Global Ecology
CHAPTER OUTLINE
Life on the Land
Rainforests
Savannas and Deciduous Tropical Forests
Deserts
Grasslands
Temperate Deciduous Forests
Temperate Mixed and Coniferous Forests
Mediterranean Scrub
The Northernmost Forests—Taiga and Boreal Forest
Arctic Tundra
I magine driving across the United States from New York
City to Los Angeles. As an astute botanist you are able to
separate the patches of natural vegetation you see from agri-
cultural fields, heavily grazed pastures, and frequently
mowed roadsides. What you observe is a gradual change in
this natural vegetation. Leaving New York City, you first
encounter deciduous forest, then pass through grasslands
where trees are mostly confined to the margins of rivers, and
eventually you enter the deserts of the arid southwest with
much bare ground, scattered shrubs, and cacti. Along the
way, you also go up and down mountains, perhaps driving
from desert up into forests and grasslands, and then back
down to desert. Crossing the coastal mountains into southern
California takes you into landscapes dominated by dense
shrubs, most of them species that you have not seen any-
(a)
32–1 Mediterranean-type vegetation Regions with
Mediterranean-type climates are characterized by hot, dry summers
and mild winters, with strongly seasonal rainfall that occurs primarily
in the cooler part of the year. The vegetation is dominated by dense,
low, mostly evergreen or summer-deciduous shrubs. There is a striking
Chapter
332
where else. Such a trip makes it clear that the natural ecosys-
tems of the eastern United States are dramatically different
from those of the west. You would also observe that when
the topography is reasonably uniform, you tend to be in one
kind of vegetation for very long stretches.
The major regional ecosystems you encounter on your
trip are called biomes. Biomes are the major subunits of the
biosphere and are characterized by a more or less homoge-
neous type of plant cover. The plants and animals that occur
in particular biomes have characteristic growth forms and
other adaptations that have evolved in relation to specific
climates. It is because of these common growth forms that
we can recognize biomes, such as grasslands or deserts,
wherever they occur, even though the individual species of a
biome in one region are often completely different from
those of the same biome in another region, having achieved
similar characteristics as a result of parallel evolution.
Biomes are shaped by climate—primarily temperature
and precipitation—and so similar biomes are found world-
wide in regions of similar climate (Figure 32–1). Temperate
deciduous forest, for example, is found in North America,
Europe, and Asia, and tropical rainforest is found in Central
and South America, Africa, Asia, and northernmost Australia.
Although it is useful to identify the major biomes, it is also
important to keep in mind that climatic, geologic, and other
factors often change gradually over space and time. Thus, on
your cross-country drive, you would have noticed that biomes
do not shift abruptly from one to the next. As with other
classification systems, opinions differ about the best way to
(b)
superficial similarity in the vegetation, whether in California, Chile,
Australia, South Africa, or the Mediterranean region, as seen here for
(a) chapparal in southern California and for (b) similar vegetation,
called maquis, on the Greek island of Corfu. Each region, however,
has its unique set of species.
32–1
32–2 CHAPTER 32 Global Ecology

CHECKPOINTS
divide up the biosphere into biomes, and many scientists
By the time you finish reading this chapter, you should be able to answer the following questions: argue that it is better not to assume that discrete units exist,
1. What is a biome, and what factors affect the distribution of but rather to consider how ecosystems change as one moves
biomes on Earth? across the gradients in climate, soils, geology, and topography.
We will adopt a middle approach, using the generally recog-
2. Given that tropical rainforests contain such a diversity of nized biomes as reference points along gradients. But in real-
species, why are tropical soils unsuitable for agriculture? ity, change is often so gradual that it is difficult to know on
the ground when one leaves one biome and enters another. In
3. How is an African savanna different from a North American
grassland? What role does fire play in grasslands?
our discussion, we will emphasize vegetation, but it must not
be forgotten that heterotrophs also play essential roles in the
4. Which best characterizes a desert—high temperature or low biomes.
precipitation? How have plants adapted to desert living?

5. How does the appearance of the temperate deciduous forest Life on the Land
biome change from one season to the next? What accounts for The scheme we present for classifying the natural vegetation
these changes? of the world is based on the work of the late A. W. Küchler,
of the University of Kansas (Figure 32–2). Study of this map

8
8
7 8
7
7
10 4
4
5
4 4
4 9 2 2

6 5 1
13
11 2
12 10 3

9
16 10
10 17
5
17
14 16
16

10
9

17
16
16

10 17

5
12 15
3 17
14

11
5
6
12

32–2 Biomes of the world The information in these maps and the 9
accompanying key was originally supplied by A.W. Küchler of the University of
Kansas, one of the leading authorities on the distribution of biomes. Because of
the global coverage of the maps, the scale is relatively small and the content is
generalized. Any given biome is not always uniform, and all of the biomes
include considerable variations in vegetation.The boundaries between the
biomes may be sharp, but more often they are blurred, consisting of broad
zones of transition from one type of vegetation to another.
 Life on the Land 32–3

1 Temperate deciduous forests

2 Temperate mixed forests
3 Subtropical mixed forests
4 Taiga
4 Northwestern coniferous forest
5 Alpine tundra and mountain forests
6 Mixed west-coast forests
7 Arctic tundra
8 Ice desert
9 Grasslands
10 Savannas
11 Mediterranean scrub
12 Deserts and semideserts
13 Juniper savanna
14 Southern woodland and scrub
15 Tropical mixed forests
16 Monsoon forests
17 Rainforests

8 8
7

7
7
4

10 4
2 1 5
1 10 4
9 9
1 10 9 2 4
2 10
5 5
11 9 5 12 12 2
11 1
5 3
5

3
12 12 17
16
16
5
10
16 5 10
16
17
17
12
17
17
5 5
17
10
15
16 9
9 16

10 17

10 12
12
12 9
11 10

17 11
11 3
10
6 6
9
32–4 CHAPTER 32 Global Ecology
reveals some broad patterns, but also complexities. In the
Northern Hemisphere, the sequence from north to south,
from ice desert to treeless Arctic tundra, to northern conifer-
ous forest (taiga), and to temperate forest or grassland is evi-
dent, with each type occupying a vast area. This sequence is
not apparent in the Southern Hemisphere because of the
small land mass in the subarctic and temperate zones. We
see also that large areas of the Earth are occupied by arid or
semiarid vegetation—desert and savanna—in both the tropi-
cal and temperate zones. Within the tropics (the zone
between the latitudes of 23.5° north and 23.5° south), one
finds the quintessential tropical ecosystem—tropical rainfor-
est—as well as areas of seasonally dry forest types, such as
monsoon forest and alpine mountain forest.
These global patterns are largely explained by the latitu-
dinal gradients in temperature and the distribution of precip-
itation influenced by the latitudinal movement of air masses
(Figure 32–3). Moisture and temperature are not the only
important factors, but they strongly restrict what can grow
in a particular area. We do not find rainforests without rain,
and desert plants that tolerate drought and intense sunlight
cannot survive along the fog-bound coasts of Ireland. The
tilt of the Earth’s axis relative to the plane in which it moves
around the sun determines the amount and seasonal varia-
tion of solar energy that reaches the land surface. The higher
latitudes, both north and south, have strongly seasonal cli-
mates with short days in winter and long days in summer. In
the regions closer to the North or South Poles, the input of
solar energy dips so low that temperatures fall below freez-
ing for extended periods. Most plants can tolerate tempera-
tures just above freezing, and some can adjust to tolerate
Green arrows:
Moist air rises
and cools—
high precipitation
60° N
Westerlies
30° N
Northeast trade winds
0°
Equatorial doldrums
Southeast trade winds
30° S
Westerlies
60° S
freezing temperatures, but only plants with special structural
and physiological adaptations, such as the timing of growth
and flowering, can survive prolonged subzero temperatures.
Evaporation by solar energy is the means by which
water enters the atmosphere to be returned to the surface as
precipitation. The worldwide distribution of precipitation,
however, is extremely variable. Regions differ markedly both
in total amounts of precipitation received and in the sea-
sonal distribution of precipitation, with profound effects on
ecosystems. Because the oceans are the primary source of
evaporated water, the central regions of the continents tend
to be dry, and in some cases extremely dry. Precipitation is
also very low in the latitudinal band between 15° and 30° in
both hemispheres (Figure 32–3). High-pressure zones domi-
nate this zone and the air is generally stable, with many
cloudless days and relative calm, explaining a traditional
name for this zone—“the doldrums.” Within the tropics,
some areas have high precipitation with little seasonal varia-
tion. Intense solar radiation causes high evaporation, a
buildup of cloud masses, and nearly daily rainfall. The
regional movement of air masses ensures a constant supply
of moisture to maintain the cycle. In other tropical regions,
however, rainfall is strongly seasonal with alternating peri-
ods of prolonged drought and intense rainfall—best typified
by the monsoon climate.
Both Elevation and Latitude Determine Ecosystem
Properties
In general, air temperature decreases with increasing height
above the ground, a consequence of the fact that it is pri-
32–3 Global air currents The Earth’s
surface is covered by belts of air currents,
which determine the major patterns of wind
and rainfall. In this diagram, the blue arrows
indicate the direction of movement of the air
within the belts. The green arrows indicate
regions of rising air, which are characterized
Direction of
Earth’s rotation by high precipitation, and the brown arrows
indicate regions of descending air, which are
Brown arrows:
Dry air descends characterized by low precipitation. The dry air
and warms— descending at latitudes of 30° north and
low precipitation
south is responsible for the great deserts of
the world. The prevailing winds on the Earth’s
surface, indicated by the black arrows, are
produced by the twisting effect of the Earth’s
rotation on the air currents within the
individual belts.

marily the absorption of short-wave energy at the Earth’s
surface that captures the heat energy of the sun. The sun-
warmed surface radiates long-wave radiation back into the
atmosphere so that the air closest to the ground is warmest.
The warming is enhanced because water vapor, carbon diox-
ide, and some other gases, as well as particulate matter,
absorb the re-radiated heat, warming the air and radiating
heat back to the surface. As a result of these processes, the
usual pattern is for the temperature of the air to decrease
with height above the ground. On average, the air tempera-
ture decreases by about 6.4°C for each 1000 meters of ele-
vation (3.5°F for each 1000 feet). This is just an average,
however—local anomalies and departures from the average
are common and are important in understanding weather
phenomena.
A result of the decrease in temperature with altitude is
that locations on the Earth that are topographically higher
will have significantly cooler temperatures, on average, than
locations at the same latitude at lower elevations. This effect
is slightly counterbalanced by the fact that the air is thinner
(because pressure also decreases with elevation) and the path
of the sunlight through the absorbing atmosphere is shorter,
so that surface heating on a clear day on a high mountain
will be greater than on a spot at sea level at the same lati-
tude. The cooling with increasing altitude has profound cli-
matic consequences when air masses move from regions of
low elevation across mountains. As the air mass is forced
upward, it is cooled. Because cool air cannot contain as
much moisture as warm air, moisture-rich air masses will
produce precipitation as they are carried upward. The high-
est rainfalls on Earth are in places where this effect is partic-
ularly strong, the best examples being places where high
mountains occur on islands or continental coasts. For exam-
ple, Mount Waialeale on Kauai, Hawaii, is such a mountain.
The average annual precipitation at the highest elevations on
the windward slope is over 1200 centimeters (40 feet). This
is extreme, but mountains typically have both cooler and
wetter climates than the adjacent lowlands, a fact well
known to anyone with an interest in downhill snow sports.
Snow-
capped
peak
Snow
lin
e
Tundra
Tree
lin
e
Taiga
Altitude →
Temperate
deciduous forest
Tropical
rainforest
Equator
Life on the Land 32–5
Descending,
warming air
Rising,
cooling air Rain shadow
Prevailing wind
32–4 Rain shadow The effect of coastal mountains on patterns of
precipitation in the Northern Hemisphere. As winds come off the
water, the air is forced upward by the contour of the land, cools, and
releases its moisture in the form of rain or snow. As the air descends
on the far side of the mountains and becomes warmer, its capacity to
hold water increases, producing a rain shadow.
The tendency of mountains to extract moisture from air
masses affects the downwind climate. As the air descends, it
will warm, and its capacity to contain moisture will corre-
spondingly increase, so that even if downwind areas provide
some moisture to the atmosphere by evapotranspiration,
there will be limited precipitation. For example, the leeward
side of Mount Waialeale receives only about 50 centimeters
of annual precipitation. This downwind dryness is the “rain-
shadow” effect (Figure 32–4). It is particularly strong along
the Pacific coast of North America, with zones of aridity or
semiaridity on the east side of mountain chains from the
northern United States down into Mexico.
The climatic changes that one observes when climbing
a mountain mimic in several important ways the changes
that occur as one travels north from the equator—mainly
with respect to temperature and the occurrence of freezing
conditions (Figure 32–5). In general, a change in mean
atmospheric temperature corresponding to an increase in
latitude of 1° occurs with each increase in elevation of
32–5 Relationship between altitude and
latitude In the Northern Hemisphere, we can
experience a similar sequence of dominant
plant life and its associated animal life by
either traveling north for hundreds of
kilometers or ascending a mountain. This
relationship between altitude and latitude
was first pointed out by Alexander von
Humboldt. To experience a similar sequence
of vegetation in the Southern Hemisphere, we
could ascend a mountain. However, by simply
Polar traveling south, we would never encounter
ice
vegetation corresponding to the taiga and
the tundra of the Northern Hemisphere. Can
Latitude → Pole you explain why?
32–6 CHAPTER 32 Global Ecology

Alexander von Humboldt

Alexander von Humboldt (1769–1859) was On leaving Latin America in 1804, Hum- new lands encouraged Jefferson’s own great
perhaps the greatest scientific traveler who boldt visited the United States for eight weeks. scheme for the exploration of the western
ever lived and was certainly one of the greatest He spent three of these weeks as Thomas Jef- United States. Thus, it is fitting that Hum-
writers and scientists of his era. A native of Ger- ferson’s guest at Monticello, talking over many boldt’s name is commemorated in the names
many, Humboldt ranged widely across the inte- matters of mutual interest. It is thought that of several counties, mountain ranges, and
rior of Latin America from 1799 to 1804 and Humboldt’s enthusiasm for learning about rivers in the American West.
climbed some of its highest mountains. Explor-
ing the region between Ecuador and central
Mexico, Humboldt was the first to recognize the
incredible diversity of tropical life and, conse-
quently, the first to realize just how vast a num-
ber of species of plants and animals there must
be in the world.
In his travels, Humboldt was impressed
with the fact that plants tend to occur in
repeatable groups, or communities, and that
wherever there are similar conditions—relat-
ing to climate, soil, or biological interac-
tions—similar groupings of plants appear. He
also discovered a second major principle—
the relationship between altitude and lati-
tude. He found that climbing a mountain in
the tropics is analogous to traveling farther
north (or south) from the equator. Humboldt
illustrated this point with his well-known dia-
gram of the zones of vegetation on Mount
Chimborazo in Ecuador, which he climbed. On
this mountain, he reached the highest eleva-
tion on record attained by any human being
up to that date.

approximately 100 meters. As a result, climbing a tall moun-
tain can take you through a sequence of ecosystem types that
will resemble what might occur if you went north toward
the Arctic. For example, in the southwestern United States,
the highest mountains are topped by conifer-dominated
forests, then, at higher elevations, by tundra-like meadows
and scree fields, and eventually by a zone of perpetual snow
cover. In the tropics, the climatic changes are just as dra-
matic, but the vegetation of the cold upper elevations does
not necessarily look like anything you will ever see in Alaska
or Siberia. This does not refute the generality, but shows only
that seasonality is an important aspect of climate.
Soils and Fire Influence Regional Patterns
of Distribution
The sketch we have just given of the major factors shaping
biome types omits many other important influences on plant
growth. Of these, two deserve mention because of their
importance at regional levels—soils and fire. Soil, as the
medium of plant growth, determines how much water and
what amounts of necessary nutrient elements are available to
plants. The kind of soil present within a region is deter-
mined, in part, by climate but also by the “parent material”
in which the soil forms and by the length of time that a
region has been stable and therefore has had surfaces ex-
posed to the processes of soil weathering. Australia, for
example, has been geologically stable over millions of years,
and largely because of this, it has many areas that are nutri-
ent poor, with deficiencies of essential elements and espe-
cially phosphorus. It is thought that these deficiencies partly
explain unique features of Australian vegetation, such as
the abundance of sclerophyllous (leathery-leaved) species,
most notably the many species of Eucalyptus. In contrast,
the continental glaciers of the Pleistocene that covered
much of northern North America and Eurasia renewed the
topsoil with fresh material, as bedrock was ground up by
the glacier and redistributed in the form of glacial till and
as windborne dust (loess) was carried far beyond the edges
of the glaciers. Geologically speaking, such glacier-derived
or glacier-affected soils are very young, so they have high
levels of essential elements.
Fire is particularly important for biomes that have alter-
nating wet and dry seasons, such as grasslands, open savan-
nas, or the chaparral of California, and much less so in
evergreen, moist areas, such as tropical forests. In some
 Life on the Land 32–7

regions, fires determine the line between two vegetation releases the seeds at a time most suitable for their germina-
types, as between Eucalyptus woods or savannas and moist tion and establishment. The huge fire in Yellowstone
tropical forests in eastern Australia. Here the area of moist National Park in the summer of 1988 was basically an
forest has progressively been reduced as indigenous people example of a natural fire cycle in the park, burning mainly
have set fires to improve grazing. For at least 1.4 million through lodgepole pines (Figure 32–6). Subsequent examina-
years, human beings have had the ability to start and keep tions showed that a similar cycle of fires had occurred
artificial fires, which they have done extensively in some around the year 1700.
areas, especially during the last 20,000 years or so. In Fires have many significant ecological effects. They
regions that have burned, most species of organisms survive, release stored nutrients, such as phosphorus, calcium, and
and in fact they often depend on fires for the renewal of potassium, from the woody or perennial vegetation back
their populations. into the soil. This stimulates the regrowth of plants, which
Wildfires, which are set by lightning, generally consume temporarily have access to large amounts of nutrients. Some
masses of vegetation that have built up in the moist season, plants exist as seeds stored in the soil, and they germinate
and these fires may burn each dry season. In regions where and flower only after a fire. Certain perennials grow luxuri-
lightning is particularly frequent, such as the state of Florida, antly and flower only after fire releases necessary amounts of
there may be many wildfires. Along with fires set acciden- nutrients, and the plants have sufficient light. Particularly on
tally or sometimes even deliberately by humans, wildfires in relatively infertile soils, fires may be essential for the renewal
the western United States may burn hundreds of thousands and regrowth of the vegetation. At the same time, soil ero-
of square miles of fire-prone forests, woodlands, brushlands, sion may be accelerated, which is why management agencies
and grasslands each year. Sometimes there are disastrous are often anxious to sow cover crops on burned slopes in
consequences for cities, towns, or individual houses or struc- the wake of a fire. Fires not only remove vegetation, but the
tures isolated in areas that burn periodically, and even the intense heat associated with fires can cause soil particles to
loss of lives. Hundreds of fires may break out in a single day become water repellent so that water runs off rapidly, accel-
over the region, greatly taxing the ability of firefighters to erating erosion still further.
battle them effectively. As people have built homes in areas
where fires are a normal part of regeneration and renewal,
the tendency is to suppress fires. This suppression can lead
eventually to the accumulation of large amounts of dry vege-
tation that act as tinder when they are ignited. When these
fires occur along the edges of cities, their effects are greatly
magnified, and they may cause great loss to human life and
to property.
Better methods of managing vegetation, either by con-
trolled burns or other methods of removing the accumulated
biomass, will be necessary if these fires are to be limited in
number and scope in the future. Controlled burning by set-
ting prescribed fires works best in areas characterized by
low-intensity fires. In areas subject to high-intensity fires,
such burns can rage out of control, as happened around
Los Alamos, New Mexico, in 2000. All methods of sup-
pressing fires affect both the appearance of the forests or
other vegetation types and the well-being of the biodiversity
they contain. The application of fire-suppression methods
has become the subject of arguments, often fierce, among
proponents of different approaches to the problem.
Understanding the frequency of fires in a particular area
provides facts useful in predicting and managing future fires.
This information can be obtained by examining fire scars on
trees; analyzing sediments in lakes, ponds, or other wet
areas; or studying the written history of particular areas.
Thus, we have learned that lodgepole pines (Pinus contorta),
which occur frequently at higher elevations throughout the
western United States, usually burn at intervals of 250 to
400 years. Following a fire, these pines can then germinate
and grow well in the bare soil, outcompeting other trees
locally. Lodgepole pines also have cones that remain closed 32–6 Lodgepole pines Following the disastrous 1988 fire in
and attached to the branches until they are heated. The heat Yellowstone National Park, lodgepole pine (Pinus contorta) seedlings
melts the thick, waxy substance coating the cones and became established at the burn site.
32–8 CHAPTER 32 Global Ecology
Fire and its consequences in particular areas will be dis-
cussed further in the biome descriptions that follow.
Rainforests
It makes sense to begin our exploration of biomes by asking
what kind of system is found where the fewest factors are
limiting—where temperatures are relatively constant and
never below freezing, where rainfall is consistent and there-
fore soil moisture ample throughout the year, and where
geology and topography allow deep soil formation and
retention of at least an adequate supply of nutrients. These
conditions favor high and consistent photosynthesis and
therefore maximum plant growth. Plants can expand their
canopies rapidly and, without the constraints of freezing and
drought, can have large or thin leaves for maximum absorp-
tion of light. The result is strong competition for light.
Plants that fall behind in capturing sunlight are quickly over-
topped. Over evolutionary time, this has favored a prolifera-
tion of woody species capable of obtaining their place in the
sun by one means or another. The wet lowland tropics
therefore are dominated by tropical rainforests—dense
forests consisting of species that are active all year round.
The plants are either evergreen (that is, their leaves are
always actively photosynthesizing) or leafless for only very
brief periods.
The dominance of trees influences all other aspects of the
rainforest. Much of the sunlight is absorbed by the dense
canopies, leaving relatively little light to penetrate to the for-
est floor. As a result, undisturbed tropical rainforest usually
has only a sparse, patchy herbaceous understory consisting of
species with special adaptations for dealing with low light
intensity. Seedlings and saplings of shade-tolerant trees are
often the most abundant plants in the ground layer.
Because of the dominance of trees in the rainforest, the
biomass in the system that is of value to herbivores is mostly
in the tree canopies. This, and the favorable climate,
explains the abundance of arboreal (tree-dwelling) animals,
such as gibbons and howler monkeys, which exhibit a way
of life highly specialized to exploit the fruits, insects, and
nutritious new growth of the treetop environment.
Trees in all regions (unless subject to air pollution) have
algae, mosses, and lichens on their branches and bark, but in
the wet tropics, conditions favor an abundance of plants
specially adapted to grow on trees. These plants, known as
epiphytes (meaning “on plants”), grow on the branches of
other plants that grow in the illuminated zone far above the
forest floor (Figure 32–7). Epiphytes can survive without soil
by collecting rain as well as nutrients from animal drop-
pings, plant litter, dust, and dead insects. There are epiphytes
from many plant families, but two well-known groups are
the orchids and the bromeliads; there are also many epi-
phytic ferns. Lianas, or woody vines also known as climbers,
represent another strategy for exploiting trees for support.
Lianas are rooted in the soil, but grow up along the trunks
of the trees into the canopy. Some lianas start as epiphytes
and drop stems to the soil where they take root. Strangler
figs (Ficus spp.) begin growth as epiphytes, dropping roots
to the ground and growing to envelop their host trees. The
fig eventually becomes an independent, hollow tree, while its
original host often dies.
Tropical rainforests have extraordinary species richness.
A typical hectare (2.47 acres, or about 2.5 football fields) of
boreal coniferous forest will have 1 to 3 tree species, a rich
temperate deciduous forest 10 to 20 species, but tropical
rainforests commonly have 100 species, and some have 200
or more. Putting this in more concrete terms: in a typical
temperate forest, there is a very good chance that if you
select a tree, the tree closest to it will be the same species; in
tropical rainforest, the tree closest to your selected tree is
more likely to be some other species. Ecologists are still
searching for a full explanation of why tropical rainforests
are so diverse. It seems reasonably certain that uninterrupted
32–7 Epiphytes By collecting and storing
water and nutrients from the surrounding
air, rain, and dust, epiphytes create little
patches of soil from accumulated debris.
Bromeliads, seen here as leafy tufts growing
along the tree branches, are among the
most common of the epiphytes. The leaves
of bromeliads merge at their bases to form
watertight tanks that, in the larger species,
can hold as much as 45 liters of rainwater.
These pools of water are microcosms of
bacteria, protozoa, larvae, insects, and
insect-eaters. Many rainforest mosquitoes
breed exclusively in bromeliad tanks. The
bromeliads absorb water from their built-in
reservoirs and are also supplied with
nutrients from the debris.

evolution is part of the reason, so that over time more
species accumulate. This could be more or less chance, or it
could represent the action of evolutionary pressures for the
subtle differentiation of species to fill different ecological
roles. Other theories stress that seed predators and
pathogens might act disproportionately on the most com-
mon species, or environmental stability may make the ex-
tinction of smaller populations less likely.
Like all natural systems, rainforests are dynamic and
constantly changing. Because of the abundant rainfall, nat-
ural fires are rare. But tree fall, usually due to wind, is com-
mon and is the main natural mechanism by which rainforest
is disturbed. Those tropical rainforests that lie within the
(a)
(b)
Rainforests 32–9
reach of hurricanes can suffer widespread canopy loss, but
such catastrophes are not essential for tree fall. Although
rainforest trees are well protected against wind damage by a
variety of mechanisms, including rot-resistant wood and
buttressing (Figure 32–8), sooner or later they succumb.
Even if it is only initially a single tree that falls, it often
brings down other trees around it by direct impact, uproot-
ing, or because the canopy of the falling tree is connected to
others by lianas. The result is a large gap in the canopy.
Light can then penetrate to the forest floor, and a strong
pulse of growth by herbs, shrubs, seedling trees, and
resprouted trees follows. Any sapling trees that survived the
falling of their large neighbors can experience a growth
(c)
32–8 Tropical rainforest (a) The interior
of a rainforest in Costa Rica. The broad-
leaved plants with red flowers are Heliconia
irrasa. (b) The diversity of the trees in the
forest, which may reach several hundred
species per hectare, is revealed when
individual trees burst into bloom, such as
these trees in the coastal rainforest of Brazil.
(c) A buttressed tree in the rainforest of
Ecuador. Note the woody vines known as
lianas on the trunk.
32–10 CHAPTER 32 Global Ecology
spurt that allows them to extend up into the canopy. Some
species of tropical rainforest trees require such sunny
patches to become established.
Temperate areas with cooler climates also can have cli-
mates with high rainfall. These rainforests of higher lati-
tudes, like those along the eastern side of Australia, in
Tasmania, and in parts of New Zealand, have much lower
species diversity but share the quality of dense tree growth.
Rainforests Are Rapidly Being Destroyed
Tropical rainforests, which already have been cut back to
about half of their original extent, are being lost rapidly as a
result of human activity. Since they are estimated to contain
half of the world’s species of plants and other organisms, this
loss is extremely serious. A traditional pattern of human use,
known as shifting cultivation, mimics the natural dynamics
of the rainforest by opening small gaps in the canopy, culti-
vating crops for a few years, and then abandoning them.
These areas then undergo succession and return to natural
forest cover. This kind of land use is thought to be sustain-
able, but only so long as population levels are low relative
to the land being exploited. With increasing populations,
rotation time is shortened. But the greatest threat to the
tropical rainforest is the conversion of forest to cropland
and open pastures. In areas less suitable for agriculture,
forestry practices often involve removing more trees than
can be replaced by natural succession. Creating large gaps
in the tropical rainforest makes it susceptible to fire to a
degree unprecedented in history, as was tragically demon-
strated by the destructive fires of 1997–1998 in the Kali-
mantan region of Indonesia, where over 5 million hectares
of forest were destroyed. Destruction of rainforest of this
kind will lead to the loss of many species of plants, animals,
and microorganisms.
The destruction of tropical ecosystems would be more
defensible if the results were always highly productive agri-
cultural lands. But experience shows otherwise. Tropical
soils often prove to be fragile. Tropical vegetation is lush,
but in large areas of the tropics, upland soils tend to lose fer-
tility rapidly when cleared for agriculture. Much of the fer-
tility in tropical rainforest ecosystems is tied up in the
vegetation. Soils are generally acidic and deficient in cal-
cium, phosphorus, and other nutrients. Plant roots tend to
spread out in a thin layer no more than a few centimeters
below the soil surface, and they rapidly transfer the nutrients
released by the decomposition of fallen leaves and branches
back into living plants. Below this thin layer of topsoil,
which is easily destroyed during the process of clearing away
the trees, there is virtually no organic matter. Fertilization
may not help much because tropical soils often fix the phos-
phorus into insoluble forms. For all of these reasons, the cul-
tivation of tropical soils presents formidable problems.
Nevertheless, the tropical forests of the world are being cut
and burned at an ever-increasing rate, often to produce fields
that become completely useless to agriculture within a few
years (Figure 32–9). It is estimated that by the middle of this
century, only about 5 percent of the area originally covered
32–9 Erosion Of the native vegetation
that once covered Madagascar, an island in
the Indian Ocean 420 kilometers east of the
African mainland, only 10 percent remains,
and that is fast disappearing. On
Madagascar, as in other tropical areas, the
soils exposed by clearing vegetation are
often not suitable for sustainable
agriculture. The abundant rains rapidly
wash the topsoil off increasingly barren
landscapes, choking the rivers with silt, as
seen here, and sometimes causing floods
and landslides. The sea around Madagascar
is often rust-red from the loss of the island’s
weathered red soil, and it is difficult to
produce enough food to sustain the people
of the world's fourth-poorest country.
 Savannas and Deciduous Tropical Forests 32–11

by tropical rainforests will survive, and those relatively small

patches will be extremely vulnerable to further harm.

Savannas and Deciduous Tropical Forests

Not all areas in the tropics have high or continuous rainfall.
In regions in which there is a prolonged dry season alternat-
ing with a rainy period, tropical rainforest gives way to
other forest types tolerant of drought. These forests can be
grouped under the general term of tropical seasonal forests.
For example, in central and southern Africa, the Indian sub-
continent, and Southeast Asia, there are extensive areas of
monsoon forest, and similar climates and forests are found
in northern Yucatán Peninsula in Mexico, in northern
Colombia and Venezuela, in Ecuador, and in eastern Brazil
(Figure 32–2). These climates produce enough rain to per-
mit vigorous tree growth and dense and productive forests,
but the dominant trees are deciduous, losing their leaves
during the dry season. Elsewhere in areas of the tropics
with seasonal rainfall, there are other types of dry forests,
such as the short-tree forests of tropical Mexico and Cen- 32–10 Subtropical mixed forest Slash pine (Pinus elliottii) is one of the
tral America, or the biologically distinct dry forests of widespread evergreen conifers that occurs in the subtropical mixed
Madagascar. Tropical mixed forests, in which both ever- forests of the southeastern United States. Conifers tend to dominate on
green trees and shrubs are present, occur locally in eastern soils that are either nutrient-poor or seasonally flooded, or both.
and southern Brazil and northern Australia (Figure 32–2).
The disappearing tropical rainforest has caused alarm
among conservationists, but dry tropical forests also have
extraordinarily high levels of biodiversity and are equally canopy. Trees and shrubs are scattered individually or grow
deserving of protection. in groups in a grassland matrix, and, in places, woody plants
Although pine trees are often associated with northern nearly disappear. Such areas are called savannas, and they
forests, pines also occur in the tropics and subtropics. Most show many variations worldwide. Savannas usually have
of Florida and extensive areas throughout the southeastern much lower annual rainfall than tropical rainforests—fre-
United States are covered by subtropical mixed forests. In quently in the range of 90 to 150 centimeters a year. There is
these forests, pines (Figure 32–10) and some evergreen also a wider range in average monthly temperatures, due to
broadleaf trees are mixed with deciduous trees; the precipi- the seasonal drought and the sparse covering of vegetation.
tation occurs mainly in the summer. In the tropics, savanna trees are broad-leaved deciduous
With further decreases in rainfall, a point is reached at and evergreen trees, which may occur singly or in groves,
which moisture is not sufficient to sustain a continuous tree and some savannas are dominated by shrubs (Figure 32–11).

32–11 Savanna A savanna in

East Africa with giraffes surrounded
by a herd of impala. The
transitional nature of this biome,
relative to the characteristic
vegetation of the tropical rainforest
biome and the desert biome, is
evident in the grasses, shrubs, and
short trees (acacias) seen here.
32–12 CHAPTER 32 Global Ecology
Savannas cover large areas of East Africa and are also found
on all continents on the margins of rainforests, where the
rainfall is seasonal and limiting (Figure 32–2). The thorn
forest (cerrado) that covers wide areas in Brazil belongs to
the savanna biome.
In savannas, because of the scattered distribution of
taller woody plants, the ground is generally well illuminated,
and perennial herbs (mostly grasses) are common. Bulbous
plants, which are able to withstand periodic burning, can be
abundant. Because of the dense cover of perennial herbs
made possible by the high seasonal rainfall, there are few
annual herbs. Epiphytes are also rare because of the stress of
the prolonged dry season.
Trees that occur in savannas often have thick bark as
protection against fire. They are well branched, but they are
seldom more than 15 meters tall. Many trees, such as the
acacias, are protected by stout spines to discourage grazing
(Figure 31–7a). Their leaves are generally smaller than those
of the evergreen trees of the rainforest, and so they lose less
water by transpiration.
Fire, which is common in all tropical savannas, is a
strong influence on the relative abundance of trees, shrubs,
and grass. Where soils and climate allow dense grass cover
to develop, fire can be frequent, sometimes occurring annu-
ally. With such high fire frequencies, trees are limited to pro-
tected situations, such as along rivers or in rocky areas.
Thus a climate already challenging for trees can favor fire,
which will further restrict and perhaps even completely elim-
inate trees over large areas.
Savannas and similar seasonally deciduous tropical and
subtropical plant communities grade into tropical rainforests
as the rainfall becomes higher and its seasonal distribution
more even, and corridors of tropical rainforest (called gallery
forest) extend out into the savannas along rivers. At their
poleward margins, savannas and related plant communities
grade into deserts.
Savannas extend into temperate areas, occurring in the
region between the prairies and temperate deciduous forests,
as well as in the region between prairies and taiga in North
America and over much of eastern Mexico, Cuba, and south-
ernmost Florida. Savannas in these cooler, moister climates
are thought to be dependent on fire for their maintenance.
Many believe that the savanna vegetation was greatly ex-
panded in pre-Colombian times because of burning by the
indigenous people. Deliberate burning was also practiced in
other parts of the world, often to improve grazing for domes-
tic stock, a practice that continues today in many places.
Deserts
With further decreases in rainfall, a point is reached at
which extended drought prevents the plant cover from being
continuous in time or space or both, resulting in deserts.
Bare patches of soil appear, and dense vegetation is conspic-
uously restricted to drainages, or areas where local runoff
accumulates, such as at the base of rocky cliffs, or around
the rare springs or seeps, as in the desert palm oases. The
most important factor creating the dry conditions is the rela-
tive stability of the atmosphere in desert regions. The great
deserts of the world are all located in the zones of atmo-
spheric high pressure that flank the tropics at about 30°
north latitude and 30° south latitude, and they extend pole-
ward in the interior of the large continents (Figure 32–2).
Many deserts receive less than 20 centimeters of rainfall per
year. In the Atacama Desert of coastal Peru and northern
Chile, the average rainfall is less than 2 centimeters per year.
Averages, however, do not tell the whole story, because as
average rainfall decreases, the variability from year to year
increases. In a humid region, a severe drought would be
50 percent less rain than average, but in a desert a drought
can be no precipitation at all—sometimes for several years.
The plants and animals of extremely arid regions must be
able to tolerate these periods of extreme dryness and to
exploit the good years when they occur.
Extensive deserts are located in North Africa and in the
southern part of the African continent, where the Namib
Desert is inhabited by some of the world’s most extraor-
dinary organisms, including Welwitschia (Figure 18–38).
Other deserts occur in the Near East, in southeast Mongo-
lia and northern China, in western North America and
western and southern South America, and in Australia. The
Sahara, which extends all the way from the Atlantic coast
of Africa to the Arabian peninsula, is the largest desert in
the world. Seventy percent of the entire continent of Aus-
tralia is covered in semiarid or arid areas. Less than 5 per-
cent of North America is desert, and much of that is not
extreme (Figure 32–12).
The temperatures in many deserts—the hot deserts—are
very high because the skies are usually completely clear and
little heat is taken up in evaporation and transpiration. Sum-
mer temperatures of more than 36°C are common in some
deserts. The same conditions that make deserts hot in the
day cause them to cool rapidly at night. With little or no
cloud cover, generally clear air because of limited moisture,
and no tree canopies to trap outgoing radiation, the heat
stored during the day is lost through re-radiation, and tem-
peratures drop precipitously as soon as the sun sets. Human
beings can die from exposure in deserts, not only from
extreme heat but also from cold. Although all deserts are
dry, not all of them are characterized by continuously high
daytime temperatures. There are also cold deserts. An exam-
ple is the Great Basin Desert of western North America,
which lies between the Sierra Nevada–Cascade Mountain
system and the Rocky Mountains (Figure 32–12d). Here
there are only a few weeks of high temperatures each year,
and winters can be bitterly cold, with some of the scant pre-
cipitation falling as snow.
The annual distribution of rainfall in deserts generally
reflects that of the adjacent areas. On the equatorial side, it
rains in the summer; on the poleward side, it rains in the
winter. Between the two, as in the lowlands of Arizona,
there may be two annual peaks of precipitation. As a result,
such an area generally has two periods of active plant
growth—one in the winter and one in the summer—and dif-
ferent plants are active in each period. In general, the pat-
terns of activity of desert plants reflect the origins of the
plants. Characteristically, plant species that have migrated

(a)
(c)
32–12 Desert Shown here are some representative plants of the
principal deserts of North America. (a) The Sonoran Desert stretches
from southern California to western Arizona and south into Mexico. A
dominant plant, the giant saguaro cactus (Carnegiea gigantea), is often
as much as 15 meters high, with a widespreading network of shallow
roots.Water is stored in a thickened stem, which expands, accordionlike,
after a rainfall. (b) To the east of the Sonoran is the Chihuahuan Desert;
one of its principal plants is the agave, or century plant (Agave deserti),
a monocot. (c) North of the Sonoran is the Mojave Desert, with its
characteristic plant, the Joshua tree (Yucca brevifolia). This plant was
from areas where they grew actively in the winter continue
doing so in the desert. Similarly, plant species that have
migrated into the deserts, from areas where they grew dur-
ing periods of summer rainfall, continue to grow actively in
the summer in the desert, wherever a sufficient amount of
summer rainfall makes this possible.
Desert Plants Are Adapted to Low Precipitation
and Extremes of Temperature
High variability in moisture conditions means that desert
plants must be able either to survive the dry season or dry
Deserts 32–13
(b)
(d)
named by early Mormon colonists who thought that its form resembled
that of a bearded patriarch gesticulating in prayer.The Mojave contains
Death Valley, the lowest point on the continent (90 meters below sea
level), only 130 kilometers from Mount Whitney, with an elevation of
more than 4000 meters. (d) The Mojave blends into the Great Basin
Desert, a cold desert bounded by the Sierra Nevada to the west and the
Rockies to the east. It is the largest and bleakest of the American deserts.
The dominant plant is sagebrush (Artemisia tridentata), shown here
with the snow-covered Sierra Nevada in the background.
periods in a dormant condition or to have some mechanism
of averaging out variable moisture conditions. The life his-
tory of annual plants is ideally suited to survival through
droughts of unpredictable length. It therefore makes sense
that annuals are better represented, both in number and in
kind, in the deserts and semiarid regions of the world than
in any other biome. Annuals can germinate and complete
their life cycles in the open areas of ground during the lim-
ited periods when water is available. The seeds of these
plants are able to survive extended droughts in the soil.
Then, when rains thoroughly wet the soil, the seeds can ger-
minate rapidly, flower, and set seeds before the soil moisture
32–14 CHAPTER 32 Global Ecology
is depleted. At the other extreme are species of plants, the
succulents, that can endure the dry periods by storing water.
Succulent plants are found in a wide range of plant families,
but two groups are notable for having many succulent
species—the cacti of the New World, essentially all of which
are succulents, and the succulent euphorbs (Euphorbiaceae)
of the Old World. Succulents are designed to absorb water
while it is available and store it in special tissues.
CAM photosynthesis provides another means of con-
serving the stored water. CAM plants absorb carbon dioxide
only at night; their stomata remain closed during the day. By
this means, they are able to take up carbon dioxide with
much reduced water loss. Another feature of succulents is a
morphology that allows them to change volume without
damage (see the essay “How Does a Cactus Function?”).
The success of the succulent life history is evidenced by the
fact that succulents often are among the tallest and most
robust plants of deserts—the tall saguaro cactus of the Sono-
ran Desert of the southwestern United States being a well-
known example (Figure 32–12a).
Other plants—creosote bushes are an example—do not
store large amounts of water, but they have the ability to tap
into deep-water reserves by means of extensive root systems
(Figure 32–13). Because deep water is accumulated by slow
downward percolation, it tends to vary much less than sur-
face water and to be available during dry seasons and, in
some situations, through many dry years. Tapping deep
water would not work, however, if the plants were not able
to use water efficiently. Adaptations, such as small, leathery
leaves with relatively few stomata and stems resistant to the
loss of conductivity due to embolisms, restrict water loss and
allow the plant to resist wilting when under extreme mois-
ture stress. C4 photosynthesis is likewise more common
among the plants of deserts and other seasonally dry, warm
habitats than it is elsewhere. Such plants also have the
32–13 Creosote bushes One of the most
characteristic plants of the Mojave, Sonoran, and
Chihuahuan deserts of North America is the creosote
bush (Larrea divaricata), which has small, leathery
water-conserving leaves. Creosote bushes in the
Mojave Desert may form circular or elliptical clones
because of new branch production at the periphery
of stem crowns and the segmentation and death of
older stem segments, resulting in a ring of satellite
bushes around a central bare area.The latter usually
accumulates a mound of sand, which may reach a
depth of about half a meter. Some clones may attain
extreme ages: the King Clone, shown here, is
estimated to be nearly 12,000 years old. Such
ancient clones started from seeds that germinated
near the end of the last glacial expansion.
capacity to drop leaves and even entire branches in dry peri-
ods so that they can maintain their water balance. Then,
when better conditions return, they can regenerate their
canopies. Other plants have green stems rich in chlorophyll
and thus are able to photosynthesize even when leafless.
Perennial herbs face difficult challenges in the desert
because of the extremely dry conditions, but grasses are
found in many deserts, usually as widely scattered, dense
bunches known as tussocks. Some plants have deeply buried
bulbs that break their domancy when stimulated by rain,
while others occur along arroyos or washes, where moisture
is available near the surface. The amazing capacity of plants
to adapt to desert conditions is exemplified by the gray-
white wooly shrub known as Arizona honeysweet (Tidestro-
mia oblongifolia), a C4 perennial herb common in the Death
Valley region of California and Nevada. The maximum pho-
tosynthetic rate for this species is achieved at temperatures
between 45° and 50°C in full sunlight in midsummer—tem-
peratures that would be damaging or even fatal to plants of
regions with higher moisture.
As we have discussed, deserts grade into savanna-type
communities. One such community is the widespread juniper
(Juniperus spp.) woodlands of western North America (Fig-
ure 32–14). Juniper woodlands occur in places that are usu-
ally drier than will support prairie but wetter and cooler than
desert areas. In many places in the western United States,
juniper, often found with pinyon pine, grows in areas adja-
cent to deserts but at higher elevations. Studies of pollen and
macrofossils preserved in pack-rat middens show the degree
to which vegetation types that seem like permanent features
of the landscape actually are quite dynamic over time. Dur-
ing the pluvial periods of the Pleistocene epoch, at the times
of maximum expansion of the continental glaciers, juniper
woodland communities moved down into the lowland areas
that presently support juniper-free deserts.
 Deserts 32–15

How Does a Cactus Function?

The barrel cactus Ferocactus acanthodes, fold and shrink. When rain finally returned, the transpired to mass of CO2 fixed is only about
which grows along the northwestern edge of shallow root systems (with a mean depth of 70:1 for the entire year. This is considerably
the Sonoran Desert in southern California, gets only about 8 centimeters) took in water so lower than for a typical C3 plant, which requires
its name because it is shaped like a barrel. It rapidly that the stomata were fully functional larger quantities of water to fix an equivalent
looks as if its stem has been folded like a fan, again within 24 hours of rainfall. amount of carbon but has a higher maximum
and it is covered with spines like a porcupine. But these are not the only mechanisms the rate of photosynthesis.
Why should this desert plant present such an barrel cactus uses for conserving water. Like Because seedlings of the barrel cactus,
unusual and formidable appearance? many other desert plants, the barrel cactus unlike their parents, cannot tolerate extremely
When the barrel cactus is full of water, the exhibits crassulacean acid metabolism (CAM). It high temperatures and prolonged drought,
folds swell and are barely visible, but when the opens its stomata only at night and so under- they survive only in certain years and in pro-
plant dries, the folds are deep and the stem is goes gas exchange with cooler air, which can tected microhabitats. By the age of 26 years,
able to contract without crushing the cells. This hold less water than warmer air. Consequently, these plants have usually grown to only about
ridge-and-valley folding of the stem has other the plant loses less water to the atmosphere 34 centimeters tall, adding about 10 percent
advantages, too. Deep inside the valleys be- through transpiration. Its ratio of mass of water to the mass of their stems each year.
tween the folds are the stomata. The spines
help to break up the wind currents, and thus
the valleys serve as protected retreats where
dry desert winds cannot easily reach to carry
moist air away from the vicinity of the stomatal
chambers.These formidable spines also help to
protect the cactus from the rodents and birds
that are in constant search of water, even
going so far as to steal it from a succulent stem.
In a study conducted by Park Nobel, of the Uni-
versity of California, Los Angeles, barrel cacti
stored enough water in their succulent stems to
permit them to open their stomata for about
40 days after the soil became too dry to furnish
them with any additional water. Under such
conditions, many of the fine roots are sloughed
off, to prevent water loss to the soil. After seven
months of drought, stomatal activity ceased,
and the osmotic potential of the stem was
more than double the value it had been during
wet periods, despite the ability of the stem to Barrel cactus in a hydrated state (left), and in a dehydrated state (right).

(a) (b)

32–14 Juniper woodlands (a) Juniper (Juniperus osteosperma) are characteristic of the pinyon-juniper savannas and woodlands of
savanna in the Great Basin near Wellington, Utah. (b) Cold winters the Great Basin, as shown here south of Moab, Utah.
32–16 CHAPTER 32 Global Ecology
Grasslands
Grasslands occur where the amount of rainfall is less than is
needed to support vigorous tree growth but great enough
to allow herbaceous plants, especially grasses, to dominate.
There are many kinds of grasslands; the major variations re-
late to rainfall. Grasslands with the highest rainfall are domi-
nated by tall grasses in closely spaced clumps or dense sods.
Grasses in drier areas occur in widely spaced clumps, with
sparse or virtually no vegetation between them. Grasslands
occur from the tropics to the edges of the boreal regions and
from sea level to high mountain meadows. They grade into
savanna and desert and occur as patches in woodland and
forest. They are most extensive, however, in the middle lati-
tudes. These grasslands are characterized by cold winters and
warm summers, with moderate to low rainfall and occasional
periods of extreme dryness.
Grasses, which regenerate at the beginning of each grow-
ing season from buds at or below ground level, have many
fine roots. Over long periods of time, grasslands tend to build
soils that are deep and rich in organic matter. Such soils are
ideal for agriculture, and vast areas of grassland have been
converted to cropland. In areas too dry for agriculture, grass-
lands have value as grazing lands (Figure 29–5).
Grasslands generally occur over large areas in the interior
portions of the continents, most notably in North America
and across Eurasia (Figure 32–2). In North America, there is
a transition from the more desertlike, western shortgrass
prairie (the Great Plains), through the moister, richer, tallgrass
prairie (the Corn Belt), to the eastern temperate deciduous
forest (Figure 32–15). Grasslands become progressively drier
(a)
32–15 Grasslands The grasslands of North America include large
regions of shortgrass and tallgrass prairie. (a) A female bison nursing
her calf in the shortgrass prairie of Custer State Park, South Dakota.
with increasing distance from the Atlantic Ocean and the Gulf
of Mexico, which are the major sources of moisture-bearing
winds in the eastern half of the North American continent.
Farther north, grasslands become moister again as evapora-
tion decreases at relatively cool temperatures. “Typical” short-
grass prairie occurs under near-drought conditions. In years
with greater moisture, taller grasses (but not the very tall
ones of the tallgrass prairie) tend to predominate. Grasslands
also occur as patches within forested biomes when soil con-
ditions, such as shallow soil over bedrock, limit tree growth.
In the central and eastern United States, such grassy patches
are often called glades, and these are typically dominated by
prairie plants (Figure 32–16).
Perennial bunchgrasses and sod-forming grasses domi-
nate, but other perennial herbs are common. Although the
growth of grassland plants is seasonal, there is little room
for the development of annual herbs, and these are mostly
limited to disturbed areas, such as prairie-dog towns or bad-
ger diggings. Annuals and introduced weeds are common in
areas disturbed by humans, such as near buildings and along
roadsides and railroads.
Because most grasslands have more or less continuous
cover and die back for part of the year, they carry fire read-
ily. Where woody plants and grasses coexist, fires carried in
grasses can top kill or otherwise injure trees and shrubs. As
a result, fire can push back woody plants and increase the
dominance of grasses. Though such patterns are natural,
humans can increase the frequency with which fires occur. It
is thought that burning of grasslands by Native Americans,
for example, may have increased savanna vegetation.
(b)
(b) A June day on a tallgrass prairie in North Dakota. The cottonwood
grove by the prairie creek is characteristic of this biome.

32–16 Glade Coreopsis dominating a glade in temperate deciduous
forest near St. Louis, Missouri. Such openings in the forest, which usually
occur in areas of shallow soil, are often dominated by prairie plants that
form continuous stands beyond the borders of the forest.
When disturbed, grasslands have often changed into
either forests or deserts. Thus, the grasslands that once
stretched from southern Arizona to western Texas, which
were encountered by early European settlers, were largely
overgrazed and converted to deserts or shrublands during
the past century. The failure of farmers and ranchers to
manage grasslands properly led to the “dust bowl” disasters
of the central United States in the 1930s (Figure 32–17).
All of the great grasslands of the world were once
inhabited by herds of grazing mammals associated with
large predators. Such herds were widespread in the periods
of maximum expansion of the glaciers during the Pleistocene
Temperate Deciduous Forests 32–17
epoch, when they were widely hunted by our ancestors.
Many of these grazing mammals, such as the American
bison, have since been hunted almost to extinction. They
survive today mainly in refuges, having given way to herds
of domestic animals and cultivated fields.
Temperate Deciduous Forests
As we move poleward from the tropics, seasonality in-
creases, and eventually, at about 35 degrees latitude, we
encounter climates having long periods with temperatures
below freezing. If rainfall is well distributed through the
growing season, plant growth can flourish, and the tree
form, which can compete successfully for sunlight, is fa-
vored for the same reasons that it is in the rainforests. The
kinds of trees and the structure and function of the forests
are, however, quite different. On fertile soils in these re-
gions, the dominant trees lose their leaves at the end of the
growing season and remain leafless through the winter.
From an energetic and physical standpoint, this pattern can
be explained by the fact that it requires less energy and
involves fewer risks to produce a new leaf than to try to
maintain an evergreen leaf through the rigors of the cold
season. Even if it were possible to sustain photosynthetic
function at a reasonable level, water is unavailable because
the ground is frozen for much of the year, a situation some
describe as physiological drought.
Despite the fact that the climates in these regions vary
between tropical heat and moisture in the summer and Arc-
tic cold in the winter, it is traditional to describe them as
“temperate” and the deciduous forests that are found there
as temperate deciduous forests. Such forests were present
over large areas in the Northern Hemisphere, but primarily
because of the limited land area in the appropriate latitudi-
nal bands they are almost absent in the Southern Hemi-
sphere (Figure 32–2). Temperate deciduous forests are best
developed in areas with warm summers and relatively cool
32–17 Dust bowl The prairie soils were once so
bound together with the roots of grasses that
they could not be cultivated until adequate
plows were developed. But once the plants were
removed by overgrazing or careless cultivation,
prairie soils rapidly deteriorated and were blown
away by the wind. This photograph of a badly
eroded farmyard, taken in Oklahoma in 1937,
vividly recalls the “dust bowl” conditions that led
many thousands of people to migrate away from
the central United States. John Steinbeck’s novel
The Grapes of Wrath was based on the
experiences of these migrants.
32–18 CHAPTER 32 Global Ecology
(a)
winters (Figure 32–18). Annual precipitation generally ranges
from about 75 to 250 centimeters and is either distributed
evenly throughout the year or concentrated somewhat dur-
ing the summer months.
The temperature-driven annual cycle of growth in tem-
perate forests is a salient feature of temperate deciduous
forests (Figure 32–19). In winter, the trees are leafless, and
their metabolic activity is greatly reduced. In spring, a vari-
ety of herbaceous plants burst forth in profusion on the
well-illuminated forest floor (Figure 32–18a). Some species
(the spring ephemerals) have leaves that emerge fully formed
from bulbs or rhizomes and complete their period of activity
in sunlight before the trees leaf out overhead. Others (for
example, early and late summer species, and evergreen spe-
cies) emerge more slowly and sustain photosynthetic activity
longer and later during the shady conditions prevailing in
summer. Summer-active species generally have leaves that
are broader but thinner in cross section than those active
only in spring, and they usually have smaller storage organs
than species with shorter growing periods. Forest herbs
vary in height from a few centimeters to more than one
meter. Variation in leaf height appears to be related to the
typical density of the competing foliage that different
species encounter in their microenvironments. Most of the
species that ripen their seeds in spring are dispersed by ants
(page 469), which are active when few other dispersers are
present. Most of the species that ripen seeds in the fall, how-
ever, are dispersed by birds, at a season that coincides with
the height of fall migration. Very few annual plants occur in
deciduous forests, probably because the dense plant cover in
the moist understory puts seedlings at a strong competitive
disadvantage.
The soils in regions occupied by temperate deciduous
forest are usually acidic, and they tend to contain moderate
amounts of nutrients. For these reasons, such soils are easily
depleted of nutrients after the forests are cleared, and they
may become highly infertile following years of intensive cul-
32–18 Temperate deciduous
forest Representative plants of
the temperate deciduous forests of
North America. (a) A beech and
maple forest in Michigan,
photographed in the spring. The
forest floor is carpeted with large-
flowered trillium (Trillium
grandiflorum). (b) In the fall, as
seen here in a forest in the
southern Appalachian mountains,
the leaves of the maples and
eastern sourwoods (Oxydendrum)
turn a beautiful scarlet.
(b)
tivation. Prairie soils are far more fertile and have character-
istics much more favorable to sustained cultivation.
One of the striking features of the temperate deciduous
forest is the similarity of plants found in each of its three
main regions in the Northern Hemisphere, often represent-
ing related species within the same genus. For example, the
herbaceous plants of the deciduous forests of China and
Japan resemble those of eastern North America more closely
than either group resembles those of western North Amer-
ica. This was less true some millions of years ago, when a
band of deciduous forest existed across North America.
Temperate Mixed and Coniferous Forests
Bordering the deciduous forests on the north (as noted
above, the temperate forest is limited in the Southern Hemi-
sphere) are mixed forests, in which conifers form an impor-
tant element along with the deciduous trees. Such temperate
mixed forests (Figure 32–20) are characteristic of the Great
Lakes–Saint Lawrence River region, much of the southeast-
ern United States, eastern Europe, the northern and eastern
border regions of Manchuria (in northeast China) and ad-
jacent Siberia, eastern Korea, and northern Japan (Figure
32–2). They can be considered to be the intermediate condi-
tion between temperate deciduous forests to the south and
the taiga to the north. Temperate mixed forests occur in
areas with colder winters and more reliable snow cover than
are found in areas of temperate deciduous forest. The conif-
erous component of deciduous or subtropical broadleaf
forest also increases in areas with nutrient-poor soils or sea-
sonally saturated soils to form a variant of mixed forest that
occurs closer to the equator (Figure 32–10).
The absence of extensive deciduous and mixed temper-
ate forests in western North America (excluding Mexico) is
striking. Study of the fossil record shows, however, that the
relative absence of deciduous and mixed forest genera and
species is a “recent” phenomenon—in the earlier Cenozoic
era, these genera were abundant. Most of the deciduous

32–19 Annual growth cycle The four seasons in a temperate
deciduous forest in Illinois.The trees leaf out early in spring and begin to
manufacture food; they lose their leaves in autumn and enter an
essentially dormant state, in which they pass the unfavorable growing
trees and associated herbs were eliminated in western North
America during the latter half of the Cenozoic era as the
amount of summer rainfall was greatly reduced. In their
place now grow the mountain coniferous forests and mixed
west-coast forests of western North America, which contain
such trees as the coast redwood (Sequoia sempervirens; Fig-
ure 32–21), the big tree or giant sequoia (Sequoiadendron
giganteum), the Douglas fir (Pseudotsuga menziesii), and the
sugar pine (Pinus lambertiana; Figure 31–13b), all of which
Temperate Mixed and Coniferous Forests 32–19
conditions of winter. Many herbs grow under the trees (see Figure 32–18),
and a number of them flower very early in spring, before the tree leaves
reach full size and shade the forest floor. In spring, most of the trees
produce abundant pollen, which is carried by the wind.
were much more widely distributed in the past. Similar
kinds of vegetation are found in areas with similar climatic
characteristics in Scandinavia, central Europe, the Pyrenees,
the Caucasus, the Urals, southern Tibet, and the Himalayas
northward to eastern Siberia. Vegetation types that resemble
these are also found in areas in western South America, cen-
tral New Guinea, southwestern New Zealand, the southern
Arabian peninsula, Ethiopia, and the mountains of Central
Africa (Figure 32–2). At higher elevations in these regions
32–20 CHAPTER 32 Global Ecology
32–20 Temperate mixed forest In this forest in southern Ontario,
the evergreen conifers appear dark green and the deciduous trees are
showing their fall colors.
32–21 Mixed west-coast forest Redwoods (Sequoia
sempervirens) are a prominent feature of the mixed west-coast
forests of California. Watered by the frequent fogs of the region
during the dry summers and protected from freezing temperatures by
their proximity to the ocean, redwoods often form spectacular groves,
many of which, like the one shown here in Muir Woods near San
Francisco, are now protected in parks and reserves.
32–22 Alpine tundra As seen here on the Olympic Peninsula in
Washington State, alpine tundra is comparable in many respects to the
Arctic tundra found hundreds of miles to the north. In this area,
however, forested slopes are found within a hundred meters or so of the
alpine meadows.
are found the open grassland communities called alpine tun-
dra (Figure 32–22), which in North America are intermixed
with mountain forests from the Brooks Range in southern
Alaska southward into the Rocky Mountains, the Cascades,
and the Sierra Nevada.
Mediterranean Scrub
Highly distinctive scrub communities have evolved from
mixed deciduous-evergreen forests in areas with Mediter-
ranean climates, which are characterized by cool, moist win-
ters and hot, dry summers (Figure 32–23). Such climates are
found along the shores of the Mediterranean Sea, over a
large part of California (and for a short distance to the
north and south of that state), in central Chile, on the south-
ern coast of Africa, and along portions of the coast of south-
ern and southwestern Australia (Figure 32–2). The plants in
these areas—often evergreen or summer-deciduous trees and
shrubs—have relatively short growing seasons that are
restricted to the cool part of the year, when moisture is rela-
tively abundant. They may lock up nutrients efficiently in
their evergreen leaves. In Mediterranean climates, the luxuri-
ant growth of spring is followed by drought and dormancy
during the summer. Shrublands cover vast areas in all the
Mediterranean climate regions. These shrublands have a
similar appearance when viewed at a distance, but each
region has its own unique species, and therefore deserves its
own name. In California and elsewhere in western North
America, chaparral is the word for these mostly evergreen

and typically very dense shrublands. The equivalent vegeta-
tion around the Mediterranean Sea is called maquis (Figure
32–1), in Chile it is known as matorral, in South Africa, fyn-
bos, and in Australia, mallee (kwongan is used in western
Australia).
The prolonged summer drought and the dense growth
make the chaparral-type shrublands very susceptible to fire,
and fire is a prominent ecological factor in Mediterranean-
type vegetation, both at present and in the past before the
appearance of humans. But what was once a “natural dis-
turbance” is today often seen as a catastrophe. The main
problem is the intrusion of cities and towns into areas of
highly flammable natural vegetation. This problem is partic-
ularly severe in southern California, where homes extend
far up into the chaparral and the interval between fires is
long. Turnover in homeownership is high, and while a dis-
astrous fire will stimulate much discussion at the time, the
talk slowly ebbs, the matter remains unresolved, and a new
generation of homeowners experiences the next major con-
flagration.
As is true of the deserts of the world, each area of
Mediterranean climate is geographically isolated, and each
has its own distinctive assemblage of plants and animals.
The degree of ecological convergence is high, however. Sea-
sonal drought enhances the importance of edaphic (soil-
related) and biotic variation, and small differences in
precipitation often have profound effects on the vegetation
and animal life present in the area. Hence, these areas often
have high proportions of extremely local species of plants
and animals, many of them now in great danger of extinc-
tion. In their modern form, these areas have already been
profoundly changed by people. Much of their vegetation is
now in a very different condition from that occurring before
people occupied the areas with their grazing animals—for
The Northernmost Forests—Taiga and Boreal Forest 32–21
32–23 Mediterranean scrub Chaparral in the
mountains near Los Angeles, California. This sort of
plant community, consisting of broad-leaved,
drought-resistant, and often spiny evergreen shrubs,
occurs only in limited areas of the world, but it has
evolved independently on five continents in areas
having Mediterranean (summer-dry) climates. (See
also Figure 32–1.)
example, today’s areas might be inhabited by more shrubs
and fewer trees than formerly, or by more spiny and poison-
ous plants.
The Northernmost Forests—
Taiga and Boreal Forest
Moving poleward from the mixed coniferous forest, the sea-
sonal contrast becomes more pronounced, with very short
and extremely cold days in winter and a short growing sea-
son of long days. In these conditions, trees still dominate,
because during the relatively brief summer, growth condi-
tions are favorable, and evergreen conifers are abundant.
But the shortness of the growing season makes the energetics
of the deciduous habit less favorable. It is evidently more
efficient, during the short season, to retain an evergreen nee-
dle that can begin photosynthesis as soon as conditions
allow rather than lose valuable time growing leaves from
buds. Deciduous trees do occur in the most northern forests,
but they often are restricted to particular situations, such as
the margins of streams where frost does not penetrate so
deeply into the ground. And a deciduous conifer, larch
(Larix spp.), is abundant in some taiga areas. These north-
ern coniferous forests are often referred to by the Russian
name taiga; in North America they are also called the boreal
forest. In both cases, these forests are characterized by a per-
sistent cover of snow in the winter. In the southern reaches
of the taiga, the trees are taller and more luxuriant, often
reaching 75 meters or more in height (Figure 32–24). In its
main, northern area, however, the trees are shorter, and
thousands of square kilometers are covered by this uniform
forest, with relatively few species of plants and animals (Fig-
ure 32–25). Taiga extends over much of Russia, Scandi-
navia, and northern North America (Figure 32–2).
32–22 CHAPTER 32 Global Ecology
32–24 Pacific coast rainforest Douglas firs (Pseudotsuga
menziesii) growing on the Olympic Peninsula in Washington State.
Epiphytic mosses, liverworts, Selaginella, and lichens often grow
luxuriantly on the trees.
Taiga occurs in the interior of large continental masses
at the appropriate latitudes. In such regions, extreme tem-
peratures range from –50° to 35°C. Taiga is flanked on the
south by montane forests (as in western North America),
deciduous forests, savannas, or grassland, depending on the
amount of precipitation in the region. Because continental
masses do not occur at the appropriate latitudes in the
Southern Hemisphere, taiga is absent there. Due to the influ-
ence of the prevailing westerlies blowing over relatively
warm ocean currents between 40° and 50° north latitude,
the western portions of North America and Eurasia are
characterized by milder climates than their eastern portions.
Consequently, taiga is found somewhat farther north toward
the Pacific coast than it is along the Atlantic coast in North
America, and the same is true of the individual distributions
of many kinds of plants and animals that inhabit the biome.
The northern limits of taiga are ultimately determined by the
severity of the Arctic climate, reaching a limit where the
maximum monthly temperature is approximately 10°C.
In the extensive northern reaches of the taiga, most of
the precipitation falls in the summer; the cold winter air in
these regions has a very low moisture content. The annual
total precipitation usually amounts to less than 30 centime-
ters. The rate of evaporation is low, however, and lakes,
bogs, and marshes are common (Figure 32–25). More than
three-quarters of the northern reaches of the taiga is under-
lain by permanent ice, or permafrost, usually within less than
a meter of the surface; the permafrost tends to trap surface
water and form lakes. Fires are common in the taiga, and
they result in generally warmer, more productive sites for at
least 10 to 20 years afterward, due to the local melting of the
permafrost. In general, taiga soils are highly acidic, very low
in nutrients, and poorly suited for agriculture.
Species of a few genera of trees are common in the
northern taiga, including spruce (Picea), larch (Larix), fir
(Abies), and poplar (Populus). Among the more common
shrubs are dewberries (Rubus), Labrador tea (Rhododen-
dron), willows (Salix), birches (Betula), and alders (Alnus).
Pines (Pinus) can be common, usually in drier areas. The
members of all these genera of trees and shrubs are ectomy-
corrhizal (Figure 29–1), and they occur in dense stands con-
32–25 Northern taiga The northern taiga,
which covers hundreds of thousands of
square kilometers in the cooler part of the
north temperate zone, is dominated by white
spruce (Picea glauca) and larch (Larix). This
photograph was taken in northern Manitoba,
Canada. In the more northern part of their
range, the trees are smaller than seen here.

sisting of only one or a very few species. Perennial herbs are
common, and mosses and lichens are especially prevalent,
often forming luxuriant masses. Annual plants, except in
areas where there is human disturbance, are essentially
absent.
At its northern limits, taiga grades unevenly into tundra.
In both these biomes, the days are long during the relatively
short growing season; north of the Arctic Circle, the sun
does not set during at least part of the summer. Because of
the seasonally abundant light and favorable temperatures,
cool-season cultivated plants, such as cabbages (Brassica
oleracea var. capitata), may grow rapidly in cleared areas in
the taiga, attaining large sizes in a remarkably short period
of time. Yet the infertile, highly leached soils of the taiga do
not allow most forms of agriculture.
Coniferous forests extend far to the south of taiga, along
the Pacific coast of North America. These magnificent ever-
green coniferous forests (Figure 32–2) occur where there is a
pronounced summer drought, but high and persistent rainfall
during the cooler seasons. Because photosynthesis is limited
by lack of moisture during the warm season, deciduous trees
are at a disadvantage and are usually found only along
stream banks. The evergreen conifers, however, can synthe-
size carbohydrates all year round and, because of their mas-
sive size, can store water and nutrients for use during the dry
season. Their thick barks and high crowns protect them from
all but the most intense fires characteristic of the region.
Arctic Tundra
The Arctic tundra is a treeless biome that extends to the far-
thest northern limits of plant growth (Figure 32–26). It
occupies an enormous area: fully one-fifth of the Earth’s
(a)
32–26 Arctic tundra (a) Arctic wet coastal tundra near Prudhoe
Bay, Alaska, in late summer. The reddish brown plants are the grass
Arctophila fulva, the green ones are the sedge Carex aquatilis. Note
that the water table is above the surface because of the presence of
permafrost; such conditions are characteristic of the Arctic tundra.
Arctic Tundra 32–23
land surface (Figure 32–2). The majority of tundra can be
found in the Arctic, mostly above the Arctic Circle, although
it extends farther south along the eastern sides of the conti-
nents than along the western sides. The Arctic tundra essen-
tially constitutes one huge band across Eurasia and North
America, with alpine tundra, more closely related to the
adjacent mountain forests, extending southward in the
mountains (Figure 32–22; see also Figure 32–5). Some
species of plants that occur in the Arctic tundra have wide
circumpolar ranges.
Toward the poles, the winter season lengthens relative
to that of summer. Although daylength in summer increases
—north of the Arctic Circle the sun does not set for days—
this does not compensate for the long period of little or no
sun. This portion of the Earth operates at a strong solar
energy deficit, radiating more energy back to space than is
received in direct sunlight. Frosts can occur throughout the
year, and winters are bitterly cold, with high winds that will
desiccate any plant that rises above the snow level. At low
temperatures, blowing snow acts as an abrasive that will
ruthlessly attack any exposed tissues.
The limited supply of solar energy is insufficient to thaw
more than a shallow layer of soil. The result is permafrost—
permanently frozen soil. The annual freezing and thawing of
the surface has many effects unknown in milder climates,
such as landscapes broken into polygonal blocks ranging
from 3 to 30 meters across, and small hills, or pingos, up to
50 meters high, formed with a core of ice. The soils are
acidic to neutral, low in nutrients, and generally poor for
agriculture. Even though precipitation is usually less than
25 centimeters per year, much of it is held near the surface
by underlying permafrost, and the ground is usually wet.
(b)
(b) Arctic tundra at Barrow, Alaska. A clone of the sedge Eriophrum
angustifolium is growing out into a solid patch of another species of
the same genus, Eriophrum scheuchzeri. Vegetative reproduction,
like that in E. angustifolium illustrated here, is characteristic of many
tundra plants.
32–24 CHAPTER 32 Global Ecology
Fixed nitrogen is generally in very short supply, and there
are only a few species of legumes and other plants with sym-
biotic bacteria that fix atmospheric nitrogen. The evapora-
tion rate is low because of the relatively high moisture
content of the air and low temperatures. In contrast, some
tundra areas are so dry as to constitute true polar deserts.
Most of the land north of 75° north latitude is a desert or
semidesert, with few plants taller than 5 centimeters. This is
a result of the very low precipitation in both summer and
winter and the very cold winter temperatures.
For plant growth to occur at all, the mean temperature
must be above freezing for at least one month of the year.
The growing season (the time from one killing frost to the
next) in many areas of Arctic tundra is less than two months.
Several genera of low shrubs, including birch (Betula), willow
(Salix), blueberry (Vaccinium), and Labrador tea (Rhododen-
dron), are common in the tundra, and there are a number of
genera of perennial herbs but only one native annual species
(Koenigia islandica). Many of the plants that grow in the
Arctic tundra, including a number of the grasses and sedges,
are evergreen, thus able to initiate photosynthesis soon after
adequate light, moisture, and temperature become available.
Plant height is controlled primarily by snow depth in winter;
large woody plants are absent because of their relatively high
energy allocation to unproductive stem tissue, which cannot
be supported given the short, cool growing season. A number
of tundra plants have relatively large, showy flowers, the pro-
duction of which costs the plants considerable quantities of
energy. Such flowers hold energy-rich rewards for their polli-
nators—necessary rewards, given the low temperatures that
prevail at high latitudes. Vegetative propagation is character-
istic of many of the perennials, and this may be correlated
with the uncertainties of setting seed during the brief Arctic
summer. Much of the biomass of tundra plants—from half
to as much as 98 percent—is underground, consisting not
only of roots but also of rhizomes and other kinds of under-
ground stems.
North of the Arctic tundra is ice desert, where physical
conditions are even more extreme and vegetation is absent. Ice
desert is characteristic of the interior of Greenland; Svalbard,
a small group of islands off Norway; and Novaya Zemlya,
two islands off the north coast of Siberia. Much of Antarctica,
not mapped in Figure 32–2, is also covered by ice.
SUMMARY
Biomes Are Terrestrial Ecosystems Characterized by
Distinctive Vegetation
The distribution of biomes is a result of complex interactions
among the distribution of heat from the sun, air-circulation
patterns, and geologic features. These factors cause wide dif-
ferences in temperature and precipitation from place to place
and season to season. In addition to climate, differences in
the surfaces of the continents, such as soil composition and
altitude, affect the kinds of plant and animal life found in the
various biomes on Earth.
Tropical Rainforests Have a Great Diversity
of Species, with Few Individuals per Species
Tropical rainforest, in which neither water nor low tempera-
ture is a limiting factor, is by far the richest biome in terms of
number of species. The trees are evergreen and characterized
by medium-sized leathery leaves. A poorly developed layer of
herbs grows on the forest floor, but there are many vines and
epiphytes at higher levels. Tropical soils are often acidic and
very poor in nutrients; such soils lose their fertility rapidly
when the forest is cleared.
Savannas and Deciduous Tropical Forests Occur
Where Rainfall Is Seasonal
Mostly tropical and subtropical communities that are charac-
terized by a seasonal drought are termed savannas, subtropi-
cal mixed forests, monsoon forests, tropical mixed forests,
and southern woodland and scrub. The trees and shrubs of
these communities are wholly or partly deciduous, shedding
their leaves during times of drought. Herbaceous perennials
are common. Savannas also occur between the prairies and
temperate deciduous forests and between the prairies and the
taiga in North America. Subtropical mixed forests cover
most of Florida and the Coastal Plain of the southeastern
United States. In these forests, pines and other evergreen
trees grow intermixed with deciduous trees.
Desert Plants Are Adapted to Low Precipitation
and Extremes of Temperature
Away from the equator, tropical and subtropical communi-
ties grade into deserts and semideserts, which are character-
ized by low precipitation and often by high daytime
temperatures during at least part of the year. Succulent plants
and annual herbs are common in deserts.
Grasslands Occur Where Precipitation and Fire
Interact to Support Grasses but Not Trees
Grasslands, which intergrade with savannas, deserts, and
temperate forests, are characterized by a general lack of trees,
except along streams. The most productive soils for temper-
ate agriculture are grassland soils.
Temperate Deciduous Forests Are Made up of Leaf-
Shedding Trees and Many Types of Perennial Herbs
In the temperate deciduous forests, most of the trees lose
their leaves during the cold (usually snowy) winters, when
moisture may be unavailable for growth. Many genera are
common to the temperate deciduous forests of eastern North
America and eastern Asia. Temperate deciduous forests are
bordered by temperate mixed and coniferous forests to the
north, in which conifers play an important role.
Mediterranean Scrub Is Characterized by Evergreen,
Drought-Resistant Shrubs or Trees That Form
Thickets
Distinctive scrub communities, called chaparral in North
America and maquis in the Mediterranean region, have
evolved in the five widely separated areas of the world with a

SUMMARY TABLE Some Characteristics of the Earth’s Principal Biomes
BIOME T E M P E R AT U R E A N D P R E C I P I TAT I O N
Rainforests High temperature and high rainfall year round.
Savannas and High temperature and seasonal drought.
deciduous
tropical
forests
Deserts Precipitation generally very low except for
occasional peaks; maximum temperature
varies with the type of desert.
Grasslands Moderately low precipitation; cold winters
and warm summers.
Temperate Moderate precipitation evenly distributed;
deciduous cool winters and warm summers.
forests
Temperate Moderately low precipitation and moderately
mixed and cold winters.
coniferous
forests
Mediterranean Cool, moist winters and hot, dry summers.
scrub
Taiga Moderately low precipitation and cold winters,
although in the Pacific Northwest the winters
are very wet.
Arctic tundra Very low precipitation in both summer and
winter; very cold winters.
Mediterranean climate—a dry summer and a cool, rainy
winter growing season. Such communities occur in western
North and South America, around the Mediterranean, in the
Cape Region of South Africa, and in southwestern Australia.
The Taiga Is Characterized by Forests of Evergreen
Trees
The taiga is a vast northern coniferous forest that extends in
unbroken bands across Eurasia and North America and
down the Pacific coast to northern California. In its southern
reaches, the taiga is dominated by tall trees with a lush
growth of bryophytes and lichens; northward, it consists of
vast monotonous stretches of forest with very few tree
species.
The Arctic Tundra Has Low-Lying Shrubs and Grasses
but No Trees
North of the taiga is the tundra, a treeless region that also
extends around the Northern Hemisphere, mostly above the
Arctic Circle, in a band that is broken only by bodies of
water. Both the northern reaches of the taiga and all of the
tundra are underlain by permafrost. Because of this perma-
frost, and especially because of the low rates of evapotranspi-
Questions 32–25
CHARACTERISTIC PLANTS M I S C E L L A N E O U S F E AT U R E S
Broad-leaved evergreen trees, epiphytes, and The biome with the greatest diversity of species.
lianas. Infertile soils.
Grasslands with scattered broad-leaved Periodic burning is an important aspect.
deciduous or evergreen trees or shrubs.
Succulents such as cacti; annual herbs. Adaptations include small leaves, thick cuticles,
and photosynthetic rates with high maximum
temperatures.
Perennial bunchgrasses and sod-forming Heavily exploited for agriculture.
grasses.
Deciduous trees and many perennial herbs. The dominant herbaceous plants vary with the
seasons.
Mixtures of deciduous trees and conifers. Occur as a transition zone north of the deciduous
forest. Also found in areas with nutrient-poor soils
or with less seasonal environments.
Evergreen or summer-deciduous, drought- Called chaparral in California and maquis around
resistant trees and shrubs in dense thickets. the Mediterranean Sea.
Forest of evergreen trees. Soils are highly acidic and very low in nutrients.
Permafrost may be present.
Low shrubs, grasses, sedges, and lichens. Permafrost present throughout. Much of the
biomass is underground.
ration, tundra and taiga soils are relatively moist and highly
leached of nutrients.
QUESTIONS
1. Describe the influence of latitude and altitude on the
distribution of organisms on Earth.
2. Describe the effect of mountains on local precipitation.
3. Compare tropical and temperate forests in terms of the
numbers of species found in each, and in the size and
appearance of the trees.
4. Explain why annual plants are better represented, both in
number and in kind, in the deserts and semiarid regions of
the world than anywhere else.
5. Compare the relative amounts of nutrients found in forest
and grassland soils.
6. How are the evergreen conifers of the Pacific Northwest of
the United States and Canada adapted to the winter-wet/
summer-dry environment of that region?
7. What are the principal differences between taiga and
tundra? What role does permafrost play in these biomes?