). Most typical is a kobresia and carex sedge
Figure 3.14 Lush meadows in the Tien Shan Mountains, Kyrgyzstan, are surrounded
by high, snowy peaks. (Photo by author.)
127
community that includes 75 flowering plant species in several associations. Shag-
spine pea-shrub and alpine bluegrass dominate. Many fruticose and foliose lichens
and several mosses grow on the soil surface. Where frost polygons limit vegetation
cover, plants are androsace cushions, buttercups, snow lotus, and crustose lichens.
These extensive wet sites are the basis for a rather complex ecosystem of plants,
herbivores, and carnivores.
The highest alpine zone below permanent ice and snow, occurring at elevations
of 13,000 ft (4,000 m), is an herb tundra and cushion plant community similar to
that found in the Arctic. Plants do not form a continuous cover on the shallow,
stony soil, but grow in patterns similar to frost boil tundra. Cushion plants of Dry-
adanthe tetranda dominate but cover only 1030% of the ground. Most other flower-
ing plants and mosses grow within the cushions.
Bog communities can be found at any elevation in the alpine zone where ground-
water reaches the surface. Soils are peaty glei with organic content up to 60%. All
bogs are characterized by sedge communities dominated by carex species. Forbs
include lousewort, viviparous bistort, and buttercup. Mosses grow on the soil surface.
Although many plant species are restricted to particular communities, animals
in the Tien Shan tend to be more widespread, occupying several elevation zones
and habitats. Gray marmots and Tolei hares are common in all communities
except the highest moist meadows and herb tundra. Marmots are abundant wher-
ever it is rugged and rocky. Gray hamsters occur
only in lower elevations, in desert to dry steppe.
Large-eared pika, southern mole vole, and silvery
mountain vole, limited to higher elevations, live
in habitats from dry steppe to moist meadows.
Voles and marmots are the most characteristic
mammals of the moist meadows. Ungulates that
graze the meadows are Siberian ibex and the
argali or Kyzylkum sheep. Carnivores include
weasels, fox, and an occasional brown bear or
wolf, especially at the higher elevations. Birds,
including Horned Lark, Lesser Sand Plover, Isa-
belline Wheatear, and Snow Finch, are abundant
in dry communities. White-winged Redstart and
Alpine Chough are also common. In bogs, voles
are the only permanent inhabitants and Water Pi-
pet is the only nesting bird, although other ani-
mals and birds feed in bogs.
Himalaya Mountains. At approximately 30 N,
the isolated Himalayan mountain range contains
the southernmost and highest alpine ecosystem of
the world. In a southward curving arc, the range
.................................................
Speciation in the Himalayas
Because of their subtropical latitude, the Hima-
layas have little floristic relationship with cir-
cumpolar tundra. No woody perennials are
common to both regions, but the Himalayas
have 20 herbs of circumpolar distribution,
including viviparous bistort, a carex sedge, fire-
weed, kobresia, Iceland purslane, and alpine
sorrel. The cryptogam flora is less known. Fruti-
cose species of Iceland lichen, reindeer lichen,
and white-worm lichen dominate but do not
form extensive carpets. The range supports a
distinctive diversity of alpine plants. Only six
species of saxifrage occur in the Arctic, but 100
grow in the Himalayas, Similarly, one primrose
grows in the Arctic compared with 90 species
in the Himalayas. The Arctic has only two spe-
cies of rhododendron and six of saussurea,
while the Himalayas have 40 rhododendrons
and 60 saussureas.
.................................................
128
covers a large east-west extent from the Indus Gorge in northern Pakistan to the
Tsang Po Gorge in northeastern India. Most of the Himalayas are composed of
acidic rocks, such as granite and gneiss, and soils are young regosols. At elevations
below 13,000 ft (4,000 m), the landscape is characterized by U-shaped glacial val-
leys with moraines on higher slopes. Valley floors are filled with muddy outwash
material and alluvial fans that developed on steep side walls after valley glaciers
retreated. Solifluction occurs above 13,000 ft (4,000 m). Above 14,750 ft (4,500 m),
debris-covered glaciers fill valley floors between steep slopes (aretes) that rise above
the ice. Because of steep slopes and frost action, scree slopes are plentiful, provid-
ing habitat for pioneer plants. The lower alpine zone is characterized by closed veg-
etation on all slopes. In the higher alpine zone, sedge mats and cushions exist in
the shelter of large rocks or on patterned ground. The highest plants in the world
grow in isolated protected sites on the slopes of Mount Everest and surrounding
peaks.
The alpine environment varies according to geographic location and aspect
north-facing and south-facing slopes are significantly different. Treeline elevation,
as well as species, is highly variable but generally 12,000 ft (3,600 m). On shady
slopes in the northwest, treeline species are fir, spruce, and pine. On sunny slopes
in the same locality, sea wormwood dwarf shrublands and Grecian juniper wood-
lands merge with subalpine meadows. In the Central Himalayas, cloud forests of
east Himalayan fir, birch, and drooping juniper at the upper treeline are replaced
by subalpine thickets of rhododendron on shady slopes and juniper on sunny
slopes. In the drier Inner Himalaya, because it is more sheltered from monsoon
rain, closed dwarf-shrub thickets and mats are replaced by the open dwarf shrub-
lands and steppes of the Tibetan Plateau.
No long-term weather stations exist, and the climate of the Himalayas is not
well known. Vegetation and data from short-term weather measurements indicate
that the entire range is humid, but especially so in the southeast. Even in the drier
northwestern part, dwarf sage shrubs form a closed vegetation cover with many
forbs. From northwest to southeast, summer rains increase while winter and spring
rains decrease. The northwest gets winter precipitation from cyclonic storms mov-
ing in from the west; only one-quarter of its annual total falls in summer. Plants
there grow best under or around winter snowbeds, while others depend on light
summer rain. The southeast, dry in the winter due to monsoon winds from the
Asian interior, receives most of its precipitation in the summer from southerly
monsoon flow. The highest amount of precipitation falls in the cloud forest below
the alpine zone. Although it receives lower precipitation, the alpine belt has high
humidity in the summer, which promotes growth of foliose lichens and diversity in
floristic composition. A pronounced difference in microclimate is evident on the
north and south slopes in the eastern Himalayas during the dry winter. South-
facing slopes are hot, while north-facing slopes are shady and snowy until the end
of May. The difference diminishes in the summer when both slopes are moist due
to high humidity.
129
The lower alpine zone, near 14,000 ft (4,250 m), gets mostly rain, and a long-
lasting cloud cover suppresses diurnal differences in temperature and humidity. In
the upper alpine, at 15,500 ft (4,750 m), weather is dominated by drizzling mists
alternating with sunshine. This causes short-term changes in temperature and rela-
tive humidity. Surfaces are always damp and covered by more lichens than flower-
ing plants.
At the lower limit of the alpine zone, more than 200 days have a mean tempera-
ture around freezing. The upper alpine zone is more continental, with more
extreme temperatures, especially during the wet summer monsoon when the cloud
layer is either thin or well below this high zone. Temperatures in the upper alpine
regularly fall below freezing during the wet summer monsoon season, causing nee-
dle ice. Between the rainy season and the first snowfall, some plants like gentians,
snow lotus, and larkspur can withstand temperatures down to 5 F (15 C) at
night. According to estimates from existing data, the mean summer temperature at
the upper limit of the alpine belt is 36.5 F (2.5 C). The extent of permafrost is not
known, but extensive arctic-size areas of patterned ground can be seen above
16,500 ft (5,000 m). The elevation of the open alpine mats at the upper limit of the
alpine zone may coincide with the lower limit of permafrost.
Both wind and duration of snow-cover affect plant communities. Sheltered
areas have less temperature variation, while windy ridges are more extreme. High
ridges exposed to cold, dry winter winds may be completely blown free of snow.
Evergreen woody perennials like dwarf-shrub rhododendrons are restricted to
snow-protected slopes.
Because the Himalayas cover such a large area and have a wide range in tem-
perature, precipitation patterns, glaciation, substrate, slope steepness, and slope as-
pect, there are many plant communities depending on geographic location.
Although the growthforms of the basic communities are similar throughout the
range, species composition varies.
The two major communities of the lower alpine zone are both dwarf-shrub
thickets (see Figure 3.15). Rhododendrons grow as cushion-forming dwarf shrubs
on both sunny and shady slopes up to 14,750 ft (4,500 m) where they are protected
by winter snow. Tall forbs such as lousewort, monkshood, and kobresia sedge
grow throughout the woody evergreen thickets. The humid conditions support
abundant lichen growth in the shelter of the rhododendrons. Several junipers form
a second set of dwarf shrublands in the lower alpine, some restricted according to
geographic area. Single-seed juniper is found throughout the Himalayas, common
juniper in the northwest, and Grecian juniper on the driest rocky outcrops. Some
form flag trees due to strong upslope winds. Juniper roots stabilize soil on steep
slopes better than rhododendrons, even in areas affected by solifluction. Between
the juniper patches are mat-forming meadows of Nepal kobresia sedges and flat,
silvery-colored cushions of everlasting and edelweiss. Mosses and lichens are rare.
Rhododendron-kobresia mosaics are the major vegetation in the upper alpine
zone at 15,000 ft (4,600 m). Dwarf shrubs, common in the lower alpine, are limited
130
to protected warmer habitats in the upper alpine. Other plants in the kobresia turf
include plants with long taproots, both rosettes and cushions, such as rock prim-
rose. Except for their flowering stalks, plants are less than 2 in (5 cm) high, and the
leaves of the cushions or rosettes are tight to the ground. On drier and windier sites
susceptible to disruption by solifluction, only small patches of flowering plants
such as taprooted cushions of snow lotus, sandwort, and sibbaldia grow. Mosses
and lichens outnumber flowering plants. Crustose and foliose lichens cover dead
kobresia plants, and fruticose lichens grow on woody parts of rhododendrons.
Gravels between plants are also covered with lichens. Open scree slopes in the
northern part of the range have cushions and rosettes with long roots that anchor
plants in the unstable ground. The drier north slope is an alpine short-grass steppe
with needlegrass, carex sedges, and flat cushions of rock primrose and locoweed.
Two major plant communities depend on variations in soil. Snowbeds, similar
to those found in the European Alps and other mountains in North America and
Eurasia with winter precipitation, are more common in the upper alpine zone in
the northwestern Himalayas. In this summer-dry region, melting snow provides a
needed source of moisture in summer. Bistort, buttercup, and mosses dominate.
Vascular plants are rare on unstable boulder slopes with little soil development,
and crustose lichens cover the rocks. Young glacial moraines in the upper alpine
that have only recently been exposed can be found in several stages of plant
succession.
The Himalayan alpine is home to several large ungulates, including goats, ibex,
and tahr, and blue sheep or bharal. The tahr, related to wild goats, has a double
coat for warmth and hooves well suited to gripping rocks. Bharal is neither sheep
Figure 3.15 Vegetation varies with elevation as climate and environment changes in
the Himalayas. (Illustration by Jeff Dixon.)
131
nor goat but is a larger mammal with characteristics of each. High meadows in both
the Himalayas and Tibet are grazed by yaks. This large animal, males weighing
about 1,000 lb (450 kg), survives by grazing grasses, forbs, and lichens. Like the
muskox of the Arctic, the yak has a dense undercoat covered by longer guard hairs to
give it insulation during cold winters. A large lung capacity and numerous red blood
cells enable it to cope with low oxygen at high elevations. Yaks conserve energy
when traveling over snow in winter by walking single file and carefully stepping in
the footprints of the lead animal. Usually congregating in herds of 1030, yaks were
formerly more widespread. Numbers have decreased as a consequence of both
uncontrolled hunting and conversion of grazing grounds to pasture for domesticated
animals. The only horse native to alpine regions anywhere in the world is the Tibetan
wild ass, or kiang. Smaller herbivores include marmots, pikas, hares, and voles.
Large carnivores include the elusive snow leopard, less than 4 ft (1.2 m) long
but with a 3 ft (1 m) tail that provides balance in rugged terrain (see Plate XII). The
snow leopard has the thickest coat of any cat and also has thick fur on the soles of
its feet. Enlarged nasal cavities help it breathe thin air. Brown bears are occasion-
ally seen, and the Himalayan black bear wanders into the alpine from the forest
below. Like their Arctic or North American counterparts, both bears are omnivo-
rous. Red fox, Tibetan wolf, and Siberian weasel are smaller predators.
Counterparts of the ptarmigan in North American mountains are Snow Par-
tridge, Himalayan Snow Cock, and Tibetan Snow Cock, all members of the pheas-
ant family. Raptors include Golden Eagle, Lammergeier or Bearded Vulture, and
Himalayan Griffon Vulture.
Few locations in the Himalayas have not been influenced by human activity.
Destruction of the upper forest belt by fire has had the most effect, allowing mat-
forming, nongrazed alpine plants to extend their distribution to lower elevations.
Cattle and goats graze up to the highest alpine zone, even to 17,400 ft (5,300 m) in
Tibet. Secondary growth is rich in flowering plants such as species of rock prim-
rose, gentians, iris, and cinquefoil. After grazing has opened the natural vegetation,
woody weeds appear. Rhododendrons and junipers, which are not grazed, are peri-
odically removed by fire to maintain plants suitable for forage, such as kobresia
sedges. Forests on shady, humid slopes are less susceptible to fire and are removed
only in patches.
Southern Hemisphere
Gradients along slopes from exposed rocky ridges to valley bottoms are similar to
those in the Northern Hemisphere, but they control a different flora. Very small
dwarf shrubs are the most characteristic plants in all alpine habitats.
Southern Africa. Although the high plateau of Lesotho, 29 S, is a treeless zone
at 11,000 ft (3,300 m), temperatures are higher than in most alpine areas (see
Figure 3.16). The plateau consists of the Thaba-Putso Range, Central Range, and
132
.......................................................................................................
Southern Andes
The high-elevation region in central Argentina and part of Chile at 2739 S can be considered ei-
ther alpine tundra or grassland because it contains elements of both. Elevations drop from 14,000 ft
(4,250 m) in the north to about 5,500 (1,700 m) in the south. Less than 25 in (630 mm) of precipita-
tion falls annually throughout the region. January (summer) means of 55 F (13 C) are warmer than
in most alpine regions, but these high elevations are covered by snow for several months in winter.
A diverse flora with characteristic genera exists, and many endemic species have adaptations to
extreme dry, cold, and windy conditions with possibility of frost year-round. A treeline of tall shrubs
gives way to small shrubs, needlegrass tussocks, and cushion plants. In the subnival zone, small forbs,
rosettes, and small grasses dominate, but cushions are common. Conspicuous endemic shrubs are
Adesmia pinifolia, Chuquiraga ruscifolia, and C. echegarayi. Endemic cushions include Adesmia subter-
ranea, Azorella cryptantha, Laretia acaulis, and Oxychloe bisexualis. Although most animal life is
related to Andean dry puna (see Chapter 4) and Patagonian steppe, some animals, such as the nearly
extinct wild chinchilla, Patagonian chinchilla mouse, and Andean field mouse, are endemic.
......................................................................................................
Figure 3.16 Alpine tundra in southern Africa is limited to the high plateau of Lesotho.
(Map by Bernd Kuennecke.)
133
Maloti Mountains and is bounded by the Drakensberg Mountains (the east Africa
escarpment) on the east. The highest peak on the plateau is Thabana Ntlenyana
(Pretty Little Mountain) at 11,430 ft (3,484 m). The plateau is broad and dissected,
with boggy areas at the headwaters of the Orange River system. Geology consists
of basaltic lava up to 5,250 ft (1,600 m) thick covered by thin to no soils and rock
rubble. During summers, the region is wet and the ground is waterlogged. Winters
bring nightly freezing and daily thawing, formation of needle ice, solifluction, and
patterned ground, all characteristics of alpine or tundra landscapes.
Low temperature combined with low precipitation in winter is stressful for vegeta-
tion. Snow falls about eight times per year in winter (April to September), but usually
melts quickly so there is no long-lying snow cover. Although the frost-free season is
about six months, the growing season is still short. Mean temperature of the warmest
month (January) is 52 F (11 C), but temperatures can be extreme. The highest tem-
perature recorded was 88 F (31 C). The lowest was 5 F (21 C). The climate is
generally humid, and the mean annual precipitation varies from 25 in (630 mm) to 63
in (1,600 mm). Highest precipitation occurs on the eastern side, part of the Drakens-
berg Mountains, with rainshadow on the western side of the escarpment. Relative hu-
midity varies 1872%, according to the wet summer season or dry winter season.
High winds in late winter and spring are especially detrimental to plants because soils
are already dry at that time. Fire has been part of the ecology of the plateau, both
induced by humans to increase pasture and caused by natural lightning.
Vegetation is primarily heath communities dominated by low woody species of
erica shrubs and strawflower cushions scattered in extensive grasslands. Alpine
heath is the dominant vegetation, with dwarf shrubs less than 24 in (60 cm) high.
However, much has been cut for fuel. Boulder fields support a taller heath, 24 ft
(0.51.2 m) high, with aster-like plants and strawflower.
Outcrops of horizontal basalt layers have many pioneer species of lichens and
mosses. A succulent euphorbia called lions spoor forms cushions on sunny and
rocky north-facing slopes. Wiregrass tussocks and a dwarf bulrush are found in
wetter areas. Three semiwoody shrubs, euryops daisy, strawflower, and a member
of the milkwort family, all form cushions. Alpine grassland ranges from a low turf
to tall plants in an open landscape, interrupted by mud patches. Three temperate
genera of grassesfescue, wiregrass, and Pentaschistisdominate, with wiregrass
being most important. All are tufted, drought tolerant, and dormant in winter.
With enough water, wiregrass tussocks can be up to 3.3 ft (1 m) tall. Other grasses
include tufted hairgrass and lovegrass according to aspect. Flowering forbs are
numerous in spring and autumn.
Bogs develop in swales, where aquatic plants such as aponogeton, a succulent
crassula, oxygen weed, mudwort, and algae grow in open pools. Pool edges may
have pipewort, poker plants, bluegrass, buttercups, and bladderworts. Bog carpets
without open water have hummocks of bush tea and everlasting, with many
mosses and a variety of higher plants. Drier soil at the edges of the bogs are pre-
dominantly carex sedge meadows.
134
Due to climate and hunting, vertebrate fauna is species poor. Larger mammals
such as antelope are no longer present. Major small mammals are the ice rat that
digs up soil and peat in bogs, and the common mole rat. Insects, especially flies,
beetles, and bees, are still numerous. The area has long been used as pasture for
domestic animals and is overgrazed and degraded, with serious trampling of the
bogs that constitute important headwaters.
New Zealand. Most of the alpine region in New Zealand is on South Island at
4147 S latitude (see Figure 3.17). Treeline is roughly 3,300 ft (1,000 m), and
Figure 3.17 Characteristics of the alpine zone in New Zealand change from west to
east because of the southern Alps location in the path of westerly winds. (Map by Bernd
Kuennecke.)
135
permanent snow begins at 6,550 ft (2,000 m). The indigenous plants and animals
of both islands are largely endemic. Human influences during the last 100 years,
however, have changed alpine communities. Treeline in many places has been
destroyed by burning, allowing tussock grasses to extend farther downslope.
Today, the upper limit of fescue tussocks coincides with natural treeline, and cli-
mate factorssuch as average summer temperatures of less than 50 F (10 C)or
indicator species, rather than vegetation, define treeline and the alpine zone. New
Zealands mountains are young, and North Islands volcanoes are still active, so
lava and loose scoria are common. South Islands geology is more varied and
includes igneous, metamorphic, and sedimentary rock. Extensive moraines and
outwash from alpine glaciation on South Island provide parent material for unsta-
ble soils, such as poorly developed inceptisols and lithosols. Freeze-thaw cycles are
active in moist soils, and small-scale patterned ground such as stone polygons,
stripes, and solifluction terraces and lobes are present.
Climate is generally maritime. The westcoast location of mountains combined
with prevailing westerly winds and storm tracks result in a distinct west to east cli-
mate gradient, particularly on South Island. Precipitation on the windward side is
more than 160 in (4,000 mm) per year and may reach 470 in (12,000 mm) on some
slopes. Lee sites receive 50 in (1,250 mm), with less than 14 in (350 mm) in the dri-
est valleys. Gradients also exist for solar radiation, temperature, and humidity, but
overall, the climate is moist with no soil moisture deficit. Even when warming
from downslope winds increases temperature, cool nights raise relative humidity
to saturation. The moderate temperatures reflect the maritime influence felt
throughout the islands. Mean summer temperature at the upper limit of the low-
alpine zone is 40 F (5 C), decreasing to 32 F (0 C) in the high alpine. Tempera-
ture extremes in the low-alpine zone are 75 F (24 C) in the summer and 2 F
(19 C) in the winter; frost is common almost nightly. Needle ice may occur on
summer nights. The longest recorded frost-free period was only 13 days. Winters
are much warmer than in continental mountains and rarely drop below 30 F
(1 C) in the low alpine, or below 7 F (14 C) in the high alpine. Seasonal soil
temperatures vary less than air temperatures, and even in the high-alpine soil only
freezes 4 in (10 cm) deep. Winds redistribute snow cover, blowing it into drifts and
insulating the soil. Summer days can be sunny even with the common occurrence
of fog and low clouds. If surfaces become parched by sun during the day, they will
be relieved come nightfall.
Although growing under conditions less severe than either the arctic or mid-lat-
itude alpine zones, the alpine flora of the Southern Alps has evolved many typical
tundra vegetation formscushions, rosettes, mats, succulents, perennials, tap-
roots, and hairy or wooly coverings. Many plants have close relatives (that is, coge-
ners) in other parts of the world, while many others are native only to New
Zealand or to New Zealand and a few islands off Antarctica.
One peculiarity of this alpine region is the abundance of fleshy succulent plants
found particularly on sunny, dry scree slopes. Fleshy structures preserve moisture
136
during the day, a distinct advantage since groundwater is unavailable on extremely
well-drained talus.
The most distinctive plants of the New Zealand alpine flora are two endemic
genera called vegetable sheepso named because, at a distance, a group of them
resembles a flock of sheep. The genera, Haastii and Raoulia, are both members of
the sunflower family, but they are not closely related. Haastii forms hummocks of
either tight rosettes or loose branches, covered with fine wooly or wispy hairs.
Haastii are found in dry mountain areas where in summer sunshine alternates with
cold, foggy winds. By experimentation, it was found that the hairs prevent water
loss via excessive transpiration and condense every possible bit of moisture in the
air during droughts to keep the interior tissues moist. Two species of Raoulia con-
sist of a hard, rounded surface that makes a strong crust around peaty humus. The
surface is covered with fine white hairs that shed water easily. Because the purpose
of the hairs is to shed excess moisture rather than absorb the maximum air mois-
ture possible, Raoulia are adapted to and found in wet mountain areas. This use for
plant hairs may be unique to the vegetable sheep of New Zealand.
The maritime climate allows a long growing season compared with most other
alpine areas. For snow-tussock grasses, vegetative growth takes place for about
eight months at the lower limit of the alpine zone and only decreases to five
months in the high alpine. High-alpine cushion plants produce flowers over an
eight-month time span (October to May), although different species bloom at dif-
ferent times. Despite the long growing season, most flower buds preform the sea-
son prior to blooming.
Frequently no krummholz forms at treeline in New Zealand, and the change
from forest to alpine tundra is abrupt. Erect southern beech trees give way to low
shrubs of various species. These low shrubs are not environmentally stunted forms
of forest species, but are genetically short. Only under severe conditions of snow
and winter desiccation does treeline beech grow as krummholz, with semiprostrate
stems up to 10 ft (3 m) and sometimes forming flag trees.
The low-alpine zone, a narrow elevational belt, is dominated by tall evergreen
snow tussock grasses, shrubs, and forbs. Grass tussocks grow as tall as 4 ft (1.2 m),
unusual in alpine regions outside the Tropics, but probably made possible by the mild
climate. Large forbs and shrubs are conspicuous on the wetter western mountains,
especially on rocky ridges and sunny aspects where snow fails to accumulate. A vari-
ety of plants sort themselves according to soil and microhabitat conditions, especially
snow cover. Evergreen shrubs might be Dracophyllum and Olearia; common herba-
ceous plants are alpine daisy and New Zealand flax. Most widespread in high rain
areas, bogs with underlying peat occupy poorly drained depressions with high water
tables. Plants include several cushion plants such as alpine daisy along with Oreobolus
sedges. Several low, trailing subshrubs and herbs grow in bogs, where insectivorous
sundews, bladderworts, fruticose lichens, sedges, and rushes are also common.
The high-alpine zone has a discontinuous cover of shorter grasses, forbs, and
shrubs, with several distinct communities. Fellfields of stable rock are sparsely
137
vegetated. The drier areas are characterized by cushion plants, particularly vegeta-
ble sheep, and small Hebe shrubs, strawflower, and cushion speargrass. Wet fell-
fields have more diversity, with more small herbs and grasses. Since talus or scree
of loose, angular stones on steep slopes can be mobile, especially during spring
snowmelt, such areas support only a sparse cover of summer-green herbs. Plants
such as fireweed and buttercup have strong roots penetrating into the deep soils
beneath the rocks that enable them to withstand scree movement.
Cushion fields on broad plateau summits with strong winds, cold summers,
and freeze-thaw cycles have dwarf cushions, mats, and prostrate plants less than
0.75 in (2 cm) tall, vegetation similar to that found in arctic tundra. Where abraded
or scoured by strong wind, they may assume asymmetrical shapes. Dracophyllum
muscoides is a prevalent prostrate shrub. Patterned ground contributes to a mosaic
on the landscape. Snowbanks, which persist until mid or late summer in depres-
sions and areas sheltered from prevailing winds, are generally well vegetated. The
smallest snow tussock grass is dominant, forming dense turf 4 in (10 cm) high, but
many other plants grow in this microenvironment as well.
The nival zone, with lichens on rock faces too steep to maintain snow cover, is
extensive in the Southern Alps. Three flowering plants grow above 8,200 ft (2,500
m) on Mt. Cook, a buttercup and subshrubs Hebe haastii and Parahebe birleyi.
Except for two species of bat, no mammals are indigenous to the alpine zone.
Birds, however, are represented by many indigenous and endemic breeding spe-
cies. Several are rare and endangered. A distinctively alpine bird is the Flightless
Rail or Takahe, now limited to Fiordland National Park. It feeds in low-alpine tus-
sock grasslands, taking the tillers of the plants. Today, the introduced red deer
offers serious competition. The endemic Alpine Parrot, the Kea, is widespread and
common in the Southern Alps, but spends winters below treeline (see Figure 3.18).
This noisy, olive-green and red bird is omnivorous and, in spite of being only 12 in
(30 cm) tall, has successfully attacked and killed sheep. The widespread New Zea-
land Falcon is a strict carnivore. The larger raptor, Australasian Harrier, soars on
thermals over grasslands, occasionally rising as high as the alpine zone. The only
bird restricted to the alpine environment of South Island is the small Rock Wren,
which feeds on insects and fruit and spends the winter in rock crevices beneath the
snow. Some coastal birds, such as the South Island Pied Oystercatcher and Banded
Dotterel, migrate to the alpine for breeding and nesting in cushion fields. The
Southern Black-backed Gull, as well as other gulls, nest in bogs. The New Zealand
Pipit may also nest in alpine zones. Few seed eaters are found in the alpine zone
because the seed supply is unreliable. Reptiles are few; two skinks and two geckos
are known to live at high elevations.
Several vertebrates have been introduced. Red deer are widely distributed
throughout the mountains and are most damaging by virtue of their heavy grazing
and trampling of the soil. Widespread hares also detrimentally affect the alpine
zone. Goats, wapiti, chamois, and tahr are more limited in extent but do damage
where they occur.
138
Invertebrate fauna is rich in all habitats free of snow in summer up to 9,800 ft
(3,000 m). Insects are important contributors to the alpine ecosystem, pollinating
some plants, defoliating others, and consuming litter. Conspicuous in size and
number of species are wetas, grasshoppers, beetles, moths, butterflies, cicadas, and
flies, especially black flies and blow flies. Characteristic of alpine New Zealand are
several genera of giant weta (Orthoptera: Stenopelmatidae), large beetles that live
under rock slabs. With bodies up to 2.75 in (70 mm long), the insect weighs up to
2.5 oz (70 g). More than 40% of New Zealands moths and butterflies are alpine in
distribution; some are endemic. Several diurnal wolf spiders and dark-colored
jumping spiders overwinter beneath the snow. They are most diverse on South
Island. Because of the short growing season, insects have a short active period. It
may take more than one year to complete the generation cycle, with larvae waiting
out unfavorable periods concealed by host plants. Common adaptations include
dark color, long and dark body hair, and a large body.
Further Readings
Blumstein, Daniel T. n.d. Marmot Lore 101: Class Notes. Marmotburrow. http://
www.marmotburrow.ucla.edu/lore.html.
Bowen, Ezra. 1972. The High Sierra, The American Wilderness. New York: Time-Life.
Figure 3.18 The Kea is a alpine parrot native to New Zealand. (Courtesy of Shutterstock.
Copyright: Dario Diament.)
139
European Environmental Agency. n.d. Mountain Distribution of Plants. Epaedia, Envi-
ronment Explained. http://epaedia.eea.europa.eu/page.php?pid341.
Great Himalayan National Park. n.d. Great Himalayan National Park. http://www.
greathimalayannationalpark.com/GHNP_home.htm.
Price, Larry W. 1981. Mountains and Man: A Study of Process and Environment. Berkeley: Uni-
versity of California Press.
Ross, Graham. n.d. The Alpine Plants of New Zealand. http://alpine-plants-new-zealand.
110mb.com.
Zwinger, Ann H., and Beatrice E. Willard. 1972. Land Above the Trees: A Guide to American
Alpine Tundra. New York: Harper & Row.
140
Appendix
Biota of the Mid-Latitude Alpine Tundra Biome
North American Alpine Tundra
Some Characteristic Plants of the Coastal Ranges
Treeline and krummholz
Mountain hemlock Tsuga mertensiana
Subalpine fir Abies lasiocarpa
Alaska cedar Chamaecyparis nootkatensis
Low shrubs or subshrubs
Resin birch Betula glandulosa
Tea-leaved willow Salix planifolia
Mountain heather Phyllodoce spp.
Graminoids
Mountain hare sedge Carex phaeocephala
Altai fescue grass Festuca altaica
Forbs
Cushions or mats
Spreading phlox Phlox diffusa
Creeping sibbaldia Sibbaldia procumbens
Leafy or rosettes
Alpine bittercress Cardamine bellidifolia
Chiming bells Mertensia paniculata
Fireweed Epilobium spp.
141
Cryptogams
Jewel lichen (Crustose) Caloplaca elegans
Sphagnum moss Sphagnum ssp.
Some Characteristic Animals of the Coastal Ranges
Herbivores
Mountain goat Oreamnos americana (Introduced)
Dalls sheep Ovis dalli (Alaska and Canada only)
Olympic marmot Marmota olympus (Endemic to
Olympics)
Hoary marmot Marmota caligata (Alaska and
Canada only)
American pika Ochotona princeps
Mazama pocket gopher Thomomys mazama
Deer mouse Peromyscus maniculatus
Meadow vole Microtus pennsylvanicus
Carnivores
Dusky shrew Sorex monticolus
Brown or Grizzly bear Ursus arctos
Birds
White-tailed Ptarmigan Lagopus leucurus
Horned Lark Eremophila alpestris
Water Pipet Anthus spinoletta
Gray-crowned Rosy Finch Leucosticte arctoa tephrocotis
Red-tailed Hawk Buteo jamaicensis
Stellars Jay Cyanocitta stelleri
Some Characteristic Plants of the Cascade Range
Whitebark pine Pinus albicaula
Engelmann spruce Picea engelmannii
Alpine larch Larix lyallii
Partridge foot Luetkea pectinata
Kinnikinnik or Bear-berry Arctostaphylos uva-ursi
Western moss heather Cassiope mertensiana
Pink mountain heather Phyllodoce empetriformis
142
Graminoids
Mountain hairgrass Deschampsia atropurpurea
Kobresia sedge Kobresia myosuroides
Blackish sedge Carex nigricans
Forbs
Cushions or mats
Tolmie saxifrage Saxifraga tolmiei
Leafy or rosettes
Cliff Indian paintbrush Castilleja rupicola
Lemmons rockcress Arabis lemmonii
Lyall lupine Lupinus lyallii
Alpine aster Aster alpigenus
Bistort Polygonum bistortoides
Newberry knotweed Polygonum newberryi
Pussypaws Calyptridium umbellatum
Cryptogams
Jewell lichen (Crustose) Caloplaca elegans
Some Characteristic Animals of the Cascade Range
Herbivores
Mountain goat Oreamnos americana (Introduced)
Yellow-bellied marmot Marmota flaviventris
Heather vole Phenacomys intermedius
Mazama pocket gopher Thomomys mazama
American water vole Microtus richardsonii
Carnivores
Birds
Gray-crowned Rosy Finch Leucosticte arctoa tephrocotis
Rock Wren Salpinctes obsoletus
Clarks Nutcracker Nucifraga columbiana
White-tailed Ptarmigan Ptarmigan leucurus
143
Some Characteristic Plants of the Sierra Nevada
Timberline and krummholz
Whitebark pine Pinus albicaulis
Sierra lodgepole pine Pinus contorta var. murrayana
Foxtail pine Pinus balfouriana
Dwarf bilberry Vaccinium caespitosum
Timberline sagebrush Artemisia rothrockii
Graminoids
Shorthair reedgrass Calamagrostis breweri
Short-grass sedge Carex exserta
Forbs
Cushions or mats
Sierra saxifrage Saxifraga aprica
Alpine pussytoes Antennaria alpina
Clubmoss ivesia Ivesia lycopodioides
Tufted phlox Phlox caespitosa
Davidsons penstemon Penstemon davidsonii
Leafy or rosettes
Nutalls sandwort Arenaria nuttallii
Dwarf knotweed Polygonum minimum
Bottlebrush squirreltail Sitanion hystrix
Great Basin violet Viola beckwithii
Oval-leaved buckwheat Eriogonum ovalifolium
Prickly gilia Leptodactylon pungens
Pussypaws Calyptridium umbellatum
Cryptogams
Some Characteristic Animals of the Sierra Nevada
Herbivores
Alpine chipmunk Tamias alpinus (Endemic)
Sierra pocket gopher Thomomys monticola
Bighorn sheep Ovis canadensis
144
Carnivores
Birds
White-tailed Ptarmigan Lagopus leucurus (Introduced)
Rosy Finch Leucosticte arctoa
Some Characteristic Plants of the Great Basin
Limber pine Pinus contorta
Big sagebrush Artemisia tridentata
Bristlecone pine Pinus longaeva
Graminoids
Alpine reedgrass Calamagrostis purpurascens
Alpine fescue grass Festuca brachyphylla
Spiked trisetum grass Trisetum spicatum
Sheep fescue grass Festuca ovina
Elk sedge Carex elynoides
Rocky Mountain sedge Carex scopulorum
Forbs
Cushions or mats
Covilles phlox Phlox covillei
Alpine avens Geum rossii
Leafy or rosettes
Mono clover Trifolium monoense
Bottlebrush squirreltail Sitanion hystrix
Broad-podded phoenicaulis Phoenicaulis eurycarpa
(Continued)
145
Masons Jacobs ladder Polemonium chartaceum
Rambling fleabane Erigeron vagus
Marsh marigold Caltha leptosepala
American bistort Polygonum bistortoides
Cryptogams
Some Characteristic Animals of the Great Basin
Herbivores
Bighorn sheep Ovis canadensis (Rare)
Northern pocket gopher Thomomys talpoides
Carnivores
Birds
Some Characteristic Plants of the Rocky Mountains
Krummholz and treeline
Limber pine Pinus contorta
Alpine larch Larix lyallii
a
Tamarack Larix laricina
a
Western hemlock Tsuga heterophylla
a
White spruce Picea glauca
a
a
a
Sibbaldia Sibbaldia procumbens
a
a
Mountain heather Phylladoce spp.
a
146
Marsh Labrador tea Ledum palustre
a
a
Graminoids
Elk sedge Carex elynoides
Cottongrass sedge Eriophorum angustifolium
a
a
Rocky Mountain sedge Carex scopulorum
Drummonds rush Juncus drummondii
a
Forbs
Cushions or mats
Saxifrage Saxifraga spp.
Alpine phlox Phlox condensata
Alpine candytuft Thlaspi alpestre
Dwarf clover Trifolium nanum
One-flowered harebell Campanula uniflora
American bistort Polygonum bistortoides
Sky pilot Polemonium viscosum
Alpine avens Geum rossii
Alpine sage Artemisia borealis
Alpine locoweed Oxytropis podocarpa
a
Dotted saxifrage Saxifraga bronchialis
a
Leafy or rosettes
Alpine primrose Primula angustifolia
Milkvetch Astralagus plumbeus
Draba Draba spp.
Ragwort Senecio spp.
Arctic gentian Gentiana algida
Snow buttercup Ranunculus adoneus
Kings crown Sedum rosea
Marsh marigold Caltha leptosepala
Cryptogams
Iceland lichen (Fruticose) Cetraria islandica
Snow lichen (Fruticose) Cetraria nivalis
(Continued)
147
Knob lichen (Fruticose) Dactylina madreporiformis
Worm lichen (Fruticose) Thamnolia vermicularis
Reindeer lichen (Fruticose) Cladonia rangifera
a
Rock clubmoss Selaginella densa
a
Note:
a
Northern Rocky Mountains, Alaska, and Yukon only.
Some Characteristic Animals of the Rocky Mountains
Herbivores
Elk Cervus elaphus
Mountain bighorn sheep Ovis canadensis
b
Dalls sheep Ovis dallii
a
Mountain goat Oreamnos americana
a
Northern pocket gopher Thomomys talpoides
b
b
Collared pika Ochotona collaris
a
b
Hoary marmot Marmota caligata
a
Alaska marmot Marmota broweri
a
Water vole Microtus richardsoni
Carnivores
Brown or Grizzly bear Ursos arctos
a
Birds
White-tailed Ptarmigan Lagopus leucurus
Water Pipit Anthus spinoletta
Rosy Finch Leucosticte arctoa
b
Notes:
a
Northern Rocky Mountains, Alaska, and Yukon only;
b
Southern and Central Rocky Mountains
only.
148
Some Characteristic Plants of Mount Washington
Balsam fir Abies balsamea
Paper birch Betula papyrifera
Mountain heather Phyllodoce caerulea
Mountain cranberry Vaccinium vitis-idaea
Sweet blueberry Vaccinium angustifolium
Three-toothed cinquefoil or Sibbaldiopsis Potentilla tridentata
Bearberry willow Salix uva-ursi
Mountain alder Alnus biridus ssp. crispa
Labrador tea Ledum groenlandicum
Graminoids
Forbs
Cushions or mats
Greenland sandwort Arenaria groenlandica
Lapland diapensia Diapensia lapponica
Leafy or rosettes
Cryptogams
Clubmoss Lycopodium spp.
Some Characteristic Animals of Mount Washington
Herbivores
Groundhog Marmota monax
Carnivores
Masked shrew Sorex cinereus
149
Birds
Slate-colored Junco Junco hyemalis
White-throated Sparrow Zonotrichia albicaulis
Eurasian Alpine Tundra
Some Characteristic Plants of the European Alps
Treeline
Norway spruce Picea abies
Prostrate pine Pinus mugo
European larch Larix decidua
Stone pine Pinus cembra
Mountain pine Pinus uncinata
Lapland rosebay Rhododendron lapponicum
Alpenrose Rhododendron ferrugineum
Creeping azalea Loiseleuria procumbens
Green alder Alnus viridus
Net-leaved willow Salix reticulata
Graminoids
Fescue grass Festuca hallari
Fescue grass Festuca varia
Curly sedge Carex rupestris.
Scorched alpine sedge Carex atrofusca
Forbs
Cushions or mats
Rock jasmine Androsace helvetica
Two-flowered saxifrage Saxifraga biflora
Leafy or rosettes
Glacier buttercup Ranunculus glacialis
Edelweiss Leontopodium brachyactis
Houseleek or Liveforever Sempervivum montanum
Cryptogams
Norwegian haircap moss Polytrichum norvegicum
150
Some Characteristic Animals of the European Alps
Herbivores
Ibex Capra ibex
Red deer Cervus elaphus
Chamois Rupricapra rupricapra
Snow vole Arvicola nivalis
Alpine marmot Marmota marmota
Carnivores
Lynx Lynx lynx
a
Brown bear Ursus arctos
a
Wildcat Felis silvestris
Alpine shrew Sorex alpinus
Birds
Lammergeier or Bearded
Vulture
Gypaetus barbatus
a
a
Note:
a
Extinct or very rare.
Some Characteristic Plants of the Tien Shan Mountains
Treeline
Central Asian spruce Picea schrenkiana
Shrubs
Sagebrush Artemisia rhodantha
In the Chenopod family Kalidium schrenkianum
Shagspine pea-shrub Caragana jubata
Graminoids
Fescue grass Festuca sulcata
False needlegrass Ptilagrostis subsessiliflora
Needlegrass Stipa caucasica
Spike fescue grass Leucopoa olgae
Reedgrass Calamagrostis tianshanicus
Tien Shan fescue Festuca tianshanica
Alpine bluegrass Poa alpina
Kobresia sedge Kobresia capilliformis
Carex sedge Carex melanantha
Forbs
Cushions and mats
Rose family Dryadanthe tetranda
Androsace Androsace sericea
151
Leafy or rosettes
Sea lavender Limonium hoeltzeri
Snow lotus Saussurea leucophylla
Locoweed Oxytropis chionobaia
Aster Aster spp.
Gentian Gentiana spp.
Dandelion Taraxacum spp.
In the Buttercup family Callianthemum alatavicum
Lousewort Pedicularis rhinanthoides
Buttercup Ranunculus albertii
Cryptogams
Parmelia (Foliose lichen) Parmelia spp.
Aspicilia (Crustose lichen) Aspicilia spp.
Twisted moss Tortula ruralis
Wideleaf stegonia moss Stegonia latifolia
Some Characteristic Animals of the Tien Shan Mountains
Herbivores
Siberian ibex Capra sibirica
Argali or Kyzylkum sheep Ovis ammon
Gray marmot Marmota baibacina
Tolei hare Lepus tolei
Gray hamster Cricetulus migratorius
Silvery mountain vole Alticola argentata
Large-eared pika Ochotona macrotis
Southern mountain vole Ellobius talpinus
Carnivores
Wolf Canis lupus
Steppe polecat Mustela eversmannii
Birds
Lesser Sand Plover Charadrius mongolus
Isabelline Wheatear Oenanthe isabellina
Snow Finch Montifrigilla nivalis
Alpine Chough Pyrrhocorax graculus
White-winged Redstart Phoenicurus erythrogaster
152
Some Characteristic Plants of the Himalayas
Western Himalayan fir Abies pindrow
Western Himalayan spruce Picea smithiana
Himalayan pine Pinus wallichiana
Eastern Himalayan fir Abies spectabilis
Drooping juniper Juniperus recurva
Shrubs
Western Himalayan birch Betula utilis
Sea wormwood Artemisia maritima
Rhododendron Rhododendron spp.
Meyer single-seed juniper Juniperus squamata
Common juniper Juniperus communis
Grecian juniper Juniperus excelsa
Graminoids
Needlegrass Stipa purpurea
Mount Everest sedge Carex montis-everestii
Nepal kobresia sedge Kobresia nepalensis
Forbs
Cushions or mats
Snow lotus Saussurea gnaphalodes
Sandwort Arenaria bryophylla
Everlasting Anaphalis triplinervis
Edelweiss Leontopodium brachyactis
Sibbaldia Sibbaldia purpurea
Rock primrose or Jasmine Androsace lehmannii
Locoweed Oxytropis spp.
Leafy or rosettes
Fireweed Epilobium angustifolium
Iceland purslane Koeniga islandica
Alpine sorrel Oxyria digynia
Monkshood Aconitum spp.
Buttercup Ranunculus spp.
Cryptogams
Reindeer lichen (Fruticose) Cladonia rangiferina
White-worm lichen (Fruticose) Thamnolia vermicularis
Umbilicaria (Foliose lichen) Umbilicaria indica
Ochrolechia (Crustose lichen) Ochrolechia glacialis
153
Some Characteristic Animals of the Himalayas
Herbivores
Blue sheep or Bharal Pseudois nayur
Yak Bos grunniens
Himalayan tahr Hemitragus jemlahicus
Ibex Capra ibex
Tibetan wild ass or Kiang Equus kiang
Himalayan marmot Marmota himalayana
Himalayan pika Ochotona himalayana
Silvery mountain vole Alticola argentata
Tolai hare Lepus tolai
Carnivores
Snow leopard Panthera uncia
Himalayan black bear Ursus thibetanus
Tibetan wolf Canis lupus chanku
Siberian weasel Mustela sibirica
Birds
Snow Partridge Lerwa lerwa
Himalayan Snow Cock Tetraogallus himalayensis
Tibetan Snow Cock Tetraogallus tibetanus
Lammergeier or Bearded Vulture Gyptaetus barbatus
Himalayan Griffon Vulture Gyps himalayensis
Southern Hemisphere Alpine Tundra
Some Characteristic Plants of the Lesotho Plateau
Shrubs
Erica heath Erica dominans
Erica heath Erica glaphyra
Strawflower or Everlasting Helichrysum trilineatum
Graminoids
Goat fescue grass Festuca caprina
Wiregrass Merxmuellera drakensbergensis
Copper wiregrass Merxmuellera disticha
In the Bluegrass family Pentaschistis oreadoxa
Bog bluegrass Poa binata
Lovegrass Eragrostis caesia
Carex sedge Carex spp.
Dwarf bulrush Scirpus ficinioides
154
Aquatic
Aponogeton Aponogeton junceus
Crassula Crassula inanis
Oxygen weed Lagarosiphon muscoidas
Cape mudwort Limosella capensis
Forbs
Cushions or mats
Euryops daisy Euryops decumbens
In the Milkwort family Muraltia saxicola
Lions spoor Euphorbia clavarioides
Leafy or rosettes
In the Aster family Eumorphia sericea
Bush tea Athrixia fontana
Poker plants Kniphofia caulescens
Buttercup Ranunculus meyeri
Pipewort Eriocaulon dregei
Bladderwort Utricularia spp.
Some Characteristic Animals of the Lesotho Plateau
Herbivores
Ice rat Otomys sloggetti
Mole rat Cryptomys hottentotus
Some Characteristic Plants of the Southern Alps, New Zealand
Treeline
Southern beech Nothofagus menziesii
Southern beech Nothofagus solandri
Shrubs
Vegetable sheep Haastii spp.
Vegetable sheep Raoulia spp.
In the Aster family Olearia spp.
In the Figwort family Hebe spp.
Strawflower or Everlasting Helichrysum spp.
In the Heath family Dracophyllum muscoides
In the Figwort family Parahebe birleyi
Graminoids
Red snow tussock grass Chionochloa rubra
Snow tussock grass Chionochloa oreophila
Bentgrass Agrostis spp.
Oreobolus sedge Oreobolus pectinatus
Rush Carpha alpina
Rush Juncus spp.
155
Forbs
Cushions or mats
Alpine daisy Celmisia argentea
In the Figwort family Chionohebe spp.
Leafy or rosettes
Succulent in the Bellflower family Lobelia roughii
Succulent in the Bluegrass family Poa sclerophylla
Succulent in the Gentian family Gentiana divisa
New Zealand flax Phormium tenax
In the Madder family Coprosma perpusilla
Sundew Drosera spp.
Bladderwort Utricularia monanthos
Cushion speargrass Aciphylla dobsonii
Fireweed Epilobium spp.
Cryptogams
Cladina (Fruticose lichen) Cladina spp.
Cladonia (Fruticose lichen) Cladonia spp.
Some Characteristic Animals of the Southern Alps, New Zealand
Herbivores
Red deer Cervus elaphus
a
Chamois Rupicapra rupicapra
a
Tahr Hemitragus jemlabicus
a
Goat Capra hircus
a
Wapiti or Elk Cervus canadensis
a
European hare Lepus europaeus
a
Birds
Flightless Rail or Takahe Porphyrio mantelli
Alpine Parrot or Kea Nestor notablis
New Zealand Falcon Falco novaeseelandiae
Australasian Harrier Circus approximans
Rock Wren Xenicus gilviventris
Pied Oyster Catcher Haematopus ostralegus finschi
Banded Dotterel Charadrius bicinctus
Southern Black-backed Gull Larus dominicanus
Reptiles and amphibians
Skink Leiolopisma spp.
Gecko Hoplodactylus spp.
Note:
a
Introduced.
156
4
Tropical Alpine Biome
High-elevation tropical ecosystems, generally between 23
1
=
2
N and 23
1
=
2
S, are
the least known of all tropical biomes. Geographically isolated, they are like cold
islands surrounded by warm wet rainforests. Tropical alpine zones are distinct
from their arctic and mid-latitude counterparts in both vegetation and climate, and
the use of the term tundra is misleading because of numerous differences. Most
areas have unique geographic names but are too small to be shown on a world
map. The tropical alpine environment in the northern Andes, straddling the Equa-
tor from Venezuela to northern Peru with a few outliers in Costa Rica, is called the
paramo, while farther south in the central Andes, the altiplano is also known as
puna. The puna is the largest area of tropical alpine habitat in the world; it ranges
from central and southern Peru and Bolivia to northern Chile and Argentina. In
East Africa, it is called the afroalpine zone, and in Indonesia and New Guinea it is
simply called tropical-alpine. High zones on Hawaiian volcanoes are also above
tree growth. Unless a particular geographic region is specified, the term tropical
alpine zone is used here to refer to all high-elevation ecosystems on tropical moun-
tains. The lower limit of the zone is often difficult to define because there may not
be a recognizable treeline. The alpine zone is above rainforest or cloud forest and
treeline may consist of clumps of shrubs or stunted trees draped with epiphytes sur-
rounded by grassland. On dry slopes where there is no forest, shrubs and succulents
give way to typical alpine vegetation.
Although taxa on tropical mountains vary considerably from place to place
because the high mountains are isolated from one another, similarities in climate,
growthforms, and general environment exist. The paramo is similar to the afroalpine
157
in terms of plant growthforms. Like the isolated
mountains in Africa and Indonesia, the paramo is
not continuous but consists of smaller isolated
patches in Venezuela, Colombia, Ecuador, and
northern Peru. The puna has no Old World equi-
valent. In terms of growthforms, it is most similar
to the mid-latitude alpine region of Tibet, but in
terms of continuous extent, the puna is most simi-
lar to the Ethiopian Plateau. High mountain grass-
lands dominated by bunchgrasses with a scattering
of other plants, especially shrubs and rosettes in
the sunflower family, are typical of all tropical al-
pine areas. Some localities are purely grassland,
while others have ground covered by giant rosettes
and little grass. Locally important families with sig-
nificant genera include pineapple (genus Puya) in
South America, bellflower (genus Lobelia) in east-
ern Africa, sunflower (genus Senecio) also in eastern
Africa, rose (genus Polylepis) in the Andes, and tree
fern (genus Cyathea) in New Guinea. Frost poly-
gons and stripes may be found at the highest elevations. Ground cover varies from
barren with occasional mats or cushions to a spongy mat of mosses.
Tropical mountains are geologically young, building primarily since the late
Cenozoic. Therefore most of the biota has only recently evolved, and most species
were derived from lowlands as mountains uplifted. Of about 300 alpine plants on
East African volcanoes, 80 percent are endemic, having evolved from surrounding
lowland vegetation. Similar development has taken place in other tropical moun-
tains such as the Andes.
Climate
Except for wet and dry seasons, tropical mountains display daily changes rather
than seasonal ones (see Figure 4.1). Because of their tropical locations, daylength
and temperature vary little throughout the year. Day and night are each consis-
tently 12 hours long. No seasonal change in sun angle like that experienced in the
mid-latitudes occurs, but sun angle differs according to slope aspect and steepness.
Because the sun crosses the Equator in its path between the Tropics of Cancer and
Capricorn, either the northern or southern slope can receive direct rays while the
other is in shadow, depending on time of year. No seasonal variation in tempera-
ture and no long period of cold and dormancy occur. However, diurnal tempera-
ture changes are pronounced. During the day, tropical alpine sites are bombarded
with solar radiation, intensified by the thin atmosphere at high elevation. Much
heat is lost at night as energy radiates back to space. The difference in temperature
.................................................
Island Biogeography
Within the discipline of Geography, a subfield
called Island Biogeography strives to explain
the distribution of species on oceanic islands.
Because of limited habitat and long isolation
on oceanic islands, a unique biota often devel-
ops through the adaptive radiation of the few
organisms that can make it far out to sea. The
Galapagos and Hawaiian Islands are prime
examples of places where plant and animal
immigrants evolved to fill vacant niches. The
same concept can be applied to island-like
areas, such as high-elevation mountain top
environments that have never been connected.
They rely on similar types of long-distance dis-
persal and evolution and often have many
endemic species.
.................................................
158
Figure 4.1 Tropical alpine climates have similar temperatures but may vary in total
and seasonality of precipitation. (a) Cuzco, Peru, is in the dry puna, but (b) closer to
the Equator, the meteorological station on Mount Kenya receives more precipitation.
(Illustration by Jeff Dixon.)
from day to night in tropical alpine areas is much greater than the difference
between winter and summer. Consequently, plants must able to withstand temper-
ature extremes every day of the year.
Mean annual temperature for all tropical alpine environments is about 50 F
(10 C) at about 9,800 ft (3,000 m), the same temperature value that is often used to
define the boundary between boreal forest and arctic tundra. The boundary also
roughly coincides with the lowest elevation occurrence of freezing temperatures
on wet slopes. Frequency of frost increases from a few days a year at 9,800 ft
(3,000 m) to 100 days at 14,700 ft (4,500 m) and daily at the level of permanent
snow above 15,400 ft (4,700 m). On dry mountains, frost begins slightly lower, but
permanent snow is higher, at 19,700 ft (6,000 m), probably due to less precipita-
tion. As temperatures decline with increasing latitude such as on volcanic moun-
tains in Mexico at 19 N, the frost zone and treeline both begin lower. The
coincidence of freezing temperatures with treeline in tropical mountains is signifi-
cant for life. Plants must not only be adapted to temperatures that are consistently
low, but also must be permanently frost resistant. Unlike plants in mid-latitude
environmentsalpine or notthat have different frost tolerances in summer than
in fall and must winter-harden before they are able to withstand freezing condi-
tions, tropical alpine plants may be subjected to freezing on any night of the year
and must always be prepared. Plants must also photosynthesize under both low
temperatures and low light conditions because days are frequently cloudy.
Tropical mountains experience rainy seasons and less rainy or even dry seasons
lasting up to six months. In equatorial areas where precipitation is controlled by
convection, rainy seasons occur twice a year during the equinoxes, when the sun
and equatorial low pressure belt pass overhead. The dry seasons coincide with the
solstices when high pressure dominates. Climates dependent on trade winds to
bring moisture usually have only one rainy period related to seasonal shifts in the
low pressure belt. Most precipitation is due to thunderstorms.
While temperature predictably decreases with elevation, changes in precipita-
tion are more complex. In the Tropics, precipitation increases to a maximum at
mid-level elevations where cloud forests occur, but then steadily decreases toward
the mountain summits. Stable air above the cloud forests prevents storms from ris-
ing farther upslope. Therefore, alpine areas are considerably drier than lowlands in
the same latitude. The zone of maximum precipitation is generally between 3,000 ft
(900 m) and 4,600 ft (1,400 m), but varies according to geographic location. Maxi-
mum rainfall is reached at 2,500 ft (750 m) in Hawaii, 3,150 ft (950 m) in the tropi-
cal Andes, and up to 5,000 ft (1,500 m) in eastern Africa. After the maximum,
precipitation decreases with elevation at a rate of 4 in per 330 ft (100 mm per 100 m).
It is difficult to generalize about precipitation totals for tropical alpine zones
because several variables such as slope aspect and position with regard to equato-
rial low pressure or trade winds create different patterns of precipitation and rain-
shadow. Such variations contribute to diversity and influence the distribution of
environments and plant communities.
160
Tropical alpine zones may have seasonal temperature regimes that coincide
with the rainy and dry seasons. Periods of rain and clouds depress diurnal tempera-
ture changes, because less solar radiation enters during the day but also because
more infrared radiation (heat) is trapped at night. Under clear skies, both high and
low temperatures are more extreme. Most frosts occur during the drier season
because with less cloud cover more infrared radiation is lost at night, causing tem-
peratures to drop. Minimum temperaturesespecially those below freezingand
maximum temperatures are more important to plant life than are daily, monthly,
or annual means.
Tropical mountains rarely experience the strong winds common to mid-latitude
mountains under the jet stream and cyclonic storms. It may be windy on exposed
ridges, but not everywhere.
Soil
Tropical alpine soils are similar to those in other alpine regions: young and poorly
developed, usually bearing chemical resemblance to parent material. Steep slopes
tend to be covered by scree and talus, while more level areas may have an accumu-
lation of sediment and organic matter from plants. Surface soils may be dry, but
deeper moisture is usually available. Landscapes in the high Andes south of Ecua-
dor appear very dry with only a thin plant cover, but plants extend roots more than
3.3 ft (1 m) into the soil and well-established plants have access to soil moisture.
Coarse and dry cinders on volcanic mountains prevent evaporation from moist
sand or soil beneath. Surfaces are subjected to temperature extremes on a daily
basis, but temperatures at depth are more stable. Soil temperatures 3 in (8 cm) deep
on Mount Kenya are constant at about 37 F (3 C). The surface, however, is sub-
ject to frost-heave. Unlike arctic tundra and mid-latitude alpine environments,
tropical alpine areas have no permafrost, but they do experience small-scale soli-
fluction where nightly freezing disrupts soil development. In bare soil, needle ice
forms nightly in the afroalpine and other tropical alpine zones, disturbing the soil
and making it difficult for seedlings to survive.
Plant Adaptations
Tropical alpine areas support five growthforms, each dominating a different envi-
ronment largely based on soil and moisture availabilitytussock grasses, cushion
plants, sclerophyllous shrubs, ground-level rosettes, and giant stalked rosettes
(see Figure 4.2). Many growthforms are the same as in arctic tundra or in mid-
latitude alpine regions, but others have evolved unique ways to endure the extreme
temperature range.
Low-stature plants, such as stalk-less rosettes, tussock grasses, and cushion
plants, create a mild microclimate within their foliage at ground level, thereby
avoiding harsher air conditions. Grass tussocks have a layer of dead vegetation that
161
both insulates and holds water. After a fire, flowering and regrowth quickly follow
to replenish this necessary insulation. The masses of stems in tussock grasses also
support the plant against wind. Hairiness is a common characteristic and serves
several purposes. In addition to shielding the plant from intense sun, hairs provide
insulation at night, which helps retain heat. Air trapped by the hairs maintains its
humidity, thereby reducing the transpiration loss during the hot, dry day. Tropical
alpine plants, especially large ones, often have a silvery color that reflects excessive
solar radiation. Shrubs with leathery or waxy (sclerophyllous) leaves are well
adapted to periodic droughts and intense radiation in alpine zones. Small leaves
present less surface area to the cold, and a waxy cuticle is able to withstand dry air
conditions. Many high-elevation plants, such as giant lobelias in the afroalpine,
have a red pigment that protects against strong radiation. The number of plants
that are either hairy or have red coloration increases with elevation. Many plant
adaptations, such as hairiness or waxy leaves, make then unpalatable to herbivores,
providing an additional survival trait.
Little information is available about reproduction by seed among tropical
alpine plants. Flowering takes place all year on Mount Kenya, but most occurs in
January and July following the rainy seasons. In the Andean paramo, however,
Figure 4.2 Major growthforms in tropical alpine regions include (a) giant rosettes,
(b) tussock grasses, (c) ground-level rosettes, (d) cushions, and (e) shrubs. (Illustration by
Jeff Dixon.)
162
seasonal drought limits flowering time. Flies, bumblebees, and wind are important
pollinators, although two species of hummingbirds are known to pollinate espele-
tias. Several plants require cross-pollination. Espeletia seeds need a cold period of
30 days at 35 F (2 C) before germination. Giant rosettes in the Tropics are long-
lived plants. Silversword in Hawaii does not flower until it reaches age 20 and it
lives to be 90 years old. One of the lobelias (Lobelia telekii) on Mount Kenya flowers
and dies at 4070 years, while the much larger giant groundsel may live for several
hundred years.
Giant Rosettes
Although not present in every habitat, a conspicuous growthform unique to tropi-
cal alpine tundra is giant rosettes. Giant rosettes include espeletias and puyas in the
Andes; lobelias, giant senecios, and giant groundsels in Africa; tree fern in New
Guinea; and silversword in Hawaii (see Figure 4.3). Giant rosettes, whether
ground-hugging or stalked, have unique ways of coping with the tropical alpine cli-
mate. Stalks or inflorescences can be 330 ft (110 m) tall. Plants usually have a
sheath of dead leaves, 412 in (1030 cm) thick, which insulates the stem from
cold, traps water, and provides humus. Temperature inside the stem remains above
Figure 4.3 Although not related, giant rosettes have similar growthforms: (left) giant
groundsel (Dendrosenecio keniodendron) in Equatorial Africa and (right) espeletia in
the Andean paramo in El Angel Ecological Reserve, Ecuador. (Left, courtesy of Rainer
W. Bussmann, Missouri Botanical Garden, and right, Susan L. Woodward.)
163
the freezing point. The chemical composition of the leaves changes, indicating that
nutrients are transferred from dead leaves to new ones, which is an efficient way to
recycle scarce soil resources. In taller rosettes in East Africa, the outer leaves
always enclose the central growing point. These leaves close at night. The closer
temperatures are to freezing, the more tightly the leaves close; on warm cloudy
nights, the rosettes only loosely enclose the growing point. Flower buds may be
protected the same way but not only at night. Leaves may curl over the bud when-
ever weather turns cloudy and cool. One of the giants (Senecio brassica) in the upper
alpine zone of Mount Kenya has an extra advantage. It supplements its insulation
with a dense wooly covering on the underside of its leaves. When a rosette closes,
this wool provides extra warmth. The flowers on tall inflorescences, which are
exposed to air temperatures farther above ground level, are protected from
extremes of temperature by insulating bracts or hairs, or they are sunken into the
stalk. Buds on puyas in the Andean puna are protected deep within a hairy
inflorescence.
A unique method of preventing freezing has developed among the giant lobe-
lias on Mount Kenya. The broad leaf base catches water. The plant secretes a fluid
that lowers the freezing point of the water, keeping it liquid. The lobelia requires a
wet habitat so the prevention of freezing is necessary. A small fly takes advantage
of the warmer site and lays its eggs in the water, thereby protecting its larvae from
freezing.
Stems of both Andean espeletias and the African giant rosettes contain a pith
that stores water for use during times of low rainfall or when soil water is unavail-
able because the ground is frozen. So much water can be held that the mountain
gorilla of southwestern Uganda will snap off rosettes of giant groundsels to
obtain the watery pith. Plants from wetter habitats or warmer lower elevations
have less pith.
Giant rosettes also grow in tropical alpine zones where the surface soil may be
quite dry. The plants must be able to pull water up and through tall stalks. Even
though dead leaves keep the plant warmer than
outside air temperatures, the xylem (tissue that
transports water from roots to leaves) is suscepti-
ble to freezing at night. When night temperatures
are above freezing, giant groundsels and tall lobe-
lias on Mount Kenya do not alter transpiration
the next day. After a hard frost, however, some
do decrease transpiration, perhaps because the
xylem is too cold or frozen, causing a temporary
physiological drought. Many giant rosettes have
short stems so the water has less chance of freez-
ing overnight. Giant rosettes, however, rarely
suffer moisture stress because deep roots tap deep
soil moisture.
.................................................
Nature or Nurture?
Intense solar radiation or temperature changes
may be the reason for rosette growthform in
Tropical Alpine Biomes. Alpine plants were
moved from 13,700 ft (4,175 m) on Mount
Kenya to a lab in Nairobi at an elevation of
5,327 ft (1,624 m) and kept at 75 F (24 C).
After only 1027 days, leaves grewtwice as large
and stems elongated, destroying the compact
rosette form. Plants subjected to cold night
temperatures, however, did not change.
.................................................
164
Treeline Flora
Treeline flora, as well as alpine flora, in the Tropics is diverse because mountains
are isolated with little to no past or present connections to each other. Treeline in
Old World tropical mountains is dominated by members of the heath family
evergreen shrubs with tough, leathery leaves able to regenerate after fires. Tree heather
and Philippia heath species form shrublands in Equatorial East Africa. Shrubby Polyle-
pis, in the rose family, grows at high elevations of the tropical Andes, and true trees
can be found at 13,800 ft (4,200 m) or even higher. Evergreen alder ranges from Mex-
ico to northern Argentina and forms treeline where Polylepis is absent.
It is difficult to define treeline in tropical alpine mountains because giant
groundsels in eastern Africa and espeletias in South America can be called trees
because of their erect woody stems. In some regions, such as the west slope of the
Andes, desert conditions extend high into the mountains, and where there is no
forest, there can be no treeline. The lower limit of the alpine zone is often better
designated by climate conditions and plant growthforms.
Animal Adaptations
Faunas of tropical alpine environments are usually dominated by only a few fami-
lies or genera. Individuals are difficult to see in the field because they are most
often small and have dull colors, unlike lowland animals in the Tropics. Most are
also secretive, hiding in burrows or among rocks.
In spite of the decrease in both air density and oxygen, birds do not appear to
have special adaptations to high-elevation environments. They are already physio-
logically adapted to maximize intake of oxygen in their breathing mechanisms.
Lower air density does, however, mean that less lift is available and more energy
needs to be expended to fly. Birds with great wing area or smaller body size are at
an advantage. So in a seeming paradox, very small birdssuch as New World
hummingbirds and Old World sunbirdsand very large soaring birdssuch as
condors and Old World vulturesare the most common birds at high elevations.
Hummingbirds have another advantage in that they can survive a lowered body
temperature, called torpor, and achieve a corresponding decrease in metabolism.
A bird will go into torpor only on those nights when energy and food supplies are
low. Nectar-feeding sunbirds in the Old World and hummingbirds in the New
World are common pollinators in their respective tropical alpine areas.
In the Andean paramo, temperature can fluctuate up to 45 F (25 C) each day,
and nightly lows reach 20 F (6.5 C). Tropical alpine insects must survive or
avoid both nightly freezing and daily high temperatures with possible desiccation.
Insects shelter in rock scree or in vegetation, especially in dead leaves of espeletia
rosettes where temperatures remain above freezing. As many as 130,000 insects
have been estimated to live on a 4 ft (1.2 m) stem, with different species occupying
165
different parts of the plant. Most paramo insects cannot supercool and would
freeze without shelter. However, some grasshoppers stay in open vegetation
between rosettes and survive low ground night temperatures because they are toler-
ant of freezing. Similar patterns of insects occur in and around giant senecios on
Mount Kenya.
Regional Expressions of the Tropical Alpine Biome
South America
Treeline in the Andes Mountains is about 11,000 ft (3,250 m), above which is the
tropical alpine zone (see Figure 4.4). High-elevation landscapes in the northern
Andes, 10 N to about 5 S, are called paramo. From northern Peru through
Bolivia to northern Chile and Argentina, 522 S, high inland basins are called
puna or altiplano. Although both are basically grasslands, major differences include
amount of rainfall, height of grasses, associated plants, and animal life. With up to
75 in (1,900 mm) of precipitation, paramo is high-elevation wet grassland, while
puna with less than 27 in (685 mm) is high-elevation desert grassland. Giant
Figure 4.4 Alpine environments in tropical South America. (Map by Bernd Kuennecke.)
166
espeletia rosettes characterize parts of the paramo but are absent from the puna.
Large animals such as guanaco and vicuna graze the puna but not the paramo.
Landform differences are evident as well. In contrast to rugged alpine glaciation
and snow-capped volcanoes typical of the paramo in the northern Andes, the puna
is characterized by broad intermontane basins or altiplanos.
Paramo. The southern border of the paramo is the great Huancabamba Depres-
sion where the upper Mara~ n on River cuts deeply through the Andes in northern
Peru. While creating a corridor for species passing between the Amazon and the
Pacific coast, the low elevation is a significant geographic barrier for passage of
high Andean flora and fauna between north and south.
The paramo from Venezuela through Colombia to Ecuador is a moist grass-
land, an herbaceous ecosystem lying between forest at elevations below 9,850 ft
(3,000 m) and permanent snowline above 15,000 ft (4,500 m). It generally has
low mean temperatures with night and morning frosts and high humidity. The
Spanish name paramo, referring to sterile and uninhabited lands, reflects its tree-
less nature. It is an area of steep mountains, with typical alpine glacial landforms.
In Venezuela, much of the paramo occurs as isolated islands above cloud forest.
The ice layer in the soil is not deep like arctic permafrost, and melts daily, saturat-
ing the soil and causing solifluction. No seasonal changes occur, so the growing
season lasts all year. The paramo is dominated by grasses and rosette plants. With
its tall bunchgrasses or tussock grasses and giant rosettes, the Andean paramo is
similar to the alpine zones of Mount Kenya, especially in the resemblance in
appearance of some of the giant rosette plants.
The paramo climate is generally moist, with almost daily presence of mists.
Precipitation at 9,850 ft (3,000 m) varies from 27 in (685 mm) to 75 in (1,900 mm)
depending on location with respect to the Equator and position on the windward
or lee side of the Andes. It is only 36 in (900 mm), however, at higher elevations.
There may be either one or two rainfall seasons, but the driest months are January
and February north of the Equator and June and July south of the Equator. The
rainy seasons, April through June and August through November, respectively,
coincide with times of the year when the equatorial low pressure is near the Equa-
tor. Low precipitation in the dry season, however, makes parts of the paramo like
deserts for four or more months due to both evaporation and runoff on steep
mountains.
Temperature on the paramo can change dramatically throughout the day. At
night, temperatures drop below freezing, but they can rise to 73 F (23 C) by mid-
day. Many days, however, have mean temperatures around freezing. Glaciers still
cover several peaks, but they have been retreating rapidly in the last decade. With
little difference in daylength throughout the year because of its equatorial location,
the amount of sunlight received does not vary from season to season. Clear skies
versus cloud cover, however, affect the amount of incoming radiation reaching the
surface during the dry season and in the morning before clouds form. Upslope
167
winds cause cloud formation by early afternoon, blocking the intense equatorial
sun and chilling the air. Plants must be ready to withstand variations in solar radia-
tion, extreme heat, and freezing temperatures on a daily basis.
Temperature varies according to height above the ground and cloud cover.
During the dry season, temperatures 5 ft (1.5 m) above the ground average 27 F
(2.8 C) at night compared with 52 F (11 C) during the day, and temperatures
remain below freezing for 13 hours, from 8 PM to 9 AM. Temperature variation
even 4 in (10 cm) above the ground is similar, but variation in soil surface tempera-
tures is extreme, ranging from 14 F (10 C) at night to sometimes as high as 104 F
(40 C) during the day. During the wet season when it is cloudy, all temperature
variations are less extreme.
Soils are shallow with few nutrients and low water-holding capacity. Above
13,100 ft (4,000 m), in what is known as the superparamo, a daily freeze-thaw cycle
leads to solifluction, resulting in unstable soils. At slightly lower elevations, nightly
frosts create needle ice. Most soils are poorly developed and are well-drained enti-
sols or inceptisols. However, some areas are inundated with water all year, result-
ing in bogs with sphagnum moss. Most soils are acidic.
Vegetation. Even though by definition the paramo is treeless, high Andean wood-
land groves do occur. Polylepis trees grow in small stands above treeline, forming
vegetation islands where protected microclimates are warmer. In these areas, wet
scrublands consisting of reedgrass, bentgrass, St. Johns wort, as well as Polylepis
trees develop.
The flora is rich. Some 420 kinds of flowering plants occur in the Venezuelan
paramo, and more than 230 are in the superparamo. With no direct connection
now or ever with the Arctic, few arctic plants have reached the Andes, and most
species, some of which are unique to a particular
peak or cluster of peaks, derive from local low-
land floras. A major exception is Iceland pur-
slane, which dispersed from the Arctic south
via the Rockies and Andes all the way to Tierra
del Fuego. This plant may have had a dispersal
advantage because it is an annual and its seeds
can be transported long distances by wind or
migrating birds. Some forbs, such as geranium,
gentian, paintbrush, buttercup, and others, are
represented at the genus level in both the Arctic
and mid-latitudes.
The paramo contains several plant com-
munities. The most common growthform is
bunchgrass. The vegetation has two layers, an
upperstory of bunchgrasses and an understory
of mats, cushions, rosettes, creepers, subshrubs,
.................................................
Frailejones
Several rosettes in the sunflower family, espe-
cially espeletiascalled frailejones in Spanish,
meaning gray friarsare prominent in the
paramo and superparamo of Venezuela,
Colombia, and northern Ecuador. Some are tree
size, up to 10 ft (3 m) tall, while others are only
a few inches high. Most espeletias, grow at ele-
vations between 9,800 and 13,100 ft (3,000
4,000 m), but some may be found as low as
7,200 ft (2,200 m). Espeletia has a tall stem or
trunk with leaves at the top. Old leaves die and
drape down over the stalk. Plants depend on
insects, birds, and wind for pollination.
.................................................
168
lichens, and mosses. Grasses are diverse, with 30 species in the Venezuelan
paramo. Most are bentgrasses, bromes, reedgrasses, fescues, and needlegrasses.
Mosses, liverworts, and lichens are abundant and diverse. In spite of the moist cli-
mate, the paramo always looks as though it were autumn because dry vegetation
among new growth lends a brownish tinge to the landscape (see Figure 4.5). Wet
grasslands with tall bunchgrasses or tussock grasses often include giant rosette
espeletias and puyas. Boggy or swampy areas support large cushion plants, while
forbs, including alpine orchids, occupy open rocky areas. Dwarfed bamboo, shrubs
from several families (sunflower, mahogany, and St. Johns wort), and sedges also
occupy these areas. The moist ground may be covered with a dense mat of mosses,
lichens, and cushion plants. Wet, marshy soils, with rocky outcrops of volcanics
and other rock types, are the headwaters of streams. Notable flora in marshy areas
includes quinine shrubs.
Above 12,800 ft (3,900 m), in the desert paramo, short cushion plants become
more prominent. Around 14,000 ft (4,275 m), grasses decrease in abundance and
low shrubs, mats, and rosettes begin to dominate. Sandwort cushions form dense
colonies only a few inches high. In some places, large cushions of endemic Azorella
crenata and A. julianii grow with tall espeletia rosettes. With age, azorella cushions
can grow to 3.3 ft (1 m) high. The Andean desert paramo is primarily found in
the Cordillera de Merida in Venezuela, from 13,100 ft (4,000 m) up to the edge of
Figure 4.5 The paramo in the northern Andes is predominantly tussock grasslands.
(Photo by author.)
169
glaciers at 15,100 ft (4,600 m). Extreme habitats at this elevation are called perigla-
cial, characterized by daily freeze-thaw cycles, small-scale patterned ground, and
solifluction. These highest desert periglacial zones have sparse vegetation.
Three plant associations with giant espeletia rosettes occupy periglacial habi-
tats, each with a different species. One, up to 10 ft (3 m) tall and topped by an inflo-
rescence of 40 in (100 cm), is most widely distributed above 13,000 ft (4,000 m), on
steep slopes with unstable fine scree. Colonization stabilizes the slope, and plants
may live for 170 years. A different, smaller espeletia grows on rocky sites such as
aretes and cirque walls where large boulders retain heat, allowing the plants to sur-
vive at higher elevations. This species grows only 3 ft (1 m) tall and has a short
inflorescence; it also has the shortest life span of the three, 70 years. A medium-size
espeletia grows 7 ft (2 m) tall on gentle slopes with medium-size gravels. Its inflo-
rescence is 30 in (75 cm) long and the plant lives for 130 years. Long life is an
advantage because these three species only reproduce by seeds. Each plant pro-
duces 1 million seeds in its lifetime. Two other espeletias have prostrate stems
forming giant cushions or shorter rosettes, respectively. These two are more impor-
tant in the lower Andean zones but grow up to 13,800 ft (4,200 m) in wetter parts
of the desert paramo.
The three high-elevation paramo espeletias continue to grow after flowering.
Their inflorescences sprout from between young leaves near the top of the rosette,
even as the terminal bud continues to grow upward. Espeletias that die after flower-
ing because the terminal bud becomes the flower stalk are found only below 13,000 ft
(4,000 m).
Many plants at elevations of 14,000 ft (4,275 m) or higher have dense, wooly
hairs. One type of espeletia, a large rosette of thick, wooly, white leaves on a stalk
draped with the old dead leaves, grows at this elevation. This plant, the largest
espeletia species, can grow up to 15 ft (4.5 m) tall and is similar in appearance to
the giant groundsels of Africa. It grows tallest in wet areas, near streams, lakes, and
peat bogs and makes a kind of savanna with the grasses.
Other habitats in the high alpine zone include loose sand and gravel. Two
shrubs, chuquiragua (or flower of the Andes) and jata (or candlebush) are pioneers
on this substrate, growing until sufficient soil has developed for grasses to take root.
Occasional rock scree in the upper alpine within the permanent rock and snow at
13,50015,750 ft (4,1004,800 m) supports only scattered small rosette plants.
Animals of the paramo. The few mammals inhabiting the paramo are primarily
small herbivores, including paramo rabbits and several rodents. Rabbits are found
mostly under cover of St. Johns wort. Their burrows are in two sections, a small
one for refuge or escape, and a larger one used as the nesting chamber. Nests are
constructed of dry grass, moss, and hair, which insulates against cold temperatures.
Adaptations to the paramo include heavy fur, long gestation period, small litters,
and few teats. Grasses, especially bromes, are the rabbits main food. Rabbits are
170
preyed on by Black-chested Buzzard Eagle, White-rumped Hawk, owls, and an
occasional puma and paramo wildcat as well as by feral dogs.
Larger mammals of note include mountain tapir, little red brocket deer, north-
ern pudu, and the omnivorous spectacled bear. The mountain tapir is the only tapir
that does not live in tropical forests. Little red brocket deer and northern pudu are
both small ungulates that usually live in montane forests but, because of their small
size, can also take shelter in grassland. Brocket deer measures 28 in (70 cm) tall at
the shoulder, and pudu, the worlds smallest deer, is only 14 in (35 cm) high.
White-tailed deer are also found all along the Andes chain, living on valley floors
in small family groups of up to five individuals. Although they prefer Polylepis
stands, they also inhabit open and shrubby paramo. The only other large herbi-
vores are domesticated llamas and alpacas.
Thomas small-eared shrew is significant in the paramo ecosystem, building a
complex system of nests and tunnels beneath espeletias and St. Johns wort. It preys
on young rodents, eggs and chicks of ground-nesting birds, lizards, and insects, and
is itself prey for opossum, weasel, eagles, hawks, and owls. Long-tailed weasel is
another major small predator. Puma and paramo wildcat occasionally may be seen.
The most commonly seen animals are small birds. The high elevation of the
paramo makes it a convenient resting stop for migratory birds. Among the breeding
birds, nectar-seeking hummingbirds are important pollinators and have the advan-
tage of inducing torpor when temperatures drop. Alpine birds generally build larger
nests than do their lowland counterparts, and also choose sites that are more stable
in temperature. With few predatory snakes and lizards and no trees (except scat-
tered Polylepis groves) in the paramo, nests are built on the ground or in low bushes.
Small insect-eating birds are most numerous. Ground-feeding insectivores include
Paramo Pipit and Streak-backed Canastero, while Merida Wren and Sedge Wren
glean insects from small bushes. Fruit eaters, large seed eaters, and scavengers are
not common because fleshy fruits, large seeds, and large carcasses are rare. Eagles
are present but scarce, as are Andean Condors. Condors, extinct since the 1950s in
Venezuela, are being reintroduced, but there is concern that too few large animals
are available for food.
Many insects, birds, and even a small rice rat live in the sheath of dead leaves
that cover giant rosette stems. Simultaneous September to December flowering of
most giant rosettes attracts many pollinators. More then 1,000 insects were col-
lected around espeletia flowers in the Venezuelan paramo. Most abundant were
flies and midges, but bees and wasps were also well represented. Butterflies are con-
spicuous, and beetles are important both in numbers and species diversity, with
about 25 families represented in the Andean highlands. Carabid beetles, in particu-
lar, have many species in the paramo.
Although espeletia has chemical toxins to deter grazing by native animals, non-
native grazers are not affected. Much of the paramo has been converted to agricul-
tural land and is subject to overgrazing, a significant threat to the ecosystem.
171
Puna. From the Huancabamba Depression south to western Bolivia and northern
Chile and Argentina is the puna, at a general elevation of 9,85016,400 ft (3,000
5,000 m). The high ranges to the east face the easterly trade winds, and the puna in
their rainshadow is dry. Puna is cold grassland with drought-adapted bunch
grasses. Needlegrass is prominent, along with shrubs of the sunflower family.
Other common plants are fescue and reedgrass, with prostrate rosettes. The eastern
puna is more moist and has a closed grassy vegetation, while the central and west-
ern parts are dry and shrubby. Precipitation and plant growth patterns reflect the
occurrence of a single rainy season and an intense winter dry season. The wetter
puna in the east receives 30 in (750 mm) of rainfall. Conditions become drier both
to the west and south, until extreme drought is reached in the Atacama Desert at
the border of Bolivia, Chile, and Argentina. The decrease in precipitation is espe-
cially noticeable south of 15 S. Seasonality also increases toward the south, in the
form of a longer dry season. About 85% of annual precipitation falls in the rainy
summer, with almost none during the dry winter. Temperatures also vary accord-
ing to latitude. Toward the south, increasingly greater seasonality and colder
winter temperatures are experienced. Temperatures of 5 F (15 C) have been
recorded in the altiplano in southern Peru and 22 F (30 C) in the Bolivian
puna. The southern limit of the puna at 20 S lies where the annual temperature
range increases to 18 F (10 C), and winters are more severe. The altiplano has
been affected by livestock grazing for centuries, and it is questionable as to whether
the current vegetation represents natural landscape or managed pasture.
The wet puna, located between 12,100 and 13,800 ft (3,700 and 4,200 m) from
northern Peru to northern Bolivia, is covered with tussock grassland and shrubs
(see Figure 4.6). The Andean mountains in this area divide into two ranges, the
Cordillera Occidental and Cordillera Oriental. A central large plateau called the
Altiplano lies between them. The high peaks and valleys above the Altiplano are
covered with ice; glacial tongues may advance as low as 9,800 ft (3,000 m). Bent-
grass, reedgrass, fescue, and needlegrass (ichu grass) are the most conspicuous
grasses, with mountain bamboo and pampas grass in more humid areas. Poorly
drained regions in this zone support sedges, rushes, and bulrushes.
The high Andean puna, above 13,800 ft (4,200 m) on the Altiplano, has extreme
diurnal temperature changes because of the high elevation. Temperatures drop
below freezing nightly. Annual precipitation frequently falls as snow or hail instead
of rain. Grasses are fescue, Peruvian feathergrass, and reedgrass. Tussocks can be
3 ft (1 m) in diameter and just as tall. Prostrate or rosette plants include gosmore
and azorella cushions (see Plate XIII). Cushion bogs called bofedales are wet areas
above 13,000 ft (4,000 m), with large cushion plants submerged or floating in water.
The dry puna occurs in western Bolivia and northern Chile and Argentina. This
is a vegetation of primarily tropical alpine herbs and dwarf shrubs rather than
grasses. Characterized by low precipitation, 216 in (50400 mm) yearly, the dry
season here lasts eight months. This shrubland steppe has drought-resistant shrubs
such as Adesmia, broom rape, and tolillar, or grassy steppe with reedgrass, fescue,
172
and needlegrass. Shrubs may be 8 ft (2.5 m) tall. A vast inland sea on the Altiplano
in this region left behind extensive salt flats, called salares, including Coipasa,
Uyuni, Chalviri, and Arizaro (see Figure 4.7). Halophytic plants more commonly
found in desert areas such as saltbush, saltgrass, pickleweed, and seepweed grow in
this environment. Salt concentration in the tissue of saltbush is higher than that in
the soil, which allows water to move from the salty soil into the plant. To prevent
......................................................................................................
Puya
Although the genus is the only member of the pineapple family to grow in the high Andes, several
species of puya are common giant rosettes in both the paramo and puna (see Plate XIV). Two of
the most common are Puya hamata, which is an indicator species of the high paramo, and Puya
clava-herculis. Rosettes of both species are generally small, less than 2 ft (0.6 m) in diameter, but
the inflorescence can be 5 ft (1.5 m) tall. In contrast, the endangered Puya raimondii, which only
grows at 13,100 ft (4,000 m) or above, can reach 10 ft (3 m) in diameter. After 80100 years, this rare
plant flowers once, producing an inflorescence 15 ft (4.5 m) tall before dying. Up to 20,000 small
flowers on the stalk are protected from freezing by hairy bracts. Except for the giant P. raimondii,
puyas resemble in size and form the lobelias of the afroalpine zone in East Africa.
......................................................................................................
Figure 4.6 The Altiplano, seen here in southern Peru, is a high-elevation plain between
high mountains. Ampato Volcano is in the background. (Copyright C Dr. James S. Kus.)
173
even more salt buildup inside the plant
as evaporation takes place, the plant exudes salt
onto the surface of its leaves. The salty surface
gives the plant a grayish look and also renders it
inedible to most herbivores. Old Man of the
Andes cactus, named for its long white hairs,
grows on islands in the Uyuni salt flats. Bofedales
with large floating cushion plants also occur.
Animals of the puna. The most representative
native mammals are vicuna, guanaco, chinchilla,
and vizcacha. Members of the camel family such as
guanaco, vicuna, and the domesticated relatives al-
paca and llama have strong incisors that enable
them to crop plants with high silica content (see
Figure 4.8). Guanacos, probable ancestors to
domesticated llamas and alpacas, are wide-ranging,
but vicunas are restricted to high elevations above
.................................................
San Pedro Cactus
Growing in the mountains of Ecuador and Peru
at 5,0009,000 ft (1,5002,700 m), the San
Pedro cactus can withstand cold conditions. It
is a multistemmed columnar giant up to 20 ft
(6 m) tall, with arms that can spread 6 ft (2 m)
wide. Because its smooth green flesh contains
small quantities of mescaline and other hallu-
cinogenic compounds, it has a long history of
ritual use in pre-Columbian times. Peruvian
shamans, or curanderos, still drink a mixture
made from the plant. They believe it helps
them diagnose a spiritual or subconscious
basis for illness. The plant is legal to grow in
most countries as an ornamental, but, with few
exceptions, is illegal to use.
.................................................
Figure 4.7 The vizcacha rat is adapted to feed on salty plants around salt flats in the
Altiplano. (Courtesy of Susan L. Woodward.)
174
11,500 ft (3,500 m). The golden vizcacha is a
rodent well adapted to browsing on saltbush. Like
other rodents, it has both upper and lower inci-
sors. It also has an extra upper pair of teeth that
are actually made of stiffened, bristly hair, which
allows the animal to scrape salt off saltbush leaves
before consuming them. The animals kidneys are
specialized to excrete excess salt.
The Andean hairy armadillo, 12 in (30 cm)
long with a tail half again its size, primarily feeds
on small vertebrates and insects, but it will also
eat roots or tubers. Hairs between its plates help
keep it warm. Puma and Andean fox, are typical
predators.
Darwins Rhea, an ostrich-like bird standing
3 ft (1 m) high, is endemic to South America and
found on the puna as well as on lower-elevation
grasslands. Its diet consists of plants, seeds, roots, insects, and small vertebrates.
Except during breeding season, the birds live in groups of up to 30 individuals. The
male becomes very territorial when he incubates the eggs and cares for the young,
.................................................
Reed Boats
Bolivian Indians have traditionally built boats
from reeds growing around Lake Titicaca,
although they are now replacing them with
longer-lasting wooden boats. When Thor
Heyerdahl first attempted to cross the Atlantic
Ocean in a reed boat, Ra I, it failed just short of
his goal in the Caribbean. A year later, in 1970,
with an improved design and assistance from
four Aymara Indians in Bolivia, Ra II was built.
The journey from Morocco to Barbados was
successful, covering 4,000 miles (6,450 km) in
57 days. The boat remained intact and sea-
worthy to the end.
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Figure 4.8 Vicuna, in Salinas-Aguado Blanca Reserve north of Arequipa, Peru, is fre-
quently seen grazing the puna. (Copyright C Dr. James S. Kus.)
175
often adopting orphaned chicks into his flock.
Flightless, rheas are fast runners, attaining speeds
up to 37 mph (60 km/h). Rheas escape predation
by running a zigzag path or suddenly squatting in
brush and flattening their bodies against the
ground. The Puna Tinamou is another common
ground bird. Many endemic birds, including Ash-
breasted Tit-tyrant, Royal Cinclodes, three species
of canastero, Olivaceaous Thornbill, and Gray-bel-
lied Flower-piercer are widespread. Other endemic
bird species are more locally distributed. Polylepis
forests and scrub provide habitat for most of the
endemic birds, found in both wet and dry puna.
Large flocks of flamingo are found around the salt
lakes.
Afroalpine
Equatorial East Africa refers to Kenya, Uganda,
Tanzania, and adjacent countries, while the term
East Africa also includes Ethiopia, which has
some dissimilarity in climate and landscape (see
Figure 4.9). Afroalpine applies to all mountains
within 10 to 15 N and S of the Equator that rise
above treeline. With two exceptions, the Rwen-
zori Range and the Ethiopian Plateau, the afroal-
pine region consists of geologically young,
isolated volcanic mountains that arose along the
Eastern African Rift zone from Lake Malawi in
the south to the Red Sea in the north. The largest
volcanoes are the Virunga group along the west-
ern rift, and Aberdare, Mount Kenya, Mount Kili-
manjaro, and Mount Meru on the eastern rift.
Age varies from more than 15 million years for
Mount Elgon and 2 million years for Mount
Kenya to less than 200,000 years for Mount Meru,
a factor affecting colonization and current vegeta-
tion. The Rwenzori Mountains consist of older,
Precambrian intrusive igneous and metamorphic
rocks. The Ethiopian highlands are a lava plateau
broken by the East African Rift zone. At least 10 mountains exceed 13,000 ft
(4,000 m). Mount Kenya at 17,058 ft (5,199 m) and Mount Kilimanjaro at 19,340
ft (5,895 m) are the highest. Glaciation has sculpted most peaks, and glaciers still
exist today on the Rwenzoris, Mount Kenya, and Mount Kilimanjaro.
.................................................
Chinchillas
Due to demand from the fur industry for their
soft, furry coats, these rodents were hunted
almost to extinction, until the practice became
illegal in 1929. Captive-raised animals that pro-
duce superior pelts minimized pressure on wild
populations, but decimation of wild animals
was so severe that chinchillas are still considered
endangered or threatened. It is unknown how
many remain in the wild. The Andean Mountain
cat, the size of a domestic cat, is equally rare.
Although its habits are largely unknown, it pre-
fers high rocky mountains where chinchillas
take shelter under rocks. It is likely that the cat
may have depended on chinchilla for food.
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Flamingos
Three of the worlds six flamingo species are
found in the high Andes. In spite of frigid winter
temperatures, hot springs that moderate water
temperature help make the Altiplano salt lakes
prime flamingo habitat. The attraction is an
abundance of algae, brine shrimp, and other
aquatic invertebrates. The rare and endangered
James and Andean Flamingos, which live year-
round in the Altiplano, are joined by the more
abundant Chilean Flamingo during breeding
season. With their beaks almost constantly mov-
ing through salt water while feeding, flamingos
also ingest salt, which is excreted by special
glands in their nostrils. The birds obtain fresh
water from springs, puddles, or from their own
feathers saturated with rain water.
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176
The afroalpine flora is much less diverse than that of the Andes. Because high
peaks are isolated and not a continuous chain, unique species occur on each moun-
tain. More than 80% of the flora is endemic to the high mountains of East Africa,
but not all species are confined to the afroalpine. With no corridors from the Arctic
or from Central Asian highlands, few northern plants other than saxifrages and
primroses are found. The flora instead has affinities with several other floral
regions, including South African, Northern Hemisphere temperate, Mediterra-
nean, and Himalayan.
Climate. Few records exist, but differences in vegetation and zonation imply dif-
ferences in climate. Two rainy and two dry seasons are typical. WinterDecember
through March in the Northern Hemisphere and June through August in the
Southern Hemisphereis driest. The majority of precipitation falls in the summer,
April through May and November through December, respectively. Precipitation
Figure 4.9 Alpine environments in Tropical Africa. (Map by Bernd Kuennecke.)
177
varies with exposure, and because moisture-laden winds come from the south and
southeast, those sides of all peaks and mountain chains receive more rainfall.
Total rainfall and lengths of wet and dry seasons vary considerably. The Rwenzo-
ris are always moist, Mount Kenya has one month of drier conditions, and Mount
Kilimanjaro has 411 months of drought. Annual rainfall in the alpine zone of
Mount Kenya, more than 12,000 ft (3,650 m), is 35 in (900 mm), with less rainfall
received at higher elevations. Mount Kilimanjaro is drier, receiving only 30 in
(750 mm) at 12,500 ft (3,800 m), and only 5 in (125 mm) above 13,000 ft (4,000
m). The upper alpine zone of Mount Kilimanjaro is more barren than Mount
Kenya because no sheltered valleys occur and the more porous substrate holds lit-
tle water.
Although mean annual temperature is similar to that of both arctic tundra
and mid-latitude alpine, major differences in the temperature regime exist. Major
contrasts include a year-long growing season and a daily freeze-thaw cycle.
Because of its equatorial location, the growing season is not limited by cold win-
ters. As is true for other tropical environments, afroalpine zones are characterized
by small seasonal changes and extreme daily temperature changes. On Mount
Kenya at both 10,000 ft (3,000 m) and 17,000 ft (5,200 m), mean monthly tempera-
tures vary only 3 F (1.5 C) throughout the year. Temperature decreases with ele-
vation, from an annual mean of 45 F (7 C) in the forest at 10,000 ft (3,000 m) to
35 F (1.5 C) in tussock grassland at 13,750 ft (4,200 m), to 18 F (7.5 C) in
rocky areas at 15,650 ft (4,750 m) with glaciers nearby.
Due to both high elevation and equatorial location, intense solar radiation
received during the day is lost as infrared radiation at night and temperatures
drop below freezing. Rain and cloudiness affect temperature. Mount Rwenzori,
with more clouds and rain, experiences only an 11 F (6 C) daily temperature
range, while on Mount Kenya, where cloudy conditions are less common, the
day-to-night temperature change is 22 to 32 F (12 to 18 C). On the more conti-
nental Sanetti Plateau in the Bale Mountains of Ethiopia, diurnal temperature
changes are even more extreme. All these measurements, however, indicate the
air temperature, not the microclimate at ground level. When air temperature at
14,800 ft (4,500 m) is 38 F (3.5 C), temperature at ground level or inside vegeta-
tion clumps can be a much warmer 67 F (19.4 C). Extreme differences between
ground and air temperatures can be 56 F (31 C) at midday. Ground tempera-
tures change rapidly in response to sunny or cloudy conditions. Under a cloud
cover or in shaded sites, air and ground temperatures are similar. The lowest
ground temperature measured in the lower alpine on Mount Kenya was 16 F
(9 C). Although such low temperatures are infrequent, heavy frost may occur
nightly. Topography contributes to microclimate conditions, especially cold air
drainage onto valley floors where more extensive solifluction disrupts soils and
prevents colonization by plants.
Because of daily fluctuation in temperature, the relative humidity changes are
extreme. Relative humidity of the air on Mount Kenya drops from 90% in the early
178
morning to 20% or less at midday. At ground level, however, it remains fairly con-
stant, at about 70%. Because of increasing relative humidity at night, moisture
from dew and frost is significant. Wind has a local effect, and plants tend to grow
in the lee of boulders near ridgetops and glaciers.
Soils. Soils are derived almost solely from volcanic material but vary according to
rock chemistry, ash deposits, slope, and precipitation. All are subject to serious ero-
sion, often accelerated by overgrazing and agriculture. Fresh moraines are com-
mon parent material, coarse at valley sides and fine on valley floors where they
retain water, leading to the formation of bogs. Because it is too cold for much
chemical activity, soils are acid. Bare, steep slopes on valley sides are subject to sol-
ifluction. Small sorted polygons 5 in (13 cm) diameter on ridgetops above 14,000 ft
(4,250 m) are the result of either frost action due to daily temperature fluctuations
or to desiccation cracking. Finer material concentrates in the center while the edges
are outlined by gravel. Mosses first take hold in the gravel edges, then are suc-
ceeded by grasses. Below 14,000 ft (4,250 m) where more tussock vegetation covers
and insulates the soil, frost action is less common. Finer sediment on flat valley
floors over 12,000 ft (3,650 m), however, is disturbed by needle ice, especially in
bare soil between grass tussocks or on dried lake beds, preventing the establishment
of plants.
Vegetation. Altitudinal belts of montane forest, heath, and afroalpine vegetation
are largely intact in Equatorial Africa; however, human-induced fires on Mount
Elgon and Mount Kenya, have modified the afroalpine environment. Alpine zones
in Ethiopia are highly disturbed by fire, wood collection, grazing, and agriculture.
Montane forest, depending on location, primarily is composed of hardwood trees
with a few conifers and often a bamboo zone. The heath vegetation is a distinct
treeline zone, with trees and shrubs of tree heather and two Philippia species.
The alpine zone extends from the upper edge of the heath zone, about 12,000 ft
(3,650 m) to the nival zone at 15,000 ft (4,500 m). Although no clear demarcation
exists between the forest and heath belts and the alpine zone, few plants are able to
survive the harsh afroalpine environment, resulting in a flora that differs from adja-
cent lowlands. Heath shrubs diminish in number and are replaced by giant rosettes
and tussock grasslands as elevation increases. The upper limit of vascular plants,
marked by an occasional strawflower, is 16,000 ft (4,875 m), but lichens are found
on summits above 17,000 ft (5,200 m).
Three growth layers define afroalpine vegetation. An aerial level contains the
giant rosette tree senecios, below which is an herb or shrub layer, usually of straw-
flower and grass tussocks. The ground layer consists of low rosettes and prostrate
plants. Although the alpine zone is characterized by tussock grasses, the most
obvious plants are lobelias and seneciosboth giant rosettesand ladys mantle
and strawflower (see Plate XV). Many interesting alpine species are endemic to
Mount Kenya or to East African tropical mountains in general.
179
All lobelias are low, compact rosettes with inflorescences up to 6 ft (2 m) tall.
Woody, creeping stems remain prostrate along the ground. While Lobelia keniensis
rosettes, endemic to Mount Kenya, grow from a single prostrate stem in the alpine
zone, related species in the montane forest below have several erect branches. The
growing points of alpine lobelias are suppressed by cold, allowing only one stem to
grow. The rosettes close their leaves into tight buds at night to protect the growing
point from cold. The stems and inflorescences of alpine lobelias are hollow, provid-
ing yet more insulation. While air temperatures drop to 23 F (5 C), the core of
the plant remains above freezing at 35 F (1.7 C).
L. keniensis commonly grows in groups connected by shallow underground
stems. Leaves are broad without hairs, and the inflorescence grows up to 5 ft
(1.5 m). The plants grow in moist areas and reach maximum density in the middle
alpine elevations. A different lobelia, Lobelia telekii, endemic to Mount Kenya,
Aberdare, and Mount Elgon, grows on well-drained, coarse substrate, primarily in
the high alpine. It is differentiated by its narrow leaves with a waxy covering. Hairy
bracts hide the flowers on the 6 ft (1.8 m) inflorescence.
Giant groundsels (genus Dendrosenecio) and giant senecios (genus Senecio) vary in
growthform according to elevation, aspect, and microclimate, and dominant species
vary with habitat. All continue to grow after flowering and all possess insulating
layers of dead leaves around their stems. In several instances, people have used these
leaf girdles as a ready source of fuel, killing the stripped plants. Giant groundsels are
treelike with woody stems. Not all giant rosettes are upright. Senecio brassica, a giant
senecio endemic to Mount Kenya and growing at lower elevations in the alpine
zone than any other large senecio, has a low rosette of leaves, densely hairy on the
underside, topping the end of a woody stem that creeps along the ground surface.
The tall inflorescence, up to 3 ft (1 m), has bright yellow flowers. Some senecio spe-
cies also occur as low, creeping herbs that can form mats up to 10 in (25 cm) high.
Of the several plant communities found in the afroalpine zone, the five most
important are unevenly distributed. Although defined as above treeline, the afroal-
pine has distinctive woodlands or forests of tree-like giant groundsels, usually
growing on deep soil with access to underground water. These giant rosettes occur
on all high mountains except Mount Meru, where it may be too dry, and are one of
the plants found at the highest elevations. The genus (Dendrosenecio) is endemic to
Equatorial East Africa and absent from Ethiopia. Different but related species have
evolved on each of the isolated mountains. Strawflower scrub also has different
species in different locations. Found on rocky ground in all mountains, including
Ethiopia, it is most impressive on the Rwenzoris, where it forms a dense scrub up
to 6.5 ft (2 m) tall. Ladys mantle scrub is found on gently sloping, well-drained
ground, and different mountains have different dominant species (see Figure 4.10).
Dull brown tussock grassland dominated by fescue, but also containing bentgrass,
bluestem, hairgrass, and Pentaschistis, is found on all mountains, but it is less com-
mon in wetter areas such as on the Virunga volcanoes and in the Rwenzoris. Grass-
land grows on well-drained, somewhat steep soils and replaces ladys mantle scrub
180
where the latter has been burned. Carex bogs form on flat or gentle slopes with
poor drainage and usually create peat. Although found in the afroalpine, bogs are
more important in the heath zone and are absent from Mount Meru. On wetter
mountains, epiphytic mosses, lichens, and liverworts grow on the trunks of giant
groundsels, especially where the leaf shield has been destroyed. Lichens predomi-
nate on drier mountains.
Figure 4.10 Ladys mantle is a common shrub in East African alpine environments.
(Courtesy of Rainer W. Bussmann, Missouri Botanical Garden.)
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Case Study: Altitudinal Zonation of Mount Kenya
Mount Kenya is an extinct volcano on the Equator, with many other volcanic cones dotting its
north and northeast slopes. Deep valleys, glaciated and U-shaped, radiate from its peak. Teleki Val-
ley is one of the largest with smooth walls and craggy aretes. Steep valley walls, however, are not
subject to erosion because surface scree allows water to drain through. Moraines are common.
Mount Kenyas craggy summit and glaciated landscape with steep valleys contrasts with the
smooth and rounded features of Mount Kilimanjaro.
181
The heath zone forms a less distinctive belt on Mount Kenya than it does on other East African
mountains (see Figure 4.11), in part because it has been burned by human activities and only
stunted shrubs remain. Composition varies due to different climate on different aspects. Protea
may form closed communities up to 13 ft (4 m) tall, with undergrowth according to habitat. Tree
heather and Philippia species up to 12 ft (36 m) tall occupy rocky ridges and moraines, where the
ground is covered with a rich growth of mosses. Usnea lichens may be draped over heath plants.
On boggy ground, fescue grass and carex sedge tussocks dominate, with ladys mantle growing
between the clumps. Because the tussocks cover most of the ground, the general appearance is
grassland with occasional giant rosettes of Lobelia keniensis.
The treeless environment can be divided into lower alpine, upper alpine, and nival zones,
based on dominance of different giant rosettes, even though considerable overlap of species and
interdigitation of communities occur, depending on slope, aspect, drainage, and other local char-
acteristics. (Zones are more precisely defined on mountains not subjected to glaciation, such as
Mount Kilimanjaro, where aspect and slope are less variable.)
The lower alpine zone, roughly 12,000 ft (3,650 m), is marked by the abundance of the ground-
level giant senecios (Senecio brassica) and lack of the tree-like giant groundsels (Dendrosenecio
keniodendron). Two basic habitats, meadows and ridgetops, are encountered. The flat or gently
sloping meadows and valley bottoms are usually wet and characterized by tussock grasslands, an
almost continuous cover of fescue with few or no shrubs. Fescue grass forms clumps to more than
3 ft (1 m) high, and smaller carex sedge tussocks also occur. Mosses and smaller rosettes grow in
the dark humus soil between tussocks. Scattered groups of lobelia rosettes, 1 ft (0.3 m) high and
18 in (45 cm) in diameter, occupy semibogs where soil is rich in humus and kept damp by seepage
or standing water. Tussocks of nodding hairgrass with patches of prostrate senecios grow where
meadows border rocky areas.
Weathered and eroded ridgetops and sometimes old eroded moraines offer a rockier, more
exposed, and drier habitat with plant communities that continue into the upper alpine zone.
Figure 4.11 Vegetational zonation on Mount Kenya. (Illustration by Jeff Dixon.)
182
East African alpine animals. Equatorial East African mountains are not well
inventoried, but the more conspicuous species are known. Mammalian herbivores
on Mount Kenya include rock hyrax, groove-toothed rat, and the common duiker.
Moraines or crags adjacent to tussock grassland and water constitute their habitat,
each animal occupying a microenvironment. Hyrax live among the rocks, rats
dig burrows at the bases of giant senecios or grass tussocks, and duikers prefer
giant groundsel forest. Hyrax is commonly seen in the afroalpine, with different
subspecies on different mountains (see Figure 4.12). They eat mosses and grasses,
types varying according to proximity to their burrows. Fur on the mountain hyrax
is over 2 in (5 cm) long compared with only 0.5 in (1.3 cm) on lowland savanna
species, and the alpine form has a larger body size. Also unlike the lowland hyrax,
they drink water, often trampling trails to lakes or streams. Although their burrows
are just deep holes between rocks, temperatures inside are above freezing and
A cover of nodding hairgrass and other grasses extends up to 15,000 ft (4,500 m), the grasses
becoming shorter with increased exposure. Few rosettes grow between the grass tussocks
because drainage is excessive and soils become dry, and frost-heaving is too severe for soils to
remain stable. Creeping plants and woody plants such as strawflower are found only in sheltered
spots. Scattered giant groundsels, more common at higher elevations, grow on well-drained
ridges, indicating a somewhat harsher environment and transition into the upper alpine zone.
Dung accumulations near hyrax burrows support dense growth of an endemic succulent.
Characterized by the appearance of giant groundsels in distinct communities, the upper alpine
zone lies at elevations of about 14,000 ft (4,250 m). Valley walls, the steeply sloping sides of glaci-
ated valleys, where soil is damp but well-drained, are typically covered by a pattern of a groundsel
and a senecio, according to moisture requirement. Higher on the slope, tall tree-like giant ground-
sels are found on the well-drained, shallow soils with access to abundant subsurface water, while
the prostrate giant senecios grow lower down the valley wall on waterlogged finer soils and
around lakes. Thick masses of ladys mantle shrubs grow at the base of the giant groundsels. Pro-
tected areas enclosed by steep cliffs allow heath vegetation to grow at higher-than-usual eleva-
tions. Solifluction terraces and damp, flat ground on valley floors support fescue or bentgrass
tussocks. Ridgetops are plagued by wind and intensive frost. Sparse grass tussocks exist up to
14,500 ft (4,400 m) on smooth ridges, but almost none occur over 15,000 ft (4,500 m). Ladys man-
tle stands occupy ground just below the ridgetops, along with woody strawflowers. Rocky out-
crops are more varied, but include herbaceous senecios in the lee of boulders in windy areas. The
small Senecio purtschelleri is common around hyrax colonies.
Rather than permanent snow and ice, the nival zone on Mount Kenya refers to elevations
above 15,000 ft (4,500 m) from which glaciers have recently retreated. Stunted plants grow in shel-
tered locations, and pioneering species such as herbaceous senecios (Senecio keniophytum and
S. purtschelleri) can be found just 25 ft (7.5 m) from the ice. Browns strawflower, a hardy shrub and
the highest plant on the mountain, is found sheltering in rock cracks with little soil at elevations
up to 16,000 ft (4,875 m) that have long been ice-free.
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183
considerably warmer than the outside air. Plants growing at burrow entrances are
not eaten, which helps to conceal burrow locations.
Long-furred groove-toothed rats are commonly found on tussock grassland.
They burrow into the base of giant groundsels and then up into the plant as far
as the base of the leaves where temperatures remain more constant. The rats feed
on plant roots and seeds. Fewer in number, duikers live in the heath zone or in
giant groundsel forest where they browse on ladys mantle and other woody
plants. Other small mammals include a shrew, the harsh-furred mouse, and giant
mole rat.
Predators, feeding mainly on hyrax, include an occasional leopard, wild dog,
and several raptors such as Mackinders Owl, Augur Buzzard, Verreaux Eagle,
and Lammergeier.
The three most common birds on Mount Kenya are the Scarlet-tufted
Malachite Sunbird, the Hill Chat, and Streaky Seed Eater; all live in leaf girdles of
giant groundsels. The sunbird, which breeds all year in the alpine zone, takes good
advantage of giant rosettes. It uses its long bill to get nectar from deep inside lobelia
flowers and also feeds on insects, especially flies that live in the inflorescences. It
also eats midges that breed in the pools of water that collect in lobelia rosettes, and
strips the hairy covering from senecio leaves to line its nest. The Hill Chat, common
in both heath and alpine environments, feeds mainly on carabid beetles, weevils,
and spiders from the ground and from the leaves of senecios and lobelias. It does
Figure 4.12 The herbivorous hyrax lives in burrows between rocks in the afroalpine.
(Courtesy of Shutterstock. Copyright: Steffen Foerster Photography.)
184
not, however, touch insects on the lobelia inflorescences used by the sunbird. The
seed eater of course eats seeds.
Insects also take advantage of the microclimates in plants. Two moths living in
fescue tussocks build silk tubes from the base to the outer leaves. The insects move
through the tube in response to diurnal temperature fluctuations, spending the hot-
test part of the day and cold nights at the base of the plant and only venturing to
the outer leaves in early morning and evening when air temperatures are more
moderate. Insects take shelter from the cold in other animal nests, under rocks,
rosettes of lobelias or senecios, or in dead leaves on giant groundsel stems. Midges
breed in lobelia rosettes and shelter in giant groundsels. Many specialized insects,
especially beetles, are restricted to the environment of mole rat burrows. Some spe-
cies are apparently limited to one mountain or habitat, but there has been little
study of afroalpine invertebrates to confirm this. However, many species have
evolved; in one small area on Mount Elgon, 22 different species and subspecies of
the carabid beetle genus Trechus were counted.
The only resident reptile is the alpine meadow lizard, which lives in tussocks
and under stones.
Ethiopian Montane Moorlands
The Ethiopian plateau is a large expanse of land more than 6,500 ft (2,000 m) in
elevation with mountain chains and peaks more than 9,850 ft (3,000 m) high.
Although the Simen Mountains in the north are the highest, with Ras Dejen at
15,158 ft (4,620 m), the Bale Mountains in the south make up the largest continu-
ous afroalpine area. In contrast to the mountains of East Africa, which are sur-
rounded by wetter climates, the Ethiopian highlands rise from dry lands. The
mountain climates are largely unknown, but change according to elevation.
Annual rainfall varies considerably, from 100 in (2,500 mm) in the southwest to
40 in (1,000 mm) in the north. Depending on locale, the dry season can be as short
as two months or as long as 10. Frost occurs all year, but especially in winter from
November to March. Diurnal temperature ranges can be extreme, from 5 F (15 C)
at night to 79 F (26 C) during the day.
About 80% of the land above 9,850 ft (3,000 m) in Africa is in Ethiopia. The
lower part of the alpine zone is a montane forest or grassland now degraded by
grazing. Several alpine lakes add interest to the landscape. The flora here has affin-
ities with both Palaearctic and Afrotropical plant realms, and high levels of endem-
ism are found in this isolated volcanic upland. The Bale Mountains are a center of
endemism, with 20% of their wild mammals unique to the range. Development of
the African Rift Valley divided the plateau and its biota into two parts. The highest
areas were glaciated during the Pleistocene and were uncovered and colonized
only a few thousand years ago.
Adaptations to the high-altitude environment include gigantism, such as giant
lobelias, tree heather, and giant St. Johns wort. Many St. Johns worts are herba-
ceous or shrubby, but in Ethiopia, the species grows up to 40 ft (12 m) tall with a
185
trunk 10 in (25 cm) in diameter (see Figure 4.13). Some adaptations serve to reduce
transpiration. Leaves of lobelias are thick and leathery, and many species have a
small leaf surface area. The dry, paper-like flowers on many perennial plants are
able to withstand harsh winds.
Vegetation is predominantly a grassland with shrubs or a moorland domi-
nated by strawflower scrub and tussock grasses. The dominant treeline scrub con-
sists of Philippia, tree heather, and other shrubs. Smaller plants in the primarily
bare soil include strawflower, ladys mantle, Junegrass, and hairgrass. Wetter
areas are dominated by carex sedges. Above 12,150 ft (3,700 m), carex sedge
and fescue grasslands with giant lobelias dominate. A single species of lobelia
is the only giant rosette in Ethiopia. In the highest regions, a fescue and straw-
flower scrub community extends all the way to mountain summits, although
high-elevation cliffs and rocky slopes have little vegetation. The Ethiopian
Figure 4.13 St. Johns wort grows to unusually large sizes in the Ethiopian highlands.
(Courtesy of Rainer W. Bussmann, Missouri Botanical Garden.)
186
highlands differ from other afroalpine mountains in that they have few shrubby
ladys mantles and no giant groundsels. They also have African rose and yellow
primrose, typical genera of Europe or Asian alpine areas. Lichens often cover
woody trees or shrubs.
The Ethiopian highlands is a unique area in terms of animals and has several
endemic and threatened mammals. Simen National Park was named a World
Heritage Site in 1978. Significant is the extremely rare Ethiopian wolf, also called
the Simen fox. It lives on the open moorlands above 9,800 ft (3,000 m) and eats
burrowing rodents such as the giant mole rat. Evolutionarily related to the Euro-
pean grey wolf, it provides a major example of prior connections with Eurasia. The
giant mole rat, a large endemic rodent, also has a European origin. Other rodents
in the wolfs diet include diurnally active grass rats, and Starcks hare. Speciation
among rodents has been high. Other rodents endemic to the Ethiopian alpine are
the giant climbing mouse, narrow-headed rat, and black-clawed brush-furred rat.
Near-endemic animals include Walia ibex, mountain nyala, and gelada baboon.
The ibex, nyala, and baboon, along with klipspringer and rock hyrax found in
rocky habitats, point to relationships with desert ecosystems.
Alpine lakes and streams provide excellent avian habitat, particularly to over-
wintering Palaearctic birds. Ducks that can be seen in the thousands include
wigeon and shovelers. Wading birds such as Ruffs, and Greenshanks are also
abundant. Golden Eagle, Chough, and Ruddy Shelduck, all Palaearctic species,
breed in the Ethiopian alpine, their only known breeding site outside the Northern
Hemisphere temperate zone.
Good rainfall has long attracted agricultural peoples to the Ethiopian Plateau
and continues to foster increasing human populations. The Ethiopian highlands
have been cultivated for centuries and much of the area is degraded. The more in-
hospitable environments of most of the afroalpine zone have made it easier to pre-
serve. The montane and heath zones in the mountains of Equatorial East Africa,
however, are affected by human activities.
Oceania: Hawaiian Islands
Some 2,500 mi (4,000 km) from North America and 2,000 mi (3,200 km) from the
nearest islands of comparable height in French Polynesia, Hawaii is one of the most
isolated groups of islands in the world. Mauna Kea, on the big island of Hawaii, is
highest at 13,802 ft (4,207 m), closely followed by nearby Mauna Loa at 13,681 ft
(4,170 m). Haleakala on Maui is only 10,026 ft (3,056 m). Because these volcanoes
built up from the seafloor and have never been connected with a continent, many
continental lifeforms such as ants, coniferous trees, most land birds, and all mam-
mals except a bat, were unable to reach the islands. Long-distance dispersal, by
means of wind, rafting on the ocean, or attached to feathers or feet of birds was infre-
quent, and plants or animals that did arrive evolved in isolation. The diversity of
Hawaiis biota stems from adaptive radiation, meaning that immigrant plants and
187
animals evolved into many different species to occupy vacant niches or habitats. Pol-
ynesians colonized the islands about 300 years ago, bringing with them pigs, jungle
fowl, dogs, and Polynesian rats, and (unintentionally) geckos, skinks, and snails.
Many food plants such as taro were also introduced. Cattle, goats, and sheep were
introduced by Europeans at a later date.
Even though Hawaii lies at or near tropical latitudes (20 N), summits of the
higher mountains are cold. Temperatures on Mauna Kea average 39 F (4 C), and
the mountain was glaciated in the Pleistocene. Snow frequently falls on Mauna
Kea and Mauna Loa at elevations as low as 8,200 ft (2,500 m), but only occasion-
ally on Haleakala. Fresh lava or areas of cinders and ash have seen little soil
development.
Because Hawaii is under the influence of both the trade winds and subtropical
high pressure, an inversion layer develops. Warm moist trade wind air becomes
cooler as it rises, while the descending air in the high-pressure cell becomes warmer
as it sinks. This results in warm air overlying cool air at an altitude between 5,000
and 7,000 ft (1,5002,150 m), called an inversion, because the normal trend of tem-
peratures becoming cooler with elevation is turned upside down or inverted. The
inversion layer prevents trade wind air from rising any higher, blocking moisture
from the alpine zone. Haleakalas summit gets less than 30 in (750 mm) of precipi-
tation annually, and Mauna Kea is even drier, with less than 15 in (380 mm). The
dry, clear skies on Mauna Kea provide ideal conditions for astronomical observa-
tions, and one of the worlds major observatories sits on its summit.
Closed forest extends up to about 6,500 ft (2,000 m), which coincides with the
level of trade wind inversions. Treeline generally occurs about 9,000 ft (2,750 m)
and is characterized by shrublands or open parkland with small ohia trees. The
change from forest to alpine scrub above treeline is abrupt due to the inversions
effect on precipitation. Not only is the alpine zone much drier than the lowlands, it
also receives intense radiation due to the absence of cloud cover.
Three sparsely vegetated life zones occur above treeline: alpine scrub, alpine
desert, and the aeolian zone. Around 8,200 ft (2,500 m) elevation, an alpine scrub-
land dominated by prostrate shrubs of pukeawe, Hawaiian blueberry, mamame,
pilo, cranesbill, and tarweed typifies the change from wet forest to dry subalpine.
The lower stature of plants is caused by low precipitation at this elevation, most of
it falling as winter snow. Nightly frost is common. On drier slopes the alpine scrub
extends to lower elevations. At 8,000 ft (2,450 m) on the northeastern side of
Haleakala, an open grassland with endemic hairgrass bunches dominates. At the
highest levels of plant growth, at about 11,300 ft (3,450 m) and just before the
alpine desert, prostrate shrubs of pukeawe and Hawaiian blueberry grow with sev-
eral bunchgrasses, such as hairgrass, panicum, and bentgrass, carex sedges, and
ferns. Two conspicuous alien (nonnative) species in the alpine scrub are gosmore,
a plant that resembles a dandelion, and wooly mullein. Except for a few hardy indi-
viduals, the alpine scrub level is the zone where silverswords, the giant rosettes of
Hawaii, grow.
188
Above the alpine scrub zone to the upper limit of plant life at about 11,300 ft
(3,450 m) is an alpine desert of fresh ash, cinders, and lava flows extending from
the summits of Mauna Kea, Mauna Loa, and Haleakala. Frost occurs almost every
night. Permanent ice can be found less than 3.3 ft (1 m) beneath the cinder surface
on the summit cones of Mauna Kea, and Haleakala has some active patterned
ground such as stone stripes. Plant life is limited to lichens and mosses.
The highest part of the mountains, above the alpine desert, is the aeolian zone.
Life is limited to native arthropods adapted to the extreme cold and barren environ-
ment and surviving on plant and animal material carried on upslope winds. Cater-
pillars of a flightless moth on Haleakala survive by eating leaves trapped in their
webs. At least one spider eats insects blown upslope. The Mauna Kea wekiu bug
and its close relative the Mauna Loa wekiu bug are also flightless. Instead of a veg-
etable diet like their low-elevation relatives, they suck the body fluids of cold or
dead insects. A change in their blood that prevents freezing in spite of low tempera-
tures is so powerful that the bugs are extremely sensitive to heat, even that of a
human hand.
Most other animal life in the Hawaiian alpine consists of introduced animals
that have obliterated native species. Pigs, rats, dogs, and jungle fowl dominate.
Both grassland and shrubland in Haleakala National Park incurred considerable
damage from pigs before the 1980s when they were eliminated from the park.
While digging for preferred food of gosmore and bracken fern, they disrupted
native plants, which allowed exotics to spread and displace even more natives.
Goats, sheep, and cattle were also detrimental to native vegetation.
Silverswords. Silverswords (ahinahina in Hawaiian), endemic to the islands of
Hawaii and Maui, are rosettes of stiff succulent leaves that grow in the alpine scrub
regions of Mauna Kea, Mauna Loa, and Haleakala (see Plate XVI). Although they
appear silvery white, the long, pointed leaves are actually lime green but densely
covered with silky silver hairs. Silversword habitat receives some of the highest
solar radiation ever measured, and the silver hairs, easily damaged by touching,
reflect both visible and ultraviolet light.
Silverswords and greenswords are among three genera and 30 species in a
group of related plants called the Silversword Alliance that includes herbs, vines,
shrubs, trees, and rosette plants, all of which are endemic to Hawaii. The alliance
is a good example of adaptive radiation and convergent evolution because all the
species evolved from a North American tarweed in the sunflower family that
arrived in Hawaii about 5.2 million years ago. The taxonomy of silverswords is in
flux, with questions as to whether some are species or subspecies.
Of the 23 species of Hawaiian tarweeds, only some with succulent leaves grow
in the alpine zone. The three species of silverswords and two species of Maui green-
swords are all in the genus Argyroxiphium. They only grow in localized distributions
above 4,900 ft (1,500 m) on the islands of Maui and Hawaii. Two silverswords
grow in the alpine scrub of Haleakala, Mauna Kea, and Mauna Loa, and like
189
desert plants, they are adapted to intense sun and dry conditions. The third is eke
(A. caliginis), a tiny silversword only 6 in (15 cm) in diameter with a 1.5 ft (0.5 m)
flower stalk, which occupies bogs at the transition between the forest and alpine
zones in western Maui. The two Maui greenswords have few silver hairs on their
green leaves and grow in misty sites. Two greenswords on Kauai belong to a differ-
ent genus (Wilkesia), and are known as iliau in Hawaiian. They are endemic to that
island and do not grow in an alpine environment.
Leaves of silverswords are long and narrow, 616 in (1540 cm long) and 0.20.6
in (0.51.5 cm) wide. The genus has evolved a growthform similar to the giant
rosettes in tropical Africa and in the tropical Andes, a rosette that protects the interior
growing point and from which a tall flower stalk emerges. Water and nutrients stored
in the leaves are used to produce the tall inflorescence. The plant grows as a ground-
hugging rosette or with a very short stem. Two subspecies vary in the shape of the in-
florescence and geographic location: one is found in Haleakala crater, the other on
Mauna Kea. Both occupy dry alpine cinder habitats between 7,000 ft (2,150 m) and
12,300 ft (3,750 m) on the volcanoes. A third species, Mauna Loa silversword, is lim-
ited to that volcano and its locale is protected in Hawaii Volcanoes National Park. A
short, shrubby tarweed found on exposed windy slopes hybridizes freely with silver-
swords in nature. More than 30 identified crosses indicate their recent evolution.
A silversword flowers only once in its lifetime. It takes 1550 years for the plant
to mature from seed to flowering stage. It remains a compact rosette up to 2 ft
(0.6 m) in diameter until it sends up a flower stalk. Flowering occurs mid-June to
November, and after the seed ripens three months later, the plant dies. The inflores-
cence, which carries 50600 pink, white, or red daisy-like flowers, grows up to 10 ft
(3 m tall) and 30 in (75 cm) wide. Each plant produces thousands of seeds. Because
they require cross-pollination, all plants in an area generally flower at the same time.
Sticky hairs on its leaves, stems, and central stalk trap crawling insects and prevent
them from accessing the pollen, allowing flying pollinators such the native yellow-
faced bee, to carry pollen to other plants. Occasionally small rosettes will branch out
from the main body, a benefit because only the rosette with the flower stalk will die.
Early in the twentieth century, silversword populations were highly endangered
because of vandalism and browsing by goats and cattle. Root systems can be easily
crushed in loose cinders. After vandalism was curtailed and cattle were eliminated
from Haleakala National Park in the 1930s and goats in the 1980s, the population
came back from a low of 2,0004,000 plants to more than 60,000 now.
Further Readings
Coe, Malcolm James. 1967. The Ecology of the Alpine Zone of Mount Kenya. The Hague:
Dr. W. Junk Publishers.
Line, Les. December 2003January 2004. Farewell to Flamingos? National Wildlife Maga-
zine. http://www.nwf.org/nationalwildlife/article.cfm?issueID65&articleID875.
190
Palomar College. n.d. The Silversword Alliance. http://waynesword.palomar.edu/
ww0903b.htm.
Public Broadcasting Service. n.d. Salt Flat Living. Andes: The Dragons Back. http://
www.pbs.org/wnet/nature/andes/saltflat.html.
Wheeler, Jane C. n.d. Evolution and Present Situation of the South American Camelidae.
http://www.conopa.org/camelidos/historia.php.
191
Appendix
Biota of Tropical Alpine Tundra Biome
South American Alpine Tundra
Some Characteristic Plants of the Andean Paramo
Trees and treeline
Polylepis (in the Rose family) Polylepis sericea
Chachacomo Escalonia spp.
Shrubs
Chuquiragua or Flower of the Andes Chuquiragua spp.
Jata or Candlebush Locicaria ferruginea
In the Mahogany family Schmardaea microphylla
Quinine or Cedro colorado Chichona officinallis
St. Johns wort Hypericum laricifolium
Graminoids
Reedgrass Calamagrostis spp.
Fescue grass Festuca spp.
Dwarf bamboo Chusquea spp.
Brome grass Bromus catharticus
Needlegrass Stipa spp.
Forbs
Giant rosettes
Large espeletia Espeletia timotensis
Small espeletia Espeletia moritziana
Midsize espeletia Espeletia spicata
Prostrate stem espeletia Espeletia semiglobulata
Short espeletia Espeletia schultzii
Wooly espeletia Espeletia hartwegiana
192
Puya Puya clava-herculis
Puya Puya raimondii
Puya Puya hamata
Cushions
Azorella (in the Aralia family) Azorella crenata
Azorella (in the Aralia family) Azorella julianii
Sandwort Arenaria spp.
Ragwort Senecio spp.
Leafy
Iceland purslane Koeniga islandica
Geranium Geranium spp.
Gentian Gentiana spp.
Paintbrush Castilleja spp.
Buttercup Ranunculus spp.
Cryptogams
Sphagnum moss Sphagnum
Some Characteristic Animals of the Andean Paramo
Herbivores
Mountain tapir Tapirus pinchaque
Little red brocket deer Mazama rufina
Northern pudu Pudu mephistophiles
Rice rat Microryzomys minutus
Paramo rabbit Sylvilagus brasiliensis.
White-tailed deer Odocoileus virginianus
Carnivores
Thomas small-eared shrew Cryptotis thomasi
White-eared opposum Didelphis albiventris.
Long-tailed weasel Mustela frenata
Paramo wildcat Felis tigrina
Puma Puma concolor
Birds
Andean Condor Vultur gryphus
Paramo Pipit Anthus bogotensis
Streak-backed Canastero Asthenes wyatti
Merida Wren Cistothorus meridae.
Sedge Wren Cistothorus platensis
Black-chested Buzzard Eagle Geranoactus melanoleucus
White-rumped Hawk Buteo leucorrhous
193
Some Characteristic Plants of the Andean Puna
Trees and treeline
Butterfly bush or Colle Buddleja coriaceae
Polylepis or Quenoa (in the Rose family) Polylepis tomentella
Polylepis or Quenoa (in the Rose family) Polylepis tarapacana
Shrubs
In the Pea family Adesmia spp
Broom rape Baccharis incarum
Broom rape or Chijua Baccharis boliviensis
Tolillar Fabiana densa
Atacama saltbush Atriplex atacamensis
Graminoids
Sedge Oreobolus spp.
Needlegrass Stipa jehu
Peruvian feathergrass or Ichu Stipa ichu
Fescue grass Festuca spp.
Reedgrass Calamagrostis spp.
Mountain bamboo Chusquea spp.
Pampas grass Cortaderia spp.
Saltgrass Distichlis humilis
Rush Juncus spp.
Bulrush Scirpus spp.
Cushion rush Distichia muscoides
Cushion rush Oxychloe andina
Cacti
Old man of the Andes Oreocereus spp.
San Pedro cactus Trichocereus pachanoi
Forbs
Giant rosettes
Puya Puya raimondii
Puya Puya hamata
Cushions
Azorella (in the Aralia family) Azorella yarita
Azorella (in the Aralia family) Azorella compacta
Cushion plantain Plantago rigida
Leafy
Gosmore Hypochoeris spp.
In the Rose family Lachemilla spp.
In the Sunflower family Culcitium spp.
194
Pickleweed Salicornia pulvinata
Seepweed Suaeda foliosa
Some Characteristic Animals of the Andean Puna
Herbivores
Vicuna Vicugna vicugna
Guanaco Lama guanicoe
Llama Lama glama
Alpaca Lama pacos
Chinchilla Chinchilla brevicaudata
b
Golden vizcacha rat Pipanococtomys aureus
a
Carnivores
Puma Felis concolor
Andean fox Pseudalopex culpaeus
Andean Mountain cat Oreailurus jacobita
b
Andean hairy armadillo Caetophractus nationi
Birds
Ash-breasted Tit-tyrant Anairetes alpinus
a
Royal Cinclodes Cinclodes aricomae
a
Berlepschs Canastero Asthenes berlepschi
a
Line-fronted Canastero Asthenes urubambensis
a
Olivaceaous Thornbill Chalcostigma olivaceum
a
Scribble-tailed Canastero Asthenes maculicauda
a
Gray-bellied Flower Piercer Diglosa carbonaria
a
Darwins Rhea or Suri Pterocnemia pennata
Puna Tinamou Tinamotis pentlandii
James Flamingo Phoenicopterus jamesi
b
Andean Flamingo Phoenicopterus andinus
b
Chilean Flamingo Phoenicopterus chilensis
Notes:
a
Endemic;
b
Rare and endangered.
African Alpine Environments
Some Characteristic Plants of Equatorial East Africa
Treeline
Bamboo Arundinaria alpina
Protea Protea kilimandscharica
Tree heather Erica arborea
Philippia heath Philippia excelsa
Philippia heath Philippia keniensis
(Continued )
195
Low or subshrubs
Strawflower or Everlasting Helichrysum stublmannii
Kilimanjaro strawflower Helichrysum kilimanjari
Johnstons ladys mantle Alchemilla johnstonii
Ladys mantle Alchemilla argyrophylla
Browns strawflower or
Everlasting
Helichrysum brownei
Graminoids
Carex sedge Carex monostachya
Fescue grass Festuca pilgeri
Bentgrass Agrostis trachyphylla
Bluestem grass Andropogon spp.
Hairgrass Koeleria spp.
Pentaschistis grass Pentaschistis minor
Forbs
Giant rosettes
Giant groundsel Dendrosenecio keniodendron
Giant groundsel Dendrosenecio johnstonii
Giant groundsel Dendrosenecio brassiciformis
Giant groundsel Dendrosenecio keniensis
Giant lobelia Lobelia telekii
Giant lobelia Lobelia keniensis
Giant senecio or Giant
groundsel
Senecio brassica
Giant senecio or Giant
groundsel
Senecio battescombie
Leafy
Ladys mantle Alchemilla cyclophylla
Sedum (succulent) Sedum ruwenzoriense
Senecio Senecio purtschelleri
Senecio Senecio keniophytum
Cryptogams
Usnea (Fruticose lichen) Usnea
Some Characteristic Animals of Equatorial East Africa
Herbivores
Rock hyrax Procavia johnstoni
Groove-toothed rat Otomys orestes
Common duiker Sylvicapra grimmia altivallis
Harsh-furred mouse Laphuromys a. aquilas
Giant mole rat Tachyryctes rex
196
Carnivores
Leopard Felis pardus
Shrew Crocidura alex alpina
Wild dog Lycaon pictus lupinus
Birds
Mackinders Owl Bubo capensis
Augur Buzzard Buteo rufofuscus augur
Verreaux Eagle Aquila verreauxii
Lammergeier or Bearded Vulture Gyptaetus barbatus
Scarlet-tufted Malachite Sunbird Nectarinia johnstoni johnstoni
Hill Chat Pinarochroa sordida ernesti
Streaky Seed Eater Serinus striolatus stridatus
Reptiles
Alpine meadow lizard Algyroides alleni
Some Characteristic Plants of the Ethiopian Plateau
Treeline
Tree heather Erica arborea
Heath Erica trimera
Giant St. Johns wort Hypericum revolutum
Philippia heath Philippia spp.
Low shrubs
Ladys mantle Alchemilla haumannii
Strawflower Helichrysum citrispinum
Graminoids
Carex sedge Carex monostachya
Junegrass Koeleria spp.
Hairgrass Aira spp.
Fescue grass Festuca macrophylla
Fescue grass Festuca abyssinica
Forbs
Giant rosettes
Giant lobelia Lobelia rynchopetalum
Leafy
African rose Rosa abyssinica
Yellow primrose Primula verticillata
Cryptogams
Usnea (Fruticose Lichen) Usnea spp.
197
Some Characteristic Animals of the Ethiopian Plateau
Herbivores
Walia ibex Capra ibex walie
Mountain nyali Tragelaphus buxtoni
Klipspringer Oreotragus oreotragus
Giant mole rat Tachyoryctes macrocephalus
a
Grass rat Lophuromys melanonyx
Starks hare Lepus starcki
Giant climbing mouse Megadendromus nikolausi
a
Narrow-headed rat Stenocephalemys albocaudata
a
Black-clawed brush-furred rat Lophuromys melanoyx
a
Rock hyrax Procavia capensis
Carnivores
Ethiopian wolf or Simen fox Canis simensis
a
Gelada baboon Theropithecus gelada
Birds
Widgeon Anas penelope
Shoveler Anas clypeata
Ruff Philomachus pugnax
Greenshanks Tringa nebularia
b
Chough Pyrrhocorax pyrrhocorax
b
Ruddy Shelduck Tadorna ferruginea
b
Notes:
a
Endemic;
b
Migratory breeder.
Oceania Alpine Tundra
Some Characteristic Plants of Hawaii Alpine
Treeline
Ohia (in the Myrtle family) Metrosideros polymorpha
Low or subshrubs
Pukeawe (in the Heath family) Styphelia tameiameiae
Hawaiian blueberry Vaccinium reticulatum
Mamame (in the Pea family) Sophora chrysophylla
Pilo (in the Madder family) Coprosma montana
Cranesbill Geranium cuneatum
Tarweed or Naenae (in the Sunflower family) Dubautia menziesii.
Graminoids
Hairgrass Deschampsia nubigena
Panicum grass Panicum spp.
Bentgrass Agrostis sandwicensis
198
Forbs
Giant rosettes
Haleakala silversword Argyroxiphium sandwicense ssp.
macrocephalum
Mauna Kea silversword Argyroxiphium sandwicense ssp.
sandwicense
Mauna Loa silversword Argyroxiphium kauense
Maui greensword Argyroxiphium kai
Maui greensword Argyroxiphium virescens
Eke Argyroxiphium caliginis
Leafy
Gosmore or Hairy cats ear Hypochoeris radicata
a
Wooly mullein Verbascum thapsus
a
Bracken fern Pteridium aquilinum
Fern Asplenium spp.
Note:
a
Introduced.
Some Characteristic Animals of Hawaii Alpine
Herbivores
Pig Sus scrofa
a
Polynesian rat Rattus exulans
a
Carnivores
Dog Canis familiaris
a
Birds
Jungle Fowl Gallus gallus
a
Insects
Haleakala flightless moth Thryocopa apatela
Mauna Kea wekiu bug Nysius wekiuicola
Mauna Loa wekiu bug Nysius aa
Note:
a
Introduced.
199
Glossary
Ablation. Means by which ice is lost from a glacier, including melting, calving, and
sublimation.
Aeolian. The zone on mountains above the limit of plant life, where wind-blown debris
and insects may accumulate.
Albedo. Reflectivity of a surface, refers to how much solar radiation is reflected.
Alpine. The zone on mountains above the region of trees.
Alpine Glacier. Ice that moves downslope in mountain valleys. Also called valley gla-
ciers because the ice is confined to the valley and does not cover the entire
landscape.
Annual. A plant that completes its life cycle in one year or one growing season.
Antarctic. Strictly, the latitudes between the Antarctic Circle at 66
1
=
2
S and the South
Pole. Can refer to the general region near or south of the Antarctic Circle.
Arctic. Strictly, the latitudes between the Arctic Circle at 66
1
=
2
N and the North Pole.
Can refer to the general region near or north of the Arctic Circle.
Arete. A narrow, sharp-edged ridge between two valley glaciers or glaciated valleys.
Aspect. Direction toward which a slope faces. Also known as exposure.
Behavioral. Activities or behavior of animals.
Biome. A large region with similar vegetation, animal life, and environmental
conditions.
Biota. The combined flora and fauna, including all the plants and animals.
Boulder Clay. Unsorted jumble of rocks and smaller particles deposited by a glacier.
Bryophyte. Group of plants that includes mosses.
Bulbil. Small plant-like shoot produced on a flower stalk.
Calving. Large pieces of ice breaking off a glacier.
201
CAM (Crassulacean Acid Metabolism). A form of photosynthesis used by plants to
conserve water.
Canopy. The uppermost layer of foliage in vegetation.
Cenozoic. A recent geologic time period, roughly 65 million years ago to the present.
Chamaephytes (Raunkiaer). Plants that hold their regenerating buds just above soil
level.
Circumpolar. Distribution that encircles the North Pole.
Climate. Typical weather (especially temperature and precipitation) patterns during a
normal year that are experienced over decades or centuries. Weather refers to the
conditions of the atmosphere at any given moment.
Cogener. Plants in the same genus.
Community. The plants and animals assembled in a given area. Sometimes refers to
only a subset of these organisms, such as the plant community or the bird
community.
Conifer. A cone-bearing plant such as pine and spruce.
Continental Glacier. Ice that covers a large part of a continent, such as Greenland.
Also called ice sheet.
Continentality. The effect a large land mass has on seasonal temperature variations.
Continental areas are warmer in summer and colder in winter. Compare with
Maritime.
Cover. The proportion of a surface on which vegetation occurs, usually measured as a
percent.
Cretaceous. A geological time period from roughly 145 million years ago to 65 million
years ago.
Crustose (lichen). Crustlike. Compare with foliose and fruticose.
Cryptogam. Plant group that includes algae, fungi, mosses, lichens, and ferns, which
lack flowers. Most are nonvascular plants and have no differentiated tissue to
transport water or nutrients.
Cushion Plant. A low-growing, multistemmed plant that grows as a dense mound.
Cyanobacteria. Also known as blue-green algae. Found in soil and water and are able
to fix nitrogen and photosynthesize.
Cyclonic Storm. Type of weather that results when cold air comes in contact with
warm air in the mid-latitudes.
Deciduous. Refers to plants, usually trees and shrubs, that drop their leaves during non-
growing seasons.
Diurnal. Change from day to night, such as temperature.
Dwarf Shrub. Small shrub with branches less than 12 in (30 cm) high.
Ecotone. A zone between two biomes or plant communities where type of climate and
plants changes gradually.
Ecotype. A plant or animal that has not quite evolved to the species level but that has
variations according to environment.
Endemic. Originating in and restricted to a particular geographic area.
Energy Balance. Comparison of the amount of energy received from the sun to the
amount lost from Earth. Can be positive or negative. Also called radiation balance.
Entisol. Recently developed soils that have no differentiated horizons or layers.
Epiderm. Outer layer of leaf cells.
202
Glossary
Equable. Refers to climate. Little variation between winter and summer temperatures.
Ericaceous. Refers to a member of the Ericaceae, a plant family consisting of heathers,
bilberries, Labrador tea, rhododendrons, and so on.
Evapotranspiration. The combined processes of adding water vapor to the atmosphere
through evaporation from the soil and bodies of water and from the passage of
water out of plants through the stomata in their leaves.
Evergreen. Refers to plants, usually trees or shrubs, which maintain leaves all year.
Leaves may be replaced throughout the year or in a flush during a single season,
but the plant is never without live foliage.
Fauna. All the animal species in a given area.
Flora. All the plant species in a given area.
Floristic Province. A region with related plants that evolved in that area.
F ohn. A warm downslope wind.
Foliose (lichen). Leaflike in appearance. Compare with crustose and fruticose.
Forb. A broad-leaved, green-stemmed, nonwoody plant. One type of herb.
Fruticose (lichen). Shrubby, upright lichens. Compare with crustose and foliose.
Genus (plural genera). A taxonomic unit composed of one or more closely related
species.
Giant Rosette. Rosettes in tropical alpine environments that grow to large size.
Gigantism. Trait of some species to grow larger than normal.
Glacial Drift. Any rock and sediment carried and deposited by a glacier. See moraine,
outwash, and boulder clay.
Gleization. The process of soil formation in bogs where undecayed plant material
accumulates under conditions of poor drainage and cold temperatures. Forms a
peaty surface underlain by clay.
Graminoid. Refers to narrow-leaved grass-like plants, including grasses, sedges, and
rushes.
Growing Season. The length of time plants are able to grow, generally between the last
frost of winter and first frost of fall, but may be related to seasonality of precipitation.
Growthform. The appearance or morphology of a plant that is adapted to particular
environmental conditions. Examples include tree, shrub, and forb.
Habitat. The place where a species lives and the local environmental conditions of that
place.
Heath. Small-leaved shrubs such as heathers and bilberries, which are members of the
Ericaceae. Leaves are often drought adapted.
Hemicryptophytes (Raunkiaer). Plants that hold regenerating buds at the surface of
the ground.
Herb. A nonwoody or soft, green-stemmed plant that dies down each year. May be an-
nual or perennial. Broad-leaved herbs are called forbs. Grasses and sedges are
called graminoids.
Herbaceous. Nonwoody plant that dies back every year.
Hibernation. When an animals body temperature is reduced to that of the environ-
ment, resulting in a decrease in metabolism and need for energy. Used by animals
to avoid a cold season.
High Arctic. Refers to arctic latitudes with high numbers, usually with a more severe
climate than Low Arctic.
Glossary
203
Histosol. Soils that are predominantly organic, usually formed by the gleization proc-
ess and common in bogs.
Humus. Partially decayed plant and animal matter that occurs as a dark brown
organic substance in soils. Important for conserving water and some plant
nutrients.
Inceptisol. In humid climates, developing soil that has at least one distinctive horizon.
Indicator Plant. A plant typical of a biome that can be used to delimit the biomes
extent in the absence of climate data.
Inflorescence. Flower stalk.
Infrared radiation. Energy that comes from the Earth. Also called longwave radiation
or terrestrial radiation.
Introduced (species). A species transported accidentally or deliberately by humans
beyond its natural distribution area. Also called alien or non-native.
Inversion. Air is warmer with increasing elevation, instead of the normal lapse rate of
cooler temperatures.
Koeppen. A climate classification system.
Krummholz. Stunted trees at treeline, deformed by harsh climatic conditions.
Lapse Rate. Decrease in temperature as elevation increases. Averages 3.5 F per
1,000 ft (6.5 C per 1,000 m), but the rate is variable.
Latitude. Distance north or south of the equator, measured in degrees. The equator is
0 latitude. Low latitudes lie between 0 and 30 north and south; mid-latitudes lie
between 30 and 60, and high latitudes between 60 and 90.
Lichen. A form of life composed of a fungus and an alga joined in a symbiotic relation-
ship and classified as a single organism.
Lifeform (Raunkiaers). A category of plant life based on morphology and the position
of the renewal bud.
Low Arctic. Refers to arctic latitudes with lower numbers, usually with a less severe
climate that High Arctic.
Maritime. The effect large bodies of water have on moderating seasonal temperature
variations. Maritime climates do not have extremes of temperature. Compare with
continentality.
Mat Plant. A low-growing, multistemmed plant that roots from stems that creep along
the ground surface. Forms a dense cover.
Mesophyll. Interior leaf cells.
Microclimate. A small area with climatic conditions that are different from the general
climate of the area.
Microhabitat. A tiny nook or habitat with specific environmental conditions that differ
from that of the larger habitat in which it occurs.
Moraine (glacial). Unsorted accumulation of rocks and sediment deposited by a gla-
cier, usually as hills. A type of till.
Morphology. Form and structure, size and shape of an organism. General appearance
of an organism.
Needle ice. Ice that freezes and expands in soil, disrupting the surface.
Nival. Zone of permanent snow and ice on mountains. Snowline marks its lower limit.
Nivation Hollow. Depression in the ground surface where snow accumulates and the
soil is moist.
204
Glossary
Nunatak. A mountain peak that extends or extended above glacial ice, providing a
place for plant or animal life during glaciation. May be refugia.
Outwash (glacial). Sediment that is sorted by size as it is deposited by water melting
from a glacier.
Paludification. The process by which acidic, waterlogged conditions are produced and
bogs are expanded because of the growth of mosses.
Parent Material. Sediment or rock from which soil is developed. Contributes to the
mineral component of the soil.
Perennial. A plant that lives for several years and undergoes active growth each year.
Periglacial. Refers to cold areas subject to intense frost action and solifluction, often
adjacent to glaciers.
Permafrost. A condition of permanently frozen ground.
Phanerophyte (Raunkiaer). A plant that holds its regenerating buds high, such as
trees.
Photoperiod. The number of hours of daylight. Depending on latitude, it may change
with the seasons.
Photosynthesis. The process by which green plants convert oxygen and carbon dioxide
in the presence of sunlight to sugars. Energy in the form of visible light is trans-
formed into chemical energy that can be used by living organisms.
Physiology. The metabolic functions and processes of organisms.
Pleistocene. Geologic time period when glaciers covered much of North America and
Eurasia, fromapproximately 1.6 millionto 10,000 years ago. Alsocalled the ice age.
Polar Desert. The coldest and driest part of the Arctic or Antarctic tundra. Polar semi-
desert is not quite as dry or as cold.
Polyploidy. Refers to more than the normal two sets of chromosomes.
Refugia. Areas that were ice free during the Pleistocene. Plants and animals could
maintain populations in refugia.
Regenerating bud(s). Point(s) where new growth will form in the growing season.
Rhizome. A horizontal root structure that lies just below the surface.
Rosette. A growthform characterized by a basal whorl of leaves around a central stem
or renewal bud. Can be flat to the ground or taller.
Sclerophyllous. Referring to leaves that are thick or waxy to prevent water loss.
Scree. Unstable slopes of small-size rocks, subject to downslope movement.
Scrub. A vegetation type characterized by sparse, small shrubs or stunted trees.
Seasonality. Difference in temperature or precipitation between winter and summer.
Sexual Reproduction. The formation of new individuals by the fusion of gametes (ova)
and pollen in plants; egg and sperm in animals.
Soil. The uppermost layer on land. Composed of a mixture of mineral and organic
materials in which plants grow.
Soil Horizon. A layer within the soil that is fairly distinct in terms of its chemistry, tex-
ture, and color.
Solar Radiation. Energy that comes from the sun, also called shortwave radiation or
insolation.
Solifluction. Process by which water-logged soil slowly flows downslope.
Species. A group of sexually reproducing individuals that can produce viable offspring.
The fundamental unit of classification in taxonomy.
Glossary
205
Spodosol. Soil that usually forms beneath forest vegetation. Has distinctive horizons,
especially a light-colored sandy surface with a darker, clay layer beneath.
Stomata. The leaf pores through which plants exchange gases with the atmosphere.
Subalpine. The zone slightly lower in elevation than alpine tundra, but with some char-
acteristics of the alpine environment.
Subarctic. The latitude just south of the Arctic Circle, but with some characteristics of
the Arctic. Subantarctic refers to just north of the Antarctic Circle in the Southern
Hemisphere.
Sublimation. When ice or snow does not melt but skips the liquid state and goes
directly into the gas form in the air.
Substrate. Surface material in which plants grow, including rock, soil, or sediments.
Succulent. A growthform that has specialized tissue in the stem, leaves, or an under-
ground organ for the storage of water.
Sun Angle. How high the sun is in the sky during the day and a major factor in temper-
ature. A higher sun angle imparts more warmth; a lower sun angle, less.
Supercool. Ability of a plant or animal to reduce its body temperature below freezing
with no harm to the tissues.
Talus. Unstable steep slopes of large rocks, subject to downslope movement.
Taproot. A root that extends deep into the ground that provides access to water.
Tarn. Alpine lake occupying a cirque in an alpine glaciated environment.
Taxonomy. The science of describing, classifying, and naming organisms.
Temperate. Refers generally to the temperature patterns of the mid-latitudes, the tem-
perate zone, where summers are warm to hot and winters are mild to cool. Not too
cold or too hot.
Tertiary. A recent geologic time period, from approximately 1.6 million to 65 million
years ago.
Thallus. The body of a nonvascular plant that is not differentiated into stem, root, or
leaf.
Tolerance Limits. Refers to the extremes of environmental factors, such as cold,
heat, drought, and snow depth, beyond which an individual species cannot
survive.
Torpor. Reduction in an animals body temperature for a short period of time, allowing
the animal to avoid an unfavorable environmental period. The animal becomes
torpid. It is not true hibernation because the body temperature is not reduced to
that of the environment.
Treeline. The transition from forest to nonforested zones.
Tropical Alpine. Treeless zones in high mountains near the Equator or in the
Tropics.
Tropics. The parts of Earth that lie between 23
1
=
2
N and 23
1
=
2
S or between the Tropic
of Cancer and the Tropic of Capricorn.
Tundra. Refers to treeless zones in the Arctic, Antarctic, or alpine.
Tussock. A growthform of grasses and sedges in which individuals grow as tufts or
clumps and form conspicuous hummocks on the ground.
Ungulate. Any hoofed mammal, such as a cow, elk, or ibex.
Vascular plant. Any plant with conducting vessels that move nutrients and water
between roots and leaves. Includes flowering plants and ferns.
206
Glossary
Vegetation. The general plant cover of an area, defined according to the appearance of
the plants (grass, forest, and shrubs) rather than the actual species present. Com-
pare with flora.
Vegetative Reproduction. The formation of new plants from pieces of the parent plant
such as fragments of leaves, roots, stems, or rhizomes. Also called asexual repro-
duction. Includes cloning.
Viviparous. Refers to plants that produce plantlets or seed-like shoots instead of seeds.
Zonation. The occurrence of particular forms of life in distinct belts. These may be
determined by latitude or elevation.
Glossary
207
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211
Index
The letter f following a page number denotes a figure. The letter t following a page number
denotes a table.
Active layer, 34
Adaptations. See Animal adaptations; Plant
adaptations
Aeolian zone, 6, 30, 189. See also
Nival zone
Afroalpine, 15758, 17687
Air: density, 84, 165; pressure, 84
Alaska, 35, 38, 40, 5051, 117
Alaska pipeline, 35
Albedo, 37
Alpine parrot, 138, 139f
Alpine tundra: boundaries, 56; differences
from arctic, 2t, 45, 83; similarities with
arctic, 1, 2t, 4, 83
Alpine tundra, mid-latitude. See Mid-
latitude alpine
Alpine tundra, tropical. See Tropical
alpine
Alps, European, 15, 101f, 103, 12125
Alps, Southern, 13539
Altiplano, 17276, 173f. See also Puna
Amphibians, 31, 63, 118
Andes Mountains, 102, 133, 16676
Animal adaptations: arctic, 37; behavioral,
28; mid-latitude alpine, 9394;
morphological, 26; physiological, 2627,
27f ; tropical alpine, 16566
Animal life, 2526
Animal life, antarctic, 7071
Animal life, arctic, 4246, 44f, 5455, 58,
62, 65, 67
Animal life, mid-latitude alpine: Eurasia,
124, 128, 13132; North America,
1068, 108f, 110, 111, 113, 114,
11718, 120; Southern Hemisphere,
133, 135, 138
Animal life, tropical alpine: afroalpine,
18384, 187; South America, 17071,
17475, 174f, 175f
Animals: and snow, 2829; effects on
vegetation, 4748
Annuals, 23, 90, 106
Antarctic tundra, 33, 6771, 68f ; animals,
7071; climate, 3638, 6870;
polar comparisons, 3334; vegetation,
6970
Appalachian Mountains, 103, 11820,
121
Arctic fox, 48, 5455
Arctic ground squirrel, 28
213
Arctic tundra, 3367; animals, 4248, 44f,
5455; boundaries, 56; climate, 3640,
39f, 4951, 5556, 59, 6364; differences
from alpine, 2t, 45; energy budget, 37;
polar comparisons, 3334; regional
expressions, 4971; similarities with
alpine, 1, 2t, 4; soils, 4041; vegetation,
4142, 41f, 5154, 51f, 5859, 6062,
61f, 6467, 66f
Arctic tundra, Eurasia, 61f ; Fennoscandia,
5963; Russia, 6367
Arctic tundra, North America, 4959, 49f,
50t; Greenland, 5559
Azorella cushions, 16970
Bering Strait, 26, 56, 103
Bird life, antarctic, 7071
Bird life, arctic, 4647; Eurasia, 62, 65, 66;
North America, 55f, 5657, 58
Bird life, mid-latitude alpine: Eurasia, 125,
128, 132; North America, 1089, 110,
111, 113, 11415, 118, 120; Southern
Hemisphere, 13839, 139f
Bird life, tropical alpine, 165, 171, 17576,
18485, 187
Birds: raptors, 62, 109f ; sea, 70, 189; water,
29, 46, 55f, 60, 62, 65, 138, 176, 187
Bharal, 131
Bogs, 41, 65, 8990, 128, 134, 137; cushion,
172
Boulder, Colorado, 115
Boulder field, 98f
Bristlecone pine, 114
Brown bear, 43, 54
Bulbils, 25, 57
Cactus, 89, 174
Cairngorm Mountains, 125
Camouflage, 26
Canada, 38, 39f, 40, 49, 50, 5154. See also
Churchill, Manitoba
Caribou, 28, 42, 43, 52, 54, 57, 58. See also
Reindeer
Carpathian Mountains, 126
Cascade Range, 84, 104, 105, 11011
Chinchilla, 176
Churchill, Manitoba, 41f, 45
Climate, 14
Climate, antarctic, 6870
Climate, arctic, 3640, 39f, 5556; Eurasia,
59, 6364, 63t; North America, 4951,
50t, 56
Climate, mid-latitude alpine, 8488;
Eurasia, 12627, 12930; North America,
8488; Southern Hemisphere, 133,
134, 136
Climate, tropical alpine, 15861, 159f,
16668, 172, 17779, 188
Cloning, 2425, 24f, 93
Coal mining, 60
Coastal Ranges, 104, 10910
Cold air drainage, 38, 87
Continentality, 3738, 63
Cottongrass, 52
Cryptoendolithic lifeforms, 70
Cryptogams, 16, 54
Cushion plants, 1618, 17f, 18f ; effect on
microclimate, 12; tropical alpine,
16970
Cyanobacteria, 47, 54, 70
Drakensberg Mountains, 134
Drought stress, 15, 22, 89
Ecotype, 14, 15, 23, 24
Edelweiss, 14, 123
Energy budget, 4; Arctic and Antarctic, 37;
mid-latitude alpine, 86; sun versus shade,
4; tropical, 158
Entisol, 12, 89
Espeletia, 16364, 163f, 165, 168, 170,
171
Ethiopian Plateau, 158, 176, 18587
Evergreen, 16, 22, 94, 97
Fellfield, 98f, 99
Felsenmeer, 98
Fennoscandia, 39f, 46, 5963,
61f, 101
Fire, 134, 136, 179, 182
Flag trees, 97, 119, 130
Flamingos, 176
214
Index
Frailejones. See Espeletia
Frost hardening, 20
Germination, 2324, 9293
Giant groundsel, 16364, 163f, 180,
18283
Giant rosettes, 16364, 163f. See also
Espeletia; Giant groundsel; Giant
senecio; Lobelias; Puya; Silversword
Giant senecio, 16364, 163f, 180, 18283
Gigantism, 186
Glacial deposits, 10
Glacial movement, 9
Glaciation, alpine, 1011
Glaciation, continental, 910
Global warming, 45, 49
Graminoid plants, 1618, 17f, 18f
Great Basin, 104, 105, 11315
Greenland, 41, 53, 5559
Grizzly bear, 43
Groundhog, 107, 120. See also Marmot
Growing season, 3738, 51, 158
Growthform, 1520, 17f, 18f ; cryptogams
and phanerogams, 16; effect on
microclimate, 1112, 161; tropical alpine,
16164, 162f
Haastii, 137
Habitats, mid-latitude alpine, 98100
Hamster, 128
Hawaii, 18790
Heath: arctic, 52, 53, 88; mid-latitude
alpine, 100, 117, 11920, 120f, 123, 134;
tropical alpine, 165, 179, 182, 186
Heat tolerance, plant, 21
Hibernation, 28
High Arctic: Greenland, 5859; North
America, 42, 5051, 5354; plant
communities, 5354
Himalaya Mountains, 15, 30, 101, 12832,
131f
Histosol, 13, 41, 89
Hot springs, 57, 58
Huancabamba Depression, 167, 172
Human impacts, 5, 31, 125; antarctic, 71;
arctic, 13, 35, 40, 4849; agriculture and
grazing, 59, 108, 123, 125, 132, 134, 135,
171, 172, 187; hunting, 125, 135; species
introduction, 138, 188, 189, 190
Hyrax, 18384, 184f
Ibex, 124f, 128, 131
Ice fog, 40, 50
Iceland, 38, 40, 59
Inceptisol, 12, 40, 89
Insects. See Invertebrates
Institute of Arctic and Alpine Research
(INSTAAR), 115
Invertebrates, 2930, 47, 70; mid-latitude
alpine, 94, 109, 139; tropical alpine,
16566, 185, 189
Island biogeography, 158, 18788
Junipers, 130, 132
Kiang, 132
Kodiak bear, 43
Krummholz, 12, 60, 9697, 97f, 112, 137.
See also Flag trees
Lemmings, 28, 29, 54; cycles, 46, 48, 62, 65
Lesotho, 13235
Lichens, 16, 2223, 25; antarctic, 6970;
arctic, 4748, 51, 6566, 66f ; mid-latitude
alpine, 91, 98, 133
Light regime, 4, 15, 36, 83, 158
Little Ice Age, 54
Llamas, 93
Lobelias, 16364, 180, 18283, 187
Low Arctic: Greenland, 58; North America,
4142, 41f, 5053; plant communities,
5153
Maps: antarctic, 68f ; arctic, 49f, 61f ;
mid-latitude alpine, 104f, 122f, 133f, 135f ;
tropical alpine, 166f, 177f
Maritime, 38, 136. See also North Atlantic
Drift
Marmot, 28, 1078, 108f. See also
Groundhog
Marsh, 100
Mat plants, 1718, 17f, 18f
Index
215
Meadow, 98f, 99, 100, 127f
Metabolic heat, 27
Microclimate, 8486; plant effects on,
1112, 161
Mid-latitude alpine, 83140; adaptations of
lowland species, 103; animal adaptations,
9394; birds, 1089; climate, 8488, 105;
energy budget, 86; habitats and com-
munities, 98100, 98f, 99f ; plant
adaptations, 9093; plant communities,
98100, 106; plant reproduction, 9193;
Pleistocene migrations, 100103; regional
expressions, 10390; similarity of floras,
100103; soils, 8890; treeline, 9498;
wind, 87
Mid-latitude alpine, Central Asia, 122f, 125
132; Himalaya Mountains, 12832, 131f ;
Tien Shan Mountains, 12628, 127f
Mid-latitude alpine, Europe, 12025, 122f ;
Alps, 101f, 12125; Cairngorm
Mountains, 125; Carpathian Mountains,
126; Pyrenees, 125
Mid-latitude alpine, North America,
10321, 104f ; animals, 1069; Cascade
Range, 11011; Coastal Ranges, 10910;
Great Basin, 11315; Mount Washington,
11820; Rocky Mountains, 11518;
Sierra Nevada, 11113
Mid-latitude alpine, Southern Hemisphere,
13239; New Zealand, 13539;
southern Africa, 13235; southern
Andes, 133
Migrations: bird, 47, 5657; mammal, 26,
5657; plant, 100103. See also Origins,
flora and fauna
Mires, 52, 53, 60, 64
Mole rat, 135; giant, 187
Monsoons, 129, 130
Mountain goat, 2829, 110, 11718, 118f
Mountain sheep, 29; bighorn, 113, 114, 117;
Dalls, 110, 117; Kyzylkum, 128
Mount Everest, 15, 129
Mount Kenya, 159f, 164, 176, 178,
18185, 181f, 182f
Mount Kilimanjaro, 15, 30, 176
Mount Rainier, 8485, 85f, 110, 111
Mount Washington, New Hampshire,
100101, 11820, 120f, 121
Muskoxen, 40, 4243, 53, 54, 58
Needle ice, 1213, 3435, 89, 161
New Zealand, 97, 13539
Nival zone, 6, 183. See also Aeolian zone
Niwot Ridge, 8485, 85f, 115
North Atlantic Drift, 38, 60, 63. See also
Maritime
Nose bot flies, 43
Nunatak, 10, 14
Olympic Mountains, 104, 110
Origins, flora and fauna, 1314, 26, 28, 90,
100; Fennoscandia, 62; North American
alpine, 103, 1056; Pleistocene
migrations, 100103; Russia, 6566;
tropical alpine, 158, 168, 177. See also
Migrations
Oxygen, reduced, 93, 132
Palsa, 36, 65
Pamir Mountains. See Tien Shan
Mountains
Paramo, 102, 15758, 16566, 16771,
169f ; desert, 16970; superparamo, 168
Patterned ground, 3436, 36f ; Himalaya
Mountains, 130; mid-latitude alpine, 52,
8586, 89, 119
Peary Land, 23, 58
Penguins, 7071
Permafrost, 13, 3436, 38, 64, 105, 130
Phanerogams, 16
Photoperiod. See Light regime
Photosynthesis, 22; Crassulacean acid
metabolism (CAM), 15, 89
Pika, 28, 1078, 108f
Pingo, 3536, 64
Plant adaptations, 1425; cold, 2023, 160;
heat, 21, 22; mid-latitude alpine, 9093;
morphological, 20; phenology, 20;
physiological, 2122; tropical alpine,
16164; winter hardening, 20
Plant age, 23, 91, 93, 98, 114, 163, 170, 190
Plant color, 12, 90, 162, 189
216
Index
Plant communities. See Vegetation,
antarctic; Vegetation, arctic; Vegetation,
mid-latitude alpine; Vegetation, tropical
alpine
Plant hairs, 12, 1920, 19f, 90, 137
Plant temperature, 1112, 20
Pleistocene, 8, 911, 1314, 56, 100103
Polar bear, 4445
Polar comparisons, 3334
Polar desert, 6, 41, 42, 54, 56, 67;
semidesert, 5354
Pollination, 23, 92, 163, 171, 190
Polygons, 3436, 36f. See also Patterned
ground
Polylepis, 168, 176
Polyploidy, 14
Precipitation: antarctic, 69; arctic, 38,
40, 5051, 56, 59, 64; mid-latitude
alpine, 88, 105, 126, 12930, 134, 136;
tropical alpine, 160, 167, 172, 17778,
185, 188
Preformed buds, 21, 90, 91
Ptarmigan, 26, 29, 47, 55, 57, 108
Puna, 15759, 159f, 17276, 175f
Puya, 163, 173
Pyrenees Mountains, 125
Rabbits, 170, 187
Raoulia, 26, 57, 62, 93, 102, 112
Reindeer, 28, 42, 44f, 4748, 62, 67. See also
Caribou
Reproduction, bird, 29, 4647, 7071, 108,
17576; mammal, 29, 42, 43, 45, 46
Reproduction, plant: mid-latitude alpine,
9193; sexual, 2324, 9193; tropical
alpine, 16263, 190; vegetative, 2425,
24f, 93
Reptiles, 31, 109, 185
Rhea, 175
Rhododendrons, 123, 129, 130, 132
Rocky Mountains, 1012, 104, 105,
11518; climate, 105; northern Rockies,
11618; southern Rockies, 11516
Rodents, 29; arctic, 4546; burrowing, 62,
9394, 100; Eurasia, 124, 128; North
America, 100, 1067, 110, 111, 113,
11718; tropical alpine, 175, 184, 187. See
also Lemmings
Roots, 19, 21, 22
Rosettes, 1718, 17f, 18f. See also Giant
rosettes
Russia, 3840, 6367
Sagebrush, 113, 127
Salinity, 41, 57, 70, 127, 173, 175
Sclerophylous, 12, 162
Scree, 98f
Sea ice, 33, 38, 45, 56, 69
Siberia, 38, 40, 6467
Sierra Nevada, 84, 101, 104, 105, 11113,
112f
Silversword, 163, 18990
Slope: angle, 87; aspect, 87, 129, 158
Snow: and animals, 2829; as insulation,
11, 28; cover, 38, 59, 64; mid-latitude
alpine, 88
Snowbeds, 60, 9091, 99f, 131
Snow leopard, 132
Snowy Owls, 29, 46, 47, 55, 65
Soil, 1213; arctic, 4041, 57; mid-latitude
alpine, 8890; tropical alpine, 161, 168,
179
Soil nutrients, 12, 13, 90
Solifluction, 12, 34, 35, 161
Southern Africa, 13235
Southern tundra, Russia, 6465
Spodosol, 40
St. Johns wort, 186f
Succulents, 89, 123, 134, 136. See also Cactus
Supercooling, 21, 30, 166
Svalbard, 38, 60
Talus, 98f
Temperature: air, 4, 86; antarctic, 69; arctic,
3640, 50, 56; diurnal, 86, 88, 119, 160,
178; ground, 87; inversions, 38, 87; lapse
rate, 84; mid-latitude alpine, 84, 8687,
126, 130, 134, 136; tropical alpine, 160,
161, 16768, 172, 178, 185, 188
Tien Shan Mountains, 12628, 127f
Timberline, 7
Torpor (torpid), 28, 165
Index
217
Treeline, 69; as a boundary, 6; elevation,
7f ; krummholz, 9697, 97f ; mid-latitude
alpine, 9499; Russia, 64; southern
hemisphere, 97; species, 9, 64, 9495;
tropical, 157, 165; variation, 7, 94,
95t, 96
Tropical alpine, 15790; animal
adaptations, 16566; animals, 17071,
17476, 174f, 175f, 184; climate, 15861,
159f, 16668, 17779, 18588; giant
rosettes, 16364; origins, 158, 177; plant
adaptations, 160, 16164; regional
expressions, 16690; soil, 161, 168, 179;
treeline flora, 16; vegetation, 16364,
16870, 169f, 17274, 173f, 174f, 17983,
181f, 182f, 186f, 18889
Tropical alpine, afroalpine, 17685;
Ethiopian Plateau, 18587
Tropical alpine, oceania, 18790
Tropical alpine, South America, 16676,
166f ; climate, 16668; paramo,
16671; puna, 17276; superparamo,
168
Tundra, 1, 33, 51, 6367. See also Arctic
tundra; Mid-latitude alpine; Tropical
alpine
Typical tundra, Russia, 6566
Ungulates, 94, 171. See also individual species
Upper limit of plant growth, 15, 86, 91, 124,
129, 179, 183
Vegetable sheep, 137
Vegetation, antarctic, 6970
Vegetation, arctic: Eurasia, 6062, 61f,
6467, 66f ; North America, 5154, 51f,
5659
Vegetation, mid-latitude alpine: Eurasia,
12324, 12628, 127f, 129, 13031; North
America, 98101, 98f, 101f, 106, 10910,
11214, 112f, 11617, 11920, 120f ;
Southern Hemisphere, 133, 135,
13638
Vegetation, tropical alpine: afroalpine,
17983, 181f, 182f, 18687, 186f ;
Oceania, 18890; South America,
16870, 169f, 17274, 173f, 174f
Vicuna, 17475, 175f
Viviparous. See Reproduction, plant:
vegetative
Warble flies, 43
Weather data, 12
Weta, 139
White Mountains, California, 11314
White Mountains, New Hampshire, 105,
119
Wind: arctic, 38, 40, 52; arctic versus alpine,
4, 15; mid-latitude alpine, 87, 121;
tropical, 161
Winter hardening, 20
Yak, 93, 132
218
Index
About the Author
JOYCE A. QUINN is a professor in the Department of Geography at California
State UniversityFresno.

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