Bab 6

6 Ecosystems
and Evolution
THE FLORIDA EVERGLADES
T he Everglades, a “river of grass” in the

southernmost part of Florida, is a vast expanse
of predominantly sawgrass wetlands dotted with
small islands of trees. It is a haven for wildlife, including
alligators (see photograph), snakes, panthers, otters,
raccoons, and thousands of birds.
The Everglades today is about half its original size
of 1.6 million hectares (4 million acres) and suffers
from many serious environmental problems. Wading
bird populations dropped 93 percent in the mid-20th
century (see graph), with many still in decline, and the
area is now home to 50 endangered or threatened
species. Invasive predator species, including Burmese
pythons—former pets—decimate many prey
populations.
More than 70 years of engineering projects
aimed at protecting humans from
storm-related flooding have
reduced the quantity of water
flowing into the Everglades,
restricting the natural
recharging process there.
Flood-control measures created
dry spaces that were then converted to
agricultural or residential use, fragmenting wildlife
habitat and polluting the water that does enter.
The Everglades will never return completely to
its original condition because there are now too
many cities and sugar plantations in the region.
However, state and federal governments are working
together on the massive Comprehensive Everglades
Restoration Plan to eventually restore a more natural
water flow to the area, repel invasions of foreign
species, and reestablish
native species.
graphingactivity
c06_EcosystemsAndEvolution.indd 126 10/27/2016 9:19:01 PM

CHAPTER OUTLINE
Factors That Shape Biomes 128
■ Environmental InSight: How Climate Shapes
Terrestrial Biomes
Describing Earth's Major Biomes 132
• Tundra
• Boreal Forest
• Temperate Rain Forest
• Temperate Deciduous Forest
• Temperate Grassland
• Chaparral
■ EnviroDiscovery 6.1: Using Goats to Fight Fires
• Desert
• Savanna
• Tropical Rain Forest
Aquatic Ecosystems 142
• Freshwater Ecosystems
■ What a Scientist Sees 6.1: Zonation in a Large
I nterpret the D ata Lake
What was the approximate percentage decrease in • Brackish Ecosystems: Estuaries
Everglades wading bird populations between the 1940s
and 1975? Population Responses to Changing Conditions
over Time: Evolution 147
• Natural Selection
■ Environmental InSight: Evidence for Evolution
Community Responses to Changing Conditions
over Time: Succession 151
• Primary Succession
• Secondary Succession
■ Case Study 6.1: Wildfires
CHAPTER PLANNER ✓
❑ Study the picture and read the opening story.
❑ Scan the Learning Objectives in each section:
p. 128 ❑ p. 132 ❑ p. 142 ❑ p. 147 ❑ p. 151 ❑
❑ Read the text and study all figures and visuals.
Answer any questions.
Analyze key features

❑ Environmental InSight, p. 129 ❑ p. 150 ❑
❑ EnviroDiscovery 6.1, p. 138
❑ What a Scientist Sees 6.1, p. 143
❑ Process Diagram, p. 149 ❑ p. 152 ❑ p. 153 ❑
❑ Case Study 6.1, p. 154
❑ Stop: Answer the Concept Checks before you go on:
p. 128 ❑ p. 142 ❑ p. 147 ❑ p. 150 ❑ p. 153 ❑
End of Chapter
❑ Review the Summary and Key Terms.
❑ Answer What is happening in this picture?
❑ Answer the Critical and Creative Thinking Questions.
127

Factors That Shape Biomes
LEARNING OBJECTIVES
1. Define biome environments such as the rainforest floor. Other abiotic fac-
2. Discuss how biomes are related to climate. tors to which certain biomes are sensitive include extreme
temperatures as well as rapid temperature changes, fires,
floods, droughts, and strong winds.
E
arth has many different climates—long-term
Elevation also affects biomes: Changes in vegetation
weather patterns—based primarily on tem-
with increasing elevation resemble the changes in vegeta-
perature and precipitation differences. Char-
tion observed in going from warmer to colder climates.
acteristic organisms have adapted to each
These differences across biomes can be further defined
climate within large regions called biomes. Each biome
by types of vegetation present and land use patterns
encompasses many interacting ecosystems (Figure 6.1).
(Figure 6.3 on pages 130 and 131).
In terrestrial ecology, a biome is considered the next
level of ecological organization
biome A large,
above community, ecosystem, and
relatively distinct
landscape.
terrestrial region with
Near the poles, temperature
similar climate, soil,
is generally the overriding cli-
plants, and animals,
mate factor defining a biome, 1. How is it that the same biome type might be
regardless of where it found in widely different parts of the world?
occurs in the world. whereas in temperate and tropi-
cal regions, precipitation is more 2. Which climate-related factors shape biomes,
significant than temperature, as shown in Figure 6.2. and how might they affect which organisms
Light is relatively plentiful in biomes, except in certain live in a particular biome?
The world’s terrestrial biomes • Figure 6.1

Although sharp boundaries are shown in this highly simplified map, biomes actually grade together at their boundaries.
Use the legend below to identify the locations of the different biomes.
60° N
60° N
30° N 30° N
0° 0°
Based on data from World Wildlife Fund.
30° S 30° S
Tundra Desert
Boreal forest Tropical rain forest
Temperate deciduous forest Tropical dry forest
0 1000 2000 Miles
and temperate rain forest Savanna
60° S
60° S
0 1000 2000 Kilometers Temperate grassland Mountains with
Chaparral complex zonation
128 CHAPTER 6

How climate shapes
Environmental InSight terrestrial biomes • Figure 6.2
✓ THE PLANNER
Two climate factors, temperature and

precipitation, have a predominant effect on
Alaska Tundra.
biome distribution.
At higher latitudes, temperature is more important
than precipitation in shaping biomes, as mean annual
temperatures decline poleward.
Costa Rica Tropical Rain Forest.
Rod Planck/Science Source Images

In temperate and tropical zones,
precipitation is more important than
temperature in shaping biomes.
Michael Melford/NG Image Collection
Ar
ld
RICHARD NOWITZ/NG Image Collection

Co
cti
c
Tundra
U RE
AT
IN
CR
ER
EA
MP
SI
TE
NG
NG
LA
Boreal forest
I
TI
AS
California Desert.
TU
RE
Biomes differ in the

DE
C
relative amounts of
DE
precipitation they
receive and in the
Temperate Temperate Temperate Chaparral Temperate seasonal distribution
rain forest deciduous forest grassland desert of precipitation.
Tro
Tropical Tropical dry Savanna Moist tropical Dry tropical

p
t
Ho
ics
rain forest forest desert desert

Wet DECREASING PRECIPITATION Dry
Based on Holdridge, L. Life Zone Ecology. Tropical Science Center, San Jose, Costa Rica (1967).

A diversity of land cover types • Figure 6.3
Similar vegetation types can occur at many
different locations.
Brand X/SUPERSTOCK
Hemis. Fr/SUPERSTOCK
Paul Davis/The Global Land Cover Facility,

University of Maryland Institute for
Advanced Computer Studies/NG Maps
Brand X/SUPERSTOCK
Brand X/SUPERSTOCK
Brand X/SUPERSTOCK
Brand X/SUPERSTOCK
Brand X/SUPERSTOCK
Brand X/SUPERSTOCK
130 CHAPTER 6 Ecosystems and Evolution

In t e r pr e t t h e Da t a
Which land cover types appear to be
most abundant worldwide?
Ernest Manewal/SUPERSTOCK

Describing Earth’s Major Biomes
LEARNING OBJECTIVES
1. Briefly describe the nine major terrestrial Arctic tundra has long, harsh winters and short sum-
biomes, giving attention to the climate, soil, mers. Although the arctic tundra’s growing season is
and characteristic organisms of each. short, the days are long. Above the Arctic Circle, the sun
does not set at all for many days in midsummer. There
is little precipitation, and most of the yearly 10 to 25 cm
E
arth’s terrestrial ecosystems can be grouped
within nine major biomes: tundra, boreal (4 to 10 in) of rain or snow falls during summer months.
forest, temperate rain forest, temperate de- Most tundra soils formed when glaciers began
ciduous forest, temperate grassland, chapar- retreating after the last ice age, about 17,000 years ago.
ral, desert, savanna, and tropical rain forest. Here we These soils are usually nutrient poor and have little
describe characteristics of each biome and highlight ef- detritus, such as dead leaves and stems, animal droppings,
fects humans have had on them. or remains of organisms. Although the tundra’s surface
soil thaws during summer, beneath it lies a layer of perma-
frost, permanently frozen ground that varies in depth and
Tundra thickness. Permafrost impedes drainage, so the thawed
Tundra (or arctic tundra) occurs in upper zone of soil is usually waterlogged during summer.
the extreme northern latitudes tundra The treeless Limited precipitation, combined with low temperatures,
where the snow melts seasonally biome in the far flat topography (or surface features), and the layer of
(Figure 6.4). The Southern Hemi- north that consists permafrost, produces a landscape of broad, shallow lakes
sphere has no equivalent of the arc- of boggy plains and ponds, sluggish streams, and bogs.
covered by lichens
tic tundra because it has no land in Tundra has low primary productivity (see Figure 5.8)
and mosses; it has
the corresponding latitudes. A sim- and supports relatively few species compared to other
harsh, cold winters
ilar ecosystem located in the higher biomes, but the species that do occur there often exist
and extremely short
elevations of mountains, above the in great numbers. Mosses, lichens, grasses, and grasslike
summers.
tree line, is called alpine tundra. sedges are the dominant plants. Stunted trees and shrubs
Arctic tundra • Figure 6.4

Because of the tundra’s short growing season
and permafrost, only small, hardy plants grow in
the northernmost biome that encircles the Arctic
Ocean. Photographed at Alaska's Colville River.
Climate graph shows monthly temperatures (line)
and precipitation (bars) for Fort Yukon, Alaska.
20
Average monthly temperature in °C
Freezing 15
point
Michael Melford/NG Image Collection

10
5
Average monthly precipitation in cm
0
–5
–10
–15
–20
–25
–30
14
12
10
8
6
4 In t e r p r e t t h e Da t a
2 During how many months, if any, is the average
0
J FMAM J J A SOND
monthly temperature in the Alaskan tundra
below freezing?
Months
Based on data from www.worldclimate.com

grow only in sheltered locations. As a rule, tundra plants across North America and Eurasia. There is no biome in
seldom grow taller than 30 cm (12 in). the Southern Hemisphere comparable to the boreal for-
Animals adapted to live year-round in the tundra est. Winters in the boreal forest are extremely cold and
include lemmings, voles, weasels, arctic foxes, snowshoe severe, although not as harsh as those in the tundra.
hares, ptarmigan, snowy owls, and musk oxen. In summer, Boreal forest receives little precipitation, perhaps 50 cm
caribou migrate north to the tundra to graze on sedges, (20 in) per year, and its soil is typically acidic and mineral
grasses, and dwarf willow. Dozens of bird species also mi- poor, with a thick surface layer of partly decomposed pine
grate north in summer to nest and feed on abundant insects. and spruce needles. Permafrost occurs only in patches,
Mosquitoes, blackflies, and deerflies survive winter as eggs or often deep under the surface. Boreal forest has numerous
pupae and appear in great numbers during summer weeks. ponds and lakes dug by ice sheets during the last ice age.
Tundra recovers slowly from even small disturbances. Black and white spruces, balsam fir, eastern larch, and
Oil and natural gas exploration and military use have other conifers dominate the boreal forest, although decid-
caused damage to tundra likely to persist for hundreds of uous trees (trees that shed their leaves in autumn), such as
years (see Case Study 17.1). aspen and birch, may form striking stands (Figure 6.5).
Climate change is beginning to affect the arctic tundra. Conifers have many drought-resistant adaptations, such as
As the permafrost melts, conifer trees (cone-bearing ever- needle-like leaves whose minimal surface area prevents wa-
greens) are replacing tundra vegetation. The trees have ter loss by evaporation. Such an adaptation helps conifers
a lower reflectivity than snow, ice, or tundra vegetation, withstand the drought of the northern winter, when roots
causing additional warming, an example of a positive feed- cannot absorb water through the frozen ground. Being
back mechanism. In addition, the permafrost is evergreen, conifers resume photosynthesis as
melting, and boreal forest is moving northward, boreal forest A soon as warmer temperatures return.
two other results of a warming climate. region of coniferous The animal life of the boreal forest con-
forest (such as pine,
sists of some larger species such as caribou,
spruce, and fir) in the
which migrate from the tundra for winter;
Boreal Forest Northern Hemisphere;
wolves; brown and black bears; and moose.
located just south of
Just south of the tundra is the boreal forest, or However, most boreal mammals are medium
the tundra. Also called
northern coniferous forest (also called taiga, taiga. sized to small, including rodents, rabbits, and
pronounced tiéguh). Boreal forest stretches smaller predators such as lynx, sable, and mink.
Boreal forest • Figure 6.5

These coniferous forests occur in cold regions of the Northern Hemisphere adjacent to the tundra. Photographed in Saskatchewan, Canada.
Climate graph shows monthly temperatures and precipitation for Fort Smith, Northwest Territories, Canada.
RAYMOND GEHMAN/NG Image Collection

20
15
10
5
0
–5
–10
–15
–20
–25
–30
14
12
10
8
6
4
2
0
J FMAM J J A SOND
Months
I nterpret the D ata

Do changes in average monthly temperatures in this boreal forest
correspond to any noticeable changes in average monthly precipitation?
If so, what are these trends?

20

15
10
5

0
–5
–10
22

20
18
16
14
12
10
8
6
4
2
0
J FMAM J J A SOND
Months
Sarah Leen/NG Image Collection

In t e r p r e t t h e Da t a
What is the range of average monthly temperatures in
this temperate rain forest? How does this compare to the
Temperate rain forest • Figure 6.6 range of average monthly temperatures in the temperate
This temperate biome has large amounts of precipitation. Photographed deciduous forest?
in the Hoh Rain Forest in Washington State. Climate graph shows
monthly temperatures and precipitation for Estacada, Oregon.
Birds are abundant in the summer but migrate to proximity of temperate rain forest to the coastline moder-
warmer climates for winter. Insects are plentiful, but few ates its temperature so that the seasonal fluctuation is nar-
amphibians and reptiles occur except in the southern row; winters are mild, and summers are cool. Temperate
boreal forest. rain forest has relatively nutrient-poor soil, though its
Most of the boreal forest is not well suited to agricul- organic content may be high. Cool temperatures slow
ture because of its short growing season and mineral-poor the activity of bacterial and fungal decomposers. Thus,
soil. However, the boreal forest yields lumber, pulpwood needles and large fallen branches and trunks accumulate
for paper products, animal furs, and other forest prod- on the ground as litter that takes many years to decay and
ucts. Currently, boreal forest is the world’s top source of release nutrient minerals to the soil.
industrial wood and wood fiber. Extensive logging, gas The dominant vegetation in the North American
and oil exploration, mining, and farming have contrib- temperate rain forest is large evergreen trees such as west-
uted to loss of boreal forest. ern hemlock, Douglas fir, western red cedar, Sitka spruce,
and western arborvitae (Figure 6.6). Temperate rain for-
Temperate Rain Forest ests are rich in epiphytes, smaller plants that grow on the
A coniferous temperate rain forest occurs on trunks and branches of large trees. Epiphytes
the northwest coast of North America. Simi- temperate rain in this biome are mainly mosses, club mosses,
lar vegetation exists in southeastern Australia forest A coniferous lichens, and ferns, all of which also carpet the
biome with cool
and in southern South America. Annual pre- ground. Squirrels, wood rats, mule deer, elk,
weather, dense
cipitation in this biome is high—more than numerous bird species, and several species of
fog, and high
127 cm (50 in)—and is augmented by conden- amphibians and reptiles are common temper-
precipitation.
sation of water from dense coastal fogs. The ate rainforest animals.

The temperate rain forest is a rich wood producer, lose their leaves seasonally, such as oak, hickory, and
supplying lumber and pulpwood. Overharvesting the beech, dominate the temperate deciduous forests of the
original old-growth (never logged) forest can devastate northeastern and mideastern United States (Figure 6.7).
that biome because such an ecosystem takes hundreds In the southern areas of the temperate deciduous forest,
of years to develop. Once harvested, the old-growth the number of broad-leaved evergreen trees, such as mag-
forest ecosystem never has a chance to fully recover. nolia, increases.
Issues surrounding logging of old-growth temper- Temperate deciduous forests originally contained
ate rain forests of the Pacific Northwest are discussed a variety of large mammals, such as puma, wolves, and
in Case Study 13.1, “The Tongass Debate over Clear- bison, which are now absent. Other more common
Cutting.” animals include deer, bears, and many small mammals
and birds.
In Europe and North America, logging and land
Temperate Deciduous Forest clearing for farms, tree plantations, and cities de-
Hot summers and cold winters characterize the temper- stroyed much of the original temperate deciduous for-
ate deciduous forest, which occurs in temperate areas est. Where it has regenerated, temperate deciduous
where precipitation ranges from about 75 to 150 cm forest is often in a seminatural state that humans have
(30 to 60 in) annually. Typically, the soil of a temperate modified for recreation, livestock foraging, timber har-
deciduous forest consists of a topsoil rich in vest, and other uses. Many forest organisms
organic material and a deep, clay-rich lower temperate have successfully reestablished themselves in
layer. As organic materials decay, mineral ions deciduous forest these returning forests.
are released. Ions not absorbed by tree roots A forest biome that Worldwide, deciduous forests were among
occurs in temper-
leach (filter) into the clay. the first biomes converted to agricultural use.
ate areas with a
The trees of the temperate deciduous for- In Europe and Asia, many soils that originally
moderate amount of
est form a dense canopy that overlies saplings supported deciduous forests have been culti-
precipitation.
and shrubs. Broad-leaved hardwood trees that vated by traditional agricultural methods for
Temperate deciduous forest • Figure 6.7

The broad-leaved trees that dominate this biome are deciduous and shed their leaves before winter. Photographed in Germany Valley, West
Virginia. Climate graph shows monthly temperatures and precipitation for Nashville, Tennessee.
Don Johnston/Alamy
28
24
20
16
12
8
4
0
–4
14
12
10
8
6
4
2
0
J FMAM J J A SOND
Months

What is the range of average monthly
precipitation in this temperate
deciduous forest? How does this
compare to precipitation fluctuations
in the temperate rain forest?

28
24
20

© DLILLC/Corbis
16
12
8
4
0
–4
14
12
10
8
6
4
2
0
J FMAM J J ASO ND
Months
I n t e r p r e t t h e Da t a
When is the dry
season in temperate
grasslands? the wet
season?
Temperate grassland • Figure 6.8

Bison graze on mixed-grass prairie in Custer State Park, South Dakota. Climate graph shows monthly
temperatures and precipitation for Lawrence, Kansas.
thousands of years without a substantial loss in fertility. midwestern states. Several species of grasses that, under
During the 20th century, widely adopted intensive agri- favorable conditions, grow as tall as a person on horse-
cultural practices, along with overgrazing and deforesta- back dominate tallgrass prairies. The land was originally
tion, contributed to the degradation of some agricultural covered with large herds of grazing animals, such as bison,
lands. Most damage to farmland has happened since the pronghorn, and elk. The principal predators were
end of World War II. wolves, although in sparser, drier areas coyotes took their
place. Smaller animals included prairie dogs and their
predators (foxes, black-footed ferrets, and various birds
Temperate Grassland of prey), grouse, reptiles such as snakes and lizards, and
Summers are hot, winters are cold, and rainfall is often great numbers of insects.
uncertain in temperate grassland. Average annual precip- Shortgrass prairies are temperate grasslands that
itation ranges from 25 to 75 cm (10 to 30 in). receive less precipitation than moist tem-
Grassland soil has considerable organic temperate perate grasslands but more precipitation
material because the aboveground portions grassland A grassland than deserts. In the United States, short-
of many grasses die off each winter and con- with hot summers, grass prairies occur in parts of Montana,
tribute to the organic content of the soil, cold winters, and less Wyoming, South Dakota, and other mid-
while the roots and rhizomes (underground rainfall than is found western states. Grasses that grow knee high
stems) survive underground. Many grasses in the temperate or lower dominate shortgrass prairies. Plants
are sod formers—that is, their roots and deciduous forest grow less abundantly than in the moister
rhizomes form a thick, continuous under- biome. grasslands, and bare soil is occasionally ex-
ground mat. Although few trees grow except posed. Native grasses of shortgrass prairies are
near rivers and streams, grasses grow in great profusion drought-resistant.
in the deep, rich soil (Figure 6.8). Periodic wildfires The North American grassland, particularly the
help maintain grasses as the dominant vegetation in tallgrass prairie, was well suited to agriculture. More than
grasslands. 90 percent has vanished under the plow, and the re-
Moist temperate grasslands, also known as tall- maining prairie is so fragmented that almost nowhere
grass prairies, occur in the United States in parts of can you see what Native Americans experienced prior to
Illinois, Iowa, Minnesota, Nebraska, Kansas, and other the arrival of European settlers in the Midwest. Today,

the tallgrass prairie is considered North America’s rar- fertile. Wildfires occur naturally in this environment and
est biome. It is not surprising that the North American are particularly frequent in late summer and autumn.
Midwest, Ukraine, and other moist temperate grasslands Chaparral vegetation looks strikingly similar in differ-
became the breadbaskets of the world because they pro- ent parts of the world, even though the individual spe-
vide ideal growing conditions for crops such as corn and cies differ by location. A dense thicket of evergreen
wheat, which are also grasses. shrubs—often short, drought-resistant pine or scrub
oak trees that grow 1 to 3 m (3 to 10 ft) tall—usually
dominates chaparral. These plant species have evolved
Chaparral adaptations that equip them to live where precipitation
Some hilly temperate environments have mild winters is seasonal. During the rainy winter season, the envi-
with abundant rainfall combined ronment may be lush and green, and during the hot,
chaparral A biome with hot, dry summers. Such dry summer, the plants lie dormant. The hard, small,
with mild, moist Mediterranean climates, as they leathery leaves of trees and shrubs resist water loss.
winters and hot,
are called, occur not only in the Many plants are also specifically fire adapted and
dry summers;
area around the Mediterranean grow best in the months following a fire. Such growth is
vegetation is
Sea but also in the North Ameri- possible because fire releases into the soil the nutrient
typically small-leaved
can Southwest, southwestern minerals present in the aboveground parts of the plants
evergreen shrubs
and small trees.
and southern Australia, central that burned. The seeds and underground parts of plants
Chile, and southwestern South that survive fire make use of the newly availabile nutri-
Africa. On the mountain slopes of southern California, ent minerals and sprout vigorously during winter rains.
this Mediterranean-type biome is known as chaparral Mule deer, wood rats, chipmunks, lizards, and many spe-
(Figure 6.9). Chaparral soil is thin and often not very cies of birds are common animals of the chaparral. (For
Chaparral • Figure 6.9

Chaparral vegetation consists mainly of drought-resistant evergreen shrubs and small trees. Hot, dry summers and mild, rainy winters
characterize the chaparral. Photographed in California. Climate graph shows monthly temperatures and precipitation for Culver City, California.
Earl Scott/Photo Researchers, Inc.
25
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–20
–15
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Months

Using monthly averages, what is
the approximate average annual
precipitation in the California
chaparral?
Describing Earth’s Major Biomes 137

EnviroDiscovery 6.1
Using Goats to Fight Fires
David McNew/Getty Images

California has an average of
8000 wildfires each year, and they are
becoming increasingly expensive
and dangerous to manage because
many people are building homes and
living in the fire-vulnerable chaparral.
Yet the topography of chaparral is so
steep that firefighters often cannot
use mechanized equipment but must
transport equipment to fires with
helicopters. Afraid that prescribed burns
will get out of control, local governments
are increasingly trying an effective,
low-tech method to reduce the fuel
load: During the 6-month fire season,
goats are clearing hills around Oakland,
Berkeley, Monterey, and Malibu. Interest
in using goats for fire control is growing
as California’s long-term drought has led
to more wildfires.
A herd of 350 goats can clear an
entire acre of heavy brush in about
a day, but their use entails advance
organization and suppor t. Before
goats can remove hazardous dry fuels
from surrounding hillsides, botanists
must fence off small trees and rare or
endangered plants to keep the goats Goats prefer woody and weedy species, such as those common to chaparral.
from eating those plants. Goatherds
typically use dogs to help herd the goats. Responsible management Goats are an excellent tool for fire management because they
includes grazing the goats for the optimal amount of time—long preferentially browse woody shrubs and thick undergrowth—exactly
enough to reduce the threat of fire but not so long as to cause the fuel that causes disastrous fires. Fires that have occurred in areas
unnecessary erosion in the area. after goats have browsed there are much easier to contain.
more on the role fire plays in nature and on how hu- to explore an innovative way to remove vegetation
mans have disrupted this role, see Case Study 6.1 at the effectively.)
end of the chapter.)
The fires that occur in California chaparral are quite
costly to humans when they consume expensive homes Desert
built on the hilly chaparral landscape. Unfortunately, ef- Desert consists of dry areas found
forts to prevent the naturally occurring fires sometimes in both temperate (cold des- desert A biome in
which the lack of
backfire. Denser, thicker vegetation tends to accumu- erts) and subtropical or tropical
precipitation lim-
late over several years; then, when a fire does occur, it is regions (warm deserts). The low
its plant growth;
much more severe. Removing the chaparral vegetation, water vapor content of the desert
deserts are found in
whose roots hold the soil in place, causes other prob- atmosphere results in daily tem-
both temperate and
lems; witness the mudslides that sometimes occur dur- perature extremes of heat and
tropical regions.
ing winter rains in these areas. (See EnviroDiscovery 6.1 cold, so that a major change in

28
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CORDIER Sylvain/Hemis/Corbis
Desert • Figure 6.10

This desert landscape includes tough-leaved yuccas and spine-covered prickly In t e r p r e t t he Da t a
pear cacti. Desert inhabitants are strikingly adapted to the demands of their Would you be warm year-round in this desert? Do
environment. Photographed in the Valley of the Gods, Utah. Climate graph shows you think it could snow there? Why or why not?
monthly temperatures and precipitation for Reno, Nevada.
temperature occurs in each 24-hour period. Deserts amphibians (frogs and toads) and many reptiles, such as
vary greatly depending on the amount of precipitation the desert tortoise, Gila monster, and Mojave rattlesnake.
they receive, which is generally less than 25 cm (10 in) Desert mammals in North America include rodents such
per year. As a result of sparse vegetation, desert soil is as kangaroo rats, as well as mule deer and jackrabbits.
low in organic material but is often high in mineral con- Birds of prey, especially owls, live on the rodents and jack-
tent, particularly salts. rabbits, and even scorpions. During the driest months
Plant cover is so sparse in deserts that much of of the year, many desert animals tunnel underground,
the soil is exposed. Plants in North American deserts where they remain inactive.
include cacti, yuccas, Joshua trees, and sagebrush Humans have altered North American deserts in
(Figure 6.10). Desert plants are adapted to conserve several ways. People who drive across the desert in off-
water and as a result tend to have few, small, or no road vehicles inflict environmental damage. When the
leaves. Cactus leaves are modified into spines, which top layer of desert soil is disturbed, erosion occurs more
discourage herbivores. Other desert plants shed their readily and less vegetation grows to support native ani-
leaves for most of the year, growing only during the mals. Certain cacti and desert tortoises are rare as a result
brief moist season. of poaching. Houses, factories, and farms built in desert
Desert animals are typically small. During the heat areas require vast quantities of water, which is imported
of the day, they remain under cover or return to shel- from distant areas. Increased groundwater consumption
ter periodically, emerging at night to forage or hunt. In by many desert cities has caused groundwater levels to
addition to desert-adapted insects and arachnids (such as drop, particularly in southern Arizona and southwestern
tarantulas and scorpions), there are a few desert-adapted New Mexico.

Savanna African savanna (Figure 6.11). Large predators, such as
Savanna, a tropical grassland, is found in areas of low lions and hyenas, kill and scavenge the herds. In areas of
rainfall or intense seasonal rain- seasonally varying rainfall, the herds and their predators
fall with prolonged dry periods. savanna A tropical may migrate annually.
Temperatures in savannas vary grassland with widely Savanna in many places is being converted into
little throughout the year. Precip- scattered trees or rangeland for cattle and other domesticated animals.
itation is the overriding climate
clumps of trees. The problem is particularly serious in Africa, where
factor: Annual precipitation is 85 human populations are growing rapidly.
to 150 cm (34 to 60 in). Savanna soil is somewhat low in
essential nutrient minerals, in part because it is heavily
leached during rainy periods—that is, nutrient minerals Tropical Rain Forest
filter out of the topsoil. Although the African savanna Tropical rain forest occurs where temperatures are warm
is best known, savanna also occurs in South America, throughout the year and precipi-
western India, and northern Australia. tation occurs almost daily. The tropical rain forest
A lush, species-rich
Savanna has wide expanses of grasses interrupted by annual precipitation in a tropi-
forest biome that
occasional trees like the acacia, which bristles with thorns cal rain forest is typically between
occurs where the
to provide protection against herbivores (see Figure 200 and 450 cm (80 to 180 in).
climate is warm and
5.17b). Both trees and grasses have fire-adapted features, A related biome not described moist throughout
such as extensive underground root systems, that enable here is the tropical dry forest, where the year.
them to survive seasonal droughts as well as periodic fires. temperatures are also warm year-
Spectacular herds of herbivores such as antelope, round as in tropical rain forests, but annual precipitation
giraffe, elephants, wildebeest, and zebra occur in the is lower and more seasonal.
Savanna • Figure 6.11

Tropical grasslands such as this one, with widely scattered acacia trees, support large herds of grazing

24
animals and their predators. Photographed in Kilimanjaro National Park, Tanzania, with Mount 20
Kilimanjaro in the background. Climate graph shows monthly temperatures and precipitation for 16

Lusaka, Zambia. 12
8
4
22 0
20
18
16
14
12
10
8
6
4
2
0
J FMAM J J A SOND
Months
In t e r p r e t t h e Da t a
Is the variability in average monthly
temperatures in this savanna
comparable to the variability in
average monthly precipitation?
Why or why not?
© DLILLC/Corbis

28
24

20
16
12

8
4
48 0
44
40
36
32
28
24
20
16
12
8
4
0
J FMAM J J A SOND
Months
© Frans Lanting/Corbis
Tropical rain forest • Figure 6.12 In t e r p r e t t h e Da t a

What is the average monthly precipitation during
A view of lowland tropical rain forest along a river in the Danum Valley, Borneo, this tropical rain forest’s two wettest months?
Malaysia. Except at riverbanks, tropical rain forests have a closed canopy during its two driest months?
that admits little light to the rainforest floor. Climate graph shows monthly
temperatures and precipitation for Belem, Brazil.
Tropical rain forest commonly occurs in areas with The middle story, or canopy, which reaches a height of
ancient, highly weathered, mineral-poor soil. Little 30 to 40 m (100 to 130 ft), forms a continuous layer of
organic matter accumulates in such soils; because tem- leaves that lets in very little sunlight to support the smaller
peratures are high year-round, bacteria, fungi, and plants in the sparse understory. Only 2 to 3 percent of the
detritus-feeding ants and termites decompose organic light bathing the forest canopy reaches the forest under-
litter quite rapidly. Roots quickly absorb nutrient miner- story. Tropical rainforest trees support thick woody vines
als from the decomposing material. Tropical rain forests and extensive communities of epiphytic plants such as
are found in Central and South America, Africa, and ferns, mosses, orchids, and bromeliads.
Southeast Asia. Not counting bacteria and other soil-dwelling organ-
Of all biomes, the tropical rain forest is unexcelled isms, about 90 percent of tropical rainforest organisms
in species richness and variety (Figure 6.12). No single are adapted to live in the canopy. Rain forests shelter the
species dominates this biome. The trees are typically most abundant and varied insects, birds, reptiles, and
evergreen flowering plants. A fully developed tropical amphibians on Earth. Most rainforest mammals, such as
rain forest has at least three distinct stories, or layers, of sloths and monkeys, are adapted to live only in the trees
vegetation. The topmost story, or emergent layer, con- and rarely climb down to the ground, although some
sists of the crowns of very tall trees, some 50 m (164 ft) large, ground-dwelling mammals, including elephants,
or more in height, which are exposed to direct sunlight. are also found in rain forests.

Human population growth and industrial expan-
sion in tropical countries may spell the end of tropical
rain forests during the 21st century. Biologists know that
many rainforest species will become extinct before they 1. What is a biome?
are even identified and scientifically described. (See 2. How do you distinguish between temperate
Chapter 13 for more discussion of the ecological impacts rain forest and tropical rain forest? between
of rainforest destruction.) savanna and desert?
Aquatic Ecosystems
LEARNING OBJECTIVES
1. Summarize the important environmental factors fishes, turtles, and whales. Benthos are bottom-dwelling
that affect aquatic ecosystems. organisms that fix themselves to one spot (sponges and
2. Describe the various aquatic ecosystems, oysters), burrow into the sand (worms and clams), or
simply walk about on the bottom (crabs and aquatic
giving attention to the environmental
insect larvae).
characteristics of each.
T
he most fundamental division in aquatic
ecology is probably between freshwater and Freshwater Ecosystems
saltwater environments. Salinity, which is Freshwater ecosystems include lakes and ponds (standing-
the concentration of dissolved salts (such water ecosystems), rivers and streams (flowing-water
as sodium chloride) in a body of water, affects the kinds ecosystems), and marshes and swamps (freshwater
of organisms present in aquatic ecosystems, as does the wetlands). Specific abiotic conditions and character-
amount of dissolved oxygen. Water greatly interferes istic organisms distinguish each freshwater ecosystem.
with the penetration of light, so floating aquatic organ- Although freshwater ecosystems occupy only about
isms that photosynthesize must remain near the water’s 2 percent of Earth’s surface, they play an important
surface, and vegetation anchored to lake floors or role in the hydrologic cycle: They help recycle precipi-
streambeds will grow only in relatively shallow water. tation that flows into the ocean as surface runoff. (See
In addition, low levels of essential nutrient minerals Chapter 5 for a detailed explanation of the hydrologic
limit the number and distribution of organisms in cer- cycle.) Large bodies of fresh water help moderate daily
tain aquatic environments. In this section, we discuss and seasonal temperature fluctuations on nearby land
freshwater ecosystems only; because the immense ma- regions, and freshwater habitats provide homes for
rine environment is so critical to the environmental many species.
well-being of Earth, we devote an entire chapter to it Zonation is characteristic of standing-water ecosystems.
(see Chapter 11). A large lake has three zones: the littoral, limnetic,
Aquatic ecosystems contain three main ecological and profundal zones (see What a
categories of organisms: free-floating plankton, strongly Scientist Sees 6.1). The littoral zone standing-water
swimming nekton, and bottom-dwelling benthos. is a productive, shallow-water ecosystem A
Plankton are usually small or microscopic organisms. They area along the shore of a lake body of fresh water
tend to drift or swim feebly, so, for the most part, they are or pond. Emergent vegetation, surrounded by land
carried about at the mercy of currents and waves. Plank- such as cattails and bur reeds, as and whose water does
ton include phytoplankton, photosynthetic algae and cya- well as several deeper-dwelling not flow; a lake or a
nobacteria that form the base of most aquatic food webs, aquatic plants and algae, live in pond.
and zooplankton, animal-like organisms that feed on algae the littoral zone. Animals here
and cyanobacteria and are in turn consumed by newly include frogs, turtles, worms, crayfish and other crus-
hatched fish and other small aquatic organisms. Nekton taceans, insect larvae, and many fishes. The limnetic
are larger, more strongly swimming organisms such as zone is the open water beyond the littoral zone—that is,

WHAT A SCIENTIST SEES 6.1
Zonation in a Large Lake
Kathleen Revis/NG Image Collection
The zonation in Bear Lake, in Rocky Mountain National

Park, Colorado, is not apparent to a visitor.
A lake is a standing-water ecosystem surrounded by

land. The littoral zone is the shallow-water area around
the lake’s edge. The limnetic zone is the open, sunlit
water away from the shore. The profundal zone, under
the limnetic zone, is below where light penetrates.
Limnetic zone
Littoral zone
Profundal
zone
away from the shore. The limnetic zone extends down thermal stratification and the water layers mix. In fall,
as far as sunlight penetrates to permit photosynthe- as surface water cools, its density increases, and eventu-
sis. The main organisms of the limnetic zone are mi- ally it displaces the less dense, warmer, mineral-rich wa-
croscopic plankton. Larger fishes also spend most of ter beneath. The warmer water then rises to the surface
their time in the limnetic zone, although they may visit where it, in turn, cools and sinks. This process of cool-
the littoral zone to feed and reproduce. The deepest ing and sinking continues until the lake reaches a uni-
zone, the profundal zone, is beneath the limnetic zone of form temperature throughout. In the spring, surface
a large lake; smaller lakes and ponds typically lack a ice melts and surface water again sinks to the bottom,
profundal zone. Because light does not penetrate ef- resulting in a mixing of the layers. In summer, thermal
fectively to this depth, plants and algae do not live stratification occurs once again. The mixing of deeper,
there. Detritus drifts into the profundal zone from nutrient-rich water with surface, nutrient-poor water
the littoral and limnetic zones; bacteria decompose during the fall and spring turnovers brings essential nu-
this detritus. This marked zonation is accentuated trient minerals to the surface and oxygenated water to
by thermal stratification, in which the temperature the bottom.
changes sharply with depth. Human effects on lakes and ponds include eutro-
Temperate lakes undergo fall and spring turnovers, phication, which is nutrient enrichment of a body of
when changing surface temperatures break down the water with inorganic plant and algal nutrients like nitrates

Features of a typical river • Figure 6.13
G L O BAL Many of the world’s busiest cities are located
The river begins at at river mouths or along rivers in floodplains.
the source, often Can you name examples in your region or
high in the mountains elsewhere?
and fed by melting LOCAL
snows or glaciers.
© Frans Lanting/Corbis
Headwater streams
flow downstream
rapidly, often over
rocks (as rapids) or
bluffs (as waterfalls).
Waterfall
Along the way,
tributaries feed into
Rapids the river, adding to
Meanders
the flow.
The floodplain is the

Bends called relatively flat area on
meanders form either side of the
as the river’s course river that is subject to
levels out, so that flooding.
the river flows more
slowly and winds Near the ocean, the
from side to side. river may form a salt
marsh where fresh water
Mouth from the river and salt Floodplain
water from the ocean
mix.
The delta is a fertile,

low-lying plain at the
river’s mouth that forms
from sediments that the
slow-moving river deposits
as it empties into the
Ocean ocean.
a. A river flows from its source to the ocean. b. Aerial view of meanders in the Tambopata River, Peru.
and phosphates. Although eutrophication is a natural or warmer in winter than in adjacent parts of the stream
process, human activities often accelerate it, such as or river. The kinds of organisms found in flowing water
the runoff of agricultural fertilizers and discharge of vary greatly from one stream to another, depending pri-
treated or untreated sewage. Eutrophication of lakes is marily on the strength of the current. In streams with
discussed in detail in Chapter 10. fast currents, some inhabitants have adaptations such as
Flowing-water ecosystems are highly variable. The suckers, with which they attach themselves to rocks to
surrounding environment changes greatly between a riv- prevent being swept away. Some stream inhabitants have
er’s source and its mouth (Figure 6.13). Certain parts of flattened bodies to slip under or between rocks. Other
the stream’s course are shaded by inhabitants such as fish are streamlined and muscular
flowing-water eco- forest, while other parts are ex- enough to swim in the current.
system A freshwater
posed to direct sunlight. Ground- Human activities such as pollution and dam construc-
ecosystem such as
water may well up through tion have adverse impacts on rivers and streams. These
a river or stream in
sediments on the bottom in one activities damage wildlife habitat and threaten water sup-
which water flows in a
current.
particular area, making the water plies and fisheries. (See Chapter 10 for more discussion
temperature cooler in summer of the environmental effects of dams.)

freshwater Freshwater wetlands include services. When rivers flood their ecosystem
wetlands Lands that marshes, dominated by grass- banks, wetlands are capable of services Important
shallow fresh water like plants, and swamps, domi- holding or even absorbing the environmental ben-
covers for at least part nated by woody trees or shrubs excess water, thereby helping to efits such as clean
of the year; wetlands (Figure 6.14). Wetland soils are control flooding. The floodwater air, clean water, and
have a characteris- waterlogged for variable periods then drains slowly back into the fertile soil that the
tic soil and water- and are therefore anaerobic (with- rivers, providing a steady flow of natural environment
tolerant vegetation. out oxygen). They are rich in accu- water throughout the year. Wet- provides.
mulated organic materials, partly lands also serve as groundwater
because anaerobic conditions discourage decomposition. recharging areas. One of their most important roles is
With their productive plant communities, wet- to trap and hold pollutants in the flooded soil, thereby
lands provide excellent wildlife habitat for migratory cleansing and purifying the water.
waterfowl and other bird species, as well as for beaver, Although wetlands are afforded some legal protec-
otters, muskrats, and game fish. In addition to provid- tion, they are still threatened by pollution, development,
ing unique wildlife habitat, wetlands serve other im- agriculture, and dam construction. (See Chapter 10 for
portant environmental functions, known as ecosystem more on threats to freshwater ecosystems.)
Freshwater swamp • Figure 6.14

Freshwater swamps are inland areas covered by water and dominated by trees, such as bald cypress.
Photographed in Lake Martin, at the edge of the Atchafalaya Basin, Louisiana.
© Dan Leeth/Alamy Limited

Brackish Ecosystems: Estuaries Temperate estuaries usually feature salt marshes,
Where the ocean meets the land, there may be one of shallow wetlands in which salt-tolerant grasses grow
estuary A coastal
several kinds of ecosystems: a rocky (Figure 6.15a). Salt marshes perform many ecosystem
body of water, partly
shore, a sandy beach, an inter- services, including providing biological habitats, trap-
surrounded by land, tidal mud flat, or a tidal estuary. ping sediment and pollution, supplying groundwater,
with access to the Water levels in an estuary rise and and buffering storms by absorbing their energy, which
open ocean and a fall with the tides; salinity fluctu- prevents flood damage elsewhere.
large supply of fresh ates with tidal cycles, the time of Mangrove forests, the tropical equivalent of salt
water from a river. year, and precipitation. Salinity marshes, cover perhaps 70 percent of tropical coastlines
also changes gradually within the (Figure 6.15b). Like salt marshes, mangrove forests
estuary, from fresh water at the river entrance, to brackish provide valuable ecosystem services. Their interlacing
(somewhat salty) water, to salty ocean water at the mouth roots are breeding grounds and nurseries for several
of the estuary. Because estuaries undergo significant commercially important fishes and shellfish, such as
daily, seasonal, and annual variations in physical factors mullet, spotted sea trout, crabs, and shrimp. Mangrove
such as temperature, salinity, and depth of light penetra- branches are nesting sites for many species of birds,
tion, estuarine organisms must have a high tolerance for such as pelicans, herons, egrets, and roseate spoonbills.
changing conditions. Mangrove roots stabilize the submerged soil, thereby
Estuaries are among the most productive ecosystems preventing coastal erosion and providing a barrier
in the world. Their high productivity is brought about against the ocean during storms.
by nutrient transport from land, tidal action that rapidly Both salt marsh and mangrove forest ecosystems have
circulates nutrients and helps remove waste products, a experienced significant losses due to coastal development.
high level of light that penetrates the shallow water, and Salt marshes have been polluted—by countless ongoing
the many plants that form the base of a detritus food web. sources as well as oil spills—and turned into dumping
Estuaries • Figure 6.15

vixterd/istock/Getty Images
Photoshot/Alamy Limited
a. A salt marsh near Wrightsville Beach, North Carolina. b. A mangrove forest in Risong Bay, Palau, Micronesia, with an
underwater view of the prop root system. Mangrove roots grow into
deeper water as well as into mudflats that are exposed at low tide.
Many animals live among the mangroves’ complex root system.

grounds; mangrove forests have been logged unsustain-
ably and used as aquaculture sites. Some countries, such
as the Philippines, Bangladesh, and Guinea-Bissau, have
lost 70 percent or more of their mangrove forests.
1. Which environmental factors shape flowing-
Descriptions of terrestrial biomes and aquatic eco- water ecosystems? standing-water ecosystems?
systems focus on large-scale environmental patterns. The 2. How do the characteristics of a freshwater
chapter’s final two sections focus next on how populations wetland differ from those of an estuary? How
of organisms—and communities of those populations— does a mangrove swamp differ from a salt
adapt to environmental change. marsh?
Population Responses to Changing

Conditions over Time: Evolution
LEARNING OBJECTIVES
1. Define evolution. Since that time, scientists have accumulated an enor-
2. Explain the four conditions necessary for mous body of observations and experiments that sup-
evolution by natural selection to occur. port Darwin’s theory. Although biologists still do not
agree completely on some aspects of the evolutionary
3. Provide evidence that supports evolution. process, the concept that evolution by natural selec-
tion has taken place and is still occurring is now well
S
cientists think all of Earth’s remarkable vari- documented.
ety of organisms descended from earlier
species by a process known as evolution. Evo-
lution occurs at the population level; an indi-
vidual organism does not evolve within its lifespan. The Portrait of a young Charles Darwin
concept of evolution dates back to the time of Aristotle • Figure 6.16
(384–322 b.c.e.), but Charles Darwin (1809–1882), a 19th-
century naturalist, proposed the
evolution mechanism of evolution that to-
The cumulative day’s scientific community still ac-
genetic changes in cepts (Figure 6.16). As you will
populations that occur see, the environment—including
during successive factors that shape biomes and eco-
generations. systems—plays a crucial role in
Darwin’s theory of evolution.
It occurred to Darwin that in a population, inher-
JAMES L. STANFIELD/NG Image Collection

ited traits favorable to survival in a given environment
tended to be preserved over successive generations,
whereas unfavorable traits were eliminated. The result is
adaptation, an evolutionary modification that improves
the chance of survival and reproductive success of a spe-
cies in a given environment. Eventually the accumula-
tion of many adaptive modifications might result in a
new species.
Darwin proposed the theory of evolution by natural
selection in his monumental book The Origin of Species by
Means of Natural Selection, which was published in 1859.
Population Responses to Changing Conditions over Time: Evolution 147

Natural Selection
Evolution occurs through the process of natural selection.
As favorable traits increase in fre-
natural selection quency in successive generations,
The tendency of and as unfavorable traits decrease
better-adapted in- or disappear, the collection of
dividuals—those characteristics of a given popula-
with a combination tion changes. Natural selection is
of genetic traits the process by which successful
best suited to traits are passed on to the next
environmental
generation and unsuccessful ones
conditions—to
are weeded out. It consists of four
survive and
phenomena that occur in the
reproduce, increasing
natural world, which can be con-
their proportion in the
population.
sidered conditions necessary for
© Images & Stories/Alamy

natural selection to take place:
1. High reproductive capacity. Each species produces more

offspring than will survive to maturity. Natural popu-
lations have the reproductive potential to increase
their numbers continuously over time (Figure 6.17).
2. Limits on population growth, or a struggle for existence.

Only so much food, water, light, growing space, and High reproductive capacity and limits to
so on are available to a population, and organisms population growth • Figure 6.17
compete with one another for the limited resources A jawfish incubates eggs in his mouth. If all offspring of a jawfish
available to them. Because there are more individu- pair survived and in turn reproduced, reefs would be choked with
als than the environment can support, not all of an jawfish. Yet this fish species has not overrun the ocean because
organism’s offspring will survive to reproductive age, individuals must avoid predation and compete for limited resources.
including many of the fish yet to hatch in Figure 6.17. Photographed at Dimakya Island, Philippines.
Other limits on population growth include predators
and diseases.
3. Heritable variation. The individuals in a population offspring. Over time, enough changes may accumu-
exhibit variation. Each individual has a unique com- late in geographically separated populations (often
bination of traits, such as size, color, and ability to with slightly different environments) to produce new
tolerate harsh environments. Some traits improve the species (Figure 6.18).
chances of an individual’s survival and reproductive
success, whereas others do not. It is important to One premise on which Darwin based his theory of
remember that the variation necessary for evolution evolution by natural selection is that individuals transmit
by natural selection must be inherited so that it can traits to the next generation. However, Darwin could not
be passed to offspring. explain how this occurs or why individuals within a popu-
lation vary. Beginning in the 1930s and 1940s, biologists
4. Differential reproductive success. Individuals that possess combined the principles of genetics with Darwin’s the-
the most favorable combination of characteristics ory of natural selection. The resulting unified expla-
(those that make individuals better adapted to their nation of evolution is known as the modern synthesis
environment) are more likely than others to sur- (where synthesis refers to a combination of parts of previ-
vive, reproduce, and pass their traits to the next ous theories).
generation. Sexual reproduction is the key to natu- The modern synthesis explains Darwin’s observa-
ral selection: The best-adapted individuals are those tion of variation among offspring in terms of mutation,
that reproduce most successfully, whereas less-fit indi- or changes in DNA. Mutations provide the genetic vari-
viduals die prematurely or produce fewer or inferior ability on which natural selection acts during evolution.

PROCESS
✓ THE PLANNER
PROCESSDIAGRAM
Darwin’s finches • Figure 6.18
Charles Darwin was a ship’s naturalist on a 5-year voyage around the world. During an extended stay in the
Galápagos Islands off the coast of Ecuador, he studied the plants and animals of each island, including 14
species of finches.
2
Ancestral South
J. DFigurening/VIREO
Galápagos
species 1 America
Ancestral Islands
reaches the
species begins
Galápagos
in Ecuador.
Islands.
There is only
one finch
DIAGRAM
species now in Ecuador
Ecuador.
Grassquit finch (seeds)
Pacific
Ocean
FLPA/Alamy Limited
3
Modern Galápagos Islands
species
descend
from Pinta Marchena
ancestral
species. Genovesa
Large ground finch Santiago
(hard seeds)
Fernandina Santa Cruz

Adapted from figure 14.11 on p. 428
Image Collection
in B.W. Murck, B.J. Skinner, and

TIM LAMAN/NG
Santa Fe
San Cristobal D. Mackenzie. Visualizing Geology.
Isabela Tortuga Copyright 2008. This material is
reproduced with permission of
Small ground finch John Wiley & Sons, Inc.
Santa Maria
(soft seeds) Española
Eric Hosking/Science Source Images
© Christopher Vernon-Parry/Alamy
Tierbild Okapia/Science Source Images
© Images & Stories/Alamy
Woodpecker finch Medium ground finch Warbler finch (insects) Cactus finch
fi h ((cactus)
(insects) (moderate seeds)
4
The apparently related species on the Galápagos Islands have different beak shapes and different diets. Darwin reasoned that
finches that colonized from the mainland had changed as the birds, now geographically isolated from each other, adapted to
different diets.
T hi nk C ri ti c al l y
Which of the species shown have similar
beaks? Is this reflected in their diets?
Population Responses to Changing Conditions over Time: Evolution 149

Evidence for evolution
Environmental InSight • Figure 6.19
Human Horse Cat Bat
Humerus
Radius
Ulna
Metacarpals
and
Carpals phalanges
Adapted from Figure 15.13 on p. 244, in S. A. Alters and B. Alters, Biology:

Understanding Life, Hoboken, NJ: John Wiley and Sons, Inc. (2009).
© Jonathan Blair/Corbis
a. The Fossil Record. b. Comparative Anatomy.
Fossils deposited in rock layers, which can be dated, show how organisms Similarities among organisms demonstrate how they are
evolved over time. This well-preserved snake fossil
f from the Messel Pit, related. These similarities among ffour vertebrate limbs
a significant ffossil site near the village of Messel, Germanyy, dates from illustrate that, while proportions of bones have changed
47 million years ago. The ancient snake bears both similarities to and in relation to each organism’s’ way of liffe, the forelimbs
f
differences
ff from snake species living today. have the same basic bone structure.
Some new traits may be beneficial, whereas others may they strive to better understand certain aspects of evo-
be harmful or have no effect at all. As a result of natural lution, such as the role of chance and how quickly new
selection, beneficial strategies, or traits, persist in a popu- species evolve. As discussed in Chapter 1, science is an
lation because such characteristics make the individuals ongoing process, and information obtained in the future
that possess them well suited to thrive and reproduce. may require modifications to certain parts of the theory
In contrast, characteristics that make the individuals that of evolution by natural selection.
possess them poorly suited to their environment tend to The final section switches the focus to the response
disappear in a population. of communities to environmental changes.
A vast body of evidence supports evolution, most
of which is beyond the scope of this text. This evidence
includes observations from the fossil record, com-
parative anatomy, biogeography (the study of the geo-
graphic locations of organisms), and molecular biology 1. What is evolution?
(Figure 6.19). In addition, evolutionary hypotheses are
tested experimentally. 2. What four phenomena or conditions are the
Although virtually all biologists accept the principles basis of natural selection?
of evolution by natural selection based on such evidence, 3. Which types of evidence support evolution?

✓ THE PLANNER
c. Molecular Biology.
Human
u Pig Duck Rattlesnake
e Tuna Moth Y
Yeast The organisms pictured here all share a
particular enzyme, but in the course of
evolution, mutations have resulted in
changes in the gene that codes ffor that
enzyme. This diagram shows the nucleotide
Time
base differences in this gene among humans

and other organisms. Note that organisms
thought to be more closely related to humans
have ffewer diff
fferences than organisms that
are more distantly related to humans.
0 10 20 30 40 50 60 70
Number of nucleotide base diffe
f rences
Adapted from figure 15.19 on p. 247 in S.A. Alters and B. Alters Biology: Understanding Life. Copyright 2006.
This material is reproduced with permission of John Wiley & Sons, Inc.

Based on the diagram, which organism shown is most
closely related to the duck? Which is most distantly
related?
Community Responses to Changing

Conditions over Time: Succession
LEARNING OBJECTIVES
1. Define ecological succession. The actual mechanisms that underlie succession are
2. Distinguish between primary and secondary not clear. In some cases, it may be that a resident species
succession. modified the environment in some way, thereby making
it more suitable for a later species to colonize. It is also
A
community of organisms does not possible that prior residents lived there in the
spring into existence spontane- ecological first place because there was little competi-
ously. By means of ecological succession The pro- tion from other species. Later, as more inva-
succession, a given community de- cess of community sive species arrived, the original species were
velops gradually through a sequence of species. development over displaced.
time, which involves
Certain organisms colonize an area; over time, Ecologists initially thought that succession
species in one stage
others replace them, and eventually the replace- inevitably led to a stable and persistent com-
being replaced by
ments are themselves replaced by still other spe- munity, known as a climax community, such as a
different species.
cies. Ecologists first studied succession in three forest. But more recently, this traditional view
diverse ecosystems: an abandoned field, a northern fresh- has fallen out of favor. The apparent stability of a “climax”
water bog, and sand dunes. forest is probably the result of how long trees live relative
Community Responses to Changing Conditions over Time: Succession 151

to the human life span. It is now recognized that mature break apart the rock, beginning the process of soil forma-
climax communities are not in a state of stable equilibrium tion. Over time, mosses and drought-resistant ferns may
but rather in a state of continual disturbance. Over time, a replace the lichen community, followed in turn by tough
mature community changes in species composition and in grasses and herbs. Once enough soil accumulates, low
the relative abundance of each species, despite the fact shrubs may replace the grasses and herbs; over time, for-
that it retains an overall uniform appearance. est trees in several distinct stages would replace the shrubs.
Succession is usually described in terms of the changes Primary succession on bare rock from a pioneer commu-
in the plant species growing in a given area, although each nity to a forest community often occurs in this sequence:
stage of the succession may also have its own kinds of animals lichens → mosses → grasses → shrubs → trees.
and other organisms. Ecological succession is measured on The concept of succession was developed in the 1880s
the scale of tens, hundreds, or thousands of years, not the by Henry Cowles, who studied the process as it occurred
millions of years involved in the evolutionary timescale. on sand dunes along the shores of Lake Michigan, which
has been gradually shrinking since the last ice age. The
Primary Succession shrinking lake exposed new sand dunes that displayed a
Primary succession is the change in species composi- series of stages in the colonization of the land. As in many
tion over time in a previously uninhabited environment other lake and ocean shore areas, the Lake Michigan
(Figure 6.20). No soil exists when primary succession sand dune environment is severe, with temperatures
begins. Bare rock surfaces, such as recently formed volca- ranging from high during the day to low at night. Few
nic lava and rock scraped clean by glaciers, are examples plants could tolerate these stresses and the low nutrient
of sites where primary succession may take place. content of the sand making up the dunes.
Details vary from one site to another, but on bare rock, As Cowles observed, grasses are common pioneer
lichens are often the most important element in the pioneer plants on Great Lakes dunes. As the grasses extend over
community, which is the initial community that develops the surface of a dune, their roots hold it in place, helping
during primary succession. Lichens secrete acids that help to stabilize the dune surface. Mat-forming shrubs then
✓ THE PLANNER
PROCESS DIAGRAM
Primary succession on glacial moraine • Figure 6.20

During the past 200 years, glaciers have retreated in Glacier Bay, Th in k Cr it ica llyIs it possible for spruce trees to
Alaska. Although these photos were not taken in the same area,
grow directly on the rocks deposited by glaciers? Why
they show some of the stages of primary succession on glacial or why not?
moraine (rocks, gravel, and sand that a glacier deposits).
1 After a glacier’s retreat, lichens 2 At a later date, 3 Still later,

initially colonize the barren dwarf trees and spruces
landscape, followed by mosses shrubs colonize dominate the
and small shrubs. the area. community.
Martin Shields/Science Source Charles D. Winters/Science Source Mira/Alamy

✓ THE PLANNER
PROCESS DIAGRAM
Secondary succession on
an abandoned field in
North Carolina • Figure 6.21
Th in k C ri ti c al l y
Based on this
example, what types of plants would
you expect to find in a field 10 years
after it is abandoned?
1 2 3 4 5
Annual weeds Annual and Pine seedlings Young pine Mature hardwood
(crabgrass) perennial weeds and saplings forest and forest (oaks,
(horseweed, (shortleaf pine developing hickory)
broomsedge, and loblolly understory of
ragweed, and pine) hardwoods
aster)
1 2–4 5–15 25–50 150

Years after cultivation
invade to further stabilize the dune, followed by a suc- the fires—to a young forest of lodgepole pines, with some
cession of tree species over the course of many years. Pri- Douglas fir seedlings.
mary succession on sand dunes around the Great Lakes Biologists have studied secondary succession on aban-
might proceed in this sequence: grasses → shrubs → pop- doned farmland extensively (Figure 6.21). Although it
lars (cottonwoods) → pine trees → oak trees. takes more than 100 years for secondary succession to oc-
cur at a single site, a single researcher can study old-field
Secondary Succession succession in its entirety by observing different sites under-
going succession in the same general area. The biologist
Secondary succession is the change in species composi-
may examine county tax records to determine when each
tion that takes place after some disturbance destroys the
field was abandoned. Secondary succession on abandoned
existing vegetation; soil is already present. A clear-cut for-
farmland in the southeastern United States proceeds in
est, open areas caused by a forest fire, and abandoned
this sequence: crabgrass → horseweed, broomsedge, and
farmland are common examples of sites where secondary
other weeds → pine trees → hardwood trees.
succession occurs. During the summer of 1988, wildfires
burned approximately one-third of Yellowstone National
Park, a disaster that provided a chance for biologists to
study secondary succession in areas that were once for-
ests. Secondary succession in Yellowstone has occurred 1. What is ecological succession?
rapidly, moving from ash-covered forest floor and charred 2. How does primary succession differ from
trees, to lilies and other herbs, and—by ten years after secondary succession?
Community Responses to Changing Conditions over Time: Succession 153

CASE STUDY 6.1 ✓ THE PLANNER
Wildfires
A wildfire is any unexpected—and unwanted—fire that burns
in grass, shrub, and forest areas. Whether started by lightning U.S. acres burned in 2015, compared to 2001–2010 average.
or by humans, wildfires are an important environmental force
in many geographic areas, especially places with wet seasons 10,000,000
followed by dry seasons, such as chaparral. Vegetation that
Acreage burned (number of acres)

9,000,000 2015
grows during the wet season dries to tinder during the dry
8,000,000 2001–2010
season. After fire ignites the dry organic material, wind spreads (average)
the fire through the area. 7,000,000
At the peak of the wildfire season in the American West
6,000,000
and Southwest, an area prone to wildfires, hundreds of new
wildfires can break out each day. In 2015, dry conditions 5,000,000
triggered U.S wildfires that consumed 4.1 million hectares (10.1 4,000,000
million acres), an annual total of destruction that ranked first 3,000,000
in the past 16 years (since records have been kept), whereas
the number of fires reported in 2015 was the second least for 2,000,000
that period (see graph). Alaska and the Pacific Northwest in 1,000,000
particular experienced record wildfires (see photos). 0
Fires have several effects on the environment. First, Jan Feb Mar Apr May June July Aug Sep Oct Nov Dec
combustion frees minerals locked in dry organic matter. The Month
ashes left by fire are rich in potassium, phosphorus, calcium, Courtesy of National Oceanic and Atmospheric Administration (NOAA 2015)
and other nutrient minerals essential for

plant growth. Thus, vegetation flourishes
after a fire. Second, fire removes plant cover
and exposes the soil, which stimulates the
germination of seeds that require bare soil
and the growth of shade-intolerant plants.
Third, fire increases soil erosion because
it removes plant cover, leaving soil more
vulnerable to wind and water.
Fires were a part of the natural
environment long before humans appeared,
and many terrestrial ecosystems have
adapted to fire. Grasses adapted to wildfire
Jeff Schmaltz/NASA
Jeff Schmaltz/NASA
have underground stems and buds. After fire
kills the aboveground parts, the untouched
underground parts send up new sprouts.
Fire-adapted trees such as bur oak and
ponderosa pine have thick, fire-resistant June 14, 2015 September 1, 2015
bark; others, such as jack pine, depend
on fire for successful reproduction because Wildfires burned more than 5 million acres in Alaska in 2015.
the fire’s heat opens the cones and releases These two NASA satellite images show the differences in the
the seeds. Alaskan landscape over two months, when wildfires burned more than
5 million acres. Dark-red areas mark burn scars.
Human interference also affects the
frequency and intensity of wildfires, even when the goal is
fire prevention. When fire is excluded from a fire-adapted partly responsible for the massively destructive fires that have occurred
ecosystem, organic litter accumulates. As a result, when a fire there in recent years. Prescribed burning is an ecological management tool
does occur, it burns hotter and is much more destructive than that allows for controlled burning to reduce organic litter and suppress
ecologically helpful. Decades of fire suppression in the West are fire-sensitive trees in fire-adapted areas.

Summary ✓ THE PLANNER
1 Factors That Shape Biomes 128
1. A biome is a large, relatively distinct terrestrial region with

amount of precipitation. Temperate grassland is grassland
with hot summers, cold winters, and less rainfall than is found
in the temperate deciduous forest biome. Chaparral is a biome
with mild, moist winters and hot, dry summers; vegetation is
characteristic climate, soil, plants, and animals, regardless of where
it occurs; a biome encompasses many interacting ecosystems. typically small-leafed evergreen shrubs and small trees. Desert
Near the poles, temperature is generally the overriding climate is a biome in which the lack of precipitation limits plant growth;
factor in determining biome distribution, whereas in temperate deserts are found in both temperate and tropical regions.
and tropical regions, precipitation is more significant. Savanna is tropical grassland with widely scattered trees or
clumps of trees. Tropical rain forest is a lush, species-rich
forest biome that occurs where the climate is warm and moist
throughout the year.
2 Describing Earth’s Major Biomes 132
1. Tundra is the treeless biome in the far north that consists of

boggy plains covered by lichens and small plants such as mosses;
it has harsh, very cold winters and extremely short summers.
3 Aquatic Ecosystems 142
1. In aquatic ecosystems, important environmental factors include

Boreal forest is a region of coniferous forest in the Northern salinity, amount of dissolved oxygen, and availability of light for
Hemisphere, located just south of the tundra. Temperate rain photosynthesis.
forest is a coniferous biome with cool weather, dense fog,
and high precipitation. Temperate deciduous forest is a 2. Freshwater ecosystems include standing-water, flowing-
forest biome that occurs in temperate areas with a moderate water, and freshwater wetlands. A standing-water ecosystem is
a body of fresh water surrounded by land
and whose water does not flow, such as a lake or pond. A
flowing-water ecosystem is a freshwater ecosystem such as a
river or stream in which the water flows in a current. Freshwater
wetlands are marshes and swamps—lands that are covered by
shallow fresh water at least part of the year; wetlands have a
characteristic soil and water-tolerant vegetation. 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 a river.
Water in an estuary is brackish rather than truly fresh. Temperate
estuaries usually contain salt marshes, whereas tropical estuaries
are lined with mangrove forests.
4 Population Responses to Changing

28 Conditions over Time: Evolution 147

24
20
16 1. Evolution is the cumulative genetic changes in populations that

12 occur during successive generations.
8
4 2. Natural selection is the tendency of better-adapted
0 individuals—those with a combination of genetic traits best
–4
14 suited to environmental conditions—to survive and reproduce,
12 increasing their proportion in the population. Natural selection
10 is based on four observations established by Charles Darwin:
8 (1) Each species produces more offspring than will survive
6
4 to maturity. (2) Organisms compete with one another for the
2 resources needed to survive. (3) The individuals in a population
0 exhibit heritable variation in their traits. (4) Individuals with the
J FMA MJ J A SOND
most favorable combination of traits are most likely to survive
Months
and reproduce, passing their genetic traits to the next generation.
Summary 155

Rattlesnake
5
Human Pig Duck Tuna Moth Yeast
Community Responses to Changing
Conditions over Time: Succession 151
Time
1. Ecological succession is the process of community development

over time, which involves species in one stage being replaced by
different species.
2. Primary succession is the change in species composition over
0 10 20 30 40 50 60 70
time in an environment that was not previously inhabited by
organisms; examples include bare rock surfaces, such as recently
Number of nucleotide base differences
formed volcanic lava and rock scraped clean by glaciers. Secondary
succession is the change in species composition that takes place
3. Scientific evidence supporting evolution comes from the fossil after some disturbance destroys the existing vegetation; soil is
record, comparative anatomy, biogeography, and molecular already present. Examples include abandoned farmland and open
biology. areas caused by forest fires.
Key Terms
• biome 128 • estuary 146 • standing-water ecosystem 142
• boreal forest 133 • evolution 147 • temperate deciduous forest 135
• chaparral 137 • flowing-water ecosystem 144 • temperate grassland 136
• desert 138 • freshwater wetlands 145 • temperate rain forest 134
• ecological succession 151 • natural selection 148 • tropical rain forest 140
• ecosystem services 145 • savanna 140 • tundra 132
What is happening in this picture?

This 1994 image from Yellowstone
National Park shows young lodgepole
pines growing among trees burned in
massive 1988 wildfires.
• What community process is taking

place?
• What type of biome is pictured

here? What other biomes are
susceptible to fires? How do
humans increase the fire risk in
these biomes?
Fzançois Gohier/Science Source Images
156

Critical and Creative Thinking Questions
1. What two climate factors are most important in determining an 10. Although most salamanders have four legs, the aquatic
area’s characteristic biome? salamander shown below resembles an eel. It lacks hind limbs
and has very tiny forelimbs. Propose a hypothesis to explain
S u stai nabl e C i ti z en Q ues ti on how these salamanders evolved according to Darwin’s theory of
natural selection.
2. In which biome do you live? Where would you place your
biome in the figure below? What human-caused threats
are faced by your biome, and how might you help
Joseph T. Collins/Science Source Images

reduce them? Which other biomes might be affected
by your lifestyle, such as the foods you eat or other
resources you consume? Ar
ld
Co
cti
c
ture
Inc
ra
re
pe
as
tem
ing
ing
lat
itu
as
de
cre
De
11. How could you test the hypothesis you proposed in

Tro
question 10? What type of evidence might you produce?

p
t
Ho
ics
Wet Decreasing precipitation Dry 12. Which biome discussed in this chapter is depicted
by the information in the graph below? Explain your
answer.
3. What environmental factors are most important in determining
the kinds of organisms found in aquatic environments?
4. Distinguish between freshwater wetlands and estuaries and

between flowing-water and standing-water ecosystems. High
5. Name and compare temperate and tropical estuaries. What

types of plants are characteristic of each? Medium
6. During the mating season, male giraffes slam their necks

together in fighting bouts to determine which male is stronger Low
and can therefore mate with females. Explain how the long
necks of giraffes may have evolved, using Darwin’s theory of
evolution by natural selection.
Av rec
Av p
Le ine
7. Explain why evolution, by definition, cannot take place within

ve ra
er ipit
er er
p
te
m
m
ag at
ag at
l o ls
one individual and during that individual’s life span.

e ion
e ure
fs
an
an
oi
l
nu
nu
al
al
8. Describe the process and stages of ecological succession.
9. Which type of ecological succession do you think occurred

in the region surrounding Mount St. Helens after the volcano
erupted in 1980? Explain your choice by comparing primary and
secondary succession. ✓ THE PLANNER
Critical and Creative Thinking Questions 157

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6 Ecosystems

T he Everglades, a “river of grass” in the

c06_EcosystemsAndEvolution.indd 126 10/27/2016 9:19:01 PM

Analyze key features

c06_EcosystemsAndEvolution.indd 127 10/27/2016 9:19:03 PM

The world’s terrestrial biomes • Figure 6.1

c06_EcosystemsAndEvolution.indd 128 10/27/2016 9:19:03 PM

Two climate factors, temperature and

Costa Rica Tropical Rain Forest.

Rod Planck/Science Source Images

Michael Melford/NG Image Collection

RICHARD NOWITZ/NG Image Collection

Biomes diﬀer in the

Tropical Tropical dry Savanna Moist tropical Dry tropical

rain forest forest desert desert

Factors That Shape Biomes 129

c06_EcosystemsAndEvolution.indd 129 10/27/2016 9:19:21 PM

Paul Davis/The Global Land Cover Facility,

130 CHAPTER 6 Ecosystems and Evolution

c06_EcosystemsAndEvolution.indd 130 10/27/2016 9:19:30 PM

Factors That Shape Biomes 131

c06_EcosystemsAndEvolution.indd 131 10/27/2016 9:19:36 PM

Arctic tundra • Figure 6.4

Michael Melford/NG Image Collection

132 CHAPTER 6 Ecosystems and Evolution

c06_EcosystemsAndEvolution.indd 132 10/27/2016 9:19:38 PM

Boreal forest • Figure 6.5

RAYMOND GEHMAN/NG Image Collection

I nterpret the D ata

c06_EcosystemsAndEvolution.indd 133 10/27/2016 9:19:39 PM

Average monthly temperature in °C

Average monthly precipitation in cm

Based on data from www.worldclimate.com

Sarah Leen/NG Image Collection

134 CHAPTER 6 Ecosystems and Evolution

c06_EcosystemsAndEvolution.indd 134 10/27/2016 9:19:41 PM

Temperate deciduous forest • Figure 6.7

I nterpret the D ata

c06_EcosystemsAndEvolution.indd 135 10/27/2016 9:19:44 PM

Average monthly precipitation in cm

Temperate grassland • Figure 6.8

136 CHAPTER 6 Ecosystems and Evolution

c06_EcosystemsAndEvolution.indd 136 10/27/2016 9:19:45 PM

Chaparral • Figure 6.9

I nterpret the D ata

Describing Earth’s Major Biomes 137

c06_EcosystemsAndEvolution.indd 137 10/27/2016 9:19:47 PM

David McNew/Getty Images

138 CHAPTER 6 Ecosystems and Evolution

c06_EcosystemsAndEvolution.indd 138 10/27/2016 9:19:48 PM

Average monthly precipitation in cm

Desert • Figure 6.10

Describing Earth’s Major Biomes 139

c06_EcosystemsAndEvolution.indd 139 10/27/2016 9:19:54 PM

Savanna • Figure 6.11

Average monthly temperature in °C

Average monthly precipitation in cm

c06_EcosystemsAndEvolution.indd 140 10/27/2016 9:19:56 PM

Average monthly temperature in °C