55
Ecology and the Geography of Life
K EY C ONCE P TS
Climate, particularly temperature and precipitation, affects
the distribution of Earths major biomes, such as tropical
rain forests and tundra.
Abiotic factorssuch as water salinity, amount of dis-
solved oxygen, availability of essential minerals, and water
depthinuence the distribution of organisms in aquatic
ecosystems.
Species richness varies widely among different biomes and
aquatic ecosystems.
Ecotonesareas of transition where two communities meet
and intergradeprovide diverse conditions that encourage
species richness.
Earth has six biogeographic realms, each consisting of a
major landmass separated by the ocean, mountains, or a
desert.
Black-tailed prairie dog. Prairie
dogs (Cynomys ludovicianus)
never wander far from their bur-
rows, which they use to escape
from predators.
Raymond Gehman /Corbis
E arth has many different environments. Natural selection affects
an organisms ability to survive and reproduce in a given en-
vironment. In natural selection, both abiotic (nonliving) and biotic
(living) factors eliminate the least-t individuals in a population.
Over time, succeeding generations of organisms that live in each
biome or major aquatic ecosystem become better adapted to local
environmental conditions.
Black-tailed prairie dogs are superbly adapted to their envi-
ronment. Their teeth and digestive tracts are modied to eat and
easily digest the seeds and leaves of grasses that grow in great
profusion on the Great Plains of western North America.
Prairie dogs live in large colonies of about 500 individuals. The
eyes of every individual in the colony watch for potential danger,
and when they see it, prairie dogs call out to warn the rest of the
colony (see photograph). When danger approaches, each prairie
dog dives into its underground home. Each burrow has at least two
openings and consists of an elaborate network of long tunnels with
several chambers: a nursery for the young, a sleeping chamber,
1
1189
 a toilet chamber, and a listening chamber (close to an entrance).
Piles of excavated soil surround the burrow entrances and help
prevent ooding during rainstorms.
To survive winter, black-tailed prairie dogs hibernate in their
burrows. Their metabolism slows, and they subsist on the stored fat
in their bodies. They do not hibernate deeply, however, and may
leave their burrows to look for food when the weather warms.
As we have seen for prairie dogs, structural, behavioral, and
physiological adaptations for its own particular environment and
lifestyle have evolved in each species. As we examine Earths major
terrestrial and aquatic ecosystems, including the species charac-
teristic of each, think about the variety of adaptations that natural
selection has produced in organisms in response to their particular
environments.

BIOMES
Learning Objectives Key Point
1 Dene biome, and briey describe the nine major terrestrial
biomes, giving attention to the climate, soil, and characteris- The distribution of the worlds biomes is largely the result of
tic plants and animals of each. climate patterns, which determine an areas water (measured
2 Describe at least one human effect on each of the biomes as precipitation) and energy (measured as temperature).
discussed.

A biome is a large, relatively distinct terrestrial region that has

similar climate, soil, plants, and animals regardless of where it
occurs. Because it covers such a large geographic area, a biome 400
encompasses many interacting landscapes. Recall from Chapter
52 that a landscape is a large land area (several to many square

Average annual precipitation (cm)

Tropical
kilometers) composed of interacting ecosystems. 300 rain
Biomes largely correspond to major climate zones, with tem- Temperate forest
rain
perature and precipitation being most important ( Fig. 55-1). forest
Near the poles, temperature is generally the overriding climate Tropical
200 dry
factor; whereas in tropical and temperate regions, precipitation Temperate forest
becomes more signicant than temperature ( Fig. 55-2). Other deciduous
abiotic factors to which biomes are sensitive include temperature forest
extremes, rapid temperature changes, oods, droughts, strong 100
Boreal Temperate Savanna
winds, and res (see section on res in Chapter 54). forest grassland,
We discuss nine major biomes in this chapter: tundra, boreal chaparral
Tundra Desert
forest, temperate rain forest, temperate deciduous forest, temper- 0
ate grassland, chaparral, desert, savanna, and tropical rain forest 10 0 10 20 30
( Fig. 55-3). Although we discuss each biome as a distinct entity, Average temperature (C)
biomes intergrade into one another at their boundaries.

Figure 55-1 Using precipitation and temperature

Tundra is the cold, boggy plains
of the far north
Tundra (also called arctic tundra) occurs in extreme northern
latitudes wherever snow melts seasonally ( Fig. 55-4). The South-
ern Hemisphere has no equivalent of the arctic tundra because it
has no land in the proper latitudes. A similar ecosystem located
in the higher elevations of mountains, above the tree line, is called
alpine tundra to distinguish it from arctic tundra (see Focus
On: The Distribution of Vegetation on Mountains on page 1195).
Arctic tundra has long, harsh winters and extremely short
summers. Although the growing season, with its warmer tem-
peratures, is as short as 50 days, the days are long. Above the
Arctic Circle the sun does not set at all for many days in midsum-
mer, although the amount of light at midnight is only one tenth
that at noon. There is little precipitation (10 to 25 cm, or 4 to
to identify biomes
Factors such as soil type, re, and seasonality of climate affect whether
temperate grassland or chaparral develops. (Adapted from R. H. Whit-
taker, Communities and Ecosystems, 2nd ed., Macmillan, New York,
1975.)
10 in, per year) over much of the tundra, and most of it falls dur-
ing summer months.
Tundra soils tend to be geologically young, because most were
formed only after the last Ice Age.1 These soils are usually nutrient
poor and have little organic litter (dead leaves and stems, animal
1
Glacier ice, which occupied about 30% of Earths land during the last
Ice Age, began retreating about 17,000 years ago. Today, glacier ice oc-
cupies about 10% of the land surface.

0
1190 Chapter 55 www.thomsonedu.com/biology/solomon
 28
24

Average monthly
temperature (C)
20
16
12
8
4
0
-4
14

precipitation (cm)
Average monthly
12
10
8
6
4
2

J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D
Month

(a) Temperate deciduous forest (b) Temperate grassland (c) Temperate desert
(Nashville, Tennessee) (Lawrence, Kansas) (Reno, Nevada)

Figure 55-2 Signicance of precipitation in temperate biomes

Average monthly temperature (top) is approximately the same in each plentiful; (b) grassland, where it is less plentiful and more seasonal; and
location in North America. However, precipitation (bottom) varies a (c) desert, where it is quite low.
great deal, resulting in (a) deciduous forest, where precipitation is

droppings, and remains of organisms) in the uppermost layer of
soil. Although the soil surface melts during the summer, tundra
has a layer of permanently frozen ground called permafrost that
varies in depth and thickness. Because permafrost interferes with
drainage, the thawed upper zone of soil is usually waterlogged
during the summer. Limited precipitation, combined with low
temperatures, at topography (surface features), and permafrost,
produces a landscape of broad, shallow lakes, sluggish streams,
and bogs.
Low species richness and low primary productivity charac-
terize tundra. Mosses, lichens (such as reindeer moss), grasses,
and grasslike sedges dominate tundra vegetation; most of these
short plants are herbaceous perennials that live 20 to 100 years.
No readily recognizable trees or shrubs grow except in sheltered
locations, although dwarf willows, dwarf birches, and other dwarf
trees are common.
Year-round animal life of the tundra includes voles, weasels,
arctic foxes, gray wolves, snowshoe hares, ptarmigan, snowy
owls, musk oxen, and lemmings (see the discussion of lemming
population cycles in Chapter 52). In the summer, caribou mi-
grate north to the tundra to graze on sedges, grasses, and dwarf
willow. Dozens of bird species also migrate north in summer to
nest and feed on abundant insects. Mosquitoes, blackies, and
deeries survive the winter as eggs or pupae and occur in great
numbers during summer weeks.
Tundra regenerates quite slowly after it has been disturbed.
Even casual use by hikers causes damage. Long-lasting injury,
likely to persist for hundreds of years, was done to large portions
of the arctic tundra as a result of oil exploration and military use.
Boreal forest is the evergreen
forest of the north
Just south of the tundra is the boreal forest, or taiga, which
stretches across both North America and Eurasia. Boreal forest is
the worlds largest biome, covering approximately 11% of Earths
land ( Fig. 55-5). A biome comparable to the boreal forest is not
found in the Southern Hemisphere, because it has no land at the
corresponding latitudes. Winters are extremely cold and severe,
although not as harsh as in the tundra. Boreal forest receives little
precipitation, perhaps 50 cm (20 in) per year, and its soil is typi-
cally acidic, is low in minerals (inorganic nutrients), and has a
deep layer of partly decomposed conifer needles at the surface.
(Conifers are cone-bearing evergreens.) Boreal forest contains
numerous ponds and lakes in water-lled depressions that grind-
ing ice sheets dug during the last Ice Age.
Black and white spruces, balsam r, eastern larch, and other
conifers dominate the boreal forest, but deciduous trees such as
aspen or birch, which shed their leaves in autumn, form striking
stands. Conifers have many drought-resistant adaptations, such
as needlelike leaves with a minimal surface area to reduce wa-
ter loss (see Chapter 33). Such an adaptation enables conifers to
withstand the drought of the northern winter months, when

1
Ecology and the Geography of Life 1191
 Equator

Tundra Chaparral

Boreal forest Mountains with complex zonation

Temperate deciduous forest and Savanna

temperate coniferous forest
(includes temperate rain forest) Tropical dry forest

Temperate grassland Tropical rain forest

Desert

Figure 55-3 Animated The worlds major biomes

This simplied diagram shows sharp boundaries between biomes. tains are keyed according to their predominant vegetation type,
Biomes actually intergrade at their boundaries, sometimes over large whereas mountains with variable vegetation are keyed as Mountains
areas. Note that forested mountains such as the Appalachian Moun- with complex zonation. (Based on data from the World Wildlife Fund.)

roots do not absorb water because the ground is frozen. Natural
selection also favors conifers in the boreal forest because, being
evergreen, they resume photosynthesis as soon as warmer tem-
peratures return.
Animal life of the boreal forest includes some larger species,
such as caribou (which migrate from the tundra to the boreal for-
est for winter), wolves, bears, and moose. However, most animal
life is medium-sized to small and includes rodents, rabbits, and
fur-bearing predators such as lynx, sable, and mink. Most species
of birds are seasonally abundant but migrate to warmer climates
for winter. Insects are numerous, but there are few amphibians
and reptiles except in the southern boreal forest.
Most of the boreal forest is not suitable for agriculture be-
cause of its short growing season and mineral-poor soil. The
boreal forest, which is harvested primarily by clear-cutting, is
currently the primary source of the worlds industrial wood and
wood ber. The annual loss of boreal forests in Canada, Siberia,
and Alaska is estimated to encompass an area twice as large as the
Amazonian rain forests of Brazil.
Temperate rain forest has cool weather,
dense fog, and high precipitation
Coniferous temperate rain forest grows on the northwestern
coast of North America. Similar vegetation exists in southeastern
Australia and in southwestern South America. Annual precipita-
tion in this biome is high, from 200 to 380 cm (80 to 150 in);

2
1192 Chapter 55 www.thomsonedu.com/biology/solomon
Eastcott /Momatiuk /Earth Scenes

Beth Davidow/ Visuals Unlimited

Figure 55-4 Arctic tundra Figure 55-5 Boreal forest
Because of the short growing season and permafrost in arctic tundra, Boreal forest is coniferous forest that occurs in cold regions of the
only small, hardy plants grow in the northernmost biome that encircles Northern Hemisphere adjacent to the tundra. Photographed in Yukon,
the Arctic Ocean. Photographed during autumn in Northwest Territo- Canada.
ries, Canada.

condensation of water from dense coastal fog augments the an-

nual precipitation. The proximity of temperate rain forest to the
coastline moderates the temperature so that seasonal uctuation
is narrow; winters are mild, and summers are cool. Temperate
rain forest has a relatively nutrient-poor soil, although its organic
content may be high. Cool temperatures slow the activity of bac-
terial and fungal decomposers. Thus, needles and large fallen
branches and trunks accumulate on the ground as litter that takes
many years to decay and release inorganic minerals to the soil.
The dominant vegetation type in the North American temper-
ate rain forest is large evergreen trees, such as western hemlock,
Douglas r, Sitka spruce, and western red cedar. Temperate rain
forest is rich in epiphytic vegetation, which consists of smaller
plants that grow nonparasitically on the trunks and branches
of large trees ( Fig. 55-6). Epiphytes in this biome are mainly
mosses, lichens, and ferns, all of which also carpet the ground.
Terry Donnelly/Dembinsky Photo Associates

Squirrels, wood rats, mule deer, elk, numerous bird species (such
as jays, nuthatches, and chickadees), several species of reptiles
(such as painted turtles and western terrestrial garter snakes),
and amphibians (such as Pacic giant salamanders and Pacic
treefrogs) are common temperate rainforest animals.
Temperate rain forest, one of the richest wood producers
in the world, supplies us with lumber and pulpwood. It is also
one of the most complex ecosystems in terms of species richness.
Care must be taken to avoid overharvesting original old-growth
forest, because such an ecosystem takes hundreds of years to de- Figure 55-6 Temperate rain forest
velop. When the logging industry harvests old-growth forest, it Large amounts of precipitation characterize temperate rain forest.
typically replants the area with a monoculture (a single species) Note the epiphytes hanging from the branches of coniferous trees.
of trees that it harvests in 40- to 100-year cycles. Thus, the old- Photographed in Olympic National Park in Washington State.

1
Ecology and the Geography of Life 1193

Barbara Miller/Biological Photo Service
Figure 55-7 Temperate deciduous forest
The broad-leaf trees that dominate the temperate deciduous forest
shed their leaves before winter. Photographed during autumn in
Pennsylvania.
growth forest ecosystem, once harvested, never has a chance to
redevelop. A small fraction of the original, old-growth temper-
ate rain forest in Washington, Oregon, and northern California
remains untouched. These stable forest ecosystems provide bio-
logical habitat for many organisms, including 40 endangered and
threatened species.
Temperate deciduous forest has
a canopy of broad-leaf trees
Seasonality (hot summers and cold winters) is characteristic of
temperate deciduous forest, which occurs in temperate areas
where precipitation ranges from about 75 to 126 cm (30 to 50 in)
annually. Typically, the soil of a temperate deciduous forest con-
sists of both a topsoil rich in organic material and a deep, clay-
rich lower layer. As organic materials decay, mineral ions are re-
leased. If roots of living trees do not absorb these ions, they leach
into the clay, where they may be retained.
Broad-leaf hardwood trees, such as oak, hickory, maple, and
beech, that lose their foliage annually dominate temperate decid-
uous forests of the northeastern and Mid-Atlantic United States
( Fig. 55-7). The trees of the temperate deciduous forest form a
dense canopy that overlies saplings and shrubs.
Temperate deciduous forests originally contained a variety of
large mammals such as mountain lions, wolves, bison, and other
4
1194 Chapter 55
species now regionally extinct, plus deer, bears, and many small
mammals and birds (such as wild turkeys, blue jays, and scar-
let tanagers). Both reptiles (such as box turtles and rat snakes)
and amphibians (such as spotted salamanders and wood frogs)
abounded, together with a denser and more varied insect life than
exists today.
In Europe and North America, logging and land clearing
for farms, tree plantations, and cities have removed much of the
original temperate deciduous forest. Where it has regenerated,
temperate deciduous forest is often in a seminatural state that
is, highly modied by humans for recreation, livestock foraging,
timber harvest, and other uses. Although these returning forests
do not have the biological diversity of virgin stands, many forest
organisms have successfully become re-established.
Worldwide, temperate deciduous forest was among the rst
biomes to be converted to agricultural use. In Europe and Asia,
many soils that originally supported temperate deciduous for-
est have been cultivated by traditional agricultural methods for
thousands of years without substantial loss in fertility. During
the 20th century, however, intensive agricultural practices were
adopted; these, along with overgrazing and deforestation, con-
tributed to the degradation of some agricultural lands.
Temperate grasslands occur in areas
of moderate precipitation
Summers are hot, winters are cold, res help shape the landscape,
and rainfall is often uncertain in temperate grasslands. Annual
precipitation averages 25 to 75 cm (10 to 30 in). In grasslands
with less precipitation, minerals tend to accumulate in a marked
layer just below the topsoil. These minerals tend to leach out of
the soil in areas with more precipitation. Grassland soil contains
considerable organic material because surface parts of many
grasses die off each winter and contribute to the organic content
of the soil (the roots and rhizomes survive underground). Many
grasses are sod formers: their roots and rhizomes form a thick,
continuous underground mat.
Moist temperate grasslands, also known as tallgrass prairies,
occur in the United States in Iowa, western Minnesota, east-
ern Nebraska, and parts of other Midwestern states and across
Canadas prairie provinces. Although few trees grow except near
rivers and streams, grasses, some as tall as 2 m (6.5 ft), grow
in great profusion in the deep, rich soil ( Fig. 55-8). Before
most of this area was converted to arable land, it was covered
with herds of grazing animals, particularly bison. The principal
predators were wolves, although in sparser, drier areas coyotes
took their place. Smaller fauna included prairie dogs and their
predators (foxes, black-footed ferrets, and birds of prey such as
prairie falcons), western meadowlarks, bobolinks, reptiles (such
as gopher snakes and short-horned lizards), and great numbers
of insects.
Shortgrass prairies, in which the dominant grasses are less
than 0.5 m (1.6 ft) tall, are temperate grasslands that receive less
www.thomsonedu.com/biology/solomon
 f o c u s o n

THE DISTRIBUTION OF VEGETATION ON MOUNTAINS

Hiking up a mountain is similar to travel- ELEVATION LATITUDE
ing toward the North Pole with respect ZONES ZONES
to the major life zones encountered (see High elevation North Pole
gure). This elevationlatitude similarity
Snow/ice Polar region
occurs because the temperature drops
as one climbs a mountain, just as it does
when one travels north; the temperature
drops about 6C (11F) with each 1000-m Alpine tundra Arctic tundra
increase in elevation. The types of species
growing on the mountain change as the
temperature changes.
Deciduous trees, which shed their
Subalpine coniferous forest Taiga
leaves every autumn, may cover the base
of a mountain in Colorado, for example.
At higher elevations, where the climate
is colder and more severe, a coniferous
subalpine forest resembling boreal forest
grows. Spruces and rs are the dominant Deciduous forest Temperate forest
trees here. Higher still, the forest thins, Low elevation Midlatitudes
and the trees become smaller, gnarled,
and shrublike. These twisted, shrublike
trees, called krummholz (a German word
meaning crooked wood), are found at
their elevational limit (the tree line). The
exact elevation at which the tree line oc-
curs depends on the latitude and distance
from the ocean. In the Rocky Mountains,
between 35 and 50 north latitude, the
tree line drops 100 m with each 1 degree
latitude northward.
Comparison of elevation and latitude zones.
Above the tree line, where the climate
is quite cold, a kind of tundra occurs, with The cooler temperatures at higher elevations of a mountain produce a series of ecosystems simi-
lar to those encountered when going toward the North Pole.
vegetation composed of grasses, sedges,
and small tufted plants, most of which
are hardy perennials. Some alpine plants found, similar to the nearly lifeless polar have the great extremes of day length that
(for example, buttercups) are lowland land areas. are associated with the changing seasons in
species that have adapted to the alpine Important environmental differences high-latitude biomes. The intensity of solar
environment, whereas other plants (for exist between high elevations and high radiation is greater at high elevations than
example, mountain douglasia) live ex- latitudes that affect the types of organisms at high latitudes. At high elevations, the
clusively in the mountains. This tundra is found in each place. Alpine tundra typi- suns rays pass through less atmosphere,
called alpine tundra to distinguish it from cally lacks permafrost and receives more which results in greater exposure to ultra-
arctic tundra. At the top of the mountain, precipitation than does arctic tundra. High violet radiation (less is ltered out by the
a permanent ice cap or snowcap might be elevations of temperate mountains do not atmosphere) than occurs at high latitudes.

precipitation than the moister grasslands just described but more
precipitation than deserts. In the United States, shortgrass prai-
ries occur in the eastern half of Montana, the western half of
South Dakota, and parts of other Midwestern states, as well as
western Alberta in Canada. The plants grow in less abundance
than in the moister grasslands, and occasionally some bare soil
is exposed.
The North American grassland, particularly the tallgrass
prairie, was so well suited to agriculture that little of it remains.
More than 90% has vanished under the plow, and the remain-
der is so fragmented that almost nowhere can we see even an
approximation of what European settlers saw when they settled
in the Midwest. Today, the tallgrass prairie is considered North
Americas rarest biome.

1
Ecology and the Geography of Life 1195
 Denser, thicker vegetation tends to accumulate when periodic
res are prevented; then, when a re does occur, it is much more
severe. Removing the chaparral vegetation, whose roots hold the
soil in place, also causes problems witness the mud slides that
sometimes occur during winter rains in these areas.

Deserts are arid ecosystems

Deserts are dry areas found in temperate (cold deserts) and sub-
tropical or tropical regions (warm deserts). North America has
four distinct deserts. The Great Basin Desert in Nevada, Utah,
and neighboring states is a cold desert dominated by sagebrush.
Harvey Payne

The Mojave Desert in Nevada and California is a warm desert

known for its Joshua trees. The Chihuahuan Desert, home of
century plants (agaves; see Fig. 52-7), is a warm desert found
Figure 55-8 Temperate grassland in Texas, New Mexico, and Mexico. The warm Sonoran Desert,
The Nature Conservancy owns this tallgrass prairie preserve in Okla- with its many species of cacti, is found in Arizona, California, and
homa. Like other moist temperate grasslands, it is mostly treeless but Mexico ( Fig. 55-10).
contains a profusion of grasses and other herbaceous owering plants.
As bison graze on the plants, they affect community structure and
The low water-vapor content of the desert atmosphere leads
diversity. to daily temperature extremes of heat and cold, so a major change
in temperature occurs in each 24-hour period. Deserts vary greatly
depending on the amount of precipitation they receive, which is
generally less than 25 cm (10 in) per year. A few deserts are so dry
Chaparral is a thicket of evergreen that virtually no plant life occurs in them. As a result of sparse
vegetation, desert soil is low in organic material but often high in
shrubs and small trees
mineral content, particularly the salts NaCl, CaCO3, and CaSO4.
Some hilly temperate environments have mild winters with Desert vegetation includes both perennials (cacti, yuccas,
abundant rainfall, combined with extremely dry summers. Such Joshua trees, and sagebrushes) and, after a rain, owering an-
Mediterranean climates, as they are called, occur not only in the nuals. Desert plants tend to have reduced leaves or no leaves,
area around the Mediterranean Sea but also in California, West- an adaptation that conserves water. In cacti such as the giant
ern Australia, portions of Chile, and South Africa. In southern saguaro, for example, the stem carries out photosynthesis and
California this environment is called chaparral. This vegetation also expands accordion-style to store water; the leaves are modi-
type is also known as maquis in the Mediterranean region, mallee
scrub in Australia, matorral in Chile, and Cape scrub in Africa.
Chaparral soil is thin and infertile. Frequent res occur naturally
in this environment, particularly in late summer and autumn.
Chaparral vegetation looks strikingly similar in different
areas of the world, even though the individual species are quite
different. A dense growth of evergreen shrubs, often of drought-
resistant pine or scrub oak trees, dominates chaparral ( Fig. 55-9).
During the rainy winter season the landscape may be lush and
green, but during the hot, dry summer the plants lie dormant.
Trees and shrubs often have hard, small, leathery leaves that re-
Edward Ely/Biological Photo Service

sist water loss. Many plants are also re-adapted and grow best
in the months following a re. Such growth is possible because
re releases minerals that were tied up in the plants that burned.
With the new availability of essential minerals, plants sprout vig-
orously during winter rains. Mule deer, wood rats, brush rabbits,
skinks and other lizards, and many species of birds (such as An-
nas hummingbird, scrub jay, and bushtit) are common animals
of the chaparral.
Figure 55-9 Chaparral
Fires, which occur at irregular intervals in California chapar-
Chaparral, which consists primarily of drought-resistant evergreen
ral vegetation, are often quite costly because they consume expen- shrubs and small trees, develops where hot, dry summers alternate
sive homes built on the hilly chaparral landscape. Unfortunately, with mild, rainy winters. Photographed in the Santa Lucia Mountains,
efforts to control naturally occurring res sometimes backre. California.

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1196 Chapter 55 www.thomsonedu.com/biology/solomon
ed into spines, which discourage herbivores. Other desert plants
shed their leaves for most of the year, growing only during the
brief moist season.
Desert animals tend to be small. During the heat of the day,
they remain under cover or return to shelter periodically, whereas
at night they come out to forage or hunt. In addition to desert-
adapted insects, there are many specialized desert reptiles (such
as desert iguanas, desert tortoises, and rattlesnakes) and a few
desert-adapted amphibians (such as western spadefoot toads).
Mammals include such rodents as the American kangaroo rat,
which does not need to drink water but subsists solely on the
water content of its food (primarily seeds and insects). American
deserts are also home to jackrabbits, and kangaroos live in Austra-
lian deserts. Carnivores such as the African fennec fox and some
birds of prey, especially owls, live on rodents and rabbits. During
the driest months of the year, many desert insects, amphibians,

Willard Clay/Dembinsky Photo Associates

reptiles, and mammals tunnel underground, where they remain
inactive; this period of dormancy is known as estivation.
Humans have altered North American deserts in several
ways. Off-road vehicles damage desert vegetation, which some-
times takes years to recover. When the top layer of desert soil is
disturbed, erosion occurs more readily, and less vegetation grows
to support native animals. Another problem is that certain cacti
and desert tortoises have become rare as a result of poaching.
Houses, factories, and farms built in desert areas require vast Figure 55-10 Desert
quantities of water, which must be imported from distant areas. Summer rainfall characterizes the warmer deserts of North America,
Irrigation of desert soils often causes them to become salty and such as the Sonoran Desert shown here. The Sonoran Desert contains
unt for crops or native vegetation. Increased groundwater con- many species of cacti, including the large, treelike saguaro, which
sumption by many desert cities has caused groundwater levels to grows 15 to 18 m (50 to 60 ft) tall. Photographed in Arizona.
drop. Aquifer depletion in U.S. des-
erts is particularly critical in south-
ern Arizona and southwestern New
Mexico.

Savanna is a tropical
grassland with
scattered trees
The savanna biome is a tropical
grassland with widely scattered
clumps of low trees ( Fig. 55-11).
Savanna is found in areas of rela-
tively low or seasonal rainfall with
prolonged dry periods. Tempera-
tures in savannas vary little through-
out the year, and precipitation,
not temperature as in temperate
grasslands, regulates seasons. An-

Carlyn Iverson
nual precipitation is 85 to 150 cm
(34 to 60 in). Savanna soil is low in
essential minerals, in part because
it is strongly leached. Savanna soil Figure 55-11 Savanna
is often rich in aluminum, which In this photograph of African savanna in Tanzania, the smaller animals in the foreground are Thomsons
resists leaching, and in places the gazelles (Gazella thomsonii), and the larger ones near the trees are wildebeests (Connochaetes taurinus).

1
Ecology and the Geography of Life 1197
 Mark Moffett /Minden Pictures
Frans Lanting/Minden Pictures

(a) A broad view of tropical rain forest vegetation (b) Thick, epiphyte-covered lianas grow into the
along a riverbank in Southeast Asia. Except at river canopy using a tree trunk for support. Photographed
banks, tropical rain forest has a closed canopy that in Costa Rica.
admits little light to the forest floor.

Figure 55-12 Tropical rain forest

aluminum reaches levels that are toxic to many plants. Although to savanna. Temperate grasslands and tropical dry forests can also
the African savanna is best known, savanna also occurs in South be degraded to desert.)
America and northern Australia.
Wide expanses of grasses interrupted by occasional trees There are two basic types
characterize savanna. Trees such as Acacia bristle with thorns
that provide protection against herbivores. Both trees and grasses
of tropical forests
have re-adapted features, such as extensive underground root There are many kinds of tropical forests, but ecologists generally
systems, that enable them to survive seasonal droughts as well as classify them as one of two types: tropical dry forests or tropical
periodic res that sweep through the savanna. rain forests. Tropical dry forests occur in regions with a wet sea-
The worlds greatest assemblage of hooved mammals occurs son and a dry season (usually 2 to 3 months each year). Annual
in the African savanna. Here live great herds of herbivores, in- precipitation is 150 to 200 cm (60 to 80 in). During the dry sea-
cluding wildebeests, antelopes, giraffes, zebras, and elephants. son, many tropical trees shed their leaves and remain dormant,
Large predators, such as lions and hyenas, kill and scavenge the much as temperate trees do during the winter. India, Brazil, Thai-
herds. In areas of seasonally varying rainfall, the herds and their land, and Mexico are some of the countries that have tropical dry
predators may migrate annually. forests. Tropical dry forests intergrade with savanna on their dry
Savanna is rapidly being converted to rangeland for cattle and edges and with tropical rain forests on their wet edges. Logging
other domesticated animals, which are replacing the big herds of and overgrazing by domestic animals have fragmented and de-
wild animals. The problem is particularly acute in Africa, which graded many tropical dry forests.
has the most rapidly growing human population of any conti- The annual precipitation of tropical rain forests is 200 to
nent. In some places severe overgrazing by domestic animals has 450 cm (80 to 180 in). Much of this precipitation, which occurs
contributed to the conversion of marginal savanna into desert, a almost daily, comes from locally recycled water that enters the
process known as desertication. In desertication, the reduced atmosphere by transpiration from the forests own trees. Tropical
grass cover caused by overgrazing allows wind and water to erode rain forests are often located in areas with ancient, highly weath-
the soil; erosion removes the topsoil and decreases the soils abil- ered, mineral-poor soil. Little organic matter accumulates in such
ity to support crops or livestock. (Desertication is not restricted soils. Because temperatures are high and soil moisture is abun-

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1198 Chapter 55 www.thomsonedu.com/biology/solomon
dant year-round, decay organisms and detritus-feeding ants and
termites decompose organic litter quite rapidly. Vast networks of
roots and mycorrhizae quickly absorb minerals from decompos-
ing materials. Thus, minerals of tropical rain forests are tied up
in the vegetation rather than in the soil.
Tropical rain forests are found in Central and South America,
Africa, and Southeast Asia. Tropical rain forest is very productive
despite the scarcity of minerals in the soil. Its plants, stimulated
by abundant solar energy and precipitation, capture considerable
energy by photosynthesis. Of all the biomes, the tropical rain for-
est is unrivaled in species richness.
Most trees of tropical rain forests are evergreen owering
plants ( Fig. 55-12a). A fully developed rain forest has three or
more distinct stories of vegetation. The topmost story consists
of the crowns of the oldest, tallest trees, some 50 m (164 ft) or
more in height; these trees are exposed to direct sunlight and
are subject to the warmest temperatures, lowest humidities, and
strongest winds. The middle story, which reaches a height of 30 to
40 m (100 to 130 ft), forms a continuous canopy of leaves over-
head that lets in little sunlight for the support of the sparse un-
derstory. Only 2% to 3% of the light bathing the forest canopy
reaches the forest understory, which consists of both smaller
plants specialized for life in shade and seedlings of taller trees.
Vegetation of tropical rain forests is not dense at ground level
except near stream banks or where a fallen tree has opened the
canopy.
Tropical rainforest trees support extensive epiphytic commu-
nities of smaller plants such as orchids and bromeliads. Although
epiphytes grow in crotches of branches, on bark, or even on the
leaves of their hosts, most use their host trees only for physical
support, not for nourishment.
Because little light penetrates to the understory, many
plants living there are adapted to climb already-established host
trees. Lianas (woody tropical vines), some as thick as a human
thigh, twist up through the branches of tropical rainforest trees
( Fig. 55-12b). Once in the canopy, lianas grow from the up-
per branches of one forest tree to another, connecting the tops
of the trees and providing a walkway for many of the canopys
residents.
Not counting bacteria and other soil-dwelling organisms,
about 90% of tropical rainforest organisms live in the middle and
upper canopies. Rainforest animals include the most abundant
and varied insect, reptile, and amphibian fauna on Earth. Birds,
too, are diverse, with some specialized to consume fruits (such as
parrots), some to consume nectar (such as hummingbirds and
sunbirds), and others to consume insects. Most rainforest mam-
mals, such as sloths and monkeys, live only in the trees and never
climb down to the ground surface. Some large ground-dwelling
mammals, including elephants, are also found in tropical rain
forests.
Unless strong conservation measures are initiated soon, hu-
man population growth and agricultural and industrial expan-
sion in tropical countries will spell the end of tropical rain forests
by the middle of the 22nd century. Many rainforest species may
become extinct before they are even identied and scientically
described. (Tropical rainforest destruction is discussed in detail
in Chapter 56.)
Review
What climate and soil factors produce the major biomes?
What representative organisms are found in each of these
forest biomes: (1) boreal forest, (2) temperate deciduous
forest, (3) temperate rain forest, (4) tropical rain forest?
In which biome do you live? Does it match the description
given in this text? If not, explain the discrepancy.
How does tundra compare to desert? How does temperate
grassland compare to savanna?
AQUATIC ECOSYSTEMS
Learning Objectives
3 Explain the important environmental factors that affect
aquatic ecosystems.
4 Distinguish among plankton, nekton, and benthos.
5 Briey describe the various freshwater, estuarine, and marine
ecosystems, giving attention to the environmental character-
istics and representative organisms of each.
6 Describe at least one human effect on each of the aquatic
ecosystems discussed.
Aquatic biomes do not exist, in the sense that aquatic ecologists
do not distinguish aquatic ecosystems based on the dominant
form of vegetation. Aquatic ecosystems are classied primarily
on abiotic factors, such as salinity, that help determine an aquatic
life zones boundaries. Salinity, the concentration of dissolved
salts in a body of water, affects the kinds of organisms present
in aquatic ecosystems, as does the amount of dissolved oxygen.
Water greatly interferes with the penetration of light, so oating
aquatic organisms that photosynthesize remain near the waters
surface, and vegetation attached to the bottom grows only in
shallow water. In addition, low levels of essential minerals often
limit the number and distribution of organisms in certain aquatic
environments. Other abiotic determinants of species composi-
tion in aquatic ecosystems include water depth, temperature, pH,
and presence or absence of waves and currents.
Aquatic ecosystems contain three main ecological categories
of organisms: free-oating plankton, strongly swimming nekton,
and bottom-dwelling benthos. Plankton are usually small or mi-
croscopic organisms that are relatively feeble swimmers. For the
most part, they are carried about at the mercy of currents and
waves. They are unable to swim far horizontally, but some species
are capable of large vertical migrations and are found at differ-
ent depths of water at different times of the day or at different
seasons. Plankton are generally subdivided into two major cat-
egories: phytoplankton and zooplankton. Phytoplankton (pho-
tosynthetic cyanobacteria and free-oating algae) are producers
that form the base of most aquatic food webs. Zooplankton are
nonphotosynthetic organisms that include protozoa, tiny crusta-
ceans, and the larval stages of many animals. Nekton are larger,
more strongly swimming organisms such as shes, turtles, and
whales. Benthos are bottom-dwelling organisms that x them-
selves to one spot (sponges, oysters, and barnacles), burrow into
the sand (many worms and echinoderms), or simply walk or
swim about on the bottom (craysh, aquatic insect larvae, and
brittle stars).
1
Ecology and the Geography of Life 1199
 Freshwater ecosystems are closely linked
to land and marine ecosystems
Freshwater ecosystems include streams and rivers (owing-water
ecosystems), ponds and lakes (standing-water ecosystems), and Source
marshes and swamps (freshwater wetlands). Each type of fresh-
water ecosystem has its own specic abiotic conditions and char- Headwater
stream
acteristic organisms. Although freshwater ecosystems occupy a
relatively small portion about 2% of Earths surface, they are
important in the hydrologic cycle: they assist in recycling precipi-
tation that ows as surface runoff to the ocean (see discussion of
hydrologic cycle in Chapter 54). Freshwater habitats also provide
Waterfall
homes for large numbers of species.
Tributaries
Rapids
Streams and rivers are owing-water ecosystems
Many different conditions exist along the length of a stream or
river ( Fig. 55-13). The nature of a owing-water ecosystem Floodplain
changes greatly from its source (where it begins) to its mouth
Meanders
(where it empties into another body of water). Headwater
streams (small streams that are the sources of a river) are usually
shallow, clear, cold, swiftly owing, and highly oxygenated. In
contrast, rivers downstream from the headwaters are wider and Salt
deeper, cloudy (that is, they contain suspended particulates), not marsh
as cold, slower owing, and less oxygenated. Surrounding for-
est may shade certain parts of the stream or river, whereas other Mouth
parts may be exposed to direct sunlight. Along parts of a stream
or river, groundwater wells up through sediments on the bottom;
this local input of water moderates the water temperature so that
summer temperatures are cooler and winter temperatures are
warmer than in adjacent parts of the owing-water ecosystem. Delta
The kinds of organisms in owing-water ecosystems vary
greatly from one stream to another, depending primarily on the
strength of the current. In streams with fast currents, inhabitants
have adaptations such as suckers to attach themselves to rocks
so they are not swept away. The larvae of blackies, for example, Ocean
attach themselves with suction discs located on the ends of their
abdomens. Some stream inhabitants, such as immature water-
penny beetles, have attened bodies that enable them to slip un- Figure 55-13 Features of a typical river
der or between rocks. The water-penny beetle larva gets its com- The river begins at a source, often high in mountains and fed by melt-
mon name from its attened, nearly circular shape. Alternatively, ing snows or glaciers. Headwater streams ow downstream rapidly,
inhabitants such as the brown trout are streamlined and muscu- often over rocks (as rapids) or bluffs (as waterfalls). Along the way,
tributaries feed into the river. As the rivers course levels, the river ows
lar enough to swim in the current. more slowly and winds from side to side, forming meanders. The ood-
Streams and rivers depend on land for much of their energy. In plain is the area on either side of the river that is subject to ooding.
headwater streams, up to 99% of the energy input comes from de- Near the ocean, the river may form a salt marsh where fresh water from
tritus (dead organic material such as leaves) carried from the land the river and salt water from the ocean mix. Sediments deposited by
into streams and rivers by wind or surface runoff. Downstream, the river as it empties into the ocean form the delta, a fertile, low-lying
plain at the rivers mouth.
rivers contain more producers and therefore depend slightly less
on detritus as a source of energy than do the headwaters.
Human activities have several adverse impacts on rivers and dam, the once-powerful river is reduced to a relative trickle, so
streams, including water pollution and the effects of dams built to the owing-water ecosystem is altered.
contain the water of rivers or streams. Pollution alters the physi-
cal environment of a owing-water ecosystem and changes the
biotic component downstream from the pollution source. Dams Ponds and lakes are standing-water ecosystems
change the nature of owing-water ecosystems, both upstream Zonation characterizes standing-water ecosystems. A large lake
and downstream from the dam location. A dam causes water to has three basic zones: the littoral, limnetic, and profundal zones
back up, resulting in ooding of large areas of land and forma- ( Fig. 55-14). Smaller lakes and ponds typically lack a profun-
tion of a reservoir, which destroys terrestrial habitat. Below the dal zone.
0
1200 Chapter 55 www.thomsonedu.com/biology/solomon
 Littoral zone
Limnetic zone
Profundal
zone
Figure 55-14 Animated Zonation in a large lake
A lake is a standing-water ecosystem surrounded by land. Here, vegetation around the lake is not drawn to scale.
The littoral zone is a shallow-water area along the shore of
a lake or pond. It includes rooted, emergent vegetation, such as
cattails and burreeds, plus several deeper-dwelling aquatic plants
and algae. The littoral zone is the most productive zone of the
lake. Photosynthesis is greatest in the littoral zone, in part be-
cause light is abundant and because the littoral zone receives nu-
trient inputs from surrounding land that stimulate the growth of
plants and algae. In addition, ponds and lakes like streams and
rivers depend on detritus carried from the land for much of
their energy. Animals of the littoral zone include frogs and their
tadpoles, turtles, worms, craysh and other crustaceans, insect
larvae, and many shes such as perch, carp, and bass. Surface
dwellers, such as water striders and whirligig beetles, are found
in the quieter areas.
The limnetic zone is the open water beyond the littoral zone,
that is, away from the shore; it extends down as far as sunlight
penetrates to permit photosynthesis. The main organisms of the
limnetic zone are microscopic phytoplankton and zooplankton.
Larger shes also spend some of their time in the limnetic zone,
although they may visit the littoral zone to feed and reproduce.
Because of the depth of this zone, less vegetation grows in the
limnetic zone than in the littoral zone.
Beneath the limnetic zone of a large lake is the profundal
zone. Because light does not penetrate effectively to this depth,
plants and algae do not live in this zone. Food drifts into the pro-
fundal zone from the littoral and limnetic zones. Bacteria decom-
pose dead plants and animals that reach the profundal zone, thus
liberating minerals. These minerals are not effectively recycled,
because no photosynthetic organisms are present to absorb them
and incorporate them into the food web. As a result, the profun-
dal zone tends to be both mineral rich and anaerobic (oxygen
decient), with few organisms other than anaerobic bacteria oc-
cupying it.
Thermal stratification in temperate lakes. The marked
layering of large temperate lakes caused by light penetration is
accentuated by thermal stratication, in which the temperature
changes sharply with depth ( Fig. 55-15a). Thermal stratica-
tion occurs because the summer sunlight penetrates and warms
surface water, making it less dense. (Recall from Chapter 2 that
the density of water is greatest at 4C; both above and below this
temperature, water is less dense.) In summer, cool (and there-
fore more dense) water remains at the lake bottom and is sepa-
rated from warm (and therefore less dense) water above by an
abrupt temperature transition called the thermocline. Seasonal
distribution of temperature and oxygen (more oxygen dissolves
in water at cooler temperatures) affects the distribution of sh
in the lake.
In temperate lakes, falling temperatures in autumn cause a
mixing of the lake waters called the fall turnover ( Fig. 55-15b
1 ). As surface water cools, its density increases, and it sinks and
displaces the less dense, warmer, mineral-rich water beneath.
Warmer water then rises to the surface where it, in turn, cools and
sinks. This process of cooling and sinking continues until the lake
reaches a uniform temperature throughout. In winter 2 , surface
water cools to below 4C, its temperature of greatest density. Ice,
which forms at 0C, is less dense than cold water. Thus, ice forms
on the surface, and the water on the lake bottom is warmer than
the ice on the surface. In the spring 3 , a spring turnover occurs
as ice melts and surface water reaches 4C. Surface water again
sinks to the bottom, and bottom water returns to the surface. As
summer arrives 4 , thermal stratication occurs once again.
The mixing of deeper, nutrient-rich water with surface, nu-
trient-poor water during the fall and spring turnovers brings es-
sential minerals to the surface and oxygenated water to the bot-
tom. The sudden presence of large amounts of essential minerals
in surface waters encourages the development of large algal and
cyanobacterial populations, which may form temporary blooms
in the fall and spring.
Increased nutrients and algal growth. The presence of
high levels of plant and algal nutrients such as nitrogen and
phosphorus cause enrichment, the fertilization of a body of water.
Excess amounts of these nutrients enter waterways from sewage
and from fertilizer runoff from lawns and elds. The water in an
enriched pond or lake is cloudy because of the vast numbers of
1
Ecology and the Geography of Life 1201
 Summer

5 1520C
4 Thermocline

Water depth (m)

Thermocline 410C
10
1
Spring turnover Fall turnover
15

20 Entire lake Entire lake

5 10 15 20 310C 310C
Temperature (C)
Winter
(a)
3 Ice 0C

Figure 55-15 Animated Thermal stratication 2

in a temperate lake 4C
(a) Temperature varies at different depths during the summer. There is
an abrupt temperature transition, the thermocline. (b) During fall and
spring turnovers, a mixing of upper and lower layers of water brings
oxygen to the oxygen-depleted depths of the lake and minerals to the (b)
mineral-decient surface waters. (The numbered steps are explained in
the text.)

algae and cyanobacteria that the nutrients support. The species Wetlands are valued as a wildlife habitat for migratory water-
composition is different in enriched and unenriched lakes. For fowl and many other bird species, beavers, otters, muskrats, and
example, an unenriched lake in the northeastern United States game shes. Wetlands are holding areas for excess water when
may contain pike, sturgeon, and whitesh in the deeper, colder rivers ood their banks. The oodwater stored in wetlands then
part of the lake where there is a higher concentration of dissolved drains slowly back into the rivers, providing a steady ow of wa-
oxygen. In contrast, the deeper, colder levels of water in enriched ter throughout the year. Wetlands also serve as groundwater re-
lakes are depleted of dissolved oxygen because of the greater charging areas. One of their most important roles is to trap and
amount of decomposition on the lake oor. Fishes such as pike, hold pollutants in the ooded soil, thereby cleansing and purify-
sturgeon, and whitesh die out, and shes such as catsh and
carp, which tolerate lower concentrations of dissolved oxygen,
replace them.
Enrichment is reversible and has declined in North America
since the 1970s because the passage of legislation limits the phos-
phate content of detergents and because better sewage treatment
plants have been constructed. Agriculture is the leading source of
water-quality impairment of U.S. surface waters today. Fertilizer
runoff, as well as animal wastes and plant residues in waterways,
still cause enrichment problems.
Gregory J. Dimijian /Photo Researchers, Inc.

Freshwater wetlands are transitional

between aquatic and terrestrial ecosystems
Freshwater wetlands, which are usually covered by shallow
water for at least part of the year, have characteristic soils and
water-tolerant vegetation. They include marshes, dominated by
grasslike plants, and swamps, in which woody trees or shrubs
dominate ( Fig. 55-16). Freshwater wetlands also include hard-
wood bottomland forests (lowlands along streams and rivers that
are periodically ooded), prairie potholes (small, shallow ponds Figure 55-16 Freshwater swamp
that formed when glacial ice melted at the end of the last Ice Age), Trees, such as bald cypress (shown), dominate freshwater swamps. In
and peat moss bogs (peat-accumulating wetlands where sphag- this wetland, photographed in northeastern Texas, a oating carpet of
num moss dominates). tiny aquatic plants covers the waters surface.

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1202 Chapter 55 www.thomsonedu.com/biology/solomon

Jack Dermid/Photo Researchers, Inc.
Figure 55-17 Salt marsh
Cordgrass (Spartina alterniora) is the dominant vegetation in this salt
marsh along a creek in North Carolina. This photograph was taken at
low tide, showing an exposed oyster bed and an immature little blue
heron (Egretta caerulea).
ing the water. Such important environmental functions as these
are known as ecosystem services.
At one time wetlands were considered wastelands, areas to
be lled in or drained so that farms, housing developments, and
industrial plants could be built on them. Wetlands are also breed-
ing places for mosquitoes and therefore were viewed as a men-
ace to public health. The crucial ecosystem services that wetlands
provide are widely recognized today, and wetlands have some
legal protection. Agriculture, pollution, engineering (dams),
and urban and suburban development still threaten wetlands,
however.
extensive photosynthetic carpet and also mechanically traps de-
tritus, forming the basis of detritus food webs. Most commer-
cially important shes and shellsh spend their larval stages in
estuaries among the protective tangle of decaying stems.
Temperate estuaries usually contain salt marshes, shallow
wetlands in which salt-tolerant grasses dominate ( Fig. 55-17).
Uninformed people have often seen salt marshes as worthless,
empty stretches of land. As a result, people have used them as
dumps, severely polluting them, or lled them with dredged bot-
tom material to form articial land for residential and industrial
development. A large part of the estuarine environment is lost in
this way, along with many of its ecosystem services, for example,
biological habitats, sediment and pollution trapping, ground-
water supply, and storm buffering (salt marshes absorb much of
the energy of a storm surge and therefore prevent ood damage
elsewhere).
Mangrove forests, the tropical equivalent of salt marshes,
cover perhaps 70% of tropical and subtropical coastal mudats
where tides and waves uctuate ( Fig. 55-18). Like salt marshes,
mangrove forests provide valuable ecosystem services. Mangrove
roots stabilize the sediments, preventing coastal erosion and pro-
viding a barrier against the ocean during storms. Their interlac-
ing roots are breeding grounds and nurseries for commercially
important sh and shellsh species, such as blue crabs, shrimp,
mullet, and spotted sea trout. Mangrove branches are nesting
sites for many species of birds, such as pelicans, herons, egrets,
and roseate spoonbills. Mangroves are under assault from coastal
development, including aquaculture facilities, and unsustainable

Estuaries occur where fresh water

and salt water meet
Where the ocean meets the land, there may be one of several
kinds of ecosystems: a rocky shore, a sandy beach, an intertidal
mudat, or a tidal estuary. An estuary is a coastal body of water,
partly surrounded by land, with access to the open ocean and a
large supply of fresh water from rivers. Water levels in an estu-
ary rise and fall with the tides, and salinity uctuates with tidal
cycles, the time of year, and precipitation. Salinity also changes
gradually within the estuary, from unsalty fresh water at the river
entrance to salty ocean water at the mouth of the estuary. Because
estuaries undergo marked daily, seasonal, and annual variations
Patti Murray/Earth Scenes

in temperature, salinity, and other physical properties, estuarine

organisms have a wide tolerance to such changes.
Estuaries are among the most fertile ecosystems in the world,
often having much greater productivities than the adjacent
ocean or freshwater river (see Table 54-1). This high productivity
is the result of four factors: (1) The action of tides promotes a
Figure 55-18 Mangroves
rapid circulation of nutrients and helps remove waste products.
Red mangroves (Rhizophora mangle) have stiltlike roots that support
(2) Minerals are transported from land into streams and rivers the tree. Many animals live in the complex root systems of mangrove
that empty into the estuary. (3) A high level of light penetrates forests. Photographed at low tide along the coast of Florida, near
the shallow water. (4) The presence of many plants provides an Miami.

1
Ecology and the Geography of Life 1203
 Intertidal Pelagic environment The immense, complex marine environment is subdi-
zone vided into several zones: the intertidal zone, the benthic (ocean
Neritic Oceanic oor) environment, and the pelagic (ocean water) environment
High Low
province province ( Fig. 55-19). The pelagic environment is, in turn, divided into
tide tide two provinces the neritic province and the oceanic province.

Depth The intertidal zone is transitional

200 m between land and ocean
The intertidal zone is the shoreline area between low and high
tide. Although high levels of light and nutrients, together with
an abundance of oxygen, make the intertidal zone a biologically
Euphotic
zone productive environment, it is also a stressful one. If an intertidal
beach is sandy, inhabitants must contend with a constantly shift-
ing environment that threatens to engulf them and gives them
scant protection against wave action. Consequently, most sand-
dwelling organisms, such as mole crabs, are continual and active
4000 m
Benthic environment burrowers. Because they follow the tides up and down the beach,
most do not have any notable adaptations to survive drying or
exposure.
A rocky shore provides a ne anchorage for seaweeds and
Abyssal zone
of benthic
invertebrate animals. However, it is exposed to constant wave ac-
6000 m environment tion when immersed during high tides and to drying and tem-
perature changes when exposed to the air during low tides. A
typical rocky-shore inhabitant has some way of sealing in mois-
ture, perhaps by closing its shell, if it has one, plus a powerful
means of anchoring itself to rocks. Mussels, for example, have
Hadal zone
of benthic horny, threadlike anchors, and barnacles have special cement
environment glands. Rocky-shore intertidal algae (seaweeds) usually have
thick, gummy polysaccharide coats, which dry out slowly when
exposed, and exible bodies not easily broken by wave action
( Fig. 55-20). Some rocky-shore community inhabitants hide in
burrows or crevices at low tide.

Figure 55-19 Animated Zonation in the ocean
The ocean has three main life zones: the intertidal zone, the benthic
environment, and the pelagic environment. The pelagic environment
consists of the neritic and oceanic provinces. (The slopes of the ocean
oor are not as steep as shown; they are exaggerated to save space.)
logging. Some countries, such as the Philippines, Bangladesh,
and Guinea-Bissau, have cut down more than two thirds of their
mangrove forests.
Marine ecosystems dominate Earths surface
Although lakes and the ocean are comparable in many ways, they
have many differences. Depths of even the deepest lakes do not
approach those of the ocean, which has areas that extend more
than 6 km (3.6 mi) below the sunlit surface. Tides and currents
profoundly inuence the ocean. Gravitational pulls of both sun
and moon produce two tides a day throughout the ocean, but the
height of those tides varies with season, local topography, and
phases of the moon (full moons and new moons cause the high-
est tides).
Seagrass beds, kelp forests, and coral reefs
are part of the benthic environment
The benthic environment is the ocean oor. It is divided into
zones based on distance from land, light availability, and depth.
The benthic environment consists of sediments (mostly sand
and mud) in which many marine animals, such as worms and
clams, burrow. Bacteria are common in marine sediments, and
living bacteria have been found in ocean sediments more than
500 m (1625 ft) below the ocean oor at several different sites
in the Pacic Ocean. The abyssal zone is that part of the ben-
thic environment that extends from a depth of 4000 to 6000 m
(2.5 to 3.7 mi.). (In Chapter 54, Focus On: Life without the Sun
describes some of the unusual organisms in hydrothermal vents
in the abyssal zone.) The hadal zone is that part of the benthic
environment deeper than 6000 m.
Here we describe benthic communities in shallow ocean
waters seagrass beds, kelp forests, and coral reefs. Sea grasses
are owering plants that have adapted to complete submersion in
ocean water ( Fig. 55-21). They are not true grasses. Sea grasses
live in shallow water, to depths of 10 m (33 ft), where they receive
enough light to photosynthesize efciently. Extensive beds of sea
grasses occur in quiet temperate, subtropical, and tropical waters;

4
1204 Chapter 55 www.thomsonedu.com/biology/solomon
 ally greater than 21C), shallow sea water. The living portions
of coral reefs grow in shallow waters where light penetrates.
Many coral reefs are composed principally of red coralline algae
that require light for photosynthesis. Coral animals also require
light for the large number of symbiotic dinoagellates, known
as zooxanthellae, that live and photosynthesize in their tissues
(see Fig. 53-11). Although species of coral without zooxanthellae
exist, only those with zooxanthellae build reefs. In addition to
obtaining food from the zooxanthellae living inside them, coral
animals capture food at night; they use their stinging tentacles to
paralyze small animals that drift nearby. Coral reefs grow slowly
in warm, shallow water, as coral organisms build on the calcare-
ous remains of countless organisms before them. The waters in
which coral reefs are found are often poor in nutrients. Other
factors favor high productivity, however, including the presence
of symbiotic zooxanthellae, warm temperatures, and plenty of
sunlight.
William E. Ferguson

Coral reef ecosystems are the most diverse of all marine en-
vironments and contain hundreds or even thousands of species
of shes and invertebrates, such as giant clams, sea urchins, sea
stars, sponges, brittle stars, sea fans, and shrimp ( Fig. 55-22).
The Great Barrier Reef, along the northeastern coast of Australia,
Figure 55-20 Seaweeds in a rocky intertidal zone occupies only 0.1% of the oceans surface, but 8% of the worlds
Sea palms (Postelsia), which are 50 to 75 cm (20 to 30 in) tall, are com- sh species live there. The multitude of relationships and interac-
mon on the rocky Pacic coast from Vancouver Island to California. The
tions that occur at coral reefs is comparable only to those in tropi-
bases of these brown algae are rmly attached to the rocky substrate,
enabling them to withstand heavy surf action. Photographed at low cal rain forests among terrestrial ecosystems. As in the rain forest,
tide. competition is intense, particularly for light and space to grow.
Coral reefs are ecologically important because they both pro-
vide habitat for a wide variety of marine organisms and protect
no sea grasses live in polar waters. Sea grasses have a high primary coastlines from shoreline erosion. They also provide humans
productivity and are therefore ecologically important in shallow with seafood, pharmaceuticals, and income from tourism and
marine areas. Their roots and rhizomes stabilize the sediments, recreation. Although coral formations are important ecosystems,
reducing surface erosion. Sea grasses provide food and habitat for they are being degraded and destroyed. According to the UN En-
many marine organisms. In temperate waters, ducks and geese vironment Program, 27% of the worlds coral reefs are at high
eat sea grasses; in tropical waters, manatees, green turtles, parrot
sh, sturgeon sh, and sea urchins eat them. These herbivores
consume only about 5% of the sea grasses. The remaining 95%
eventually enter the detritus food web when the sea grasses die
and bacteria decompose them. In turn, a variety of animals such
as mud shrimp, lug worms, and mullet (a type of sh) consume
the bacteria.
Kelps, which may reach lengths of 60 m (200 ft), are the larg-
est brown algae (see Fig. 25-14b). Kelps are common in cooler
temperate marine waters of both the Northern and Southern
Hemispheres. They are especially abundant in relatively shallow
waters (depths of about 25 m, or 82 ft) along rocky coastlines.
Kelps are photosynthetic and are therefore the primary food pro-
ducers for the kelp forest ecosystem. Kelp forests also provide
habitats for many marine animals. Tube worms, sponges, sea cu-
cumbers, clams, crabs, shes (such as tuna), and mammals (such
Ron Phillips

as sea otters) nd refuge in the algal blades. Some animals eat the
blades, but kelps are consumed mainly in the detritus food web.
Bacteria that decompose dead kelp provide food for sponges, tu-
Figure 55-21 Seagrass bed
nicates, worms, clams, and snails. Kelp beds support a diversity
Turtle grasses (Thalassia) have numerous invertebrates and algal epi-
of life that almost rivals that found in coral reefs. phytes attached to their leaves. These shallow underwater meadows of
Coral reefs, which are built from accumulated layers of cal- marine owering plants are ecologically important for shelter and food
cium carbonate (CaCO3), are found in warm (temperature usu- for many organisms. Photographed off the coast of Mexico.

1
Ecology and the Geography of Life 1205
 risk. Coral reefs of southeastern Asia,
which contain the most species of all
coral reefs, are the most threatened of
any region.
In some areas, silt washing down-
stream from clear-cut inland forests has
smothered reefs under a layer of sedi-
ment. Some scientists hypothesize that
high salinity resulting from the diver-
sion of fresh water to supply the grow-
ing human population is killing Florida
reefs. Overshing, pollution from sew-
age discharge and agricultural runoff,
oil spills, boat groundings, shing with

Denise Tackett / Tom Stack & Associates

dynamite or cyanide, hurricane damage,
disease, coral bleaching, land reclama-
tion, tourism, and the mining of corals
for building material are also taking a
heavy toll. (Coral bleaching is discussed
in Chapter 29.)

The neritic province consists Figure 55-22 Coral reef organisms

of shallow waters close to shore A panoramic view of a coral reef in the Indian Ocean off the coast of the Maldives shows the many
The neritic province is open ocean that animals that live on and around coral reefs.
overlies the continental shelves, that is,
the ocean oor from the shoreline to a depth of 200 m (650 ft). 75% of the oceans water. Cold temperatures, high hydrostatic
Organisms that live in the neritic province are oaters or swim- pressure, and an absence of sunlight characterize the oceanic
mers. The upper reaches of the neritic province make up the eu- province; these environmental conditions are uniform through-
photic zone, which extends from the surface to a depth of ap- out the year.
proximately 100 m (325 ft). Enough light penetrates the euphotic Most organisms of the oceanic province depend on marine
zone to support photosynthesis. snow, organic debris that drifts down into the aphotic (with-
Large numbers of phytoplankton, particularly diatoms in out light) region from the upper, lighted regions. Organ-
cooler waters and dinoagellates in warmer waters, produce food isms of this little-known realm are lter-feeders, scavengers,
by photosynthesis and are thus the base of food webs. Zooplank-
ton (including tiny crustaceans; jellysh; comb jellies; protists
such as foraminiferans; and larvae of barnacles, sea urchins,
worms, and crabs) feed on phytoplankton.
Plankton-eating nekton, such as herring, sardines, squid,
manta rays, and baleen whales, consume zooplankton ( Fig.
55-23). These, in turn, become prey for carnivorous nekton such
as sharks, tuna, dolphins, and toothed whales. Nekton are thought
to be mostly conned to the shallower neritic waters (less than
60 m, or 195 ft, deep) because that is where their food is. How-
ever, not much is known about the behavior and migration pat-
Alex Kerstich / Visuals Unlimited

terns of marine nekton. Many shes appear to be wide ranging.

For example, an individual sh tagged on one side of an ocean
may be recaptured on the other side a few months later. Whether
the sh traveled alone or in a school is not known at this time.

The oceanic province makes up most of the ocean

Figure 55-23 Fish in the open ocean
The average depth of the worlds ocean is 4000 m (2.4 mi). The
Pacic sardines (Sardinops sagax) are one of the worlds important
oceanic province is that part of the open ocean that covers the commercial shes and provide an important source of protein for
deep-ocean basin, that is, the ocean oor at depths more than the human diet. Sardines are also an important food source for other
200 m. It is the largest marine environment and contains about marine animals.

6
1206 Chapter 55 www.thomsonedu.com/biology/solomon
or predators. Many are invertebrates,
some of which attain great sizes. The
giant squid, for example, measures up
to 18 m (59 ft) in length, including its
tentacles. Fishes of the oceanic prov-
ince are strikingly adapted to dark-
ness and food scarcity. For example,
the gulper eels huge jaws enable it
to swallow large prey ( Fig. 55-24).
(An organism that encounters food in- Image not available due to copyright restrictions
frequently needs to eat as much as pos-
sible when it has the chance.) Many
animals of the oceanic province have
illuminated organs that enable them to
see one another to mate or to capture
food. Adapted to drifting or slow swim-
ming, these shes often have reduced
bone and muscle mass.

Human activities are

harming the ocean
Because the ocean is so vast, it is hard
to visualize that human activities could
affect, much less harm, it. They do,
however. Development of resorts, cities,
industries, and agriculture along coasts
alters or destroys many coastal ecosys-
tems, including mangrove forests, salt marshes, seagrass beds,
and coral reefs. Coastal and marine ecosystems receive pollution
from land, from rivers emptying into the ocean, and from at-
mospheric contaminants that enter the ocean via precipitation.
Disease-causing viruses and bacteria from human sewage con-
taminate shellsh and other seafood and sometimes threaten
public health. Millions of tons of trash, including plastic, shing
nets, and packaging materials, end up in coastal and marine eco-
systems; some of this trash entangles and kills marine organisms.
Less visible contaminants of the ocean include fertilizers, pesti-
cides, heavy metals, and synthetic chemicals from agriculture and
industry.
Offshore mining and oil drilling pollute the neritic province
with oil and other contaminants. Millions of ships dump oily bal-
last and other wastes overboard in the neritic and oceanic prov-
inces. Fishing is highly mechanized, and new technologies can
remove every single sh in a targeted area of the ocean. Scallop
dredges and shrimp trawls are dragged across the benthic en-
vironment, destroying entire communities with a single swipe.
Most countries recognize the importance of the ocean to life on
this planet, but few have the resources or programs to protect and
manage the ocean effectively.
Review
What are plankton, nekton, and benthos?
What environmental factors are most important in deter-
mining the adaptations of organisms that live in aquatic
environments?
How do you distinguish between freshwater wetlands and
estuaries? Between owing-water and standing-water
ecosystems?
What are the four main marine environments?
Which aquatic ecosystem is often compared to tropical rain
forests? Why?
ECOTONES
Learning Objective
7 Dene and describe some of the features of an ecotone.
We have discussed the various terrestrial biomes and aquatic
ecosystems as if they were distinct and separate entities, but
within landscapes, ecosystems intergrade with one another at
their boundaries. You learned in Chapter 53 that the transition
zone where two communities or biomes meet and intergrade is
called an ecotone. Ecotones range in size from quite small, such
as the area where an agricultural eld meets a woodland or where
a stream ows through a forest, to continental in scope. For ex-
ample, at the border between tundra and boreal forest an exten-
sive ecotone exists that consists of tundra vegetation interspersed
with small, scattered conifers. Such ecotones provide habitat
diversity. In fact, often a greater variety and density of organ-
isms populate ecotones than live in either adjacent ecosystem
( Fig. 55-25).

1
Ecology and the Geography of Life 1207
 Key Experiment BIOGEOGRAPHY
QUESTION: Does species richness vary among ecotones and Learning Objective
their adjoining communities? 8 Dene biogeography, and briey describe Wallaces bio-
HYPOTHESIS: Ecotones have greater species richness than the geographic realms.
communities they connect.
The study of the geographic distribution of plants and animals is
EXPERIMENT: Plant species were sampled for two communities called biogeography (see Chapter 18). Biogeographers search for
in southwestern Oregon and for the ecotone between them. The
patterns in geographic distribution and try to explain how such
communities, one with nonserpentine soil and one with serpen-
tine soil, are dened largely by soil conditions. Nonserpentine patterns arose, including where populations originated, how they
soils (samples 1 to 10) are normal soils. Serpentine soils (sam- spread, and when. Biogeographers recognize that geologic and
ples 18 to 28) contain high levels of elements such as chromium, climate changes such as mountain building, continental drift,
nickel, and magnesium that are toxic to many plants. and periods of extensive glaciation inuence the distribution of
species. Biogeography is linked to evolutionary history and pro-
Nonserpentine Serpentine vides insights into how organisms may have interacted in ancient
Ecotone
soil soil ecosystems. Studying biogeography helps us relate ancient eco-
Black oak systems to modern ones, because ancient ecosystems form a con-
Poison oak tinuum with modern ecosystems.
Iris One of the basic tenets of biogeography is that each species
Douglas fir
originated only once. The particular place where this occurred
Hawkweed
is known as the species center of origin. The center of origin is
Fescue
not a single point but the distribution of the population when
Plant species

Snakeroot
Canyon live oak the new species originated. From its center of origin, each species
Collomia spreads until a barrier of some kind halts it; examples of barriers
Ragwort include the ocean, a desert, or a mountain range; unfavorable
Yarrow climate; or the presence of organisms that compete successfully
Buck brush for food or shelter.
Fescue Most plant and animal species have characteristic geographic
Fireweed distributions. The range of a particular species is that portion of
Knotweed Earth in which it is found. The range of some species may be a
0 5 10 15 20 25 28 relatively small area. For example, wombats, the marsupial equiv-
Samples taken along transect alent of groundhogs, are found only in drier parts of southeastern
Australia and nearby islands. Such localized, native species are
RESULTS AND CONCLUSION: Shown are the various plant
said to be endemic, that is, they are not found anywhere else in
species found in the two communities (yellow and blue) and in the world. In contrast, some species have a nearly worldwide dis-
the ecotone between them (green). The ecotone had a greater tribution and occur on more than one continent or throughout
richness than either adjoining community. much of the ocean. Such species are said to be cosmopolitan.
One of the early observations of biogeographers is that the
ranges of different species do not include everywhere that they
Figure 55-25 Ecotones and species richness
could survive. Central Africa has elephants, gorillas, chimpanzees,
(Modied from C. D. White, Vegetation-Soil Chemistry Correlations
lions, antelopes, umbrella trees, and guapiruvu trees, whereas ar-
in Serpentine Ecosystems, Ph.D. dissertation, University of Oregon,
Eugene, 1971. Reprinted with permission of Dr. Charles D. White.) eas in South America with a similar climate have none of these.
These animals and plants originated in Africa after continental
drift had already separated the supercontinent Pangaea into sev-
eral landmasses. The organisms could not expand their range
Ecologists who study ecotones look for adaptations that en-
into South America because the Atlantic Ocean was an impass-
able organisms to survive there. They also examine the relation-
able barrier. Likewise, the ocean was a barrier to South American
ship between species richness and ecotones and how ecotones
monkeys, sloths, tapirs, balsa trees, and snakewood trees, none of
change over time. Long-term studies of ecotones have revealed
which are found in Africa.
they are far from static. The ecotone boundary between desert
and semiarid grassland in southern New Mexico, for example,
has moved during the past 50 years as the desert ecosystem has Land areas are divided into
expanded into the grassland.
six biogeographic realms
Review As the various continents were explored and their organisms
What is an ecotone? studied, biologists observed that the world could be divided into
Where do ecotones occur? major blocks of vegetation, such as forests, grasslands, and des-

8
1208 Chapter 55 www.thomsonedu.com/biology/solomon
erts, and that these vegetation types
corresponded to specic climates.
The relationship between animal dis-
tribution, geography, and climate was
not deduced until 1876. At that time,
Alfred Wallace, who discovered the Nearctic
same theory of evolution by natural
selection as Charles Darwin, divided
Earths land areas into six major bio-
geographic realms: the Palearctic,
Nearctic, Neotropical, Ethiopian, Ori-
ental, and Australian ( Fig. 55-26). A Neotropical
major barrier separates each of the six
biogeographic realms from the others
and helps maintain each regions bio-
logical distinctiveness.
Many biologists quickly embraced
Wallaces classication, which is still
considered valid today. However, hu-
man activities, such as the intentional Figure 55-26 Wallaces biogeographic realms
and unintentional introduction of for- Certain unique species characterize each of the six biogeographic realms.
eign species, are contributing to a ho-
mogenization of the biogeographic realms (see Chapter 56).
Refer to Figure 55-26 as we briey consider some of the
characteristic animals in each realm. The Nearctic and Palearctic
realms are more closely related than the other regions, especially
in their northern parts, where they share many animals such as
wolves, hares, and caribou. This similarity may be due to a land
bridge that has periodically connected Siberia and Alaska. This
bridge was present late in the Pleistocene epoch, about 10,000
years ago. Animals adapted to cold environments may have dis-
persed between Asia and North America along this bridge.
The Neotropical realm was almost completely isolated from
the Nearctic realm and other landmasses for most of the past
70 million years. During this time, many marsupial species
evolved. The isthmus of Panama, which formed a dry-land con-
nection about 3 million years ago, linked North and South Amer-
ica and provided a route for dispersal. Only three species, the
opossum, armadillo, and porcupine, are descendants of animals
that survived the northward dispersal from South America, but
many species, such as the tapir and llama, are descendants of ani-
mals that survived the southward dispersal from North America.
S U M M ARY WI T H K EY TERMS
Learning Objectives
1 Dene biome, and briey describe the nine major terrestrial
biomes, giving attention to the climate, soil, and characteris-
tic plants and animals of each (page 1190).
A biome is a large, relatively distinct terrestrial region with
characteristic climate, soil, plants, and animals.
A frozen layer of subsoil (permafrost) and low-growing
vegetation that is adapted to extreme cold and a short
growing season characterize tundra, the northernmost
biome.
Palearctic
Oriental
Ethiopian
Australian
Competition from these species caused many of South Americas
marsupial species to go extinct.
The Sahara Desert separates the Ethiopian realm, which con-
tains the most varied vertebrates of all six realms, from other land-
masses. Some overlap exists between the Ethiopian realm and Ori-
ental realm because a land bridge with a moist climate linked Africa
to Asia during the Miocene and Pliocene epochs. The Oriental
realm has the fewest endemic species of all the tropical realms.
The Australian realm has not had a land connection with
other regions for more than 85 million years. It has no native
placental mammals, and marsupials and monotremes, includ-
ing the duck-billed platypus and the spiny anteater, dominate it.
Adaptive radiation of the marsupials during their long period of
isolation led to species with ecological niches similar to those of
placental mammals of other realms (see Fig. 31-28).
Review
What is biogeography?
Which biogeographic realm has been separated from the
other biogeographic realms for the longest period? What
animals characterize this biogeographic realm?
Coniferous trees adapted to cold winters; a short growing
season; and acidic, mineral-poor soil dominate the boreal
forest, or taiga.
Large conifers dominate temperate rain forest, which
receives high precipitation.
Temperate deciduous forest occurs where precipita-
tion is relatively high and soils are rich in organic matter.
Broad-leaf trees that lose their leaves seasonally dominate
temperate deciduous forest.
1
Ecology and the Geography of Life 1209
 Temperate grassland typically has a deep, mineral-rich
soil and has moderate but uncertain precipitation.
Thickets of small-leaf evergreen shrubs and trees and a
climate of wet, mild winters and dry summers characterize
chaparral.
Desert, found in both temperate (cold deserts) and
subtropical or tropical regions (warm deserts) with low
levels of precipitation, contains organisms with specialized
water-conserving adaptations.
Tropical grassland, called savanna, has widely scattered
trees interspersed with grassy areas. Savanna occurs in
tropical areas with low or seasonal rainfall.
Mineral-poor soil and high rainfall that is evenly distrib-
uted throughout the year characterize tropical rain forest.
Tropical rain forest has high species richness and high
productivity.
Learn more about biomes by clicking on the
gure in ThomsonNOW.
2 Describe at least one human effect on each of the biomes
discussed (page 1190).
Oil exploration and military exercises result in long-lasting
damage to tundra. Clear-cut logging destroys boreal and
temperate rain forests. Temperate deciduous forests are
removed for logging and for use as farms, tree planta-
tions, and land development.
Human population growth and its accompanying agricul-
tural and industrial expansion threaten most of the worlds
tropical rain forests.
Farmland has replaced most temperate grasslands; savan-
nas are increasingly converted to rangeland for cattle. De-
velopment of hilly chaparral results in mud slides and costly
res. Land development in deserts reduces wildlife habitat.
3 Explain the important environmental factors that affect
aquatic ecosystems (page 1199).
In aquatic ecosystems, important environmental factors
include salinity (concentration of dissolved salts), amount
of dissolved oxygen, availability of light, levels of essential
minerals, water depth, temperature, pH, and presence or
absence of waves and currents.
4 Distinguish among plankton, nekton, and benthos
(page 1199).
Aquatic life is ecologically divided into plankton (free-
oating organisms), nekton (strongly swimming organ-
isms), and benthos (bottom-dwelling organisms).
Phytoplankton are photosynthetic algae and cyano-
bacteria that form the base of the food web in most
aquatic communities. Zooplankton are nonphotosynthetic
organisms that include protozoa, tiny crustaceans, and
the larval stages of many animals.
5 Briey describe the various freshwater, estuarine, and marine
ecosystems, giving attention to the environmental character-
istics and representative organisms of each (page 1199).
Freshwater ecosystems include owing-water ecosystems
(streams and rivers), standing water ecosystems (ponds
and lakes), and freshwater wetlands. In owing-water
ecosystems the water ows in a current. Flowing-water
ecosystems have few phytoplankton and depend on detri-
tus from the land for much of their energy.
Large standing-water ecosystems (freshwater lakes)
are divided into zones on the basis of water depth. The
marginal littoral zone contains emergent vegetation and
algae and is very productive. The limnetic zone, open
water away from the shore that extends as far down as
sunlight penetrates, contains phytoplankton, zooplankton,
and larger shes. The deep, dark profundal zone holds
little life other than bacterial decomposers.
0
1210 Chapter 55
Freshwater wetlands, lands that are transitional between
freshwater and terrestrial ecosystems, are usually covered
at least part of the year by shallow water and have charac-
teristic soils and vegetation. Freshwater wetlands perform
many valuable ecosystem services.
An estuary is a coastal body of water, partly surrounded
by land, with access to both the ocean and a large supply
of fresh water from rivers. Salinity uctuates with tidal
cycles, the time of year, and precipitation. Temperate
estuaries usually contain salt marshes, whereas mangrove
forests dominate tropical coastlines.
Four important marine environments are the intertidal
zone, the benthic environment, the neritic province, and
the oceanic province. The intertidal zone is the shore-
line area between low and high tides. Organisms of the
intertidal zone have adaptations to resist wave action and
the extremes of being covered by water (high tide) and
exposed to air (low tide).
The benthic environment is the ocean oor. Sea grasses,
kelps, and coral reefs are important benthic communities
in shallow ocean waters.
The neritic province is open ocean from the shoreline to a
depth of 200 m. Organisms that live in the neritic province
are all oaters or swimmers. Phytoplankton are the base of
the food web in the euphotic zone, where enough light
penetrates to support photosynthesis.
The oceanic province is that part of the open ocean that
is deeper than 200 m. The uniform environment is one of
darkness, cold temperature, and high pressure. Animal
inhabitants of the oceanic province are either predators
or scavengers that subsist on marine snow, detritus that
drifts down from other areas of the ocean.
Learn more about lake and ocean zonation by
clicking on the gures in ThomsonNOW.
6 Describe at least one human effect on each of the aquatic
ecosystems discussed (page 1199).
Water pollution and dams adversely affect owing-water
ecosystems. Increased nutrients, supplied by human activi-
ties, stimulate algal growth, resulting in enriched ponds
and lakes. Agriculture, pollution, and land development
threaten wetlands and estuaries. Pollution, coastal devel-
opment, offshore mining and oil drilling, and overshing
threaten marine ecosystems.
7 Dene and describe some of the features of an ecotone
(page 1207).
An ecotone is the transition zone where two communities
or biomes meet and intergrade. Ecotones provide habitat
diversity and are often populated by a greater variety of
organisms than lives in either adjacent ecosystem.
8 Dene biogeography, and briey describe Wallaces biogeo-
graphic realms (page 1208).
Biogeography is the study of the geographic distribution
of plants and animals, including where populations came
from, how they got there, and when. Each species origi-
nated only once, at its center of origin. From its center of
origin, each species spreads until a physical, environmen-
tal, or biological barrier halts it. The range of a particular
species is that portion of the Earth in which it is found.
Alfred Wallace divided Earths land areas into six major
biogeographic realms: the Palearctic, Nearctic, Neotropi-
cal, Ethiopian, Oriental, and Australian. Each realm is
biologically distinctive because a mountain range, desert,
ocean, or other barrier separates it from the others. Today,
human activities are contributing to a homogenization of
the biogeographic realms.
www.thomsonedu.com/biology/solomon
T E ST Y OU R UN D E RS TA ND ING
1. The northernmost biome, known as , , and bottom-dwelling
typically has little precipitation, a short growing season,
and permafrost. (a) chaparral (b) boreal forest (c) tundra
(d) northern deciduous forest (e) savanna
2. South of tundra is the , which consists of
coniferous forests with many lakes. (a) chaparral (b) bo-
real forest (c) alpine tundra (d) northern deciduous forest
(e) permafrost
3. Forests of the northeastern and Mid-Atlantic United States,
which have broad-leaf hardwood trees that lose their foliage
annually, are called (a) temperate deciduous forests (b) tropi-
cal dry forests (c) boreal forests (d) temperate rain forests
(e) tropical rain forests
4. The deepest, richest soil in the world occurs in (a) temperate
rain forest (b) tropical rain forest (c) savanna (d) temperate
grassland (e) chaparral
5. This biome, with its thicket of evergreen shrubs and small
trees, is found in areas with Mediterranean climates. (a) tem-
perate rain forest (b) tropical rain forest (c) savanna (d) tem-
perate grassland (e) chaparral
6. This biome is a tropical grassland interspersed with widely
spaced trees. (a) temperate rain forest (b) tropical rain forest
(c) savanna (d) temperate grassland (e) chaparral
7. This biome has the greatest species richness. (a) temperate
rain forest (b) tropical rain forest (c) savanna (d) temperate
grassland (e) chaparral
8. Organisms in aquatic environments fall into three catego-
ries: free-oating , strongly swimming
C R I TI C AL TH I N KI NG
1. In which biomes would migration be most common? Hiber-
nation? Estivation? Explain your answers.
2. Develop a hypothesis to explain why animals adapted to the
desert are usually small. How would you test your hypothesis?
3. Why do most of the animals of the tropical rain forest live in
trees?
4. What would happen to the organisms in a river with a fast
current if a dam were built? Would there be any differences
in habitat if the dam were upstream or downstream of the
organisms in question? Explain your answers.
5. Evolution Link. When a black-tailed prairie dog or other
small animal dies, other prairie dogs bury it. Develop a hy-
pothesis to explain how this behavior may be adaptive. How
would you test this hypothesis?
.
(a) nekton; benthos; plankton (b) nekton; plankton; benthos
(c) plankton; benthos; nekton (d) plankton; nekton; benthos
(e) benthos; nekton; plankton
9. Temperate-zone lakes are thermally stratied, with warm
and cold layers separated by a transitional (a) aphotic region
(b) thermocline (c) barrier reef (d) ecotone (e) littoral zone
10. Emergent vegetation grows in the zone
of freshwater lakes. (a) littoral (b) limnetic (c) profundal
(d) neritic (e) intertidal
11. This coastal body of water has access to the open ocean and
a large supply of fresh water from rivers. (a) intertidal zone
(b) estuary (c) freshwater wetland (d) neritic province
(e) standing-water ecosystem
12. The is open ocean from the shoreline to a
depth of 200 m. (a) benthic environment (b) intertidal zone
(c) neritic province (d) oceanic province (e) aphotic region
13. Sea grasses (a) occur in shallow water of the oceans benthic
environment (b) may reach lengths of 60 m (c) contain
symbiotic algae known as zooxanthellae (d) are common on
rocky shores in the intertidal zone (e) are the main producers
in freshwater wetlands
14. The transition zone where two ecosystems or biomes meet
and intergrade is called a(an) (a) biosphere (b) aphotic region
(c) thermocline (d) biogeographic realm (e) ecotone
15. Which biogeographic realm has been separated from the
other landmasses for more than 85 million years? (a) Ethio-
pian (b) Palearctic (c) Nearctic (d) Oriental (e) Australian
6. Analyzing Data. Examine Figure 55-1. What is the lowest
average annual precipitation characteristic of tropical rain
forests? The highest? What is the range of average temperature
in tropical rain forests?
7. Analyzing Data. Examine Figure 55-25. How many of the
sampled species are found in the nonserpentine soil? In the
serpentine soil? In the ecotone? What generalization about
ecotones do these data support?
Additional questions are available in
ThomsonNOW at www.thomsonedu.com/
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Ecology and the Geography of Life 1211