Enger−Ross: Concepts in IV. Evolution and Ecology 14.

Ecosystem Organization © The McGraw−Hill

Biology, Tenth Edition and Energy Flow Companies, 2002

Ecosystem

14
Organization
and Energy Flow
CHAPTER 14

Chapter Outline
14.1 Ecology and Environment
14.2 The Organization of Ecological
Systems
14.3 The Great Pyramids: Energy,
Numbers, Biomass
The Pyramid of Energy • The Pyramid
of Numbers • The Pyramid
of Biomass
OUTLOOKS 14.1: Detritus Food Chains
14.4 Community Interactions 14.6 Succession
14.5 Types of Communities HOW SCIENCE WORKS 14.1: The Changing
Temperate Deciduous Forest • Grassland • Nature of the Climax Concept
Savanna • Desert • Boreal Coniferous Forest • 14.7 Human Use of Ecosystems
Temperate Rainforest • Tundra • Tropical
Rainforest • The Relationship Between
Elevation and Climate
OUTLOOKS 14.2: Zebra Mussels: Invaders
from Europe

Key Concepts Applications

Understand the nature of an ecosystem. • Identify biotic and abiotic environmental factors.
• Explain how energy is related to ecosystems.

Recognize the types of relationships that organisms have to each
other in an ecosystem.
• Appreciate that the relationships in an ecosystem are complex.
• Describe why plants are called producers.
• Identify the trophic levels occupied by herbivores and carnivores
and why they are called consumers.
• Appreciate the role of decomposers.

Understand that energy dissipates as it moves through • Explain why predators are more rare than herbivores.
an ecosystem.

Appreciate the difficulty of quantifying energy flow through • Understand the value of using a pyramid of numbers or a pyramid
ecosystems. of biomass as opposed to the pyramid of energy.

List characteristics of several different biomes.
• Explain why some plants and animals are found only in certain
parts of the world.
• Recognize the significance of temperature and rainfall to the kind of
biome that develops.
• Understand the concept of a climax community.

Understand the concept of succession. • Recognize that humans have converted natural climax ecosystems
to human use.
• Explain why a vacant lot becomes a tangle of plants.
• Describe what the final stages of succession will look like in a given
biome.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
14.1 Ecology and Environment
Today we hear people from all walks of life using the terms
ecology and environment. Students, homeowners, politi-
cians, planners, and union leaders speak of “environmental
issues” and “ecological concerns.” Often these terms are
interpreted in different ways, so we need to establish some
basic definitions.
Ecology is the branch of biology that studies the rela-
tionships between organisms and their environments. This is
a very simple definition for a very complex branch of sci-
ence. Most ecologists define the word environment very
broadly as anything that affects an organism during its life-
time. These environmental influences can be divided into
two categories. Other living things that affect an organism
are called biotic factors, and nonliving influences are called
abiotic factors (figure 14.1). If we consider a fish in a stream,
we can identify many environmental factors that are impor-
tant to its life. The temperature of the water is extremely
important as an abiotic factor, but it may be influenced by
the presence of trees (biotic factor) along the stream bank
that shade the stream and prevent the Sun from heating it.
Obviously, the kind and number of food organisms in the
stream are important biotic factors as well. The type of
material that makes up the stream bottom and the amount
of oxygen dissolved in the water are other important abiotic
factors, both of which are related to how rapidly the water is
flowing.
As you can see, characterizing the environment of an
organism is a complex and challenging process; everything
seems to be influenced or modified by other factors. A plant
is influenced by many different factors during its lifetime: the
Figure 14.1
Biotic and Abiotic Environmental Factors
(a) The woodpecker feeding its young in the
hole in this tree is influenced by several biotic
factors. The tree itself is a biotic factor as is the
disease that weakened it, causing conditions
that allowed the woodpecker to make a hole in
the rotting wood. (b) The irregular shape of the
trees is the result of wind and snow, both
abiotic factors. Snow driven by the prevailing
winds tends to “sandblast” one side of the tree
and prevent limb growth.
(a)
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Companies, 2002
Chapter 14 Ecosystem Organization and Energy Flow 237
types and amounts of minerals in the soil; the amount of sun-
light hitting the plant; the animals that eat the plant; and the
wind, water, and temperature. Each item on this list can be
further subdivided into other areas of study. For instance,
water is important in the life of plants, so rainfall is studied in
plant ecology. But even the study of rainfall is not simple. The
rain could come during one part of the year, or it could be
evenly distributed throughout the year. The rainfall could be
hard and driving, or it could come as gentle, misty showers of
long duration. The water could soak into the soil for later use,
or it could run off into streams and be carried away.
Temperature is also very important to the life of a
plant. For example, two areas of the world can have the
same average daily temperature of 10°C* but not have the
same plants because of different temperature extremes. In
one area, the temperature may be 13°C during the day and
7°C at night, for a 10°C average. In another area, the tem-
perature may be 20°C in the daytime and only 0°C at night,
for a 10°C average. Plants react to extremes in temperature
as well as to the daily average. Furthermore, different parts
of a plant may respond differently to temperature. Tomato
plants will grow at temperatures below 13°C but will not
begin to develop fruit below 13°C.
The animals in an area are influenced as much by abi-
otic factors as are the plants. If nonliving factors do not
favor the growth of plants, there will be little food and few
hiding places for animal life. Two types of areas that support
only small numbers of living animals are deserts and polar
regions. Near the polar regions of the earth, the low temper-
ature and short growing season inhibits growth; therefore,
* See the metric conversion chart inside the back cover for conversion to Fahrenheit.
(b)
 Enger−Ross: Concepts in IV. Evolution and Ecology
Biology, Tenth Edition
238 Part 4 Evolution and Ecology
there are relatively few species of animals with relatively
small numbers of individuals. Deserts receive little rainfall
and therefore have poor plant growth and low concentra-
tions of animals. On the other hand, tropical rainforests
have high rates of plant growth and large numbers of ani-
mals of many kinds.
As you can see, living things are themselves part of the
environment of other living things. If there are too many ani-
mals in an area, they can demand such large amounts of
food that they destroy the plant life, and the animals them-
selves will die. So far we have discussed how organisms
interact with their environments in rather general terms.
Ecologists have developed several concepts that help us
understand how biotic and abiotic factors interrelate in a
complex system.
14.2 The Organization
of Ecological Systems
Ecologists can study ecological relationships at several differ-
ent levels of organization. The smallest living unit is the indi-
vidual organism. Groups of organisms of the same species are
called populations. Interacting populations of different
species are called communities. And an ecosystem consists of
all the interacting organisms in an area and their interactions
with their abiotic surroundings. Figure 14.2 shows how these
different levels of organization are related to one another.
All living things require continuous supplies of energy
to maintain life. Therefore, many people like to organize liv-
ing systems by the energy relationships that exist among the
different kinds of organisms present. An ecosystem contains
several different kinds of organisms. Those that trap sunlight
for photosynthesis, resulting in the production of organic
material from inorganic material, are called producers.
Green plants and other photosynthetic organisms such as
algae and cyanobacteria are, in effect, converting sunlight
energy into the energy contained within the chemical bonds
of organic compounds. There is a flow of energy from the
Sun into the living matter of plants.
The energy that plants trap can be transferred through
a number of other organisms in the ecosystem. Because all of
these organisms must obtain energy in the form of organic
matter, they are called consumers. Consumers cannot cap-
ture energy from the Sun as plants do. All animals are con-
sumers. They either eat plants directly or eat other sources of
organic matter derived from plants. Each time the energy
enters a different organism, it is said to enter a different
trophic level, which is a step, or stage, in the flow of energy
through an ecosystem (figure 14.3). The plants (producers)
receive their energy directly from the Sun and are said to
occupy the first trophic level.
Various kinds of consumers can be divided into several
categories, depending on how they fit into the flow of energy
through an ecosystem. Animals that feed directly on plants
are called herbivores, or primary consumers, and occupy the
14. Ecosystem Organization © The McGraw−Hill
and Energy Flow Companies, 2002
second trophic level. Animals that eat other animals are
called carnivores, or secondary consumers, and can be subdi-
vided into different trophic levels depending on what ani-
mals they eat. Animals that feed on herbivores occupy the
third trophic level and are known as primary carnivores.
Animals that feed on the primary carnivores are known as
secondary carnivores and occupy the fourth trophic level.
For example, a human may eat a fish that ate a frog that ate
a spider that ate an insect that consumed plants for food.
This sequence of organisms feeding on one another is
known as a food chain. Figure 14.4 shows the six different
trophic levels in this food chain. Obviously, there can be
higher categories, and some organisms don’t fit neatly into
this theoretical scheme. Some animals are carnivores at some
times and herbivores at others; they are called omnivores.
They are classified into different trophic levels depending on
what they happen to be eating at the moment. If an organ-
ism dies, the energy contained within the organic compounds
of its body is finally released to the environment as heat by
organisms that decompose the dead body into carbon diox-
ide, water, ammonia, and other simple inorganic molecules.
Organisms of decay, called decomposers, are things such as
bacteria, fungi, and other organisms that use dead organisms
as sources of energy (Outlooks 14.1).
This group of organisms efficiently converts nonliving
organic matter into simple inorganic molecules that can be
used by producers in the process of trapping energy. Decom-
posers are thus very important components of ecosystems
that cause materials to be recycled. As long as the Sun sup-
plies the energy, elements are cycled through ecosystems
repeatedly. Table 14.1 summarizes the various categories of
organisms within an ecosystem. Now that we have a better
idea of how ecosystems are organized, we can look more
closely at energy flow through ecosystems.
14.3 The Great Pyramids: Energy,
Numbers, Biomass
The ancient Egyptians constructed elaborate tombs we call
pyramids. The broad base of the pyramid is necessary to
support the upper levels of the structure, which narrows to a
point at the top. This same kind of relationship exists when
we look at how the various trophic levels of ecosystems are
related to one another.
The Pyramid of Energy
A constant source of energy is needed by any living thing.
There are two fundamental physical laws of energy that are
important when looking at ecological systems from an
energy point of view. First of all, the first law of thermody-
namics states that energy is neither created nor destroyed.
That means that we should be able to describe the amounts
in each trophic level and follow energy as it flows through
successive trophic levels. The second law of thermodynamics
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

Chapter 14 Ecosystem Organization and Energy Flow 239

Populations

Communities

Ecosystems

Organism

Biosphere

Figure 14.2
Ecological Levels of Organization
Ecologists can look at the same organism from several different perspectives. Ecologists can study the individual activities of an organism, how
populations of organisms change, the interactions among populations of different species, and how communities relate to their physical
surroundings.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

240 Part 4 Evolution and Ecology

Hawk

Fourth Tertiary
trophic consumer
level Carnivore
Secondary
Third consumer Snake
trophic Carnivore
level
Primary
Second consumer
Decomposer trophic
level Herbivore
Mouse

Grass

First
trophic
Producer
level

Figure 14.3
The Organization of an Ecosystem
Organisms within ecosystems can be divided into several different trophic levels on the basis of how they obtain energy. Several different sets
of terminology are used to identify these different roles. This illustration shows how the different sets of terminology are related to one
another.

Table 14.1
ROLES IN AN ECOSYSTEM

Classification Description Examples

Producers Organisms that convert simple inorganic compounds into complex Trees, flowers, grasses, ferns, mosses,
organic compounds by photosynthesis. algae, cyanobacteria

Consumers Organisms that rely on other organisms as food.

Animals that eat plants or other animals.
Herbivore Eats plants directly. Deer, goose, cricket, vegetarian human,
many snails
Carnivore Eats meat. Wolf, pike, dragonfly
Omnivore Eats plants and meat. Rat, most humans
Scavenger Eats food left by others. Coyote, skunk, vulture, crayfish
Parasite Lives in or on another organism, using it for food. Tick, tapeworm, many insects

Decomposers Organisms that return organic compounds to inorganic compounds. Bacteria, fungi
Important components in recycling.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
Figure 14.4
Trophic Levels in a Food Chain
As one organism feeds on another organism, there is a flow of
energy from one trophic level to the next. This illustration shows six
trophic levels.
states that when energy is converted from one form to
another some energy escapes as heat. This means that as
energy passes from one trophic level to the next there will be
a reduction in the amount of energy in living things and an
increase in the amount of heat.
At the base of the energy pyramid is the producer
trophic level, which contains the largest amount of energy of
any of the trophic levels within an ecosystem. In an ecosys-
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Companies, 2002
Chapter 14 Ecosystem Organization and Energy Flow 241
tem, the total energy can be measured in several ways. The
total producer trophic level can be harvested and burned.
The number of calories of heat energy produced by burning
is equivalent to the energy content of the organic material of
the plants. Another way of determining the energy present is
to measure the rate of photosynthesis and respiration and
calculate the amount of energy being trapped in the living
material of the plants.
Because only the plants, algae, and cyanobacteria in the
producer trophic level are capable of capturing energy from
the Sun, all other organisms are directly or indirectly
dependent on the producer trophic level. The second trophic
level consists of herbivores that eat the producers. This
trophic level has significantly less energy in it for several rea-
sons. In general, there is about a 90% loss of energy as we
proceed from one trophic level to the next higher level.
Actual measurements will vary from one ecosystem to
another. Some may lose as much as 99%, while other more
efficient systems may lose only 70%, but 90% is a good rule
of thumb. This loss in energy content at the second and sub-
sequent trophic levels is primarily due to the second law of
thermodynamics. Think of any energy-converting machine; it
probably releases a great deal of heat energy. For example,
an automobile engine must have a cooling system to get rid
of the heat energy produced. An incandescent lightbulb also
produces large amounts of heat. Although living systems are
somewhat different, they must follow the same energy rules.
In addition to the loss of energy as a result of the sec-
ond law of thermodynamics, there is an additional loss
involved in the capture and processing of food material by
herbivores. Although herbivores don’t need to chase their
food, they do need to travel to where food is available, then
gather, chew, digest, and metabolize it. All these processes
require energy.
Just as the herbivore trophic level experiences a 90%
loss in energy content, the higher trophic levels of primary
carnivores, secondary carnivores, and tertiary carnivores
also experience a reduction in the energy available to them.
Figure 14.5 shows the flow of energy through an ecosys-
tem. At each trophic level, the energy content decreases by
about 90%.
The Pyramid of Numbers
Because it may be difficult to measure the amount of energy
in any one trophic level of an ecosystem, people often use
other methods to quantify the different trophic levels. One
method is to simply count the number of organisms at each
trophic level. This generally gives the same pyramid relation-
ship, called a pyramid of numbers (figure 14.6). Obviously
this is not a very good method to use if the organisms at the
different trophic levels are of greatly differing sizes. For
example, if you count all the small insects feeding on the
leaves of one large tree, you would actually get an inverted
pyramid.
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242 Part 4 Evolution and Ecology

OUTLOOKS 14.1

Detritus Food Chains

lthough most ecosystems receive energy directly from the
A Sun through the process of photosynthesis, some ecosys-
tems obtain most of their energy from a constant supply of dead
organic matter. For example, forest floors and small streams
receive a rain of leaves and other bits of material that small ani-
mals use as a food source. The small pieces of organic matter,
such as broken leaves, feces, and body parts, are known as detri-
tus. The insects, slugs, snails, earthworms, and other small ani-
mals that use detritus as food are often called detritivores. In the
process of consuming leaves, detritivores break the leaves and
other organic material into smaller particles that may be used by
other organisms for food. The smaller size also allows bacteria
and fungi to more effectively colonize the dead organic matter,
further decomposing the organic material and making it available
to still other organisms as a food source. The bacteria and fungi
are in turn eaten by other detritus feeders. Some biologists
believe that we greatly underestimate the energy flow through
detritus food chains.

Predators

Mold and
bacteria
eaters

Feces
and
smaller
particles

Bacteria
and
molds

Leaves and
other organic
material
Grazers
and
shredders
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Chapter 14 Ecosystem Organization and Energy Flow 243

Solar energy (sunlight) Figure 14.5

Energy Flow Through an Ecosystem
As energy flows from one trophic level to the
next, approximately 90% of it is lost. This means
Photosynthesis by producers that the amount of energy at the producer level
Bacteria must be ten times larger than the amount of
Algae energy at the herbivore level.
Plants

Herbivores

Primary carnivores

Secondary
carnivores

Decomposers

Heat

Figure 14.6
A Pyramid of Numbers
One of the easiest ways to quantify the various trophic levels in an ecosystem is to count the number of individuals in a small portion of the
ecosystem. As long as all the organisms are of similar size and live about the same length of time, this method gives a good picture of how
different trophic levels are related. (a) The relationship between grass and mice is a good example. However, if the organisms at one trophic
level are much larger or live much longer than those at other levels, our picture of the relationship may be distorted. (b) This is what happens
when we look at the relationship between forest trees and the insects that feed on them. A pyramid of numbers becomes inverted in this
instance.

4 mice
Thousands of
leaf-eating
insects

400
grass
plants
1
tree

(a) (b)
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244 Part 4 Evolution and Ecology

Human
50 kg

Pig
500 kg Zooplankton
40 g

Corn
5,000 kg

Algae
10 g

(a) (b)

Figure 14.7
A Pyramid of Biomass
Biomass is determined by collecting and weighing all the organisms in a small portion of an ecosystem. (a) This method of quantifying trophic
levels eliminates the problem of different-sized organisms at different trophic levels. However, it does not always give a clear picture of the
relationship between trophic levels if the organisms have widely different lengths of life. (b) For example, in aquatic ecosystems, many of the
small producers may divide several times per day. The tiny animals (zooplankton) that feed on them live much longer and tend to accumulate
biomass over time. The single-celled algae produce much more living material, but it is eaten as fast as it is produced and so is not allowed to
accumulate.

The Pyramid of Biomass
Because of the size-difference problem, many people like to
use biomass as a way of measuring ecosystems. Biomass is
usually determined by collecting all the organisms at one
trophic level and measuring their dry weight. This eliminates
the size-difference problem because all the organisms at each
trophic level are weighed. This pyramid of biomass also
shows the typical 90% loss at each trophic level. Although a
biomass pyramid is better than a pyramid of numbers in
measuring some ecosystems, it has some shortcomings.
Some organisms tend to accumulate biomass over long
periods of time, whereas others do not. Many trees live for
hundreds of years; their primary consumers, insects, gener-
ally live only one year. Likewise, a whale is a long-lived ani-
mal, whereas its food organisms are relatively short-lived.
Figure 14.7 shows two biomass pyramids.
14.4 Community Interactions
In the previous section we looked at ecological relationships
from the point of view of ecosystems and the way energy
flows through them. But we can also study relationships at
the community level and focus on the kinds of interactions
that take place among organisms.
As you know from the discussion in the previous sec-
tion, one of the ways that organisms interact is by feeding on
one another. A community includes many different food
chains and many organisms may be involved in several of the
food chains at the same time, so the food chains become
interwoven into a food web (figure 14.8). In a community,
the interacting food chains usually result in a relatively stable
combination of populations.
Although communities are relatively stable we need to
recognize that they are also dynamic collections of organ-
isms: As one population increases, another decreases. This
might occur over several years, or even in the period of one
year. This happens because most ecosystems are not con-
stant. There may be differences in rainfall throughout the
year or changes in the amount of sunlight and in the average
temperature. We should expect populations to fluctuate as
abiotic factors change. A change in the size of one popula-
tion will trigger changes in other populations as well.
Figure 14.9 shows what happens to the size of a population
of deer as the seasons change. The area can support 100 deer
from January through February, when plant food for deer is
least available. As spring arrives, plant growth increases. It is
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Chapter 14 Ecosystem Organization and Energy Flow 245

Figure 14.8
A Food Web
When many different food chains are interlocked with one another, a food web results. The arrows indicate the direction of energy flow.
Notice that some organisms are a part of several food chains—the great horned owl in particular. Because of the interlocking nature of the
food web, changing conditions may shift the way in which food flows through this system.

no accident that deer breed in the fall and give birth in the
spring. During the spring producers are increasing, and the
area has more available food to support a large deer popula-
tion. It is also no accident that wolves and other carnivores
that feed on deer give birth in the spring. The increased
available energy in the form of plants (producers) means
more food for deer (herbivores), which, in turn, means more
energy for the wolves (carnivores) at the next trophic level.
If numbers of a particular kind of organism in a com-
munity increase or decrease significantly, some adjustment
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Biology, Tenth Edition and Energy Flow Companies, 2002

246 Part 4 Evolution and Ecology

Figure 14.9 350

Annual Changes in Population Size 300

The number of organisms living in an area
varies during the year. The availability of food
250
is the primary factor determining the size of

Number of deer
the population of deer in this illustration, but
water availability, availability of soil nutrients, 200
and other factors could also be important. Birth
150 of
young
100
Lowest Increasing Highest Decreasing
50 food food food food
supply supply supply supply

0
Winter Spring Summer Fall

Figure 14.10
A Pond Community
Although a pond would seem to be an easy community to characterize, it interacts extensively with the surrounding land-based communities.
Some of the organisms associated with a pond community are always present in the water (fish, pondweeds, clams); others occasionally
venture from the water to the surrounding land (frogs, dragonflies, turtles, muskrats); still others are occasional or rare visitors (minks, heron,
ducks).

usually occurs in the populations of other organisms within
the community. For example, the populations of many kinds
of small mammals fluctuate from year to year. This results in
changes in the numbers of their predators or the predators
must switch to other prey species and impact other parts of
the community. As another example, humans have used
insecticides to control the populations of many kinds of
insects. Reduced insect populations may result in lower num-
bers of insect-eating birds and affect the predators that use
these birds as food. Furthermore the indiscriminate use of
insecticides often increases the populations of herbivorous,
pest insects because insecticides kill many beneficial predator
insects that normally feed on the pest, rather than just the
one or two target pest species.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
Polar ice cap Mediterranean scrub and woodland
Tundra Grassland
Boreal coniferous forest (taiga) Desert
Temperate deciduous forest Tropical rainforest
Figure 14.11
Biomes of the World
Major climatic differences determine the kind of vegetation that can live in a region of the world. Associated with specialized groups of plants
are particular kinds of animals. These regional ecosystems are called biomes.
Because communities are complex and interrelated, it is
helpful if we set artificial boundaries that allow us to focus
our study on a definite collection of organisms. An example
of a community with easily determined natural boundaries is
a small pond (figure 14.10). The water’s edge naturally
defines the limits of this community. You would expect to
find certain animals and plants living in the pond, such as
fish, frogs, snails, insects, algae, pondweeds, bacteria, and
fungi. But you might ask at this point, What about the plants
and animals that live right at the water’s edge? That leads us
to think about the animals that spend only part of their lives
in the water. That awkward-looking, long-legged bird wad-
ing in the shallows and darting its long beak down to spear a
fish has its nest atop some tall trees away from the water.
Should it be considered part of the pond community? Should
we also include the deer that comes to drink at dusk and
then wanders away? Small parasites could enter the body of
the deer as it drinks. The immature parasite will develop into
an adult within the deer’s body. That same parasite must
spend part of its life cycle in the body of a certain snail. Are
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Chapter 14 Ecosystem Organization and Energy Flow 247
Tropical seasonal forest
Savanna
Tropical thorn scrub and woodland
Mountain (snow and ice)
these parasites part of the pond community? Several animals
are members of more than one community. What originally
seemed to be a clear example of a community has become
less clear-cut. Although the general outlines of a community
can be arbitrarily set for the purposes of a study, we must
realize that the boundaries of a community, or any ecosys-
tem for that matter, must be considered somewhat artificial.
14.5 Types of Communities
Ponds and other small communities are parts of large
regional terrestrial communities known as biomes. Biomes
are particular communities of organisms that are adapted to
particular climate conditions. The primary climatic factors
that determine the kinds of organisms that can live in an
area are the amount and pattern of precipitation and
the temperature ranges typical for the region. The map in
figure 14.11 shows the distribution of the major biomes of
the world. Each biome can be characterized by specific
 Enger−Ross: Concepts in IV. Evolution and Ecology
Biology, Tenth Edition
248 Part 4 Evolution and Ecology
Figure 14.12
Temperate Deciduous Forest Biome
This kind of biome is found in parts of the world that have
significant rainfall (75–130 centimeters) and cold weather for a
significant part of the year when the trees are without leaves.
climate conditions, particular kinds of organisms, and char-
acteristic activities of the organisms of the region.
Temperate Deciduous Forest
The temperate deciduous forest covers a large area from the
Mississippi River to the Atlantic Coast, and from Florida to
southern Canada. This type of biome is also found in parts
of Europe and Asia. Temperate deciduous forests exist in
parts of the world that have moderate rainfall (75–130 cen-
timeters per year) spread over the entire year and a relatively
long summer growing season (130–260 days without frost).
This biome, like other land-based biomes, is named for a
major feature of the ecosystem, which in this case happens to
be the dominant vegetation. The predominant plants are
large trees that lose their leaves more or less completely dur-
ing the fall of the year and are therefore called deciduous
(figure 14.12). The trees typical of this biome are adapted to
conditions with significant precipitation and short mild win-
ters. Since the trees are the major producers and new leaves
are produced each spring, one of the primary consumers in
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Figure 14.13
Grassland (Prairie) Biome
This typical short-grass prairie of western North America is
associated with an annual rainfall of 30 to 85 centimeters. This
community contains a unique grouping of plant and animal species.
this biome consists of leaf-eating insects. These insects then
become food for a variety of birds that typically raise their
young in the forest during the summer and migrate to more
moderate climates in the fall. Many other animals like squir-
rels, some birds, and deer use the fruits of the trees as food.
Carnivores such as foxes, hawks, and owls eat many of the
small mammals and birds typical of the region. Another fea-
ture typical of the temperate deciduous forest is an abun-
dance of spring woodland wildflowers that emerge early in
the spring before the trees have leafed out. Of course,
because the region is so large and has somewhat different cli-
matic conditions in various areas, we can find some differ-
ences in the particular species of trees (and other organisms)
in this biome. For instance, in Maryland the tulip tree is one
of the state’s common large trees, while in Michigan it is so
unusual that people plant it in lawns and parks as a decora-
tive tree. Aspen, birch, cottonwood, oak, hickory, beech, and
maple are typical trees found in this geographic region. Typi-
cal animals of this biome are many kinds of leaf-eating
insects, wood-boring beetles, migratory birds, skunks, porcu-
pines, deer, frogs, opossums, owls, and mosquitoes (Out-
looks 14.2). In much of this region, the natural vegetation
has been removed to allow for agriculture, so the original
character of the biome is gone except where farming is not
practical or the original forest has been preserved.
Grassland
The biome located to the west of the temperate deciduous
forest in North America is the grassland or prairie biome
(figure 14.13). This kind of biome is also common in parts
of Eurasia, Africa, Australia, and South America. The rain-
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
OUTLOOKS 14.2
Zebra Mussels: Invaders from Europe
n the mid-1980s a clamlike organism called the zebra mussel,
I Dressenia polymorpha, was introduced into the waters of the
Great Lakes. It probably arrived in the ballast water of a ship from
Europe. Ballast water is pumped into empty ships to make them
more stable when crossing the ocean. Immature stages of the
zebra mussel were probably emptied into Lake St. Clair, near
Detroit, Michigan, when the ship discharged its ballast water to
take on cargo. This organism has since spread to many areas of
the Great Lakes and smaller inland lakes. It has also been discov-
ered in other parts of the United States including the mouth of
the Mississippi River. Zebra mussels attach to any hard surface
and reproduce rapidly. Densities of more than 20,000 individuals
per square meter have been documented in Lake Erie.
Mississippi
River
Illinois
River
New Orleans
The Spread of the Zebra Mussel
fall (30–85 centimeters per year) in grasslands is not ade-
quate to support the growth of trees and the dominant vege-
tation consists of various species of grasses. It is typical to
have long periods during the year when there is no rainfall.
Trees are common in this biome only along streams where
they can obtain sufficient water. Interspersed among the
grasses are many kinds of prairie wildflowers. The dominant
animals are those that use grasses as food; large grazing
mammals (bison and pronghorn antelope); small insects
(grasshoppers and ants); and rodents (mice and prairie dogs).
A variety of carnivores (meadowlarks, coyotes, and snakes)
feed on the herbivores. Most of the species of birds are sea-
sonal visitors to the prairie. At one time fire was a common
feature of the prairie during the dry part of the year.
© The McGraw−Hill
Companies, 2002
Chapter 14 Ecosystem Organization and Energy Flow 249
These invaders are of concern for several reasons. First,
they coat the intake pipes of municipal water plants and other
facilities requiring expensive measures to clean the pipes. Second,
they coat any suitable surface, preventing native organisms from
using the space. Third, they introduce a new organism into the
food chain. Zebra mussels filter small aquatic organisms from the
water very efficiently and may remove food organisms required
by native species. Their filtering activity has significantly increased
the clarity of the water in several areas in the Great Lakes. This
can affect the kinds of fish present, because greater clarity allows
predator fish to find prey more easily. There is concern that they
will significantly change the ecological organization of the Great
Lakes.
Lake
Superior
Upper Lake
Michigan Huron
Lower
Michigan
Lake
Michigan Lake St. Clair
Zebra mussel
introduced
Lake Erie
Today most of the original grasslands, like the temper-
ate deciduous forest, have been converted to agricultural
uses. Breaking the sod (the thick layer of grass roots) so that
wheat, corn, and other grains can be grown exposes the soil
to the wind, which may cause excessive drying and result in
soil erosion that depletes the fertility of the soil. Grasslands
that are too dry to allow for farming typically have been
used as grazing land for cattle and sheep. The grazing of
these domesticated animals has modified the natural vegeta-
tion as has farming in the moister grassland regions.
Savanna
A biome that is similar to a prairie is a savanna (figure 14.14).
Savannas are tropical biomes of central Africa, Northern
 Enger−Ross: Concepts in IV. Evolution and Ecology
Biology, Tenth Edition
250 Part 4 Evolution and Ecology
Figure 14.14
Savanna Biome
A savanna is likely to develop in areas that have a rainy season and
a dry season. During the dry season, fires are frequent. The fires kill
tree seedlings and prevent the establishment of forests.
Australia, and parts of South America that have distinct wet
and dry seasons. Although these regions may receive 100 cen-
timeters of rainfall per year there is an extended dry season of
three months or more. Because of the extended period of dry-
ness the dominant vegetation consists of grasses. In addition,
a few thorny, widely spaced drought-resistant trees dot the
landscape. Many kinds of grazing mammals are found in this
biome—various species of antelope, wildebeest, and zebras in
Africa; various kinds of kangaroos in Australia; and a large
rodent, the capybara, in South America. Another animal typi-
cal of the savanna is the termite, colonial insects that typically
build mounds above ground.
During the wet part of the season the trees produce
leaves, the grass grows rapidly, and most of the animals raise
their young. In the African savanna, seasonal migrations of
the grazing animals is typical. Many of these tropical grass-
lands have been converted to grazing for cattle and other
domesticated animals.
Desert
Very dry areas are known as deserts and are found through-
out the world wherever rainfall is low and irregular. Typi-
cally the rainfall is less than 25 centimeters per year. Some
deserts are extremely hot; others can be quite cold during
much of the year. The distinguishing characteristic of desert
biomes is low rainfall, not high temperature. Furthermore,
deserts show large daily fluctuations in air temperature.
When the Sun goes down at night, the land cools off very
rapidly because there is no insulating blanket of clouds to
keep the heat from radiating into space.
A desert biome is characterized by scattered, thorny
plants that lack leaves or have reduced leaves (figure 14.15).
14. Ecosystem Organization © The McGraw−Hill
and Energy Flow Companies, 2002
Figure 14.15
Desert Biome
The desert gets less than 25 centimeters of precipitation per year,
but it contains many kinds of living things. Cacti, sagebrush, lichens,
snakes, small mammals, birds, and insects inhabit the desert. All
deserts are dry, and the plants and animals show adaptations that
allow them to survive under these extreme conditions. In hot deserts
where daytime temperatures are high, most animals are active only
at night when the air temperature drops significantly.
Because leaves tend to lose water rapidly, the lack of leaves
is an adaptation to dry conditions. Under these conditions
the stems are green and carry on photosynthesis. Many of
the plants, like cacti, are capable of storing water in their
fleshy stems. Others store water in their roots. Although this
is a very harsh environment, many kinds of flowering plants,
insects, reptiles, and mammals can live in this biome. The
animals usually avoid the hottest part of the day by staying
in burrows or other shaded, cool areas. Staying underground
or in the shade also allows the animal to conserve water.
There are also many annual plants but the seeds only
germinate and grow following the infrequent rainstorms.
When it does rain the desert blooms.
Boreal Coniferous Forest
Through parts of southern Canada, extending southward
along the Appalachian and Rocky Mountains of the United
States, and in much of northern Europe and Asia we find com-
munities that are dominated by evergreen trees. This is the
taiga, boreal coniferous forest, or northern coniferous forest
biome (figure 14.16). The evergreen trees are especially
adapted to withstand long, cold winters with abundant snow-
fall. Typically the growing season is less than 120 days and
rainfall ranges between 40 and 100 centimeters per year. How-
ever, because of the low average temperature, evaporation is
low and the climate is humid. Most of the trees in the wetter,
colder areas are spruces and firs, but some drier, warmer areas
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
Figure 14.16
Boreal Coniferous Forest Biome
Conifers are the dominant vegetation in most of Canada, in a major
part of Russia, and at high elevations in sections of western North
America. The boreal coniferous forest biome is characterized by cold
winters with abundant snowfall.
have pines. The wetter areas generally have dense stands of
small trees intermingled with many other kinds of vegetation
and many small lakes and bogs. In the mountains of the west-
ern United States, pines trees are often widely scattered and
very large, with few branches near the ground. The area has a
parklike appearance because there is very little vegetation on
the forest floor. Characteristic animals in this biome include
mice, snowshoe hare, lynx, bears, wolves, squirrels, moose,
midges, and flies. These animals can be divided into four gen-
eral categories: those that become dormant in winter (insects
and bears); those that are specially adapted to withstand the
severe winters (snowshoe hare, lynx); those that live in pro-
tected areas (mice under the snow); and those that migrate
south in the fall (most birds).
Temperate Rainforest
The coastal areas of northern California, Oregon, Washing-
ton, British Columbia, and southern Alaska contain an
unusual set of environmental conditions that support a tem-
perate rainforest. The prevailing winds from the west bring
moisture-laden air to the coast. As the air meets the coastal
mountains and is forced to rise, it cools and the moisture
falls as rain or snow. Most of these areas receive 200 cen-
timeters (80 inches) or more precipitation per year. This
abundance of water, along with fertile soil and mild temper-
atures, results in a lush growth of plants.
Sitka spruce, Douglas fir, and western hemlock are typ-
ical evergreen coniferous trees in the temperate rainforest.
Undisturbed (old growth) forests of this region have trees as
old as 800 years that are nearly 100 meters tall. Deciduous
trees of various kinds (red alder, big leaf maple, black cot-
tonwood) also exist in open areas where they can get enough
© The McGraw−Hill
Companies, 2002
Chapter 14 Ecosystem Organization and Energy Flow 251
Figure 14.17
Tundra Biome
The tundra biome is located in northern parts of North America and
Eurasia. It is characterized by short, cool summers and long,
extremely cold winters. There is a layer of soil below the surface that
remains permanently frozen; consequently, no large trees exist in
this biome. Relatively few kinds of plants and animals can survive
this harsh environment.
light. All trees are covered with mosses, ferns, and other
plants that grow on the surface of the trees. The dominant
color is green because most surfaces have something photo-
synthetic growing on them.
When a tree dies and falls to the ground it rots in place
and often serves as a site for the establishment of new trees.
This is such a common feature of the forest that the fallen,
rotting trees are called nurse trees. The fallen tree also serves
as a food source for a variety of insects, which are food for a
variety of larger animals.
Because of the rich resource of trees, 90% of the origi-
nal temperate rainforest has already been logged. Many
areas have been protected because they are home to the
endangered northern spotted owl and marbled murrelet
(a seabird).
Tundra
North of the coniferous forest biome is an area known as the
tundra (figure 14.17). It is characterized by extremely long,
severe winters and short, cool summers. The growing season
is less than 100 days and even during the short summer
the nighttime temperatures approach 0°C. Rainfall is low
(10–25 centimeters per year). The deeper layers of the soil
remain permanently frozen, forming a layer called the
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

252 Part 4 Evolution and Ecology

permafrost. Because the deeper layers of the soil are frozen,

when the surface thaws the water forms puddles on the sur-
face. Under these conditions of low temperature and short
growing season, very few kinds of animals and plants can
survive. No trees can live in this region. Typical plants and
animals of the area are grasses, sedges, dwarf willow, and
some other shrubs, reindeer moss (actually a lichen), cari-
bou, wolves, musk oxen, fox, snowy owls, mice, and many
kinds of insects. Many kinds of birds are summer residents
only. The tundra community is relatively simple, so any
changes may have drastic and long-lasting effects. The tun-
dra is easy to injure and slow to heal; therefore we must
treat it gently. The construction of the Alaskan pipeline has
left scars that could still be there 100 years from now.

Tropical Rainforest
The tropical rainforest is at the other end of the climate spec-
trum from the tundra. Tropical rainforests are found primarily
near the equator in Central and South America, Africa, parts
of southern Asia, and some Pacific Islands (figure 14.18). The
temperature is high (averaging about 27°C), rain falls nearly
every day (typically 200–1,000 centimeters per year), and there
are thousands of species of plants in a small area. Balsa (a very
light wood), teak (used in furniture), and ferns the size of trees
are examples of plants from the tropical rainforest. Typically,
every plant has other plants growing on it. Tree trunks are
likely to be covered with orchids, many kinds of vines, and
mosses. Tree frogs, bats, lizards, birds, monkeys, and an Figure 14.18
almost infinite variety of insects inhabit the rainforest. These
forests are very dense, and little sunlight reaches the forest Tropical Rainforest Biome
The tropical rainforest is a moist, warm region of the world located
floor. When the forest is opened up (by a hurricane or the
near the equator. The growth of vegetation is extremely rapid. There
death of a large tree) and sunlight reaches the forest floor, the
are more kinds of plants and animals in this biome than in any
opened area is rapidly overgrown with vegetation. other.
Because plants grow so quickly in these forests, people
assume the soils are fertile, and many attempts have been
made to bring this land under cultivation. In reality, the soils warmest near the equator and become cooler as the poles are
are poor in nutrients. The nutrients are in the organisms, and approached. Similarly, air temperature decreases as elevation
as soon as an organism dies and decomposes its nutrients are increases. This means that even at the equator it is possible
reabsorbed by other organisms. Typical North American to have cold temperatures on the peaks of tall mountains.
agricultural methods, which require the clearing of large Therefore, as one proceeds from sea level to the tops of
areas, cannot be used with the soil and rainfall conditions of mountains, it is possible to pass through a series of biomes
the tropical rainforest. The constant rain falling on these that are similar to what one would encounter traveling from
fields quickly removes the soil’s nutrients so that heavy the equator to the North Pole (figure 14.19).
applications of fertilizer are required. Often these soils
become hardened when exposed in this way. Although most
of these forests are not suitable for agriculture, large
expanses of tropical rainforest are being cleared yearly
14.6 Succession
because of the pressure for more farmland in the highly pop- Each of the communities we have just discussed is relatively
ulated tropical countries and the desire for high-quality lum- stable over long periods of time. A relatively stable, long-
ber from many of the forest trees. lasting community is called a climax community (How Sci-
ence Works 14.1). The word climax implies the final step in
a series of events. That is just what the word means in this
The Relationship Between Elevation and Climate context because communities can go through a series of pre-
The distribution of terrestrial ecosystems is primarily related dictable, temporary stages that eventually result in a long-
to temperature and precipitation. Air temperatures are lasting stable community. The process of changing from one
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

Chapter 14 Ecosystem Organization and Energy Flow 253

are often referred to as pioneer organisms,

and the community is called a pioneer
5,500 community.
Permanent snow Lichens are frequently important in
pioneer communities. They are unusual
North Pole organisms that consist of a combination of
4,500 80° 90° algae cells and fungi cells—a combination
70° that is very hardy and is able to grow on
Tundra 65°
the surface of bare rock (figure 14.20).
60°
Boreal coniferous Because algae cells are present, the lichen
Pine Fir 3,600 forest 50° is capable of photosynthesis and can form
new organic matter. Furthermore, many
tiny consumer organisms can make use of
40° the lichens as a source of food and a shel-
Temperate
deciduous tered place to live. The action of the
forest lichens also tends to break down the rock
30°
surface upon which they grow. This frag-
25°
1,800 mentation of rock by lichens is aided by
the physical weathering processes of freez-
20°
ing and thawing, dissolution by water,
Broadleaf
evergreen
and wind erosion. Lichens also trap dust
particles, small rock particles, and the
10° dead remains of lichens and other organ-
isms that live in and on them. These
processes of breaking down rock and
0 0°
trapping particles result in the formation
Elevation Latitude Equator of a thin layer of soil.
in meters As the soil layer becomes thicker,
at equator
small plants such as mosses may become
established, increasing the rate at which energy is trapped
Figure 14.19 and adding more organic matter to the soil. Eventually, the
Relationship Between Elevation, Latitude, and Vegetation soil may be able to support larger plants that are even more
As one travels up a mountain, the climate changes. The higher the efficient at trapping sunlight, and the soil-building process
elevation, the cooler the climate. Even in the tropics tall mountains continues at a more rapid pace. Associated with each of the
can have snow on the top. Thus, it is possible to experience the producers in each successional stage is a variety of small ani-
same change in vegetation by traveling up a mountain as one would mals, fungi, and bacteria. Each change in the community
experience traveling from the equator to the North Pole. makes it more difficult for the previous group of organisms
to maintain itself. Tall plants shade the smaller ones they
replaced; consequently, the smaller organisms become less
type of community to another is called succession, and each common, and some may disappear entirely. Only shade-
intermediate stage leading to the climax community is known tolerant species will be able to grow and compete success-
as a successional stage or successional community. fully in the shade of the taller plants. As this takes place we
Two different kinds of succession are recognized: pri- can recognize that one stage has succeeded the other.
mary succession, in which a community of plants and ani- Depending on the physical environment and the avail-
mals develops where none existed previously, and secondary ability of new colonizing species, succession from this point
succession, in which a community of organisms is disturbed can lead to different kinds of climax communities. If the area
by a natural or human-related event (e.g., hurricane, vol- is dry, it might stop at a grassland stage. If it is cold and wet, a
cano, fire, forest harvest) and returned to a previous stage in coniferous forest might be the climax community. If it is
the succession. Primary succession is much more difficult to warm and wet, it may be a tropical rainforest. The rate at
observe than secondary succession because there are rela- which this successional process takes place is variable. In some
tively few places on earth that lack communities of organ- warm, moist, fertile areas the entire process might take place
isms. The tops of mountains, newly formed volcanic rock, in less than 100 years. In harsh environments, like mountain-
and rock newly exposed by erosion or glaciers can be said to tops or very dry areas, it may take thousands of years.
lack life. However, bacteria, algae, fungi, and lichens quickly Primary succession can also be observed in the progres-
begin to grow on the bare rock surface, and the process of sion from an aquatic community to a terrestrial community.
succession has begun. The first organisms to colonize an area Lakes, ponds, and slow-moving parts of rivers accumulate
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

254 Part 4 Evolution and Ecology

HOW SCIENCE WORKS 14.1

The Changing Nature of the Climax Concept

W hen European explorers traveled across the North Ameri-
can continent they saw huge expanses of land covered by
the same kinds of organisms. Deciduous forests in the East,
coniferous forests in the North, grasslands in central North
America, and deserts in the Southwest. These collections came to
be considered the steady-state or normal situation for those
parts of the world. When ecologists began to explore the way in
which ecosystems developed over time they began to think of
these ecosystems as the end point or climax of a long journey
beginning with the formation of soil and its colonization by a
variety of plants and other organisms.
As settlers removed the original forests or grasslands and
converted the land to farming, the original “climax” community
was replaced with an agricultural ecosystem. Eventually, as poor
farming practices depleted the soil, the farms were abandoned
and the land was allowed to return to its “original” condition. This
secondary succession often resulted in forests or grasslands that
resembled those that had been destroyed. However, in most
cases these successional ecosystems contained fewer species and
in some cases were entirely different kinds of communities from
the originals.
Ecologists recognized that there was not a fixed, predeter-
mined community for each part of the world and began to mod-
ify the way they looked at the concept of climax communities.
The concept today is a more plastic one. The term climax is still
used to talk about a stable stage following a period of change,
but ecologists no longer believe that land will eventually return to
a “preordained” climax condition. They have also recognized in
recent years that the type of climax community that develops
depends on many factors other than simply climate. One of these
is the availability of seeds to colonize new areas. Two areas with
very similar climate and soil characteristics may contain different
species because of the seeds available when the lands were
released from agriculture. Furthermore, we need to recognize that
the only thing that differentiates a “climax” community from a
successional one is the time scale over which change occurs. “Cli-
max” communities do not change as rapidly as successional
ones. However all communities are eventually replaced, as were
the swamps that produced coal deposits, the preglacial forests of
Europe and North America, and the pine forests of the northeast-
ern United States.
So what should we do with this concept? Although the cli-
max concept embraces a false notion that there is a specific end
point to succession, it is still important to recognize that there is
a predictable pattern of change during succession and that later
stages in succession are more stable and longer lasting than
early stages. Whether we call it a climax community is not really
important.

Small annual Perennial Grasses, shrubs,

Bare rock Lichens plants, lichens herbs, grasses shade-intolerant trees Shade-tolerant trees

Pioneer stages Intermediate stages Climax community

Hundreds of years

Figure 14.20
Primary Succession
The formation of soil is a major step in primary succession. Until soil is formed, the area is unable to support large amounts of vegetation. The
vegetation modifies the harsh environment and increases the amount of organic matter that can build up in the area. The presence of plants
eliminates the earlier pioneer stages of succession. If given enough time, a climax community may develop.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
1 2
4
3
Figure 14.21
Succession from a Pond to a Wet Meadow
A shallow pond will slowly fill with organic matter from producers in the pond. Eventually, a floating mat will form over the pond and grasses
will become established. In many areas this will be succeeded by a climax forest.
organic matter. Where the water is shallow, this organic
matter supports the development of rooted plants. In deeper
water, we find only floating plants like water lilies that send
their roots down to the mucky bottom. In shallower water,
upright rooted plants like cattails and rushes develop. The
cattail community contributes more organic matter, and the
water level becomes more shallow. Eventually, a mat of
mosses, grasses, and even small trees may develop on the
surface along the edge of the water. If this continues for per-
haps 100 to 200 years, an entire pond or lake will become
filled in. More organic matter accumulates because of the
large number of producers and because the depression that
was originally filled with water becomes drier. This will usu-
ally result in a wet grassland, which in many areas will be
replaced by the climax forest community typical of the area
(figure 14.21).
Secondary succession occurs when a climax community
or one of the successional stages leading to it is changed to
an earlier stage. For example, when land is converted to agri-
culture the original climax vegetation is removed. When
agricultural land is abandoned it returns to something like
the original climax community. One obvious difference
between primary succession and secondary succession is that
in the latter there is no need to develop a soil layer. Another
difference is that there is likely to be a reservoir of seeds
from plants that were part of the original climax community.
The seeds may have existed for years in a dormant state or
they may be transported to the disturbed site from undis-
© The McGraw−Hill
Companies, 2002
Chapter 14 Ecosystem Organization and Energy Flow 255
turbed sites that still hold the species typical of the climax
community for the region. If we begin with bare soil the first
year, it is likely to be invaded by a pioneer community of
weed species that are annual plants. Within a year or two,
perennial plants like grasses become established. Because
most of the weed species need bare soil for seed germination,
they are replaced by the perennial grasses and other plants
that live in association with grasses. The more permanent
grassland community is able to support more insects, small
mammals, and birds than the weed community could. If
rainfall is low, succession is likely to stop at this grassland
stage. If rainfall is adequate, several species of shrubs and
fast-growing trees that require lots of sunlight (e.g., birch,
aspen, juniper, hawthorn, sumac, pine, spruce, and dog-
wood) will become common. As the trees become larger, the
grasses fail to get sufficient sunlight and die out. Eventually,
shade-tolerant species of trees (e.g., beech, maple, hickory,
oak, hemlock, and cedar) will replace the shade-intolerant
species, and a climax community results (figure 14.22).
14.7 Human Use of Ecosystems
Most human use of ecosystems involves replacing the natural
climax community with an artificial early successional stage.
Agriculture involves replacing natural forest or prairie com-
munities with specialized grasses such as wheat, corn, rice,
and sorghum. This requires considerable effort on our part
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Biology, Tenth Edition and Energy Flow Companies, 2002

256 Part 4 Evolution and Ecology

Annual Grasses Shrubs Spruces Spruces Chestnut Chestnut Black Tulip

weeds and walnut poplar
other
perennials

Oak Oak
Immature oaks Maple
Pines

Previous
climax 1–2
community Plowed years 2–20 Hickory Hickory
destroyed field pioneer years Intermediate years Climax community

200 years (variable)

Figure 14.22
Secondary Succession on Land
A plowed field in the southeastern United States shows a parade of changes over time involving plant and animal associations. The general
pattern is for annual weeds to be replaced by grasses and other perennial herbs, which are replaced by shrubs, which are replaced by trees.
As the plant species change, so do the animal species.

because the natural process of succession tends toward the
original climax community. This is certainly true if remnants
of the original natural community are still locally available
to colonize agricultural land. Small woodlots in agricultural
areas of the eastern United States serve this purpose. Much
of the work and expense of farming is necessary to prevent
succession to the natural climax community. It takes a lot of
energy to fight nature.
Forestry practices often seek to simplify the forest by
planting single-species forests of the same age. This certainly
makes management and harvest practices easier and more
efficient, but these kinds of communities do not contain the
variety of plants, animals, fungi, and other organisms typi-
cally found in natural ecosystems.
Human-constructed lakes or farm ponds often have
weed problems because they are shallow and provide ideal
conditions for the normal successional processes that lead to
their being filled in. Often we do not recognize what a pow-
erful force succession is.
The extent to which humans use an ecosystem is often
tied to its productivity. Productivity is the rate at which an
ecosystem can accumulate new organic matter. Because
plants are the producers, it is their activities that are most
important. Ecosystems in which conditions are most favor-
able for plant growth are the most productive. Warm, moist,
sunny areas with high levels of nutrients in the soil are ideal.
Some areas have low productivity because one of the essen-
tial factors is missing. Deserts have low productivity because
water is scarce, arctic areas because temperature is low, and
the open ocean because nutrients are in short supply. Some
communities, such as coral reefs and tropical rainforests,
have high productivity. Marshes and estuaries are especially
productive because the waters running into them are rich in
the nutrients that aquatic photosynthesizers need. Further-
more, these aquatic systems are usually shallow so that light
can penetrate through most of the water column.
Humans have been able to make use of naturally pro-
ductive ecosystems by harvesting the food from them. How-
ever, in most cases, we have altered certain ecosystems
substantially to increase productivity for our own purposes.
In so doing, we have destroyed the original ecosystem and
replaced it with an agricultural ecosystem. For example,
nearly all of the Great Plains region of North America has
been converted to agriculture. The original ecosystem
included the Native Americans who used buffalo as a source
of food. There was much grass, many buffalo, and few
humans. Therefore, in the Native Americans’ pyramid of
energy, the base was more than ample. However, with the
exploitation and settling of America, the population in
North America increased at a rapid rate. The top of the
pyramid became larger. The food chain (prairie grass—
buffalo—human) could no longer supply the food needs of
the growing population. As the top of the pyramid grew, it
became necessary for the producer base to grow larger.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization
Biology, Tenth Edition and Energy Flow
10 kilograms
100 kilograms of grain of cow
10 kilograms
100 kilograms of grain of people
eating grain
Figure 14.23
Human Biomass Pyramids
Because approximately 90% of the energy is lost as energy passes from one
trophic level to the next, more people can be supported if they eat producers
directly than if they feed on herbivores. Much of the less-developed world is in
this position today. Rice, corn, wheat, and other producers provide the majority
of food for the world’s people.
Because wheat and corn yield more biomass for humans
than the original prairie grasses could, the settlers’ domestic
grain and cattle replaced the prairie grass and buffalo. This
was fine for the settlers, but devastating for the buffalo and
Native Americans.
In similar fashion the deciduous forests of the East
were cut down and burned to provide land for crops. The
crops were able to provide more food than did harvesting
game and plants from the forest.
Anywhere in the world where the human population
increases, natural ecosystems are replaced with agricultural
ecosystems. In many parts of the world, the human demand
for food is so large that it can be met only if humans occupy
the herbivore trophic level rather than the carnivore trophic
level. Humans are omnivores that can eat both plants and ani-
mals as food, so they have a choice. However, as the size of
the human population increases, it cannot afford the 90% loss
that occurs when plants are fed to animals that are in turn
eaten by humans. In much of the less-developed world, the
primary food is grain; therefore, the people are already at the
herbivore level. It is only in the developed countries that peo-
ple can afford to eat meat. This is true from both an energy
point of view and a monetary point of view. Figure 14.23
shows a pyramid of biomass having a producer base of
100 kilograms of grain. The second trophic level only has
10 kilograms of cattle because of the 90% loss typical when
energy is transferred from one trophic level to the next (90%
of the corn raised in the United States is used as cattle feed).
© The McGraw−Hill
Companies, 2002
Chapter 14 Ecosystem Organization and Energy Flow 257
The consumers at the third trophic level, humans
in this case, experience a similar 90% loss. There-
fore, only 1 kilogram of humans can be sustained
by the two-step energy transfer. There has been a
99% loss in energy: 100 kilograms of grain are
necessary to sustain 1 kilogram of humans.
1 kilogram
of people
Because much of the world’s population is
eating steak already feeding at the second trophic level, we
cannot expect food production to increase to the
extent that we could feed 10 times more people
than exist today.
It is unlikely that most people will be able
to fulfill all their nutritional needs by just eating
grains. In addition to calories, people need a cer-
tain amount of protein in their diets and one of
the best sources of protein is meat. Although pro-
tein is available from plants, the concentration is
greater from animal sources. Major parts of
Africa, Asia, and Latin America have diets that
are deficient in both calories and protein. These
people have very little food, and what food they
do have is mainly from plant sources. These are
also the parts of the world where human popula-
tion growth is most rapid. In other words, these
people are poorly nourished and, as the popula-
tion increases, they will probably experience
greater calorie and protein deficiency. This exam-
ple reveals that even when people live as con-
sumers at the second trophic level, they may still not get
enough food, and if they do, it may not have the protein
necessary for good health. It is important to point out that
there is currently enough food in the world to feed every-
one. The primary reasons for starvation are political and
economic. Wars and civil unrest disrupt the normal food-
raising process. People leave their homes and migrate to
areas unfamiliar to them. Poor people and poor countries
cannot afford to buy food from the countries that have a
surplus.
Many biomes, particularly the drier grasslands, cannot
support the raising of crops. However, they can still be used
as grazing land to raise livestock. Like the raising of crops,
grazing often significantly alters the original grassland
ecosystem. Some attempts have been made to harvest native
species of animals from grasslands, but the species primarily
involved are domesticated cattle, sheep, and goats. The sub-
stitution of the domesticated animals displaces the animals
that are native to the area and also alters the plant commu-
nity, particularly if too many animals are allowed to graze.
Even aquatic ecosystems have been significantly altered
by human activity. Overfishing of many areas of the ocean
has resulted in the loss of some important commercial
species. For example, the codfishing industry along the east
coast of North America has been destroyed by overfishing.
Pacific salmon species are also heavily fished and disagree-
ments among the countries that exploit these species may
cause the decline of this fishery as well.
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

258 Part 4 Evolution and Ecology

SUMMARY THINKING CRITICALLY

Ecology is the study of how organisms interact with their environ- Farmers are managers of ecosystems. Consider a cornfield in Iowa.
ment. The environment consists of biotic and abiotic components Describe five ways in which the cornfield ecosystem differs from the
that are interrelated in an ecosystem. All ecosystems must have a original prairie it replaced. What trophic level does the farmer fill?
constant input of energy from the Sun. Producer organisms are
capable of trapping the Sun’s energy and converting it into biomass.
Herbivores feed on producers and are in turn eaten by carnivores, CONCEPT MAP TERMINOLOGY
which may be eaten by other carnivores. Each level in the food
chain is known as a trophic level. Other kinds of organisms Construct two concept maps, one for each set of terms, to show
involved in food chains are omnivores, which eat both plant and relationships among the following concepts.
animal food, and decomposers, which break down dead organic
matter and waste products. biome herbivore
All ecosystems have a large producer base with successively carnivore pioneer organism
smaller amounts of energy at the herbivore, primary carnivore, and climax community primary succession
secondary carnivore trophic levels. This is because each time energy consumer producer
passes from one trophic level to the next, about 90% of the energy decomposer secondary succession
is lost from the ecosystem. A community consists of the interacting food chain trophic level
populations of organisms in an area. The organisms are interrelated food web
in many ways in food chains that interlock to create food webs.
Because of this interlocking, changes in one part of the community
can have effects elsewhere. KEY TERMS
Major land-based regional ecosystems are known as biomes.
The temperate deciduous forest, boreal coniferous forest, tropical abiotic factors omnivores
rainforest, grassland, desert, savanna, temperate rainforest, and biomass pioneer community
tundra are examples of biomes. Ecosystems go through a series of biomes pioneer organisms
predictable changes that lead to a relatively stable collection of biotic factors population
plants and animals. This stable unit is called a climax community, carnivores primary carnivores
and the process of change is called succession. climax community primary consumers
Humans use ecosystems to provide themselves with necessary community primary succession
food and raw materials. As the human population increases, most consumers producers
people will be living as herbivores at the second trophic level decomposers productivity
because they cannot afford to lose 90% of the energy by first feed- ecology secondary carnivores
ing it to a herbivore, which they then eat. Humans have converted ecosystem secondary consumers
most productive ecosystems to agricultural production and con- environment secondary succession
tinue to seek more agricultural land as population increases. food chain succession
food web successional community (stage)
herbivores trophic level

e—LEARNING CONNECTIONS www.mhhe.com/enger10

Topics Questions Media Resources

14.1 Ecology and Environment 1. Why are rainfall and temperature important in an Quick Overview
ecosystem? • Organisms and their environment
2. What is the difference between the terms ecosystem and
Key Points
environment?
• Ecology and environment
Animations and Review
• Introduction
Interactive Concept Maps
• Ecology

14.2 The Organization
of Ecological Systems
3. Describe the flow of energy through an ecosystem. Quick Overview
4. What role does each of the following play in an ecosys- • Trophic levels
tem: sunlight, plants, the second law of thermodynam-
Key Points
ics, consumers, decomposers, herbivores, carnivores,
• The organization of living systems
and omnivores?
 Enger−Ross: Concepts in IV. Evolution and Ecology 14. Ecosystem Organization © The McGraw−Hill
Biology, Tenth Edition and Energy Flow Companies, 2002

Chapter 14 Ecosystem Organization and Energy Flow 259

Topics Questions Media Resources

14.3 The Great Pyramids:
Energy, Numbers, Biomass
5. Give an example of a food chain. Quick Overview
6. What is meant by the term trophic level? • Modeling and measuring energy
7. Why is there usually a larger herbivore biomass than levels
a carnivore biomass? Key Points
8. Can energy be recycled through an ecosystem? Explain
• The great pyramids: Energy,
why or why not. numbers, biomass
Animations and Review
• Introduction
• Energy flow
Interactive Concept Maps
• Ecological pyramids

14.4 Community Interactions 9. What is the difference between an ecosystem Quick Overview
and a community? • Communities can’t stand alone
Key Points
• Community interactions

14.5 Types of Communities
10. List a predominant abiotic factor in each of the follow- Quick Overview
ing biomes: temperate deciduous forest, boreal conifer- • Biomes
ous forest, grassland, desert, tundra, temperate
Key Points
rainforest, tropical rainforest, and savanna.
• Types of communities
Animations and Review
• Introduction
• Climate
• Land biomes
• Aquatic systems
• Concept quiz
Interactive Concept Maps
• Temperature and moisture

14.6 Succession 11. How does primary succession differ from secondary Quick Overview
succession? • Predictable maturing of communities
12. How does a climax community differ from a succes-
Key Points
sional community?
• Succession
Animations and Review
• Introduction
• Organization
• Succession
• Biodiversity
• Concept quiz
Interactive Concept Maps
• Text concept map
Experience This!
• Trophic levels in the market

14.7 Human Use of Ecosystems Quick Overview

• Rolling back succession
Key Points
• Human use of ecosystems