Case Study

Mussels Muscle In
ON JANUARY 31 , 1989, the residents of Monroe,
Michigan , found themselves without water.
Monroe's schools, industries, and businesses were
closed for two days while workers labored to
resolve the problem. How can a town on the shore
of Lake Erie lack water7 The answer- the intake
pipes of its water treatment plant were clogged
with hundreds of millions of zebra mussels.
Zebra mussels be long toa large group of "two-
shelled," or "bivalve, " mollusks that includes clams
and scallops. Named for the striped pattern on their
shells, zebra mussels can range in size from barely
visible to about 2 inches long. Like other mussels,
they attach to surfaces using sticky threads.
Where did the mussels come from? About three
years earlier, a cargo ship had dumped its ballast
water in Lake St. Clair (between Lake Huron and
Lake Erie). The water, taken aboard in the Black Sea
of southeastern Europe, contained stowaways-
millions of microscopic zebra mussel larvae. In the
Great Lakes, these invaders found an ideal habitat
with plenty of food and no majar predators or
competitors. Their population exploded, spreading
to all of the Great Lakes, the Ohio and Mississippi
Rivers, and their tributaries. To make matters
worse, a few years after the zebra mussel arrived,
the quagga mussel (a clase relative of the zebra
mussel) was introduced by the same means and
began to spread in a similar manner.
8oth zebra mussels and quagga mussels
present serious ecological problems to the bodies
of water where they are introduced. As filter
feeders , the mussels remove large quantities of
phytoplankton, which are photosynthetic
microorganisms that form the basis of aquatic
food chains. As phytoplankton become scarce,
populations of the microscopic animals
(zooplankton) that feed on phytoplankton decline,
reducing the food available to small fish. lf small
fish populations decline, the larger fish that feed
on them will also become less abundant, and so
on up through the food chain. Thus, as zebra and
quagga mussels filter microscopic organisms from
the water, they alter the entire community
structure of the lake or river.
Think about the zebra and quagga mussels as
you read about the community interactions that
characterize healthy ecosystems. Why have these
unwelcome imports been so enormously
successful? What impact, if any, do they have on
one another? Can anything control them?

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 512 Behavior and Ecology

rAta Glance
Case Study Mussels Musc le In Earth Watch lnvasive Species Disrupt Community
lnteractions
27.1 Why Are Community lnteractions lmportant?
27.4 What ls Parasitism?
27.2 What ls the Relationship Between the
Scientific lnquiry A Parasite Makes Ants Berry
Ecological Niche and Competition? Appealing to Birds
The Ecologica l Nich e Defin es th e Pl ace and Ro le of Each Paras ites a nd Th eir Hosts Actas Agents ofNatural
Speci es in lts Ecosystem Selection on One Another
Competition Occurs WheneverTwo Organis ms Attempt
to Use the Sa me, Limited Resources
27.5 What ls Mutualism?
Adaptations Reduce the Overlap of Ecological Niches 27.6 How Do Keystone Species lnfluence
Among Coexisting Species Community Structure?
lnterspecific Co mpetition May Reduce th e Po pulation
Size and Dis tribution of Each Species 27.7 Succession: How Do Community
Competition Within a Species ls a Majar Factor lnteractions Cause Change over Time?
Controlling Popul ation Size Th ere Are Two Maja r Forms ofSuccess ion: Primary
Case Study Continued Musse ls Muscle In a nd Seconda ry
Succession Culminates in a Clímax Community
27.3 What Are the Results of lnteractions Between
Predators and Their Prey? So rn e Ecosystem s Are Mainta in ed in a Subclimax Stage
Clímax Communities Create Earth 's Biomes
Preda tor- Prey lnteractions Sha pe Evolutionary
Adaptations Case Study Revisited Mussels Musc le In

Case Study Continued Musse ls Muscle In

27.1 WHY ARE COMMUNITY lnteractions Amon Species

INTERACTIONS IMPORTANTI Type of Effect on Effect on
lnteraction Species A Species B
An ecological community consists of all the interacting pop-
ulations within an ecosystem. Because there are direct or in- Competition between A and B Harm s Harms

direct links between all forms of life in a given area, a Predation by A on B Benef its Harm s

community can encompass th e entire biotic, or living, por- Parasitism by A on B Be nefits Harms

tia n of an ecosystem. In Chapter 26, you learned that com- Mutualism between A and B Benefits Benefi ts

munity interactions such as competition, predation, and

parasitism can limit the size of populations. A community's
interacting web of life tends to maintain a balance between
resources and the numbers of individuals consuming them.
When populations interact and influence each other's
ability to survive and reproduce, they act as agents of natural
selection on one another. For example, in killing prey that is
easiest to catch, predators leave behind individuals with bet-
ter defenses against predatio n. These better-adapted individ-
uals produce the most offs pring. and over time, their inherited
characteristics increase within the prey population . Thus, as
community interactions limit population size, they simulta-
neously shape the bodies and behaviors of the interacting
populations. The process by which two interacting species act
as agents of natural selection on one another is called
coevolution.
The most important community interactions are com-
petitio n, predatio n, parasitism, and mu tualism. If we con-
sider these interactions as involving two different species,
they can be classified according to whether each of the
species is harmed or helped by the interacti on, as shown in
Ta ble 2 7 -1.
27.2 WHAT IS THE RELATIONSHIP
BETWEEN THE ECOLOGICAL NICHE
AND COMPETITION?
The concept of the ecological niche is im portant to our un-
derstanding of how competition within and between species
selects for adaptations in body form and behavior. Although
theword "niche" may call to mind a small cubbyhole, in ecol-
ogy, it means much more.
The Ecological Niche Defines the Place and Role
of Each Species in lts Ecosystem
Each species occupies a unique ecological niche that encom-
passes all aspects of its way oflife. One important aspect of the
ecological niche is the organism's physical home, or habitat.
The primary habitat of a white-tailed deer in the United States,
for example, is the eastern deciduous forest. In addition, an
ecological niche includes all the physical environmental con-
ditions necessary for the survival and reproduction of a given
species. These can indude nesting or denning sites, climate,

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 Community lnteractions 513

the type of nutrients the species requires, its optimal temper- P. aurelia
ature range, the amount ofwater it needs, the pH and salinity P. caudatum
of the water or soi l it m ay inhabit, and (for plants) the degree
:?;-
of sun or shade it can tolerate. Finally, the ecological niche ·¡¡; 200
e:
also encompasses the entire "role" that a given species per- Q)
150
forms within an ecosystem, including what it eats (or whether "oe:
it obtains energy from photosynthesis) and the other species ~ 100
3
with which it competes. Although different species share a. 50
o
many aspects of their niche with o thers, no two species ever a.
o
occupy exactly the same ecological niche within the same
community, as explained in the following sections.
o 2 4 6 8 1 o 12 14 16 18 20 22 24
days
(a) Grown in separate flasks
Competition Occurs Whenever Two Organisms
Attempt to Use the Same, Limited Resources
:?;-
Competition is an interaction that occurs between individuals ·¡¡; 200 t
e:
within a species or between individuals of different species as Q)

they attempt to use the same, limited resources, panicularly en- "oe: 150

ergy, nutrients, or space. The more the ecological niches of two
species overlap, the greater the amount ofcompetition between
them. Interspedfic competition refers to competitive interac-
tio ns between members of different species, such as occurs if
they feed on the same things or require similar breeding areas.
In terspecific competitio n is detrimental to all of tl1e species in-
volved because it reduces their access to resources that are in
limited supply. The degree of interspecific competitio n de-
pends o n how sim ilar the requirements of the species are.
Adaptations Reduce the Overlap of Ecological
Niches Among Coexisting Species
Just as no two organisms can occupy exactly the same physi-
cal space at the same time, no two species can inhab it exactly
the same ecological niche simultaneously and continuously.
This important concept, called the competitive exdusion
principie, was fo rmulated in 1934 by Russian biologist C. F.
Cause. This principie leads to the hypothesis that if a re-
searcher forces two species with the same niche to compete
for limited resources, inevitably, one will outcompete the
other, and the species that is less well adapted to the experi-
menta l conditio ns will die out.
Cause used two species of the protist Paramecium
(P. aurelia and P. caudatum) to demo nstrate this principie.
Crown separately in laboratory flasks, both species thrived
on the same bacteria and fed in the same regio n of their
flasks (Fig. 27-1a ) . But when Cause placed the two species
together, o ne (P. aurelia) always eliminated, o r "competi-
tively excluded," the other (P. caudatum; Fig. 27-1b). Cause
then repeated the experiment, replacing P. caudatum wi th a
different species, P. bursaria, which tended to feed in a differ-
ent pan of the flask. In that case, the two species of
Paramecium were able to coexist indefi nitely because they
occupied slightly d iffe rent niches.
Ecologist Roben MacAnhur funher explored the com-
petitive exclusio n princi pie, under natural conditi o ns, by
carefully observing five species of Nonh American warbler.
These birds all hunt fo r insects and nest in a variety of spruce
trees. Altho ugh the niches of these birds appear to overl ap
~ 100
3
a. 50
o
a.
o
o 2 4 6 8 1 o 12 14 16 18 20 22 24
days
(b) Grown in the same flask
Á FIGURE 2 7-1 Competitive exclusion (a) Raised separately
w it h a constant food su pp ly, both Paramecium aure/ia and
P. caudatum show the S-curve typ ical of a popu lation that
initially grows rapid ly and then stabi lizes. (b) Raised togethe r
and f orced to occupy the same niche, P. aurelia consistently
outcompetes P. caudatum and causes that population to die off.
Data from Cause, C.F. 1934. The Struggle for Existence. Balt imore:
Williams & Wi lkins.
QUESTION Explain how competitive exclusion could
contribute to the th reat posed by an invasive species.
considerably, MacA.rthur found that each species concentrates
its search for food in specific regions within spruce trees, em-
ploys different hunting tactics, and nests at a slightly different
time. By dividing up the resources provided by the spruce
trees they share, the warblers m inimize the overlap of their
niches and reduce interspecific competitio n ( Fig. 27-2) .
Had they not evolved in competition with one another,
each of the species of warbler that MacA.rthur observed woul d
probably have evolved to search throughout the entire spruce
tree for its spider and insect prey. With more food available,
each population would li kely have been larger. But when
species wi th similar ecological niches coexist and compete,
each species occupies a smaller niche than it would by itself.
This phenomeno n, called resource partitioning, is an evo-
lutio nary adaptation that reduces interspecific competition.
Resource partitioning is the outcome of the coevolution of
species with extensive (but not total) niche overlap. Individ-
uals with fewer competitors can utilize more resources and
leave more offspring. As a resu lt, over evolutionary time, com-
peting species evolve physical and behavioral adaptations
that reduce their competitive interactions.

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 514 Behavior and Ecology
Yellow-rumped Bay-breasted
warbler warbler
& FIGURE 27-2 Resource partitioning Each of these five insect-eating species of No rth American
warbl ers searches for food in sl ig htly different reg io ns within spruce trees. They reduce competition
by occupyi ng very sim ilar, but not ide ntical, niches . Data from MacArthur, R.H. 1958. Population ecology of
sorne warblers of Northeastern coniferous fores t. Ecology 39:599- 6 19.
Anoth er exam p le o f resource pa rtiti o ning was discov-
e red by Ch a rl es Darwin amo ng rela ted species of finches of
the Galáp agos Islands. Th e finches tha t sh ared the same is-
la nd h ad evolved different bill sizes a nd sh apes and different
feeding behaviors that reduced the competitio n am ong the m
(see p. 271).
lnterspecific Competition May Reduce the
Population Size and Distribution of Each Species
Altho ugh n atural selecti on leads toa reductio n of ni che over-
la p between di ffe re n t species, those w ith sim ila r niches still
compete for limited resources, causing bo th po pula tio ns to
be restri cted . A d assic study ofthe effects of inte rspecific com-
petitio n was performed by ecologist Joseph Connell, using
barnacles (sh elled crustacean s that a ttach perman ently to
rocks a nd othe r surfaces ).
Barnad es of th e genus Chthamalus sh are rocky ocean
shores with barnades of the genus Balanus, and their niches
overlap considerably. 8oth live in the intertidal zone, an a rea
o f the sh ore that is altemately covered and exposed by the tides.
Connell found that Chthamalus d o minates the upper shore and
Balanus do minates the lower. When he scraped off Balanus, the
Chthamalus populatio n increased, spreading d ownward into the
area that its competito r had once inhabited. Whe re the h abitat
is appropriate for both genera, Balanus conquers because it is
larger and grows faster. But Chthamalus to lerates drie r condi-
tio ns, giving it a competitive advantage on the upper sho re,
where o nly high tides submerge the bamades. As this exam p le
illustrates, inte rspecific competition can limit bo th the size and
the d istributio n of competing populatio ns.
Competition Within a Species Js a Major Factor
Controlling Population Size
lndividuals of the same species h ave th e same requirements
for resou rces an d thus occupy the same ecological niche. For
this reason, intraspedfic competition-competitio n a mo ng
individuals of the same species-is the most intense for m
of co mpetitio n. As explained in Ch apter 26, intra-specific
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Cape May Black-throated Blackburnian
warbler green warbler warbler
Case Study e o n ti n u e d
Mussels Muscle In
The zeb ra a nd q uagga mussels accide ntally impo rted from
Europe have niches that overla p exte nsive ly with those of
No rth American fresh water mu ssels and clams. Whe re the
Europea n mussels have invaded, th e indigenous
population s have declined. Both native and imported
species compete directly for space, but th e zebra and
qu agga mussels re produce far more rapidly, so th eir
populations have expanded at the ex pense of the native
mollusks.
In fact, the invasive mussel s literally cove r th e
natives; more tha n 10,000 zebra mussels have bee n found
attached to a single native mussel! Zebra and quagga
mu ssels also co mpete with nati ve biva lves for food, by
filtering the same types of mi croorga nisms from the water.
This interspecific competition has caused seve ral species of
native fres hwater cla ms to be nea rly eliminated from La ke
St. Clair and western La ke Erie.
com petitio n exerts stro ng density-d epen dent e nvironmental
resistance, limiting p opula tion size. The evolu tio n ary result
o f intraspecific co m petiti o n is t hat individuals wh o a re bet-
te r equi pped to obtain scarce resources a re more li kely to re-
produce successfully, passing their heritable traits to their
o ffsp ring.
27.3 WHAT ARE THE RESULTS OF
INTERACTIONS BETWEEN PREDATORS
AND THEIR PREY?
Predators eat oth er o rga nisms. Although we generally thin k
of pred ators as being carnivores (animals th at eat o the r ani-
mals), ecologists sometimes indude herbivores (anim als
th at eat p la n ts) in this gen eral category. He re, we define p re-
d ation in this mo re general way, to include the ze bra m ussel
th at filters microscopic photosynthetic protists fro m water,
 Community lnteractions 515

{b) Long-eared bat (e) Northern goshawk

& FIGURE 27-3 Forms of predation (a) A pi ka, whose preferred food is grass, is a small relative of
the rabbit and lives in the Rocky Mountains. (b) A long-eared bat uses a sophisticated echolocation
system to hunt moths, wh ich in turn have evolved special sound detectors and behaviors to avoid
ca pture. (e) A Northern gos hawk feasts on a smaller bird.
QUESTION Describe sorne examples of coevolutio n of predators and prey.

the grass-eating pika ( Fig. 27-3a), th e bat homing in o n a

moth ( Fig. 27-3b ), and the more familiar exa mple of a hawk
eating a smaller bird (Fig. 27-3c). Predators a re generally less
abundant than their prey; you w ill learn why in Chapter 28.
Predator-Prey lnteractions
Shape Evolutionary Adaptations
To survive, predators must feed and prey must avoid becom-
ing food . Therefo re, predator a nd prey populations exert in-
te nse selective pressure on one another, resulting in
coevolution . As prey become more difficult to catch, preda-
tors must become mo re adept at hunting. Coevolution h as
endowed the mountain !ion with tearing teeth and claws, a nd
has given the hunted fawn dappled spots that serve as camou-
fl age, as well as the behavior of lying still as it awaits its
mother's return from feed ing. Coevolution has produced the
keen eyesight of the hawk a n d owl, wh ich is countered by the
earthy colors of th eir m o use and ground squirrel p rey. Evolu-
tion under predation pressure has also produced the toxins of
the poi son dart frog, the coral sn ake, and milkweed (see
Figs. 27-7, 27-9 b, a nd 27-1 2b) .
In the following sections, we examine a few of th e evo-
lutio n ary results of predator-prey interactions. In "Earth
Watch: lnvasive Species Disrupt Community Interactions" on
p. 520, we describe w hat happe ns when natural ch ecks a nd
balances a re circumvented by transporting predatory or com-
peting o rganisms into ecological communities wh ose mem-
bers a re not ad apted to deal with the m .
Sorne Predators and Prey Have Evolved
Counteracting Behaviors
Bat and mo th adaptati o ns (see Fig. 27-3 b) provide excellent
examples of h ow both body structures a nd beh avio rs are
molded by coevolution. Most bats a re nighttime hunte rs th at
Case St udy e o n ti n u e d
Mussels Muse/e In
In 1990, a round goby was dis covered in the St. Clair River.
Like zebra and quagga mussels, this fi sh is native to
southeastern Europe, and probably arrived at the Great
Lakes in the same fashion as the mussels-in th e ballast
water of ships. Recognizing its natural prey, this 5-inch-
long predator began feasting on small zebra and quagga
mussels, rapidly expanding its range into all five of the
Great La kes.
ls the introduction of the invasive mus seis' natural
predator a solution to the mussel problem? Unfortunately,
no. The gobies ignore the largest mu ssels, which produce
the most eggs. Further, the gobies are not picky predators.
In addition to mussels, they will eat the eggs and young of
many fish, including native s mallmouth bass, walleye, and
perch.
e mit pulses of sound th at are so high pitched that people can't
hear them. Using the echoes that occur as their sounds bounce
back from n earby objects, bats create a sonar image of th eir
surroundings, which allows the m to n avigate and detect prey.
Under selection pressure from this unusual prey-locating
syste m, sorne mo ths (a favorite prey ofb ats ) have evolved ears
that are particularly sensitive to the pitches used by echol ocat-
ing ba ts. When they hear a bat, these moths take evasive action,
flying erra tically o r dro pping to the ground. The bats, in tum,
have evolved the ability to counter this defense by switching
the frequency of their sound pulses away from the moth's sen-
sitivity range. Sorne moths interfe re w ith the b ats' echolocation
by producing their own high-frequency dicks. In still another
coevolutionary adaptation, a bat that is hunting a dicking

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 & FIGURE 27-4 Camouflage by blending in (a) The sand dab is a flat , bottom-dwelling ocean fish
w ith a mottled color that closely re sembles the sand on w hich it rests. (b) Thi s nightjar bird on its
nest in Central America is barely visible among the surrounding leaf litte r.

(a) A camouflaged caterpillar {b) A camouflaged leafy sea dragon

(e) Camouflaged treehoppers (d) Camouflaged cacti

& FIGURE 27-5 Camouflage by resembling specific objects (a) A citrus swallowtail butterfly
caterpillar, whose color and shape resemble a bird dropping, sits motionless on a leaf. (b) The leafy
sea dragan (an Australian "seahorse" fish) has evo lved extensions of its body that mimi c the algae in
which it often hides. (e) Florida treehopper in sects avoid detection by resembling thorns on a branch.
(d) These cacti of the American Southwest are appropriately called "liv ing rock cacti. "
QUEST ION In genera l, how m ight such camouflage have evo lved?
. 516 . 1 . .
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 Community lnteractions 517

moth may tum off its own sound pulses temporarily (thus
avoiding detection) and follow the moth's clicks to capture it.

Camouflage Conceals 8oth Predators and Their Prey

An old saying goes that the best hiding place m ay be right out
in plain sight. Both predators and prey have evolved colors,
patterns, and shapes that resemble their surroundings. Such
disguises, called camouflage, render pla nts and animals in-
conspicuous, even when they a re in full view (Fig. 27-4).
Sorne animals closely resemble specific objects such
as leaves, twigs, seaweed1 thorns, or even bird droppings
(Figs. 27-Sa-c). Camouflaged animals tend to remain mo- (a) A camouflaged cheetah
tionless; a fleeing bird droppingwould be quite conspicuous.
Whereas many camouflaged animals resemble parts of
plants, a few d esert plants have evolved to resemble rocks,
which helps them to avoid predation by animals seeking the
water they store in their bodies (Fig. 27-Sd ).
Predators that ambush are also aided by camouflage.
For example, a spotted cheetah beco mes inconspicuou&in the
grass as it watches for grazing antelope. The frogfish closely
resembles the sponges and algae-covered rocks on which it
lurks, motionless, awaiting smaller fish to eat (Fig. 27-6 ).

Bright Colors Often Warn of Danger

Sorne animals have evolved very differently, exhibiting bright
warning coloration. These animals may be capable of in-
flicting a painful sting or may be bad-tasting and poisonous,
(b) A camouflaged frogfish
as is the poison dart frog (Fig. 27-7 ). The bright colors seem
to declare "Eat me at your own risk!" _.FIGURE 27-6 Camouflage assists predators (a) As it
waits for prey, a cheetah blends into the background of the
grass. (b) Combining camouflage and aggressive mimicry, a
Sorne Prey Organisms Gain Protection Through Mimicry frogfish waits in ambush, its camouflaged body matching the
Mimicry refers to a situation in which members of one sponge-encrusted rack on which it rests. Above its mouth
species have evolved to resemble another species. By sharing dangles a lure that closely resembles a small fish. The lure
a similar warning-color patte rn, severa! poisonous species attracts small predators , who will find themselves prey. The
frogfish can expand its mouth by a factor of l 2 in a few
may all benefit. Mimicry among different distasteful speóes thousands of a second, sucking in its prey.
is called Müllerian mimicry (after the German zoologist Jo-
h a nn Müller). For example, toxic monarch butterflies have
wing patterns strikingly similar to those of equally distasteful

Why some Wild Animals Freeze in the Midd le

of a Road as Your car Approaches?
Animals that are prey have evo lved not only camouflaged
coloring, but also behaviors that enhance their survival. In
response toa predator, they will often remain motionless
(making the camouflage more effective) and only bolt at
the last minute, when it becomes obvious that the
predator has spotted or smelled them. Your car,
resembling a very large predator, is likely to evoke both of
these instinctive behaviors. Although they work well in
natural situations, these responses increase the animal's _.FIGURE 27-7 Warning coloration The South American
chances of becoming roadkill. poison dart frog advertises its poisonous skin with bright and
contrasting color patterns.

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 518 Behavio r and Ecology

(b) Viceroy (distasteful)

A FIGURE 27-8 Müllerian mimicry Nearly ident ical warni ng coloration protect s b oth (a) t he
di stast eful m onarch and (b) t he equally d istastefu l viceroy butt erfl y.

(a) Bee (venomous) Hoverfly (nonvenomous)

(b) Coral snake (venomous) Scarlet king snake (nonvenomous)

A FIGURE 27-9 Batesi an mimicry (a) A bee , w hi ch is capable of sting ing (le ft), is m imicked b y the
sting less hoverfly (rig ht). (b) The warning colorat ion of t he ve nom ous co ral snake (left) is mimicke d
by the harmless scarlet kin g snake (r ight).

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 Community ln teractio ns 519

viceroy butterflies (Fig. 27-8) . Birds that become ill fro m con-
suming one species are more likely to avo id the other as well.
A toad that is stung while attempting to eat a bee is likely to
avoid not only bees, but other bl ack and yell ow striped in-
sects (sueh as yellow jacket wasps) without ever tasting one.
A shared color pattern thus helps all similarly colored species
avoid predation .
Once warning coloration evolved, there arose a selective
advantage fo r barmless animals to resemble venomous ones,
an adaptation called Batesian mimicry (after the English natu-
ralist Henry Bates). Through Batesian mimicry, the harmless
hoverfly avoids predation by resembling a bee (Fig. 27-9a), and
the nonvenomous scarlet king snake is protected by brilliant
warning coloration closely resembling that of the highly ven-
omous coral snake (Fig. 27-9b ).
Certain prey species use another form of mimicry:
s tartle coloration. Severa! insects and even sorne vertebrates
(such as the false-eyed frog) have evolved patterns of color
that closely resemble the eyes of a m uch larger, and possibly
dangerous, animal (Fig. 27-10) . Jf a predator gets close, the
prey suddenly flashes its eyespots, startling the predator and
allowing the prey to escape.
A sophisticated variation o n the theme of prey mim-
icking dangerous anima ls is seen in snowberry flies, which
are hun ted by te rri torial jumping spiders ( Fig. 27-11 ). When
a fly spo ts an approaching spider, it spreads its wings, mov-
ing them back and fo rth in a jerky dance. Seeing this display,
the spider is likely to flee th e fly. Why? Researchers have ob-
served that the markings on the fly's wings look like the legs
of a jumping spide r, and the fly's jerky movements mimic
the behavio r o f a jumping spider driving another spider
fro m its territory. Thus, natural selection has finely tuned
both the behavior and the ap pearance of the fl y to avo id pre-
dation by jumping spiders.

(a) False-eyed frogs (b) Peacock moth (e) Swallowtail caterpillar

& FIGURE 27-10 Startle coloration (a) When threatened , the false-eyed frog raises its rump , w hi ch
rese mbles the eyes of a large predator. (b) The peacock moth from Trinidad is well camouflaged , but
sho uld a predator app roach, it sud denly o pens its wi ngs to reveal spots rese mbling larg e eyes.
(e) Predators of t hi s cat erpill ar larva of t he Eastern tige r swallowtai l butterfly are dete rred by its
re se mblance toa snake. T he caterpill ar's head is th e "snake's" nose, and it bear s two sets of eyespot s.

(a) Jumping spider (predator) (b) Snowberry fly (prey)

& FIGURE 27-11 A prey mi mies its predator (a) When a jumping spide r approac he s, (b) t he
snowberry f ly spreads its w in gs, revealin g a pattern that rese mbles spid er legs. T he fly enhances the
effect by perform ing a jerky, side-to·s ide dance th at res emble s the leg -wavi ng display of a jumping
spider defending its territory.

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 lnvasive species are species that are introduced into an
ecosystem in wh ich they did not evolve, and that are
harmful to human health, the environment, or the economy
of a region . lnvas ive species often spread widely because
they find few effective forms of enviran mental resistance,
such as strong competitors, predators, or parasites, in their
new environment. Their unchecked population growth may
seriously damage the ecosystem as they outcompete or
prey on native species .
Not all non-native species become pests; the ones that do
typically reproduce rapidly, disperse widely, and thrive under
a relatively wide range of environmental conditions. lnvasive
plants often spread by sprouting from roots as well as seeds,
and sorne aquatic forms generate new plants from fragments .
lnvasive animals often eat a wide variety of foods. By evading
(a) Cane toad
the checks and balances imposed by millennia of coevolution,
invasive species are wreaking havoc on natural ecosystems
throughout the world. Sorne examples follow.
English house sparrows were introduced into the United
States on several occasions, starting in the 1850s, in the
hope that they wou ld control caterpillars feeding on shade
trees. In 1890, European starlings were released into Central
Park in New York City by a group attempting to introduce all
the birds mentioned in the works of Shakespeare. Both bird
species have spread throughout the continental United
States. Their success has reduced the populations of sorne
native songbirds, such as bluebirds and purple martins, with
which they compete for nesting sites. Red fire ants from
South America were accidentally introduced into Alabama on
shiploads of lumber in the 1930s and have since spread
throughout the southern United States. Fire ants kili native
ants, birds, and young reptiles. Their mounds can ruin farm
fields, and their fiery stings and aggressive temperament can
make backyards uninhabitable. lmported cane toads have
proven unstoppable as they spread through northeastern
Australia, outcompeting native frogs and ki lling potential
predators with their toxic secretions (Fig. E27-1a).
lnvasive plants also threaten natural communities. In the
1930s and 1940s, the japanese vine kudzu was planted
extensively in the southern United States to control erosion.
Today, kudzu is a majar pest, killing trees and all other
vegetation in its path, and engulfing any stationary object,
such as an abandoned house (Fig. E27 -1 b). Both water
hyacinth and purple loosestrife were introduced as
ornamental plants. Water hyacinth now clogs waterways in
the southern United States, slowing boat traffic and
displacing natural vegetation. Purple loosestrife aggressively
invades wetlands, where it outcompetes native plants and
reduces food and habitat for native animals (Fig. E27-1c).
lnvasive species rank second only to habitat destruction
(e) Purple loosestrife
in pushing endangered species toward extinction. Recently,
wildlife officials have made cautious att empts to reestablish Á FIGURE E27-1 lnvasive species (a) The cane toad in
biological checks and balances by importing predators or Australia outcompetes native toads and frogs. (b) The
parasites (called biocontrols) to attack invasive species. This Japanese vine kudzu will rapidly cover entire trees and
is fraught with danger, because new imports can have houses. (e) Purple loosestrife displaces native plants and
unpredicted and possibly disastrous effects on native reduces food and habitat for native animals in wetlands.
wildlife. The cane toad, for example, was introduced into
Australia from South America in the 1930s to control for the intended invasive species, and there have been a
introduced beetles that threatened the sugarcane crop. number of successful introductions. For example, imported
Despite the risks of imported biocontrols, there are often beetles are now among the most important ways of
few realistic alternatives, because poisons kili native and non- controlling purple loosestrife in North America. Professionals
native organisms indiscriminately. Biologists now carefully and v olunteers work together to raise and release these
screen proposed biocontrols to make sure they are specific beetles by the millions, helping to restrain this invasive weed.

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& FIGURE 27-12 Chemical warfare (a) The bombard ier beetle sprays a hot toxic brew when its leg
is pinched by a force ps. (b) A monarch caterpillar feed s on milkweed that contains a powerful toxin.
QUESTI ON Why is the caterpillar colored with bright stripes?
Predators May Use Mimicry to Attract Prey
Sorne predators have evolved aggressive mimicry, a "wolf-
in-sheep's-clothing" approach, in which they entice their
prey to come clase by resembling something attractive to
the prey. For exampl e, by using a rhythm of flashes that is
unique to each species, female fireflies attract males to mate.
But in o ne species, the females sometimes mimic the flash-
ing pattern of a different species, attracting males that they
kili and eat. The frogfish (see Fig. 27-Gb) is not o nly camou-
fl aged, but exhibits aggress ive mim icry by dangling, just
above its mo uth, a wriggling Jure that resembles a small fish.
Fish attracted by the Jure are ea ten in a split second if they
ge t too clase.
Predators and Prey May Engage in Chemical Warfare
Predators and prey use toxins for attack and defense. The
ven o m of spiders and sorne snakes, such as the coral snake ( see
Fig. 27-9b ), serves both to paralyze prey and to deter predators.
Certain mollusks (induding squid, octopus, and sorne
sea slugs) emit clouds of ink when attacked. These colo rful
chemical "smo ke screens" confuse predators and mask the
prey's escape. A dramatic example of chemical defense is seen
in the bombardier beetle. In response to the bite of an ant, the
beetle releases secretions fro m special glands into an abdom-
inal chamber. There, enzym es catalyze an explosive chemicaJ
reaction that shoots a toxic, boiling-hot spray o nto the at-
tacker ( Fig. 27-12a).
Plants have evolved a variety o f chemical adaptations
that deter their herbivorous predators. Many, such as the
mil kweed, synthesize toxic and distasteful chem icals. As
plants evolved defensive toxins, certain insects evolved in-
creasingly efficient ways to detoxify o r even use these sub-
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Community lnteractions 521
(b) Monarch caterpillar
stances. The result is that nearly every toxic plant is eaten by
at least one type of insect. For exampl e, monarch butterflies
lay their eggs on milkweed; when their larvae hatch, they con-
sume this poisonous plant ( Fig. 27-12b ). The caterpillars not
only tolerate the mi lkweed poison, but store it in their tissues
as a defense against tl1eir own predators. The stored toxin is
retained in the metamorphosed monarch butterfly (see Fig.
27-Ba). Viceroy butterfli es (see Fig. 27-Bb) use a similar strat-
egy, storing a bittercompound from willows (eaten by the lar-
vae) in the tissues of the adult.
Grasses embed tough silicon (glassy) substances in
their blades that make them difficult to chew, selecting fo r
grazing animals wi th longer, harder teeth . Over evolutionary
time, as grasses evolved tougher blades that discouraged pre-
dation, horses evolved longer teeth with thicker enamel coat-
ings th at resist wear and abrasion from the tough grasses.
27.4 WHAT IS PARASITISM?
Parasites live in or o n their prey, which are called hosts,
usually harming or weake ning them but no t immediately
killing them. Parasites are generally much smaller and more
numerous than th eir hosts. Fami liar parasites include tape-
worms, fl eas, ti cks, and the many types of disease-causing
protists, bacteri a, and viruses. Many parasites, such as the
protist that causes malaria, have com plex life cycles involv-
ing two or more hosts (see Fig. 20-1 2). The newly discovered
roundworm parasite highlighted in "Scientific Inq ui ry: A Par-
asite Makes Ants Berry Appeali ng to Birds" o n p. 522 appears
to use both ant and b ird hosts during its life cyde. There are
very few parasitic vertebrales, but the lamprey (see Fig. 24-5 ),
which attaches itselfto a host fish and sucks its blood, is o ne
example.
 522 Behavior and Ecology

Scientific lnquiry

On a field trip to the rain forest of Panama in May 2005,

ecologist Steve Yanoviak and fello w researchers were
examining a colony of tree-d welling black ants (Cephalotes
atr atus) a nd found a few with strikingly red abdomens
(Fig. E27-2). When sliced open, the abdome ns disgorged
hundreds of eggs in which tiny parasitic roundworms were
developing. Both the round worm species and its effect on
the ants were new to science, so the team ea gerly purs ued
their chance observation .
To te st the hypothesis that only adult ants with red
abdomens ha rbar round worms, the researchers di ssected
300 all-black adults and found no ne to be infected, while all
those with red abdomens were infected . Noticing that the
infected abdomens broke off readily, they measured the
force required to pluck abdomen s from infected ants versus
those of norm al a nts, a nd discovered that only infected
a bdo men s could be pulled off without dislodging th e ant
from its t wig. Although ants of this species usually bite and ~FIGURE E27-2 An ant resembles berries The a bdomen
emit foul -smelling, distasteful chemicals when handled (as of a parasitized ant closely resembles berries o n trees in
a protection against insect-eating birds), the infected ants which the ants live. Bird s eating these berries may pluck off
did not exhibit these behav io rs. Further, their red a bdomens the ant's abdomen.
strongl y re sembled, in both color and shape, berries from
trees on which the ants foraged (see Fig. E27-2).
Based on these obse rvations , Yanoviak and coworkers Based on their observations and experiments, the
hypothesized that the round worms trick berry-eating birds re searchers hypothesized that the roundworm s s pend most
into dispersing their eggs by causing the ants' abdomens to of th e ir life cycle inside the a bdomens of ants, but then
re semble berries. To investigate this, the team dissected ants rely on fruit-eating birds to disperse their eggs, a strateg y
at all stages of development, and found that the parasites that would help them to survi ve. lf the round worms only
infect larval ants and develop as the ant mature s. Once the parasitized ants , then as they spread throughout an ant
roundworms produce eggs that are ready to be released, the colony they would be likely to kili th e entire colony, and
infected ants' abdomens become enlarged and red. themselves with it. Fruit-eating birds, however, v isit many
To test w hether the eggs could survive passing through trees and leave droppings that a re collected by ants from
a bird's digestive tract, a nd thu s be di s persed in its many colonies. So, natural selection would favor
droppings, the researchers fed a n infected ant abdomen to roundworms whose larvae caused their host ants'
a chicken , and later, found hundreds of healthy roundworm abdomens to swell and mimic ripe fruit , causing birds to
eggs in its feces. But how did the larval ants become disperse them widely.
infected? The resea rchers hypothesized that ant larvae As always in science, new finding s trigger new
might eat bird droppings conta ining round worm eggs. The question s; for e xample, how does the round worm infection
tea m collected food particles that adult ants were ca rrying cause the a nt abdomen to turn red? We can be s u re that
to their colony to feed their larvae , and discovered that , future studies will ans wer this question and also raise more
indeed, many of these were bits of bird droppings. questions in the process.

Parasites and Their Hosts Actas Agents
of Natural Selection on One Another
The variety o f infectiou s b acteria and viruses and the preci-
sio n o f the immune system that counters their attacks are ev-
idence of the powerful forces of coevolutio n b etween
pa rasitic m icroo rganisms a nd their hosts.
Sorne s pecific examples o f pa rasitic interactio ns influ-
encing evolution can be o bserved today. One is a p arasitic
protist, commo n in Africa, tha t causes a disease called naga na
in cattle. Sorne breeds o f African cattle from n agana-infested
regio ns h ave evolved a pa rti al immunity to it a nd genera lly
su rvive infection, w hereas diffe rent breeds of cattle im p o rted
to Africa u su ally d ie from n agana if no t treated . Likewise, the
m a lari a p arasite, w hich sp ends pa rt of its life cycle in red
blood cells, h as exerted strong selective pressure o n human
po pulation s in m alaria -infested regio ns. A mutatio n in the
he moglo bin gen e causes red bloo d cells to becom e disto rted
a n d resistant to infectio n by this p rotist. Altho ugh individ u-
a ls w h o inhe rit two copies of this mutation w ill have sickl e-
cell ane mi a, in so rne regi o ns of Africa, 20% to 40% of the
human po pulatio n carries th e sickle-cell gene because o f the
p rotection it confers against m ala ria (see p . 190) .
27 .S WHAT IS MUTUALISM?
Mutualism refers to interactio ns between species in w hich
bo th benefit. Many mutualistic rel atio ns hips are symbiotic;

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 Community lnteractions 523

(b) Clownfish

• FIGURE 27-13 Mut ualism (a) This brig htly colored lichen growing on bare rock is a mutualistic
relationship between an alga and a fungus . (b ) The clownfish snuggles unharmed among the stinging
tentacles of the anemone.

that is, they involve a close, long-term physical association be-
tween the participating species. lf you see colored patches on
rocks, they are probably lichens, a mutualistic association be-
tween an alga anda fungus (Fig. 27-13a). The fungus provides
support and protection while obtaining food from the photo-
synthetic alga, whose bright colors are actually light-trapping
pigments. Mutualistic associations also occur in the digestive
tracts of cows and termites, where protists and bacteria find
food and shelter. The miaoorganisms break down cellulose,
making its component sugar molecules available both to them-
selves and to the animals that harbor them. In our own intes-
tines, mutualistic bacteria synthesize vitamins, such as vitamin
K, which we absorb and use. Plants called legumes benefit by
providing chambers in their roots that house nitrogen-fixing
bacteria (see Fig. 19-9). These bacteria are among the few or-
ganisms that can acquire nitrogen gas from the air and chemi-
cally modify it into a form that plants can use as a nutrient.
Many mutualistic relationships are notas intimate and
extended as those just discussed. Consider, for example, the
relationship between plants and the insects that pollinate
them. The insects fenilize the plants by carrying plant sperm
(found in pollen grains), and benefit by sipping nectar and
sometimes eating poli en. Both the bee and hoverfly shown in
Figure 27-9a are important plant pollinators. The clownfish
( coated with a !ayer of protective mucus) takes shelter among
the venomous tentacles of certain species of anemones
(Fig. 27-13b ). In this mutualistic association, the anemone
provides the clownfish with protection from predators, while
the clownfish cleans its anemone, defends it from predators,
and may bring it bits of food.
27.6 HOW DO I<EYSTONE SPECIES
INFLUENCE COMMUNITY STRUCTURE?
In sorne communities, a particular species, called a
keystone species, plays a major role in determining com-
munity structure-a role that is out of proportion to its
abundance in the community. Tf the keystone species is re-
moved from its community, normal community interac-
ti.ons are significantly altered and the relative abundance of
other species changes dramatically. lt is difficult to experi-
mentally identify keystone species because the community
must be carefully monitored for years with the species pres-
ent, and for additional years after it is removed. Nonethe-
less, ecological studies have provided evidence that
keystone species are importaotin a variety of communities.
In 1969, Robert Paine, an ecologist at the University of
Washington, removed predatory Pisaster ochraceous sea stars
(Fig. 27-14a) from sections ofWashington State's rocky inter-
tidal coast. Native mussels, a favored prey of Pisaster, became
so abundant that they outcompeted other invertebrates and
algae that oormally coexist with the mussels in intertidal
communities.
Sorne species are found to be keystone species only after
they are removed as a director indirect result of human activi-
ties. For example, the lobster may be a keystone species off the
east coast of Canada. Overfishing of the lobster allowed the
population ofits prey, sea urchins, to expand enormously. The
population explosion of sea urchins nearly eliminated certain
types of algae on which the urchins feed, leaving large expanses
of bare rack where a diverse community formerly existed.
The sea otter appears to be a keystone species along the
coast ofwestern Alaska. Starting around 1990, otter numbers
declined drastically, all owing an inaease in their sea urchin
prey. The sea urchins then overgrazed the kelp forests that
provide critica! undersea habitat for a variety of marine
species. What killed the sea otters? Killer whales, which for-
merly fed primarily oo seals and sea lions, were seen inaeas-
ingly dining on sea otters as their favored prey disappeared.
Scientists hypothesize that sea! and sea lion populations have
declined, in tum, because of overfishing by humans in the
North Pacific, depleting the food supply of these fish-eaters.
In the African savanna, the elephant is a keystone
predator. By grazing on small trees and bushes (Fig. 27-14b ) ,

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 524 Behavior and Ecology

(b) African elephant

& FIGURE 27-14 Keystone speci es (a) The sea star Pisaster ochraceous is a keystone species along
t he roc ky coast of the Paci fic Northwe st. (b) The elepha nt is a keystone s pecies o n t he African
savanna.

elephants prevent the encroachm ent of fo rests and help
maintain the grassland communi ty, along with its diverse
popul ation of grazing mamm als and their predato rs (de-
scribed in Chapter 29 ). In Chapter 30, you willlearn about
the role of anoth er keystone species, the wolf, described in
"Earth Watch: Restoring a Keysto ne Predator" on p. 594.
27.7 SUCCESSION: HOW DO
COMMUNITY INTERACTIONS
CAUSE CHANGE OVER TIME?
In a mature terrestrial ecosystem, the populations that make
up the community interact with one another and with their
nonliving environ ment in in tricate ways. But this tangled web
of life did not spring fully formed fro m bare rock or naked
soil; rather, it emerged in stages over a long period, by a
process called succession . Succession is a structural change
in a communi ty and its non living environment over time. It
is a kind of "community relay" in which assemblages of
plants and animals replace one another in a sequence that is
somewhat predictable.
Succession is preceded and started by an ecological
disturbance, an event th at disrup ts the ecosystem by alter-
ing its communi ty, its abiotic (non living) structure, or both.
The precise changes that occur during successio n are as d i-
verse as the enviro nments in which successio n occurs, but we
can recognize certain general stages. Succession starts with a
few hardy plants called pioneers. The pioneers alter the
ecosystem in ways that favo r com peting plants, which grad-
ually displace them. If allowed to continue, succession pro-
gresses toa diverse and relatively stable dimax community.
Alternatively, recurring d isturbances mainta in many com-
muni ties in earlier, or subdimax, stages of succession . Our
discussion of succession will focus on plant communities,
which domínate the landscape and provide both food and
habitat for a ni mals.
There Are Two Major Forms ofSuccession:
Primary and Secondary
Succession takes two major forms: primary and secondary.
During primary succession, a comm unity gradually forms
where there is no trace of a previous community. The d istur-
bance that sets the stage for primary succession may be a gla-
cier scouring the landscape down to bare rock, o r it may be a
volcano producing an entirely new island or covering an
ecosystem with new rock as lava hardens ( Fig. 27-1 Sa) . This
buil ding of a communi ty "from scratch " typ ically requires
thousands or even tens of thousands of years.
During secondary succession, a new community de-
velops after an existing ecosystem is disturbed in a way that
leaves significant remnants of the previous community be-
hind, such as soil and seeds. For example, beavers, land-
slides, or people may da m streams, causing marshes, ponds,
or lakes to for m. A landslide or avalanche may stri p a swath
of trees from a mountainside. When Mount St. Helens in
Washingto n State erupted in 1980, it left patchy remnants of
forest and a thick ]ayer of nutrient-rich ash that encouraged
a rapid pro liferation of new life (Fig. 27-1 Sb ). Fire is another
common distu rbance. Fires produce nutrient-rich ash, and
spare sorne trees and many h ealthy roo ts. Sorne seeds with-
stand fire and even require it in order to sprout, so fires al-
low rather rap id regeneratio n of forests and other
com mun ities (Fig. 27-15c) . In the following sections, we
look at specific exam ples of succession that illustrate the
process in more detail.

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 Community ln teractions 525

(a) Mt. Kilauea, Hawaii

(e) Yellowstone National Park, Wyoming

Á FIGURE 27-1 S Succession in progress (a) Primary succession. Left: The Hawaiian volcano Mount
Kilauea has erupted repeatedly since 1983, sending rivers of lava over the surrounding countryside.
Right: A pioneer fern takes root in a crack in hardened lava. (b) Secondary succession. Left: On
May 18, 1980, the explosion of Mount St. Helens in Washington State devastated the surrounding pine
forest ecosystem. Right: Twenty years later, life abounds on the once-barren landscape. Because
traces of the former ecosystem remained , this is an example of secondary succession. (e) Secondary
succession. Left: In the summer of 1988, extensive fires swept through the forests of Yellowstone
National Park in Wyoming. Right : Trees and flowering plants are thriv ing in the sunlight, and w ildlife
populations are rebounding as secondary succession occurs.

QUESTION People have suppressed fires for decades. What are the implications of fire suppression
for forest ecosystems and succession?

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 526 Behavior and Ecology

Primary Succession Can Begin on Bare Rock shrubs are shaded out. Within the forest, shade-tolerant
Figure 27-16 illustrates primary succession on Isle Royale, seedlings of taller or faster-growing trees thrive, including bal-
Michigan, an island in northern Lake Superior that was scraped sam fir, paper birch, and wh ite spruce. Tn time, they tower
down to bare rack by glaciers that retreated rough ly 10,000 over and replace the original trees, which are intolerant of
years ago. Weathering produces cycles of freezing and thawing shade. After a thousand years or more, a tall climax forest
that cause racks to crack, producing fissures, and erades rack thrives on what was once bare rock.
surface layers, producing small partides. Rainwater, which is
naturally acidk andan excellent solvent, dissolves so me of the An Abandoned Farm Will Undergo
minerals fram rock particles, making them available to plants. Secondary Succession
Weathered rock provides a place for lichens, which are Figure 27-17 illustrates secondary successio n on an aban-
pioneer species, to attach where there are no competitors and doned farm in the southeastern United States. The pioneer
plenty of sunlight. LiChens obtain energy thraugh photosyn- species, sun-loving, fast-growing annual plants such as crab-
thesis and acquire so me of their mineral nutrients by dissolv- grass, ragweed, and Johnson grass, take root in the rich soil.
ing rack with an acid that they secrete. As the lichens spread Such species generally produce large numbers of easily dis-
over the rack surface, species of mosses that are relatively persed seeds that help them colonize open spaces, but they
drought tolerant begin growing in the rock aacks. Fortified don't compete well against longer-lived (perennial) species
by nutrients liberated by the lichens, the mosses forma dense that gradually graw larger and shade out the pioneers.
mat that traps dust, tiny rack particles, and bits of organic de-
bris. The mosses will eventually cover and kili many of the
lichens that made their grawth possible.
As sorne mosses die each year, their bodies add nutri-
ents to a thin !ayer of new soil, while the living mosses mat
acts like a sponge, absorbing and trapping moisture. Within
the moss, seeds of larger plants, such as bluebell and yarrow,
germinate. As these plants die, their bodies contribute to the
growing !ayer of soil.
As woody shrubs such as blueberry and juniper take ad-
vantage of the newly formed soil, the mosses and remaining
lichens may be shaded out and buried by decaying leaves and
vegetation. Eventually, trees such as jack pine, black spruce,
and aspen take root in the deeper aevices, and the sun-loving

spruce-fir
climax forest:
white spruce,
juniper balsam fir,
yarrow paper birch

r~o:c:k~s:c~ra~p-e_d _:m:o:s~s:o:n:_
l
bare by a
glacier ____
bare rock
________ _._.---;~¡;.~:¡----------------------------------~1,000
time tyears)

o
• FIGURE 27-16 Primary succession Primary success io n as it occurs on bare rock exposed as
glaciers retreated from lsl e Royal e in uppe r Michigan (in th e far northern portion of the midwestern
United States). Notice that t he soil deepens over time, gradually burying the bedrock and allowing
trees to take root.

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 Community ln teractio ns 527

After a few years, perenn ial plants such as asters, gold- Succession Culminates in a Clímax Community
enrod, Queen An ne's lace, and perenni al grasses move in,
Succession ends with a relatively stable cl ímax community,
foll owed by woody shrubs such as blackberry and smooth
which perpetua tes itself if it is not disturbed by externa! forces
sum ac. These pl ants reproduce rapidly and do mínate fo r
(such as fire, parasites, introduced species, or human activi-
many years as pine and cedar trees are becoming estab-
ties). The populations within a clímax communi ty have eco-
lished . Then, roughly two decades after a field is abandoned,
logical níches that allow them to coexist wíthout replacing
an evergreen fo rest do minated by pines takes over and per-
one another. In general, clímax communities have more
sists for severa! more decades.
species and more types of communí ty interactions than do
As is common during successio n, the new forest alters
early stages of succession . The plant species that domínate clí-
conditions in ways that favo r its com petitors. The shade of the
max communities generally líve longer and tend to be larger
pine fores t inhibits the growth of its own seedlings, while fa-
than pio neer species; this trend is particularly evident in
voring the growth of h ardwood trees, whose seedlings are
ecosystems where fo rest is the clím ax communi ty.
shade tolerant. Slow-growing hardwoods such as oak and
In your travels, you have undoubtedly noticed that the
hickory, which take root beneath the pines, begin to rep lace
type of clímax community varíes dramatically from one area
them after about 70 years, as the pínes die of o ld age. Roughly
to the next. For example, if you d rive th rough Colorado or
a century after the field was abandoned, the region ís covered
Wyo ming, you will see a shortgrass prairie clímax community
by relatively stable clímax forest do minated by oak and hick-
on the eastem plains (in those rare areas where it has not been
ory, whích will persist unless the forest is disturbed, for exam-
replaced by farms), pine-spruce forests in the mountains, tun-
ple, by fire or logging.
dra on the mountain sum mits, and sagebrush-dominated
Succession Also Occurs in Ponds and lakes communities in the westem valleys. The exact nature of the
Jn freshwater ponds or lakes, succession occurs not only
through changes within the pond or lake, but also through an
influx of nu trients fro m outside the ecosystem . Sediments
and nutrients carried in by runoff fro m the surrounding land
have a particularly large impact on small freshwater lakes,
ponds, and bogs, which gradually undergo succession to dry
land ( Fig. 27-18 ). In forests, meadows may be produced by
lakes undergoing succession. As the lake fills in from the
edges, grasses colonize the newly formed soil. As the lake
shrinks and the area of meadow expands, trees will encroach
around the meadow's edges. If you return to a forest lake
20 years after your first vísít, ít ís líkely to be smaller as a re-
sult of succession.

cedar

o
• FIGURE 27-17 Secondary succession Second ary success ion as it might occur on a plowed ,
abandoned farm fi eld in North Carolina (i n the south east ern Unit ed State s). Notice that a thick layer of
soil is present from the beginning, w hich greatly speed s up the process compared to primary
succession.

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 528 Behavior and Ecology

(a) Early stage of pond succession (b) Late stage of pond succession

& FIGURE 27- 1 8 Succession in a small freshwater pond In sma ll ponds, succession is speeded by
an influx of materials from the surrounding s. (a) In this small pond , dissolved minerals carried by
runoff from the surround ing s support aquatic plants , whose seeds or spores were carried in by the
wind s or by birds and oth er anim als. (b) Over t ime, t he decaying bodies of aquat ic plants build up
soil that provides anchorage for more terrestrial plants. Eventually, the pond is entirely converted to
dry land.

climaxcommunity is determined by numerous geological and
climatic variables, including temperature, rainfall, elevation,
latitude, type of rock (which determines the type of nutrients
available), and exposure to sun and wind. Natural events such
as windstorms, avalanches, and fires started by lightning may
destroy sections of clímax forest, reinitiating secondary succes-
sion and producing a patchwork of various successional stages
within an ecosystem.
In many forests throughout the United States, rangers
now allow fi res set by lightning to run their course, recogniz-
ing that this natural process is important for the maintenance
of the en tire ecosystem. Fires 1ibera te nutrients and kili sorne
(but usually not all) of the trees. As a result, more nutrients
and sunlight reach the forest floor, encouraging the growth of
subclimax plants. The combination of clímax and subclimax
regions within the ecosystem provides habitats for a far larger
number of species.
Sorne Ecosystems Are Maintained
in a Subclimax Stage
Sorne ecosystems are maintained in a subclimax stage by fre-
quent disturbances. The tallgrass prairie that once covered
northem Missouri and lllinois is a subclimax stage of an
ecosystem whose clímax community is deciduous forest. The
subclimax prairie was maintained by periodic fires, sorne set
by lightning and others deliberately set by Native Americans
to increase grazing land for bison. Forest now encroaches,
and limited prairie preserves are maintained by carefully
managed burning.
Agriculture also depends on carefully selected subcli-
max communities. Grains are specialized grasses characteris-
tic of the early stages of succession, and farmers spend a great
deal of time, energy, and herbicides to prevent competitors,
such as other grasses, wildflowers, and woody shrubs, from
taki ng over. The suburban lawn is a painstakingly maintained
subclimax ecosystem. Mowing (a disturbance) destroys
woody colonizers, and people also use herbicides to selec-
tively kili pioneer species such as crabgrass and dandelions.
Clímax Communities Create Earth's Biomes
The cl ímax communities that form during succession are
strongly influenced by climate and geography. Extensive ar-
eas of characteristic clímax plant communities are called
biomes, and include deserts, grasslands, and a variety of
forests. These biomes domínate broad geographical regions
with similar climates. The factors that influence climate as
well as the biomes that form in tbe different di mates are de-
scribed in Chapter 29.

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Case Study revisited
Mussels Muscle In
Zebra and quagga mussels (Fig. 27-19) have all the
characteristics that make invas ive species successful in their
introduced homes. They outcompete native musse ls and clams
for both food and space, and have few predators to control
them . They have phenomenal reproductive potential-each adult
female of either species may lay severa! hundred thousand eggs
each year. Their microscopic larvae can be carried for miles in
water currents, allowing them to rapidly colonize the length of a
river. Because these mussels can survive out of water for days,
those that cling to hulls of boats can be easily portaged to other
lakes and rivers, w here they rapidly establish new colonies.
Although zebra and quagga mussels are both native to
southeastern Europe, their geographic ranges usually do not
overlap. However, the ecological niches that these closely
related species occupy overlap extensively. As a result, the
two species have begun to compete with each other in their
introduced locations. Ecologists w ho study these species
hy pothesize that the quagga may outcompete the zebra
musse l because it is better adapted toa range of conditions-
it is able to colonize deeper water, and can tolerate both lower
temperatures and lower oxygen levels. But because they both
damage freshwater ecosystems in the same way, if the quagga
outcompetes the zebra mussel, this will have few (if any)
practica! benefits.
The spread of the zebra and quagga mussels continues. In
2008, both species were reported in reservoirs in Colorado
and California, almost certainly trans ported by recreational
boaters. Despite major efforts to identify selective biocontrols,
such as selective diseases or predators, there is presently no
way to restrain the explosive growth rate of the se mu sse l
CHAPTER REVIEW
Summary of Key Concepts
27.1 Why Are Community lnteractions lmportant?
Ecological co mmunities consist of all the interacting popula-
tions within an ecosystem. Community interactions influence
population size, and the interacting populations within com-
munities act upon one another as agents of na tural selection.
Thus, community interactions shape the bodies and behaviors
of members of the interacting populations.
27.2 What ls the Relationship Between the Ecological
Niche and Competition?
The ecological nich e defines all aspects of a species' habitat and
interactions with its living and nonliving environments. Each
species occupies a unique ecological niche. Interspecific compe-
tition occurs when the nich es of two species within a commu-
nity overlap. When two species with the same niche are forced
(under laboratory conditions) to occupy the sam e ecological
ni che, one species always outcompetes the other. Species within
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Community lnteractions 529
Á FIGURE 27-19 A quagga mussel (left) and zebra mussel
(right)
populations. Some researchers suggest that lack of
phytoplankton, their primary food, may eventually slow their
proliferation in the Great Lakes. Lake Michigan , for example,
has become strikingly clearer since the mu ssels' invas ion, as
its phytoplankton population has been depleted. But because
phytoplankton directly or indirectl y support most of the
wildlife within the Great Lake s, food depletion is notan
appealing solution to the mussel problem.
Consider This
lnvas ive mussels can undermine the entire community
structure of any lake into w hich they are introduced. Because
mussel larvae settle rapidly on boat hu li s, a boat that has
been in the water for 24 hours mu st be washed with a hot-
water pressure washer or dried for at least five days before
being launched in another body of water. Do yo u think there is
any practi ca! way to re strict the spread of the se musse ls to
still more lakes and reservoirs? What measu res would be
required to accomplish this?
natural communities have evolved in ways th at avoid excessive
niche overlap, with behavioral and physical adaptations that al-
low resource partitioning. Inte rspecific co mpetition limits both
the population size and the distribution of competing species.
Tntraspecific competition also limits populations because indi-
viduals of the same species occupy the same ecological niche
and compete with one another for all of their needs.
27.3 What Are the Results of lnteractions
Between Predators and Their Prey?
Predators eat other organisms and are generally larger and less
abundant than their prey. Predators and prey act as strong
agen ts of selection on one another. Prey animals h ave evolved a
variety of protective coloratio ns that render them either incon-
spicuous ( camouflage) or startling (startle coloration) to their
predators. Sorne prey are poisonous, distasteful. o r veno mo us,
and exhibit warning coloration by which they are readily recog-
nized and avo ided by p redators, while oth ers have evolved to
resemble other, more distasteful organisms through mimicry.
8oth predators and prey h ave evolved a variety of toxic chemi-
cals for attack and defense. Plants that are preyed on have
evolved elaborate defenses, ranging from poisons to overall
toughness. These d efenses, in turn, h ave selected for p redators
that can detoxify poisons and grind down tough tissues.
 530 Behavior and Ecology

27.4 What ls Parasitism? Thinking Through the Concepts

In parasitism, the parasite feed s on a larger, less abundant host,
harming it but not killing it immediately. Parasites include Fill-in-the-Biank
d isease-causing microorganisms as well as anim als such as fleas l. O rganisms that interact serve as agents of on
and tapeworms. o ne another. This resul ts in , which is the
process by which species evolve adaptations to one
27.5 What ls Mutualism? a no ther. Four types o f communi ty in teractio ns described
Mutualism benefits both interacting species. Sorne mutualistic in this ch apter are _ _ _ _ _ _ _ _ __,
interactions are close a nd extended, such as that between cows _ _ _ _ _, and _____
and the microorgan,isms that live in their digestive tracts, which
2. Predators may be meat eaters, called or
help them to digest cellulose. Other mutualisti c in teracti on s are
plant eaters, called . Both predato rs and their
m ore temporary, such as those that occur between plants and
prey may blend in to their surroundings by using
the animals that po lhnate them .
_ _ _ _ _ . Pred ators are generally and less
27.6 How Do Keystone Species lnfluence Community _ _ _ _ _ th an their prey.
Structure? 3 . Competition occurs whenever two differen t populatio ns
Keystone species have a greater influence on com mun ity struc- within a community have overl app ing
ture than can be p redicted by their numbers. For example, if the ecological . Competition between two species
African elephant were d riven to exti nction, the African grass- is called and this may lead to evolutionary
lands it now inhabits migh t revert to forests. changes that reduce competition . This phen omenon is
27.7 Succession: How Do Community lnteractions called _ _ _ __
Cause Change over Time? 4. Fill in th e types of colo ra tio n or mimicry: U sed by a prey
Successio n is a structural ch ange in a co mmuni ty a nd its no n- to signa! th at it is dist asteful: ; used by a
living environment over time. During succession , plants alter moth with large eyespots o n its wings: _ _ _ _ _,
th e environment in ways that favor th eir competi tors, t hus pro- mi n~kry o f a poisonous animal by a no npoison ous
ducing a somewh at predictable progression of d omina nt aniQ1al: ; mim icry used by a predator to
species. Primary successio n, which m ay take th ousands o f years, attract its prey: _ _ _ __
occurs where no remna nt of a previous co mmunity existed, S. Fill in the appropriate type of communi ty interactio n:
su ch as on b are rock. Secondary succession occu rs much more Bacteria, living in the human gut, that syn thesize vitamin
rapidly beca use it builds o n the remains of a disrup ted commu- K: · bacteri a that cause illness: ·a
n ity, such as an abando ned field or remnants of a fo rest after a deer eating grass: · a tick sucking blood:
fi re. Uninterrup ted succession ends with a climax community, _ _ _ _ _; a bee pollinating a flower:_ _ _ __
which tends to be self-pe rpetuating un less acted on by outside kudzu covering trees: _ _ _ __
forces, such as fire or hum an activities. Sorne ecosystems,
including tallgrass prairie and farm fields, a re m aintained in 6. A so mewhat p redictable cha nge in community structure
relatively early, subclimax stages of successio n by periodi c over time is called . This process takes two
disruptio ns. fo rms. Which of these forms would start with bare rock?
_ _ _ _ _ Which wo uld occur after a forest fire?
_ _ _ _ _ A relatively stable community that is the end
KeyTerms p roduct of this process is called a _ _ _ __
abiotic 524 intertidal zone 514 co mmunity. A mowed lawn in suburbia is an example of a
aggressive mimicry 521 i ntraspecific _ _ _ _ _ co mmuni ty.
biome 528 competition 514
biotic 512 invasive species 520 Review Questions
camouflage 51 7 keystone species 523 l. Define an ecological communi ty, a nd list four important
carnivore 514 mimicry 517 types of community interactio ns.
clímax community 524 mutualism 522 2. Describe four very different ways in which specific plants
coevolution 512 parasite 521 a nd animals p rotect themselves from being eaten. In each ,
community 512 pioneer 524 describe an ad aptation that might evolve in predators of
competition 513 predator 514 th ese species th at would overcome their defenses.
competitive exclusion primary succession 524
3 . Defi ne pa rasitism and mutua lism and provide an example
principie 513 resource partitio ning 513
o f each.
di sturbance 524 secondary succession 524
ecological niche 512 startle coloration 519 4. Define succession . Which type of succession would occur
h erbivore 514 subclimax 524 o n a fo rest clear-cut (a region in which all the trees have
host 521 successio n 524 been removed by logging) and why?
interspecific warning coloration 517 S. List two climax and two subclimax com mun ities. How do
competitio n 5 73 they di ffe r?

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6. What is an invasive species? Why are they destructive?
What general adaptations d o invasive species possess?
Applying the Concepts
l. H erbivorous animals that eat seeds are considered by
sorne ecologists to be predators of plants, and
J)erbivorous anim als th at eat leaves are considered to be
p arasites of plants. Discuss the validity of this
classification scheme.
2 . An ecologist visiting an island finds two very closely
related species of birds, one of which has a slightly larger
bill than the othe¡;, fnterpret this finding with respect to
the competitive exclusion principie and the ecological
niche, and expJain both concepts.
3. Think about the case of the camouflaged frogfish and its
prey. As the frogfish sits camouflaged on the ocean floor,
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Community lnteractions 531
wiggling its Jure, a small fish approaches the Jure and is
eaten, while a very large predatory fish fails to notice the
frogfish . Describe all the possible types of community
interacti ons and adaptati ons that these o rganisms exhibit.
Remember that predators can also be prey and that
community interacti ons are compl ex!
4. Design an experiment to determine whether the kangaroo
is a keystone species in the AustraJian outback.
5. Why is it difficult to study succession? Suggest sorne ways
you would approach this challenge for a few different
ecosystems.
(M;¡ co to www.masteringbiology.com for practice quizzes,
......;/ activities, eText, videos, current events, and more.
 Case Study

Dying Fish Feed an Ecosystem

SOCKEYE SALMON OF THE PACIFIC NORTHWEST
have a remarkable life cycle. Hatching in shallow
depressions in the gravel bed of a swiftly flowing
stream, they follow the stream's path into ev er·
larger rivers that eventually enter the ocean.
Emerging into estuaries- wetlands where fresh
water and salt water mix-the salmon 's
remarkable physiology allows the fish to adapt to
the change to salt water before they reach the sea.
The small percentage of young salmon that evade
predators grows to adulthood, feeding on
crustaceans and smaller fish.
Years later, their bodies undergo another
transformation. As they reach se x ual maturity, a
compelling instinctive drive- still poorly
understood despite decades of research- lures
them back to fresh water, but notjust any stream
or river will do. The salmon swim along the coast
(probably navigating by sensing Earth's magnetic
field) until the unique scent of their home stream
entices them to swim inland. Battling swift
currents, leaping up small waterfalls, undulating
over shallow sandbars, and evading human traps,
they carry their precious cargo of sperm and eggs
back home to renew the cycle of life, dying after
their mission has been accomplished .
The fishes' journey back to their birthplace is
remarkable in another way. Nutrients almost
always flow downstream, carried from the land
into the ocean . But the salmon , filled with muscle
and fat acquired from feeding in the ocean, not
only battle against the flow of the current in their
upstream journey, but also rev erse the usual
movement of nutrients. What awaits the salmon at
their journey's end? How does their journey affect
the web of life upstream?

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~
Ata Glance
Case Study Dying Fish Feed an Ecosystem
28.1 How Do Energy and Nutrients Move Through
Ecosystems?
28.2 How Does Energy Flow Through Ecosystems?
Energy Enters Comm unities Primarily Through
Photosynthesis
Energy ls Passed from One Trophic Level to the Next
Net Primary Production ls a Measure ofthe Energy
Stored in Producers
Food Chains and Food Webs Describe the Feeding
Relationships Within Communities
Detritus Feeders and Decomposers Release Nutrients
for Reuse
Energy Transfer Through Trophic Levels ls lnefficient
Case Study Continued Dying Fish Feed an Ecosystem
Health Watch Food Chains Magnify Toxic Substances
28.3 How Do Nutrients Cycle Within and Among
Ecosystems?
28.1 HOW DO ENERGY ANO NUTRIENTS
MOVE THROUGH ECOSYSTEMS?
Two basic laws underlie ecosystem function: Nutrients con-
stantly cycle and recycle within and among ecosystems, while
energy moves through ecological communities (the various
populations of interacting organisms that inhabit ecosys-
tems) in a continuous one-way flow.
Nutrients are atoms and molecules that organisms
obtain from their living or nonliving environment and that
are required for survival. The same atoms have been sustain-
ing life on Earth for about 3.5 billion years. The molecules
that make up your body undoubtedly include oxygen atoms
once trapped in primordial water and liberated for the first
time by ancient photosynthetic bacteria. Sorne of your car-
bon atoms were exhaled by a dinosaur, and sorne of your ni-
trogen atoms were probably liberated by decomposer
bacteria consuming the remnants of a salmon. Although nu-
trients may be transported, redistributed, or converted to dif-
ferent molecular forms, they do not leave Earth and are
continually recycled through the nutrient cycles described in
later sections.
Energy, in contrast, is continuously replenished. The ac-
tivities of life-from the migration of salmon to the active
transpon of molecules through a cell membrane- are ulti-
mately powered by sunlight. The solar energy that continu-
ously bombards Earth is captured by photosynthetic
organisms, then transformed through the myriad chemical
reactions that energize life, and is finally converted to heat
that radiates back into space.
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How Do Ecosystems Work? 533
The Hydrologic Cycle Has lts Major Reservoir in the
Oceans
The Carbon Cycle Has Majar Reservoirs in the
Atmosph e re and Oceans
.}3io lix • The Carbon Cycle
The Nitrogen Cycle Has lts Major Reservoir in the
Atmosphere
Case Study Continued Dying Fish Feed an Ecosystem
The Phosphorus Cycle Has lts Major Reservoir in Rock
28.4 What Happens When Humans Disrupt
Nutrient Cycles?
Overloading t he Nitrogen and Phosphorus Cyc les
Damages Aquatic Ecosystems
Overloading the Sulfur and Nitrogen Cycles Causes
Acid Deposition
lnterfering with the Carbon Cycle ls Warming Earth's
Climate
Earth Watch Poles in Peril
Case Study Revisited Dying Fish Feed
an Ecosystem
28.2 HOW DOES ENERGY FLOW
THROUGH ECOSYSTE~S?
Ninety-three million miles away from Earth, thermonuclear
reactions in the sun convert hydrogen into helium, transform-
ing a relatively small amount of matter into enormous quan-
tities of energy. A tiny fraction of this energy reaches Earth in
the form of electromagnetic waves, including heat, light, and
ultraviolet energy. Of the energy that reaches Earth, much of it
is reflected by the atmosphere, clouds, and Earth's surface. Still
more is absorbed as heat by Earth and its atmosphere, leaving
only about 1% to sustain life. Because only a fraction of this
remaining solar energy is captured by photosynthetic organ-
isms, the teeming life on this planet is supported by less than
0.03% of the energy reaching Earth from the sun.
Energy Enters Communities Primarily
Through Photosynthesis
During photosynthesis, pigment molecules such as chloro-
phyll absorb specific wavelengths of sunlight (see Fig. 7-4).
This solar energy is then used in reactions that store energy in
chemical bonds. To forge new molecules, plants and other
photosynthetic organisms acquire nutrients from the abiotic
(non living) portions of ecosystems. They take up nitrogen and
phosphorus from the soil and water, absorb carbon from the
col in air, and derive oxygen from the air and from water
molecules. Using energy from the sun and these inorganic
nutrients, plants synthesize sugars, starches, proteins, nucleic
acids, and all the other biological molecules they need to
 534 Behavior and Ecology

~FI G URE 28-1 Producers During photosynthesis, producers capture

solar energy and release oxygen as a by-product. Using this energy and
inorganic nutrients from the environment, producers synthesize all the
molecules they need , including carbohydrates , fats , proteins, and nucleic
acids. These molecules, in turn, provide nearly all of the energy and most
of the nutrients for the rest of life on Earth.

energy from
sunlight

~ FIGURE 28-2 Energy flow,

nutrient cycling, and feeding
relationships in ecosystems Energy
steadily enters the producers as
sunlight, which is captured by the
producers during photosynthesi s and
stored in chemical bonds. The
chemical energy is then passed
through various trophic levels. At each
level, some energy is lost as heat.
solar energy
Nutrients are continuously recycled.
. . heat energy
QUESTIO N Why is som e energy
always lost as heat? Relate t hi s loss to
t he second law of t hermodynamics, as
discussed on p. 99. . . nutrients

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sustain life (Fig. 28-1 ). During photosynthesis, plants release
the oxygen that most organisms require for the reactions that
generate ATP. Thus, photosynthesizers seiVe as a conduit for
both energy and nutrients into ecological communities.
Energy ls Passed from One Trophic Level
to the Next
The complex interactions within biological communities,
and between communities and their abiotic environments,
determine the pathways followed by energy and nutrients as
they move through ecosystems. Figure 28-2 provides a pre-
view of our study of how ecosystems function.
Energy flows through communities from photosyn-
thetic producers through severa! levels of consumers. Each
category of organisms is called a trophic leve) (literally,
"feeding leve!"). Photosynthetic organisms, from oak trees in
a forest to single-celled diatoms in the ocean, form the first
trophic leve!, and are called producers, or autotrophs (Greek,
"self-feeders"), because they produce food for themselves using
inorganic nutrients and solar energy. In doing so, they directly
or indirectly produce food for nearly all other forms of life
as well. Organisms that cannot photosynthesize, called
consumers, or heterotrophs (Greek, "other-feeders"), must
acquire energy and most of their nutrients prepackaged in the
molecules that comprise the bodies of other organisms.
Consumers occupy severa! trophic levels. Sorne con-
sumers, called primary consumers, feed directly and exclu-
sively on producers, the most abundant living energy source
in any ecosystem. These herbivores (literally, "plant eaters"),
which include animals such as grasshoppers, mice, and ze-
bras, form the second trophic leve!. Carnivores (literally,
"meat eaters"), such as spiders, hawks, cheetahs, and salmon,
comprise the higher-level consumers. Carnivores act as
secondary consumers when they prey on herbivores. Sorne
carnivores at least occasionally eat other carnivores; when do-
open
ocean
(125)
• FIGURE 28-3 Net primary productivities of ecosystems The average net primary produ ction of
some terrestrial and aquatic ecosystems is illustrated he re, meas u red in grams of biological material
produced per sq uare meter per year. Notice t he enormous differences.
QUESTION What factors contribute to these differences in productiv ity?
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How Do Ecosystems Work? 535
ing so, they occupy the fourth trophic leve!, and are called
tertiary consumers.
Net Primary Production ls a Measure
of the Energy Stored in Producers
The amount of life that a particular ecosystem can support is
determined by the energy captured by the producers in that
ecosystem. The energy that photosynthetic organisms store
and make available to other members of the communi ty over
a given period is called net primary production ( Fig. 28-3 ).
The net primary production of an ecosystem is influ-
enced by many environmental variables, including the amount
of nutrients available to the producers, the amount of sunlight
reach ing them, the availabili ty ofwater, and the temperature.
In the desert, for example, lack of water limits production. In
the open ocean, light is a limiting factor in deep water, and lack
of nutrients limits productivity in most surfacewater. In ecosys-
tems where all resources are abundant, productivity is high; for
example, in tropical rain forests and in estuaries (coastal areas
where rivers meet the ocean, carrying nutrients washed fro m
the land), as shown in Figure 28-3.
An ecosystem's contribution to Earth's overall produc-
tivity js determined both by the ecosystem's productivity and
by the portian of Earth that it covers. Because open oceans,
whose productivity is low, cover about 65% ofEarth's surface,
they con tribute about 25% to Earth's total productivity. This
is about the same overall contribution as tropical rain forests,
whose productivity is very high but which cover only about
4% of Earth's surface.
Food Chains and Food Webs Describe the
Feeding Relationships Within Communities
To illustrate who feeds on whom in a community, it is common
to identify a representative organism within each trophic level
 536 Behavior and Ecology

that is eaten by a representative ofthe level above it. This lin-
ear feeding relationship is called a food chain. Different
ecosystems support radically different food chains. The dom-
inant producers in land-based (terrestrial) ecosystems are
plants ( Fig. 28-4a). These support the familiar plant-eating
insects, birds, and mammals, each of which m ay be preyed on
by other insects, birds, or mammals. In contrast, the domi-
nant producers in most aquatic food chains (such as in lakes
and oceans; Fig. 28-4b ) are microscopic protists and bacteria
collectively called phytoplankton (literally, "plant drifters").
These producers SUJ?POrt a diverse group of consumers called
zooplankton (literally, "animal drifters"), which consist
mainly of protists and small shrimp-like crustaceans. These
are eaten primarily by fish, who are eaten by larger fish.
Animals in natural communities, however, often do not
fit neatly into categories of primary, secondary, and tertiary con-
sumers. A food web shows many interconnecting food chains,
and more accurately describes the actual feeding relationships
within a given community (Fig. 28-5). Sorne animals, such as
raccoons, bears, rats, and humans, are omnivores (literally, "all
eating"), and they act as primary, secondary, and occasionally
tertiary consumers. A hawk for instance, is a secondary con-
sumer when it eats a m o use (an herbivore), and a tertiary
consumer when it eats a robín that feeds on insects and earth-

(a) A simple terrestrial food chain

{b) A simple marine food chain

A FIGURE 28-4 Food chains on land and sea

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 How Do Ecosystems Wo rk? 537

& FIGURE 28-5 A simplified grassland food web The animals pictured in th e foreg round include a vulture (a det ritus·
feeder), a bull snake, a ground sq uirrel, a burrowing owl, a badger, a mouse, a shrew (which looks like a small mouse but
is carnivorous), and a wolf spider. In th e middle dis tance yo u'll see a grouse, a meadow lark, a gras shopper, and a
jack rabbit. In the far distance, look for pronghorn antelope, a hawk , a wolf, and bison.
EXERCISE Using thi s figure, generate a food chain with four tro phi c levels, and identi fy each level.

worms. When digesting a spider, a camivorous plant such as the
venus flytrap can tangle the food web further by acting as both
a producer and a tertiary consurner.
Detritus Feeders and Decomposers Release
Nutrients for Reuse
Among the rnost irnportant strands in the food web are detri-
tus feeders and decom posers. Both of these groups live on the
refuse of life, such as fallen leaves and fruit, as well as the
wastes and dead bodies of other organisrns. Detritus feeders
("detritus" means "debris") are an arrny of rnostly srnall and
often unnoticed organisrns, including cen ain rnites and pro-
tists, nernatode wo rrns, earthworrns, centipedes, and sorne
insects and sn ails. A few large vertebra tes sueh as vultures also
fit this category (see Fig. 28-5 ). As they feed on organic deb ris,
detritus feeders extract sorne of the energy stored there and
excrete the rest in smaller pieces that provide food fo r other
detritus feeders or for deco rnposers.
Decomposers are pri rnarily fungi and bacteria. They dif-
fer from detritus feeders in that they secrete digestive enzyrnes
outside of their bodies. These enzymes break down nearby or-
ganic rnatter, allowing the decornposers to absorb the nutri-
ents they require, with the rernaining elements and srnall
rnolecules being released into the environment. Mushrooms
in a lawn or the blue-gray fuzz you may no ti ce on old bread are
fungal decorn posers hard at work, and the smel ly slirne on
spoiled rneat signals the presence of bacterial decornposers.
Through the activities of detritus feeders and decom-
posers, the bodies and wastes ofliving organisrns are reduced
to simple rnolecules- such as carbon dioxide, water, an d
rninerals-that return to the atmosphere, soil, and water. By
liberating nutrients for reuse, detritus feeders and decom-
posers perforrn the final stages of nutrient recycling and forrn
a vital link in the nutrient cycles of ecosystems.
What would happen if detritus feeders and decomposers
disappeared? This portion of the food web, although incon-
spicuous, is absolutely essential to life on Earth . Without it,

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 538 Behavior and Ecology
commumt1es would gradually be buried by accumulated
wastes and dead bodies. The nutrients stored in these bodies
would be unavailable to enrich the soil and water. Eventually,
plants and other photosynthetic organisms would be unable to
obtain enough nutrients to survive. With these producers elim-
inated, both energy and nutrients would cease to enter the
community, and the higher trophic levels, including humans,
would disappear as well.
Energy Transfer Through Trophic Levels
ls lnefficient
As discussed in Chapter 6, a basic law of thermodynamics is
that energy use is never completely efficient. For example, as
your car burns gasoline, about 75% o( th_e energy released is
lost as heat. This is also true in living systems: Waste heat is pro-
duced by all the biochemical reactions that keep cells alive. For
example, splitting the chemical bonds of adenosine triphos-
phate (ATP) to cause muscular contraction releases heat en-
ergy; this is why walking briskly on a cold day will warm you.
Energy transfer from one trophic leve! to the next is also
quite inefficient. When a grasshopper (a primary consumer)
eats the leaves of a plant (a producer), only sorne of the solar
energy trapped by the plant is available to the insect. Sorne
was used by the plant for cellular metabolic activities, and
more was lost as heat during these processes. Sorne energy
was converted into the chemical bonds of cellulose, which
the grasshopper cannot digest. Therefore, only a fraction o f
the energy captured by the producers of the fi rst trophic leve!
can be used by organisms in the second trophic leve!. A robín
that eats the grasshopper ( the third trophic leve!) will not ob-
tain all the energy that the insect acquired from the plant-
some will have been used up to power hopping. flying. and
the gnashing o f mouthparts. Sorne energy will have been ex-
pended to construct the grasshopper's indigestible exoskele-
ton, and much of it w ill have been lost as heat. Likewise, the
energy from the robin's body will not all be available to the
hawk that eventually consumes it.
Energy Pyramids lllustrate Energy Transfer
Between Trophic Levels
Although energy transfer among trophic levels within differ-
ent communities varíes significantly, studies of a variety of
communities indicate that the net transfer of energy between
trophic levels is roughly 10% efficient. This means that, in
general. the energy sto red in primary consumers is only about
10% of the energy stored in the bodies of producers. Jn turn,
the bodies of secondary consumers possess roughly 10% of
the energy stored in primary consumers. This inefficient en-
ergy transfer between trophic levels is called the "10% law."
An energy pyramid, which shows maximum energy avail-
able at the base and steadily diminishing amounts at higher
levels, illustrates the general energy relationships between
trophic levels ( Fig. 28-6). Ecologists sometimes measure
biomass, which is the weight of living material (usually
measured as dry weight within a given area), at each trophic
leve!. Because the dry weight of organisms' bodies at each
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tertiary
consumer
(1 calorie)
secondary
consumer
(1 O calories)
primary
consumer
(1 00 calories)
producers
(1 ,000 calories)
A FIGURE 28-6 An energy pyramid for a grassland
ecosystem The width af each rectangle is prapartianal ta the
energy stared at that traph ic level. Representative argani sms far
the first three trophic levels in a U.S. grassland ecasystem
illu strated here are a grass happer, a robin, and a red -tailed hawk.
trophic level is roughly pro,Portional to the amount of energy
stored at that leve!, a biomass pyramid for a given com muni ty
usually has the same general shape as its energy pyramid.
What does this mean for community structure? If yo u
wander through an undisturbed ecosystem, you will notice
that the predominant organisms are plants, because they
have the most energy available to them. The rnost abundant
animals will be herbivores, and carnivores will be relatively
rare, because there is far less energy avail able to suppo rt them.
The inefficiency of energy transfer also has important impli-
cations for human food production. The lower the trophic
level we utilize, the more food energy available to us; hence,
far more people can be fed on grain than on meat.
Case Study e o n ti n u e d
Dying Fish Feed an Ecosystem
Salman are carnivares , eating small fish and large
zaaplanktan . Ov erfi shing and env iranmental di sruptian
have led ta a drasti c reductian in their w ild papulatians,
w hi ch has in turn led ta an increase in salman farming
aperatians. But fa rmed salman are aften fed fish meal and
fish ail made fram smaller, wild-caught fish. Because af the
inefficiency af energy tran sfer, it takes 3 paunds af the
w ild-caught fish ta raise l paund af farmed salman. There
is cancern, therefare , that farming salman is shifting th e
fi shing pres sure aw ay f ram w ild salman anta the smaller
acean fi sh that are harv ested ta feed the salman. These
smaller fish are an impartant link in acean faad chains ,
prav iding faad far larger acean carniv ares such as
mackerel , tuna, w hales, seals, and seabirds.
 How Do Ecosystems Work? 539

An unfortunate side effect of the inefficiency of energy
transfer, coupled wi th human production and release of toxic
chemicals, is that certain persistent toxic chemicals become
increasingly concentrated in the bodies of animals that oc-
cupy increasingly high trophic levels, a process called
biological magnification. Persistent poisons, such as mer-
cury, are often taken up by producers, which are then passed
on to primary consumers (who eat large numbers of produc-
ers) and then on to carnivores, who accumulate the poison
from consumers below them. Long-lived carnivorous ani-
mals, such as swordfish, may accumulate enough mercury to
pose a potential health hazard to humans, who actas high-
level consumers when they eat these top predators (see
"Health Watch : Food Chains Magnify Toxic Substances").

Health Watch
Food Chains Magnify Toxic substances
In the United States, wildlife biologists during the 1950s
and 1960s witnessed an alarming decline in populations of
several predatory birds, especially fish-eaters such as bald
eagles, cormorants, ospreys, and brown pelicans. The
decline pushed sorne, including the brown pelican and the
bald eagle, close to extinction. What caused this? The
aquatic ecosystems supporting these birds had been
sprayed with relatively low amounts of the pesticide DDT
to control insects. In the tissues of these top predators,
scientists found concentrations of DDT up to 1 mi Ilion
times greater than in the water where their fish prey lived;
this impaired their ability to reproduce.
The birds were victims of biological magnification, the
process by which toxic substances accumulate in
increasingly high concentrations in animals occupying
higher trophic levels. Fortunately, populations of predatory
birds vulnerable to DDT have recovered significantly since
the pesticide was banned in the United States in 1973.
Substances that undergo biological magnification (such
as oon share two properties that make them dangerous.
First, decomposer organisms cannot readily break them • FIGURE E28-1 The price of pollution Deformities such
down into harmless substances-that is, they are not as the one exhibited by this frog found in Oregon have been
biodegradable. Second, they tend to be stored in living linked to bioaccumulating chemicals. Abnormalities of the
tissue (often in fat), accumulating over the years in the reproductive and immune systems are also common in many
bodies of long-lived animals. Exposure to high levels of types of organisms exposed to these pollutants. Frogs, which
pesticides and other persistent pollutants has been linked have extremely thin skin and spend much of their lives in
to sorne types of cancer, infertility, heart disease, water, are particularly vulnerable to water-borne pollutants.
suppressed immune function, and neurological damage.
Mercury contamination is a particular cause for concern,
because mercury is an extremely potent neurotoxin that that these chemicals are interfering with the reproduction and
accumulates in muscle as well as fat. lts high levels in a few development of fish (including salman), fish-eating birds such
types of long-lived ocean predators-such as swordfish, as cormorants, frogs (Fig. E28-1 ), salamanders, alligators, and
shark, and albacore tuna-have prompted the U.S. Food and many other animals. Endocrine disruptors are also suspected
Drug Administration to advise women of childbearing age of causing reduced sperm counts in humans. Endocrine
and young children to avoid or limit their consumption of disruptors and other organic pollutants bioaccumulate in
these types of fish, because the mercury levels may pose a mussels that filter large quantities of water to obtain food.
potential health hazard. In the United States, coal-fired Zebra mussels (which have invaded the Great Lakes; see
power plants are the largest single source of mercury Chapter 27) are accumulating a variety of organic pollutants,
contamination-atmospheric mercury can be wafted which endanger the fish and birds that consume them.
thousands of miles from these plants and be deposited in Understanding the workings of food webs allows us to
what should be pristine environments, such as in the Arctic. understand why bioaccumulation occurs, and why people,
lnuit natives living north of the Arctic Circle have high levels as well as wildlife, are susceptible. When we eat tuna or
of mercury and other pollutants from consuming large swordfish, for example, we actas tertiary or even
quantities of predatory marine mammals and fish. quaternary consumers, and so are vulnerable to substances
A class of organic chemicals called endocrine disruptors- that bioaccumulate. In addition, the long human life span
including sorne widely used pesticides, plasticizers (which provides more time for substances to accumulate within
make plastic flexible), and flame retardants-has become the body. Threats to human health provide additional
widespread in the environment. Like DDT, these chemicals incentive to study what happens to chemicals released into
accumulate in fat, and either mimic or interfere with the the environment, and to restrict the manufacture and
actions of animal hormones. There is compelling evidence improper disposal of substances that bioaccumulate.

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 540 Behavior and Ecology
28.3 HOW DO NUTRIENTS CYCLE WITHIN
AND AMONG ECOSYSTEMS?
In contrast to the energy of sunlight, nutrients do not flow
down onto Earth in a steady stream from above. Nutri ents are
elements and small molecules that form the chemical build-
ing blocks of life. Sorne, called macronutrients, are required
by organisms in large quantiúes. These include water, carbon,
hydrogen, oxygen, nitrogen, phospho rus, sulfur, and cal-
cium . Micronutrients, including zinc, molybdenum, iron,
selenium, and iodine, are required only in trace qua ntities.
Nutrient cycles, also called biogeochemical cycles, de-
scribe the pathways these substances follow as they move
from the abiotic portions of ecosystems through communi-
ties and back to no n living storage si tes.
The major sou rces and storage sites of nutrients are
called reservoirs, and they are almost always in the non liv-
ing, or abioúc, environment. In the following sections, we de-
scribe the cycles of water, carbon, nitrogen, and phosphorus.
You will see nutrient reservoirs indicated in whi te boxes in
each nutrient cycle figure. Events that drive the movement of
nu trients are in purple boxes, and the trophic levels through
wh ich nutrients move are in yellow boxes.
A FIGURE 28-7 The hydrologic cycle
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The Hydrologic Cycle Has lts Major Reservoir
in the Oceans
The water cyde, or hydrologic cyde (Fig. 28-7), describes the
pathway that water takes as it travels fro m its major reservoir-
the oceans-through the atrnosphere, to reservoirs in fres hwa-
ter lakes, rivers, and groundwater, and then back again to the
oceans. The hydrologic cycle differs from most o ther nutrient
cydes in that the biotic portion of ecosystems plays o nly a
small role-the fundame ntal process of the hydrologic cycle
wou ld continue even if life o n Earth were to disappear.
The hydrologic cyde is driven by solar heat energy, which
evaporates water and drives the winds that carry itas water va-
por in the atrnosphere. Gravity draws water back to Earth in the
form of precipitation ( mostly rain and snow), pulls it into the
ground, and causes it to flow in rivers that empty into the
oceans. Oceans cover about three-quarters of Earth's surface
and contain more than 97% ofEarth's water, with another 2%
of the total water trapped in ice, leaving only 1% as liquid fresh
water. Most evaporaúon occurs from the oceans, and most pre-
cipitation falls back o nto them. Of the water that falls o n land,
sorne evaporates from the soil. plants, lakes, and streams; a
portion runs back to the oceans; and a small amou nt enters

natural underground reservoirs called aquifers. Aquifers are
composed of water-permeable sediments such as silt, sand, or
grave!, which are saturated with water. They are often tapped to
supply water for irrigating crops.
Unfortunately, in many areas of the world-induding
China, India, Northem Africa, and the Midwestem United
States-groundwater aquifers are being "mined" for agricul-
ture; that js, water is being pumped out faster than it is being re-
plenished. Underground water supplies are dropping rapidly in
India, which reliesbeavily on groundwater for irrigation. Jndia's
population, whicb now exceeds 1 billion people, could suffer if
its crop yields fall as groundwater beco mes unavailable.
Because the bodies ofliving things are roughly 70% wa-
ter, a tiny portian ofthe total water involved in the overall hy-
drologic cycle enters the living communities of terrestrial
ecosystems. Sorne is absorbed by the roots of plants, and
much of this is evaporated back to the atmosphere from their
leaves through a process called transpiration. A relatively
minuscule amount ofwater participares in the chemical reac-
tions of photosynthesis, and is resynthesized and released
during cellular respiration.
The hydrologic cycle is crucial for terrestrial com m u ni ti es
because it continually restares the fresh water needed for land-
based Ji fe. Water is a solvent for all of the other nutrients, and
no nutrient can en ter or leave the cells of an organism unless it
is dissolved in water. As you study the nutrient cydes that fol-
low, keep in mind that nutrients in soil must be dissolved in
D reservoirs
D processes
D trophic levels
Á FIGURE 28-8 The carbon cycle
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How Do Ecosystems Work? 541
soil water to be taken up by the roots of plants or to be ab-
sorbed by bacteria. Plantleaves can only take up carbon diox-
ide gas after it has dissolved in a thin layer of water coating the
cells inside the lea f. The hydrologic cycle doesn't depend on ter-
restrial organisms, but they would rapidly disappear without it.
The Carbon Cycle Has Major Reservoirs
in the Atmosphere and Oceans
Chains of carbon atoms form the framework of all organic mol-
ecules, the building blocks of life. The carbon cycle (Fig. 28-8)
describes the movement of carbon from its major short-term
reservoirs in the atmosphere and oceans, through producers and
into the bodies of consumers and detritus feeders, and then back
again to its reservoirs. Carbon enters the living community when
producers capture carbon dioxide (C0 2 ) during photosynthesis.
On land, photosynthetic organisms acquire C02 from the atrno-
sphere, where it represents 0.038% of all atrnospheric gases.
Aquatic producers such as phytoplankton obtain the C0 2 they
need for photosynthesis from the water, where it is dissolved.
Producers retum sorne C0 2 to the atmosphere or water
during cellular respiration, and incorporate the rest of it into
their bodies. Burning forests return carbon dioxide from these
producers back to the atmosphere. When primary consumers
eat producers, they acquire the carbon stored in the producers'
tissues. As with producers, these herbivores and the organisms
in higher trophic Levels who consume them release C02 during
 542 Behavior and Ecology

respiration, excrete carbon compounds in their feces, and store
the rest in their tissues. Al! living things eventually die, and
their bodies are broken down by detritus feeders and decom-
posers. Cellular respiration by these organisms returns C0 2 to
the atmosphere and oceans. co2passes freely between these
two great reservoirs (see Fig. 28-8). The complementary
processes of uptake by photosynthesis and release by cellular
respiration continually transfer carbon from the abiotic to the
biotic portions of an ecosystem and back again.
Sorne carbon, ~owever, cycles much more slowly.
Much of Earth's ca~bo n is bound up in limestone rock.
formed from calcium carbonate (CaC0 3 ) deposited o n the
ocean fl oor in the shells of prehistoric phytoplankton. But
because the movement of carbon from this so urce to the at-
mosphere and back again requires millions of years, this ex.-
tremely long-term process makes very little contribution to
the carbon cycling that supports ecosystems. Another long-
term reservoir for carbon is in fossil fuels, which include
coal, oil, and natural gas. These substances were produced
over millions of years from th e remains of prehistoric organ-
isms buried deep underground and subjected to high tem-
perature and pressure. In addition to carbon, the energy of
prehistoric sunlight ( originally captured by photosynthetic
organisms) is trapped in these deposits. When humans burn
fossi l fuels to use this stored energy, co2is released into the
atmosphere, with potentially severe consequences, as de-
scribed later in this chapter.
Bi oFiix ~ The Carbon Cycle
The Nitrogen Cycle Has lts Major Reservoir
in the Atmosphere
Nitrogen is a crucial component of amino acids, proteins,
many vitamins, nucleotides (such as ATP), and nucleic acids
(such as DNA). The nitrogen cyde (Fig. 28-9) describes the

O reservoirs
D processes
D trophic levels

ammonia
and nitrates
in soil

A FIGURE 28-9 The nitrogen cycle

QUESTION What incentives cause humans to capture nitrogen from the air and pump it into the
nitrogen cycle? What are so me consequences of human augmentation of the nitrogen cycle?

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process bywhich nitrogen moves from its primary reservoir-
nitrogen gas in the atmosphere-to reservoirs of ammonia
and nitrate in soil and water, through producers and into
consumers and detritus feeders, and then back again to its
reservoirs.
The atmosphere contains about 78% nitrogen gas
(N 2), but among all forms oflife, only a few types ofbacte-
ria are able to convert N 2 into a form usable by plants and
other producers. These microorganisms provide the major
natural conduit between the atmospheric reservo ir and eco-
logical communities. In aprocess called nitrogen flxation,
nitrogen-fixing bacteria ·n soil and water break the bonds
in N 2 and combine it with hyd,rogen atoms to form ammo-
nia (NI-1 3 ). Sorne nitrogen-fixing bacteria have entered a
symbiotic association with plantsjn which the bacteria live
in special swellings on roots (see Pig. 19-9). These plants,
called legurnes (including alfalfa, soybeans, clover, and
peas), are extensively planted on farms, in part because they
release excess ammonia produced by the bacteria, fertiliz-
ing the soil. Other bacteria in soil and water convert this
ammonia to nitrate (N0 3-), which producers can also use.
Nitrates are also produced during electrical storms, when
the energy of lightning combines nitrogen and oxygen
gases to form nitrogen oxide compounds, which dissolve in
rain. When the rain falls, it enriches the soil and water with
this important nutrient.
Detritus feeders and decomposer bacteria also play
a role in the nitrogen cycle, producing ammonia from the
nitrogen-containing compounds in dead bodies and wastes.
Ammonia and nitrate are absorbed by producers and are in-
corporated into a variety of biological molecules. These are
passed through successively higher trophic levels as primary
consumers eat the producers and are themselves eaten. At
each trophic level. bodies and wastes are broken down by de-
composers, which liberate ammonia back into the reservoir
Case Study e o n ti n u e d
Dying Fish Feed an Ecosystem
Salmon bodies contain high levels of nitrogen, and
ecologists are interested in how much nitrogen their
bodies contribute to the communities that surround their
spawning streams. Nearly all nitrogen has a molecular
weight of 14 ( 14 N), but a small amount is a heavier
isotope (1 SN). The ratio of these two isotopes is different
in the ocean than it is in freshwater and terrestrial
environments. Because salmon put on about 95% of their
weight in the ocean, their bodies have the nitrogen
isotope ratio of marine animals. Ecologists can measure
the nitrogen in the vegetation around streams where
salmon come to spawn and die. The percentage of
nitrogen exhibiting the ocean ratio is assumed to come
from salmon. Using this type of analysis, ecologists have
found that the contribution of dying salmon to the plant
nitrogen near their spawning streams can be as high
as 70%.
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How Do Ecosystems Work? 543
in soil and water. The nitrogen cycle is completed by
denitrifying bacteria. These residents of wet soil. swamps,
and estuaries break down ni trate, releasing nitrogen gas back
into the atmosphere (see Fig. 28-9).
Fertilizer factories use energy from fossil fuels and N 2
from the atmosphere to synthesize ammonia, nitrate, and
urea (an organic nitrogen compound also found in urine).
The burning of fossil fuels combines atmospheric N 2 and 0 2 ,
generating nitrogen oxides that form nitrates. Nitrogen com-
pounds introduced into ecosystems by fertilizing farm fields
and burning fossil fuels now dominate the nitrogen cycle, cre-
ating serious environmental concerns, as discussed later in
this chapter.
The Phosphorus Cycle Has lts Major Reservoir
in Rock
The phosphorus cycle (Fig. 28-1 O) describes the process
by which phosphorus moves from its primary reservoir-
phosphate-rich rock-to reservoirs of phosphate in soil and
water, through producers and into consumers and detritus
feeders, and then back to its reservoirs. Phosphorus is found
iu biological molecules including nucleotides (such as ATP),
nuclei acids (such as DNA), and the phospholipids of cell
membranes. It also forms a major component of vertebra te
teeth and bones.
In contrast to carbon and nitrogen, phosphorus does
not have an atmospberic reservoir. Throughout its cycle,
phosphorus remains bound to oxygen in the form of phos-
phate (P04 3 - ). As phosphate-rich rocks are exposed by geo-
logical processes, so me of the phosphate is dissolved by rain
and flowing water, which carries it into soil. lakes, and the
ocean, forming the reservoirs of phosphorus that are di-
rectly available to ecological communities. Dissolved phos-
phate is readily absorbed by consumers, which incorporate
it into phosphate-containing biological molecules. From
these producers, phosphorus is passed through food webs;
at each level, excess phosphate is excreted. Ultimately, detri-
tus feeders and decomposers return the phosphate to the
soil and water, where it may then be reabsorbed by produc-
ers or may beco me bound to ocean sediment and eventually
reformed into rack.
So me of the phosphate dissolved in fresh water is car-
ried to the oceans. Although much of this phosphate ends
up in marine sediments, sorne is absorbed by marine pro-
ducers and is eventually incorporated into the bodies of in-
vertebrales and fish. Sorne of these, in turn, are consumed
by seabirds, which excrete large quantities of phosphorus
back onto the land. At one time, seabird excrement ( called
"guano") deposited along the western coast of South Amer-
ica was collected and provided a major source of the world's
phosphorus. As this resource was depleted, phosphorus-rich
rack was mined, primarily to produce fertilizer. Soil that
erodes from fertilized fields carries large quantities of phos-
phates into lakes, streams, and the ocean, where it stimu-
lates the growth of producers. In lakes, phosphorus-rich
runoff from land can stimulate such an excessive growth of
 544 Behavior and Ecology

D reservoirs
D processes
D trophic levels

formation of
phosphate-containing
rock

a lgae and bacteria that its natural community interactions Overloading the Nitrogen and Phosphorus
are disrupted . Cycles Damages Aquatic Ecosystems
In human-dominated ecosystems-such as farm fields, gar-
28.4 WHAT HAPPENS WHEN HUMANS dens, and suburban lawns- ammonia, nitrate, and phos-
DISRUPT NUTRIENT CYCLES? phate are supplied by chemical fertilizers to stimulate plant
Many of the environmental problems that plague modern growth. Each year, roughly 150 million tons of phosphate fer-
society have resulted fro m human interference in ecosystem tilizer are produced from mined phosphate rack. A similar
fun ction. Primitive peo ples were sustained solely by the en- amount of nitrogen-based fertilizer is manufactured using at-
ergy fl owing fro m the sun, and they produced wastes that mospheric ni trogen . These are applied to farm fields to help
were readily taken back into the nutri ent cycles. Through the satisfy the agricultura) demands of a growing population.
mid-1 800s, ho rses, oxen, and mules carri ed people, trans- Water, driven by the hydrologic cyd e, washes over the
ported their goods, and pulled farmer's plows. The energy land, dissolving and carrying away enormous quantities of
expended by these "beasts o f burden " carne from sunlight
trapped by the plants that these ani mals consumed, and
their wastes became fertilizer.
But as the populatio n grew and technology increased,
hu mans began to act mo re and mo re independently o f
these natural processes. The Industrial Revolution, which
began in earnest in the mid-nineteenth century, resulted in
a tremendous increase in our relian ce on energy stored in
fossil fuels for heat, light, transportation, industry, and agri-
culture. ln mining and transpo rting these fuels, we have ex-
posed ecosystems to a variety of substances that are foreign
and often toxic to them, such as oil spilled into bodies of
water (Fig. 28-11 ) . Our reliance o n fossil fuels has also dis-
rupted the glo bal nutrient cycles of ni trogen, phosphorus, & FIGURE 28-11 A natural substance out of placeAn oil-
sulfur, and carbon. soaked bird on the shores of the Black Sea afte r an oil s pill.

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 How Do Ecosystems Wo rk? 545

phosphate and nitrogen-based fertilizer. As the water drains

into lakes, rivers, and ultimately the oceans, these fertilizers dis-
rupt the delicate balance of food webs by overstimulating the
growth of phytoplankton. This causes phytoplankton "blooms"
that can tum reasonably clear water into an opaque green so up.
As the phytoplankton die, their bodies sink into deeper water,
where they provide a feast for decomposer bacteria. Cellular res-
piration by decomposer bacteria uses up most of the available
dissolveo oxygen. Deprived of oxygen, aquatic invertebrates
and fish either die ( contributing to the problem) or leave the
area. A dramatic example occurs each year in the Gulf of Mex-
ico off the coast ofLouisiana, where the Mississippi river dumps
enormous quantities of ni trates that have washed off fertilized
farm fields in the Midwest. Eaclvsummer, this creates a "dead
zone" covering 7,000 to 8,000 square miles, where oxygen lev-
els are so low that few types of organisms can survive. World-
wide, dead zones are increasing rapialy in both size and
number as agricultura) activities intensify.

Overloading the Sulfur and Nitrogen Cycles

Causes Acid Deposition
Although natural processes-such as the activity of nitrogen-
fixing bacteria and decomposer organisms, fires, and
lightning-produce nitrogen oxides and ammonia, about
60% of the nitrogen that is available to Earth's ecosystems now
results from human activities. Buming fossil fuels combines at-
mospheric nitrogen with oxygen, producing most of the emis-
sions of nitrogen oxides. Although sulfur is released as sulfur
dioxide (S02 ) by volcanoes, hot springs, and decomposer
organisms, human industrial activities- primarily burning
sulfur-containing fossil fuels-account for about 75% of all
the sulfur dioxide emissions worldwide. Excess production of
nitrogen oxides and sulfur dioxide was identified in the late
1960s as the cause of a growing environmental threat-"acid
rain, " more accurately called acid deposition.
When combined with water vapor in the atrnosphere, ni-
trogen oxides and sulfur dioxide are converted to nitric acid
and sulfuric acid, respectively. Days later and often hundreds
of miles from the source, these acids fall to Earth dissolved in
rainwater. Acid deposition damages fo rests, can render lakes
lifeless, and eats away at buildings and statues ( Fig. 28-12) . In
the United States, the Northeast, Mid-Atlantic, upper Midwest,
and Western regions and the state of Fiorida are the most vul-
nerable, because the rocks and soils that predominate in those
areas have little ability to neutralize acids.
Acid Deposition Damages Life in Lakes and Forests
Tn the Adirondack Mountains of New York State, downwind
from coal-burning power plants and industry in the Midwest,
acid rain has rendered many lakes and ponds too acidic to
support fis h and the food webs that sustain them. Acid dep-
osition increases the exposure of organisms to toxic metals-
such as aluminum, mercury, lead, and cadmium- all of
which are far more soluble in acidified water than in water of
neutral pi-l. Aluminum dissolved from rock may inhibit plan t
growth and kili fish.
Á FIGURE 28-12 Acid deposition is corrosive Thi s lion
statue was unve iled in 1867 outside th e Le eds tow n hall in
Yorkshire, England . lt is carved f ro m lime stone , and has been
severely erod ed by acid deposition.
On land, acid rain leeches away essential plant nutri-
ents such as calcium, magnesium, and potassium. It may also
kili decomposer microorganisms, preventing the return of
nutrients to the soil. Plants in acidified soil become weak and
more vulnerable to infection and insect attack. Since 1965,
scientists have witnessed the death of about half of the red
spruce and beech trees and one-third of the sugar maples in
the Creen Mountains of Vermont. The snow, rain, and heavy
fog that commonly cloak these mountaintops are highly
acidic. For example, at a monitoring station atop Mount
Mitchell in North Carolina, the pH of fog has been recorded
at 2.9-more acidic than vinegar ( Fig. 28-13) .
Since 1990, government regulations have resulted in
substantial reductions in emissions of both sulfur dioxide
and nitrogen oxides from U.S. power pl ants. Air quality has
improved, and rain has become less acidic. However, the to-
tal nitrogen oxide release remains high because more gaso-
line is being burned by automobiles, releasing more nitrogen
oxides fro m this source. Damaged ecosystems recover slowly.
Adirondack lakes are gradually becoming less acidic, al-
though full recovery is still decades away. In the Southeast.
sorne freshwater acid levels are still increasing. Many ecolo-
gists believe that far stricter controls on nitrogen emissions
will be required to prevent further deterioration and to allow
damaged ecosystems to recover.
lnterfering with the Carbon Cycle ls Warming
Earth's Climate
Between 345 million and 280 million years ago, huge quanti-
ties of carbon were diverted from the carbon cycle. This oc-
curred under the warm, wet conditions of the Carboniferous

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 546 Behavior and Ecology
_.FIGURE 28-13 Acid deposition can destroy forests Acid
rain and acidic fog have destroyed this forest atop Mount
Mitchell in North Carolina.
period, when bodies of prehistoric organisms were buried in
sediments, escaping decomposition. Over time, heat and pres-
sure converted their bodies (containing stored energy from sun-
light) into fossil fuels such as coal. oil. and natural gas. Since the
mid-1 800s, however, we have inaeasingly relied on energy re-
leased by burning these fuels. One researcher estimates that a
typical automobile gas tank holds the transformed remains of
100 tons of prehistoric life, largely miaoscopic phytoplankton.
As we burn fossil fuels in our power plants, factories, and cars,
we harvest the energy of prehistoric sunlight and release co2
into the atmosphere. Burning fossil fuels accounts for 80% to
85% of the co2added to the atmosphere each year.
A second source of added atmospheric C0 2 is
deforestation, which destroys tens of millions of forested
What ls Your Carbon Footprint?
Each of us impacts Earth through our personal decisions
and by the choices we make. A "carbon footprint" is a
measure of the impact that human activities have on
climate, based on the quantity of greenhouse gases
they emit.
Our carbon footprint gives usa sen se of our
individual impacts. For example, each gallon of gasoline
burned releases 19.6 pounds (8.9 kg) of co2into the air.
So, if your car gets 20 miles to the gallon, then each mi le
that you drive wi ll add about a pound of co2 to the
atmosphere.
The Web si tes of the U.S. Environmental Protection
Agency as well as for several environmental organizations
provide household emissions calculators that allow you to
estimate your carbon footprint, and provide advice on
how you can reduce it. To get started, just type "carbon
footprint" into your Internet browser.
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acres annually and accounts for about 15% of C0 2 emissions.
Deforestation is occurring principally in the Tropics, where
rain forests are rapidly being converted to agricultural land to
feed growing populations and supply the world's demand for
biofuels, such as ethanol and biodiesel. The carbon stored in
the massive trees in these forests returns to the atmosphere
when they are cut down and burned.
Collectively, human activities are estimated to release
more than 10 billion tons of carbon (in the form of C0 2 ) into
the atmosphere each year, an amount that is increasing as the
human population grows andas less-developed countries in-
crease their living standards. About half of this carbon is ab-
sorbed into the oceans, plants, and soil, with the rest
remaining in the atmosphere. As a result, since 1850 (when
people began burning large quantities of fossil fuels during
the Industrial Revolution) the C0 2 content of the atmosphere
has increased by almost 37%- from 280 parts per million
(ppm) to 383 ppm-and is growing by about 2 ppm annu-
ally (see Fig. 28-15b). Based on an analysis of gas bubbles
trapped in ancient Antarctic ice, the atmospheric co2 content
is now about 30% higher than at any time during the past
650,000 years.
Carbon Oio.xide and Other Greenhouse
Gases Trap Heat in the Atmosphere
The fate of sunlight entering Earth's atmosphere is shown in
Figure 28-14. Sorne ofthe energyfrom sunlight is reflected back
into space, bouncing off water vapor in douds and other partí-
eles in the air (Fig. 28-14 f)). Most sunlight, however, reaches
Earth, and is converted into heat (Fig. 28-14 E)) that is then ra-
diated back toward space (Fig. 28-14 0). Although most heat
is released into space (Fig. 28-14 0), carbon dioxide and sev-
era! other greenhouse gases trap sorne of this heat in the at-
mosphere (Fig. 28-14 0). This is a natural process called the
greenhouse effect, which keeps our atmosphere relatively
warm and allows life on Earth as we know it.
However, a large and growing body of evidence indi-
cates that human activities have amplified the natural green-
house effect, producing a phenomenon called global
warming. For Earth's temperature to remain constant, the to-
tal energy entering and leaving Earth's atmosphere must be
equal. As greenhouse gas levels rise, more heat is retained than
is radiated back into space, causing Earth to warm. Although
co2 accounts for most human emissions of greenhouse gases,
other important greenhouse gases indude methane (CH 4 ), re-
leased by agricultura! activities, landfills, and coa! mining; and
nitrous oxide (N2 0), released by agricultura! activities and
burning fossil fuels.
Global Temperature lncreases Have Paralleled lncreases
in Atmospheric Carbon Dioxide
The decade from 1998 to 2008 was the warmest ever recorded.
Global surface temperature data are recorded from thousands
of weather stations around the world and from satellites that
measure ocean surface temperatures. Atmospheric C02 data are
collected from air that has typically traveled for hundreds of
miles before reaching the sampling si te near the barren peak of
 How Do Ecosystems Work? 547

0 Most heat is radiated

back into space

0 Some atmospheric heat is

retained by greenhouse gases

8 Some energy
is reflected back
into space

& FIGURE 28-14 Greenhouse gases and global warming lncoming sunlight warms Earth's surface •
and is radiated back to the atmosphere. Greenhouse gases, released by natural processes and
substantially augmented by human activities (yellow rectangles), are absorbing increasing amounts of
this heat, raising global temperatures. 01)
Mauna Loa, a mountain in Hawaii. Here, numerous measure- Global Warming Will Have Many Consequences
ments are taken daily. Elaborate statistical procedures are used As geochemist James White at the University of Colorado
to eliminate readings that show high variability over a short quipped, "If the Earth hadan operating manual, the chapter on
time span, indicating that they have been influenced by nearby dimate might begin with the caveat that the system has been
C0 2 sources. 1-Iistorical and recent temperature records have re- adjusted at the factory for optimum comfort, so don't touch the
vealed a global temperature increase (Fig. 28-15a) that parallels dials." Perhaps the most visible effects of global warming are
the measured rise in atmospheric C0 2 (Fig. 28-15b ). the melting of glaciers throughout the world (Fig. 28-17). For
Any scientist will tell you that correlations between example, each year about 1.6 million people flock to Glacier
events do not prove that one event caused the other. However, National Park, Montana, named for its spectacular abundance
the evidence that human activities are driving global warm ing of glaciers. Of the approximately 150 glaciers that graced its
is so compelling that the overwhelming majority of scientists mountainsides in 1850, fewer than 30 remain.
accept this hypothesis and are convinced that urgent action is
needed to slow the process. The lntergovemmental Panel on More Extremes in Weather Are Predicted and May Already Be Oc-
Climate Change (JPCC) is a consortium involving hundreds of curring Many leading dimatologists see evidence that global
scientists as well as other experts from 130 different nations. In warming is already affecting our weather. Scientists at the Na-
their 2007 report, the IPCC predicted that even under the best- tional Center for Atmospheric Research have concluded that
case scenario in which a concerted worldwide effort is m ade to global warming has increased ocean surface temperatures be-
reduce greenhouse gas emissions, the average global tempera- yond the range of normal fluctuations, contributing, since the
ture will rise by at least 3.2 o F (1.8 oq by the year 2100; a high- mid-1960s, to an increase in severe hurricanes that generate
level emissions scenario projects an increase of7.2 ° F (4.0 °C) higher wind speeds and greater rainfall. As the world warms,
(Fig. 28-16). Small global temperature changes can have enor- experts predict an increase in heat waves. Sorne world regions
mous impacts. For example, average temperatures during the will experience more flooding, while in other areas, droughts
peak of the last lee Age (20,000 years ago) were only about 5 oC are likely to last longer and to be more severe. Since the 1970s,
lower than at present. the area ofEarth impacted by severe drought has doubled as a

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represents a copyright v iolation .
 548 Behavior and Ecology •F ·e
7.2 4.0

•F ·e Q)
(J)
<U
58.1 14.5 ~
~ o 5.4 3.0
::1

~
Q)
57.9 14.4
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a. 57.7 14.3 ::1
E
2! 57.6 14.2 ~
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57.4 14.1 2!
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<U 57.2 14.0 .o 1.8 1.0
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.o 57.0 13.9 e;,
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56.8 13.8 o
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1960 1970 1980 1990 2000 2010 ·a 0.0 0.0
year a.
(a) Temperature change since 1960

390 2000 2020 2040 2060 2080 2100

380 year
370
• FIGURE 28-16 Projected range of temperature increases
ea. 360
350
The IPCC proj ection s for the twenty-first century are based on
.e: t hree d ifferent scenarios of greenhou se gas emi ssion s. The red,
N
340 blue, and green data lines are projections based on high,
o moderate, and substantially reduced growth of greenhouse gas

~ t-
()
330
emissions, respective ly. Even under the most optimistic
320
assumptions, a continued increase in global warming is
310 predicted. Global temperature change is expressed relative to a
300 baseline of zero degrees of change. The baseline is the average
1960 1970 1980 1990 2000 2010 temperature from 1980 to 1990. Data from IPCC. 2007. Fourth
year assessment report: Summary for po/icymakers.
{b) Atmospheric e02 change since 1960

• FIGURE 28-15 Global warming parallels atmospheric

C02 increases (a) Global surface temperatures. Because global
temperature varíes considerably from year to year, this
temperature curve shows trend s by averaging each successive
year with the four years preced ing it. (b) Yearly average C02
concentrations in parts per mill ion These measurements we re
recorded at 11 ,1 55 feet (3,400 meters) above sea leve!, near
the summit of Mauna Loa, Hawaii . Data from the National
Oceanographic and Atmospheric Adm inistration (NOAA).

result of increased temperatures and local decreases in rainfall.

Agricultura! disruption resulting from such extremes in
weather could be disastrous for nations that are already strug-
(a) earroll Glacier, 1904
gling to grow enough food.

Community lnteractions May Be Disrupted The predicted rapid

rise in global temperature will exceed the rate at which natu-
ral selection can allow most organisms to adapt. The impact
of global warming on forests could be profound. As the world
warms, tree distributions will change based on their tolerance
for heat. For example, sugar maples may disappear from
northeastem U.S. forests, while sorne southeastem forests
could be replaced by grasslands. Warming is contributing to
massive destruction of evergreen trees in the western United

~ FIGURE 28-17 Glaciers are melting Photos taken from the

same vantage point in (a) 1904 and (b) 2004 document the
retreat of the Carroll Glacier in Glacier Bay, Alas ka. {b) earroll Gtacier, 2004

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represents a copyright violation.

States and Canada by the pine bark beetle, which is benefit-
ing from warmer winters and longer reproductive seasons
(see pp. 591-592). Coral reefs, already stressed by human ac-
tivities, are likely to suffer further damage from warmer wa-
ters, which drive out the symbiotic algae that provide them
with energy from photosynthesis. Further, as the oceans ab-
sorb more C0 2, their waters are becoming more acidic. Be-
cause acid dissolves limestone, this makes it more difficult for
corals to form their limestone skeletons.
Spring is arriving earlier in Euro pe and in the northeast-
ern United States. The growing season in Europe has in-
Earth Wat6A
Poles in Peril
Nowhere are the impacts of global warming more visible
than at the far ends of Earth, in our planet's unique ice-
dominated ecosystems. The Antarctic Peninsula in the
Southern Hemisphere is particularly vulnerable to global
warming because its average year-round temperature hovers
close to the freezing point. As global temperatures have
risen, temperatures on the peninsula have increased ata far
faster rate than the global average, and the flow of glacial ice
into the sea has accelerated. Huge ex pan ses of floating ice
shelves have disintegrated, with far-reaching consequences.
Sea ice creates conditions that favor abundant growth of
phytoplankton and algae, which provide food for larval
krill-shrimp-like zooplankton that are a keystone species in
the Antarctic community. Krill comprise a major portion of
the diet of seals, penguins, and several species of whales.
Researchers are concerned that the impact of krill loss may
reverberate throughout the Antarctic food web, harming all
the other organisms who feed on them.
In the Northern Hemisphere, Greenland's ice cover is
melting at record rates, and Arctic temperatures have
increased by twice the global average during the past 50
years. During the twenty-first century, loss of sea ice during
Arctic summers has repeatedly shattered records, and total
summer ice cover has decreased by roughly 30% during the
past 30 years. The loss of summer sea ice is bad news for
polar bears and other marine mammals that rely on ice floes
as nurseries for their young and as staging platforms for
hunting fish or seals. As summer ice has diminished, both
walrus and polar bear populations are moving onto land in
record numbers to give birth, and the loss of ice platforms is
reducing the expanse of ocean in which these top predators
can hunt. In 2008, in response to its declining numbers and
the projected continued loss of its habitat, the polar bear
was added to the list of threatened species-the first to be
listed primarily as a result of global warming. Complete loss
of sea ice, which climate models predict will occur within
the next century, could cause the extinction of polar bears in
the wild (Fig. E28-2).
The thawing of Earth's northern regions and the melting
of sea ice are cause for particular concern because both of
these situations generate positive feedback loops that
further accelerate global warming. Extensive areas of deep
frozen soil called permafrost underlie Siberia and Alaska.
These soils store huge deposits of organic matter from
prehistoric plants and animals, which have not broken down
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How Do Ecosystems Work? 549
creased by more than 10 days during the past three decades.
In the northern hemisphere, animals as diverse as butterflies,
birds, and whales are extending their ranges northward. The
cumulative weight of data from diverse sources worldwide
provides strong evidence that warming-related biological
changes have begun. Global warming is also predicted to in-
crease the range of tropical disease-carrying organisms, such
as malaria-transmitting mosquitoes, with negative conse-
quences for human health. Arctic and Antarctic wildlife from
polar bears to penguins are among the hardest hit, as de-
scribed in "Earth Watch: Poles in Peril."
because decomposer organisms are inhibited by the cold.
Ominously, so me of these regions are now thawing for the
first time since the last ice age ended more than 10,000
years ago. In Siberia, for example, a region of frozen peat
soil larger than Texas and Oklahoma combined is thawing,
creating giant bogs that could release enormous
quantities of C0 2 and methane gases as conditions
become favorable for decomposers. Because methane is
about 20 times as effective as carbon dioxide in trapping
atmospheric heat, this is of particular concern. In a
positive feedback loop, the greenhouse gases released by
thawing permafrost will further warm the atmosphere,
causing still more permafrost to thaw.
Sea ice reflects back most of the solar energy that hits
it, but the ocean water that is exposed when ice
disappears absorbs most solar energy, converting it to
heat. This situation is a second example of positive
feedback, because the heat absorbed by the exposed
water will further warm Earth's surface, causing more ice
to melt. Melting ice has al so opened access to areas of the
Arctic Ocean and its seabed that have been inaccessible
for centuries. This has fueled a rush by several countries
to lay claim to oil fields that are predicted to underlie the
Arctic Ocean floor. lronically, this suggests yet another
type of positive feedback: Burning this oil will further
accelerate the climate change that has now made it
accessible.
• FIGURE E28-2 Polar bears are on thin ice The polar bear
is now a threatened species as a result of global warming.
 550 Behavior and Ecology
Case Study re v i s ited
Dying Fish Feed an Ecosystem
Researchers investigating the sockeye salmon's return toan
Alaskan stream witness an awesome sight. Hundreds of brilliant
red bodies writhe in water so shallow that it barely covers them.
A female beats he r tail, excavating a shallow depression in the
gravel where she releases her coral-colored eggs; meanwhile, a
male showers them with sperm. But after their long and
strenuous migration, these adult salmon are dying. Their flesh is
tattered, their muscles wasted, and the final act of reproduction
saps the last of their energy. Soon, the stream is clogged with
dying, dead, and decomposing bodies-an abundance of
nutrients unavailable at any other time of the year. Eagles,
grizzly bears, and gulls gather to gorge themselves on the
fleeting bounty. Flies breed in the carcasses, feeding spiders,
bird s, and trout. The breeding cycles of local mink populations
have evolved around the event; females lactate at the time when
the salmon provide them with abundant food . Bears drag the
carcasses into the woods, often consuming only a part of each
fish. The remnants become food for detritus feeders and
decomposers, who liberate its nutrients to the soil where they
will be taken up by plants and reenter nutrient cycles.
Because 95% of the mass of the salmon's body was
accumulated while it lived in the ocean, the upstream
CHAPTER REVIEW
Summary of Key Concepts
28.1 How Do Energy and Nutrients Move
Through Ecosystems?
Ecosystems are sustai ned by a co nlinuo us input of energy from
sunlight and the constant recyding of nutrients. Energy enters
the biotic portio n of ecosystems w hen it is ham essed by photo-
synthelic organ isms. Nutrien ts are obtained by organis ms from
their living and non living environment and are recycled within
and amo ng ecosystems.
28.2 How Does Energy Flow Through Ecosystems?
Energy enters ecosystems through photosynthesis. Photosyn-
thetic o rga nisms act as co nduits of both energy a nd nutri ents
into biological communiti es. En ergy is passed upward through
trophic (feedi ng) levels. Autotrophs are photosynthetic a nd
are called producers, the lowest trophic level. Among het-
erotrophs, herbivores form the seco nd leve!: p rim ary co n-
su mers. Carn ivores act as secondary consumers w hen they
prey o n herbivores and as terliary or higher-level consu mers
w he n th ey eat other carnivo res. Omnivores, w hich consume
b o th plants and o ther a nim als, occupy multiple trop hi c levels.
Feeding relalionships in wh ich each troph ic leve! is repre-
sented by one o rganism are called food chains. ln natural
ecosystems, feed ing relationships are complex and are de-
scribed as food webs. Detritus feeders and decomposers
(which d iges t dead bo di es and wastes) use a nd release th e en-
ergy stored in these substances and liberate thei r nutrients,
whi ch th en reemer nutrient cycles.
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migration of salmon represents a n enormous tran sfer of
nutrients in an unusual direction: upstream. Historically,
researchers estímate that 500 mill ion pounds of salmon
migrated upstream in the U.S. Pacific Northwest each year,
contributing hundreds of thousands of pounds of nitrogen
and phosphorus to the region surrounding the Columbia River.
Now, dueto factors that include overfishing, river damming,
diversion of water for irrigation , runoff from agriculture, and
pollution of the estuaries (where several salmon species spend
a significant part of their life cycle), migratory salmon
populations in the region have declined by more than 90% in
the past century. The web of life that relied on the mighty
annual upstream flow of nutrients has been disrupted.
Consider This
Some salmon populations have been so thoroughly depleted
that they are listed as endangered or threatened under the
Endangered Species Act . Some peo ple a rgue that because these
salmon are also raised commercially in hatcheries and artificial
ponds, they should not be afforded this protection. Meanwhile,
researchers studying chinook salmon raised in hatcheries noted
a 2 5% decline in the average size of eggs of hatchery-reared
fish ove r just four generations. These eggs produce smaller
juvenile fish. Based on th is information, why is there a good
case for protecting populations of wild sal mon?
The high er the trophic )evel, the less energy available to sus-
tain it. Jn general, only about lO% of the energy cap tu red by or-
ganisms a t one trophic leve! is co nverted to the bodies of
organisms in the n ext higher leve!. As a result, plants are more
abundant than herbivores, and herbivores are more common
than carnivo res. The s torage o f ene rgy at each trophic level is il-
lustrated graphically as an energy pyram id. This inefficiency of
energy transfer through trophic levels leads to biological magni-
ficalion, the process by which toxic substances accumul ate in in-
creasingly high co ncentratio ns in o rgan isms occu pyi ng
progressively higher trophic levels.
28.3 How Do Nutrients Cycle Within and Among
Ecosystems?
A nutrient cycle depicts the movement of a particular nutrient
from its reservo ir, usually in the abio tic, or nonlivi ng, portion o f
the ecosystem; through the biotic, o r living, portion of the
ecosystem; and back to its reservo ir.
The m ajor reservo ir o f water is the ocea ns. During the hy-
drol ogic cycle, solar energy evapo rates water, which returns to
Earth as precipitation. Water flows into lakes and underground
aquifers, and is carried by rivers to the oceans. Relatively s mall
amounts ofwater are passed through food webs.
Major short-term carbon reservoirs are the oceans and the
atmosphere. Carbon enters producers through photosynthesis.
From autotrophs, it is passed through the food web and released
to the atmosphere as C02 during cellul ar respiration. Fossil fu.
els represent a long-term carbon reservoir; burning them is
changing Earth's di mate.
The major reservoir of nitrogen is the atrnosphere. Plants get
their nitrogen from nitrates and am monia. Nitrogen gas is cap-
tured by nitrogen-fixing bacteria, w hich release ammo nia. Other
bacteri a convert ammonia to ni trate, which can also be produced
by lightning. Industrial processes manufacture fertilizer containing
 How Do Ecosystems Work? 551

ammonia and nitrate. Nitrogen passes fro m producers to con- nutrient cycle 540 reservoir 540
sumers and is returned to the environment through excretion and omnivore 53 6 secondary consumer 535
the activities of detritus feeders and decomposers. Nitrogen gas is phosphorus cycle 543 tertiary consumer 535
returned to the air by denitrifying bacteria. phytoplankton 536 transpiration 541
Th e reservoir for phosphorus is in rocks, as phosphate, primary consumer 535 trophic leve! 535
which dissolves in water. Phosphate is absorbed by photosyn- producer 535 zooplankton 536
thetic organism s a nd is passed through food webs. Some phos-
p~ate is excreted, and the rest is returned to the soil and water
by decomposers. Some is carried to the oceans, where it is de-
posited in marine sediments. Humans mine phosphate-rich Thinking Through the Concepts
rock to produce fertilizer. Fill-in-the-Biank
l. Nearly alllife gets its energy from , which is
BioFii)C '" The Carbon Cycle captured by the process of . In contrast,
_ _ _ _ _ are constantly recycled during processes
28.4 What Happens When Humans called _ _ _ __
Disrupt Nutrient Cycles? 2. Photosynthetic organisms are called either or
Environmental disruption occurs when human activities interfere _ _ _ _ _ . The energy that these store and make
with the natural functioning of ecosystems. Human activities re- available to other o rganisms is called _ _ _ __
lease toxic substances and produce more nutrieiJ,ts than nutrient 3. Feeding levels within ecosystems are also called
cydes can efficiently process. The use of enormous quantities of _ _ _ _ _ . An illustratio n of these levels with only
fertilizer by agricultura! activities h as disrupted many aquatic o ne organism at each leve! is called a(n) _ _ _ __
ecosystems. Nutrient-rich water washing into lakes, rivers, and the Feeding relationships are most accurately depicted as
oceans causes excessive growth of aquatic plants and phytoplank-
ton; their subsequent death and decomposition deplete oxygen,
4. In general, o nly about percent of the energy
killing many aquatic organisms and producing "dead zones" in
available in o ne trophic leve! is captured by the leve!
nearshore coastal waters. By burning fossil fuels, we have over-
above it. This concept is based on what general physical
loaded the natural cyd es for sulfur, nitrogen, and carbon. Fossil
principie? _____
fuel combustion releases sulfur dioxide and nitrogen oxides. Tn the
atmosphere, these substances are converted to sulfuric acid and ni- S. Photosynthetic organisms are consumed by other
trie acid, which fall to Earth as acid deposition. Acidification of organisms collectively called , or
many freshwater lakes in the eastern United States has substan- _ _ _ _ _ . Organism s in higher trophic levels are
tially reduced their ability to support life. At high elevations, acid collectively called , or . Animals
deposition has sign ificantly damaged many eastern forests. and protists that feed on wastes and dead bodies are
Burning fossil fuels has substantially increased atmospheric called . The major types ofdecomposer
carbon di oxide, a greenhouse gas. This increase is correlated with organisms are and _ __ _ _
increased global temperatures, leading nearly all scientists to con- 6. During the nitrogen cyde, nitrogen gas is captured from
dude that global warming is due to human activities. Global its atmospheric reservoir by in the soil, and is
warming is causing glaciers and ice sheets to melt, and is influenc- then returned to this reservoir by . The two
ing the distribution and season al activities of wildlife. Scientists forms of nitrogen that are u sed by plants are _ _ _ __
believe global warming is beginning to have a major impact on a nd _ _ _ __
precipitation and weather pattems, with unpredictable results. 7. Two relatively short-term reservoirs for carbon are the
_ _ _ _ _ and . Carbon in these short-
KeyTerms term reservoirs is in the form of . Two long-
acid deposition 545 food chain 536 term reservoirs for carbo nare and
aquifer 541 food web 536
autotroph 535 fossil fuel 542
biodegradable 539 global warming 546 Review Questions
biogeochemical cyde 540 greenhouse effect 546 l. What makes the flow of energy through ecosystems
biological greenhouse gas 546 fundamentally differen t from the flow of nutrients?
magniflcation 539 herbivore 535
2. What is an auto troph ? What trophic leve! does it occupy,
biomass 538 heterotroph 535
and what is its importance in ecosystems?
carbon cycle 541 hydrologic cycle 540
carnivore 535 legume 543 3. Define net prirnary production. Would you predict higher
consumer 535 macronutrient 540 productivity in a farm pond oran alp ine lake? Explain
decomposer 53 7 micronutrient 540 your answer.
deforestation 546 net primary production 4. List the first three trophic levels. Among the consumers,
denitrifying bacteria 543 535 whi ch are most abundant? Why would you predict
detritus feeder 537 nitrogen cycle 542 that there will be a greater biomass of plants than
energy pyramid 538 nitrogen flxation 543 h erbivores in any ecosystem ? Relate your answer to
estuary 535 nutrient 533 the "10% law."

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 552 Behavior and Ecology
5. How do food chains and food webs differ? Which is the
more accurate representa tion of actual feeding
relationships in ecosystems?
6. Define detritus feeders and decomposers, and explain their
importance in ecosystems.
7 . Trace the movement of carbon from one of its reservoirs
through the biotic community and back to the reservoir.
How h ave human activities altered the carbon cycle, and
what are th e jml?lications for future climate?
8. Explain how nitrogen gets from the atmosphere into a
p lant's body.
9. Trace a phosphorus molecule from a phosphate-rich rock
into a carnivore. What makes the phosphorus cycle
fundamentally different from the carbon and nitrogen
cycles?
10. Trace the movement of a water molecule from the ocean,
through a plant body, and back to the ocean, d escribing
all the intermediare stages and processes.
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Applying the Concepts
l. BioEthics What could your college or university doto
reduce its contribution to global warming? Be specific, and,
if possible, offer practica! altematives to current practices.
2. Define and give an examp le of biological magnification .
What qualities are present in materials that undergo
biological magnification? In which trophic leve! are the
problems worst, and why?
3. BioEthics Discuss the contribution ofhuman
population growth to (a) acid rain and (b) global
warming.
4. Describe what would happen to a population of deer if all
predators were removed and hunting banned . Include
effects on vegetation as well as on the deer population
itself. Relate your answer to carrying capacity as discussed
in Chapter 26.
{M;jGo to www.masteringbiology.com for practice quizzes,
~ activities, eText, videos, current events, and more.