CH 04

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Biological Communities and Species Interactions

Any species of bug is an irreplaceable marvel, equal to the works of art which we religiously preserve in our museums.
Claude Levi-Strauss
OBJECTIVES
After studying this chapter, you should be able to:
• describe how environmental factors determine which species live in a LEARNING ONLINE
given ecosystem and where or how they live. Visit our website at www.mhhe.com/environmentalscience for aids
• understand how random genetic variation and natural selection lead to help you study this chapter. You’ll find practice quizzes, key term
to evolution, adaptation, niche specialization, and partitioning of flashcards, career information, case studies, current environmental
resources in biological communities. news, and regional examples of important environmental issues.
• compare and contrast interspecific predation, competition, symbiosis, You’ll also discover active links to valuable web pages including:
commensalism, mutualism, and coevolution. National Marine Mammal Laboratory
• discuss productivity, diversity, complexity, and structure of biological Cornell Lab of Ornithology
communities and how these characteristics might be connected to
Biological Resources Division, USGS
resilience and stability.
National Biological Information Infrastructure (NBII) Home Page
• explain how ecological succession results in ecosystem development
and allows one species to replace another. U.S. Long Term Ecological Research Network, Home
• give some examples of exotic species introduced into biological
communities and describe the effects such introductions can have on
indigenous species.
The bears, birds, and fish of the McNeil River, Alaska, form an interconnected biological community
together with terrestrial and aquatic plants and invertebrates. © John Warden/Stone.
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Why Trees Need Salmon

Ecologists have long known that salmon in the Pacific Northwest need these insects, and nutrients from salmon thus work through the entire
clear streams to breed, and that clear streams need healthy forests. forest ecosystem.
Surprising new evidence now indicates that forests themselves need In a separate study, ecologists Robert Naiman and James Helfield
salmon to remain healthy. found that trees along salmon-rich rivers can grow up to three times
Pacific salmon (Onchorhyncus spp.) are anadromous: they hatch as fast as trees along streams without salmon. This is important, they
in freshwater lakes and streams, spend much of their lives at sea, point out, because salmon stocks are dwindling throughout the Pacific
then return to the stream where they were born to breed and die. To Northwest. In Washington, Oregon, and California, salmon populations
reproduce successfully, these fish require clear, cold, shaded streams have fallen by 90 percent from their historic numbers. Because of this
and a clean gravel riverbed. When forests are cut, sediment washes close relationship, they argue, forest management and fish manage-
down hillsides and into streams, clogging gravel streambeds and suf- ment need to be integrated. Each population—rainforest trees and
focating eggs. Sediment also absorbs sunlight, warming water and ocean-going fish—affects the stability of the other.
reducing oxygen saturation in the water. Lower oxygen level reduces Apparently, salmon need healthy forests, and forests need healthy
survival rates of eggs and young fish. salmon populations. Stream ecosystems need standing trees to retain
Every year, as millions of fish return to spawn and die, they pro- soil and provide shade. So healthy streams depend on fish, just as the
vide a banquet for bears, eagles, and other species that gorge them- fish depend on the streams. As this case shows, links among organisms
selves on the fat-rich fish. Ecologist Thomas Riemchen has found that in an ecosystem are intricate, often subtle, and essential for ecological
bears fishing in British Columbia’s rivers can catch 500 fish in a six- stability. Relationships between apparently separate environments, such
week salmon migration season (about 12 fish per bear per day). He as rivers and forests, can be equally important. In this chapter we’ll
also estimates that a bear gets 70 percent of its annual protein intake explore some of these relationships among organisms and between
from fish. But bears also drag tons of fish up on shore and leave half- organisms and their environment.
eaten carcasses strewn about the forest floor. Riemchen calculates
that these scattered fish fertilize the forest at a rate of about 120 kg To read more:
of nitrogen per acre. British Columbia’s rainforests, with at least Reimchen, T. E., D. Mathewson, M. D. Hocking, J. Moran, and D. Harris. 2003.
30,000 fishing bears, may receive 60 million kg of salmon each year. Isotopic evidence for enrichment of salmon-derived nutrients in vegetation,
Nitrogen is often a limiting nutrient for rainforest vegetation. Between soil and insects in riparian zones in coastal British Columbia. American
one-quarter and one-half of the nitrogen in a towering Sitka Spruce Fisheries Society Symposium 34:59–69.
or Douglas Fir may derive from salmon carcasses. Helfield J. M., and R. J. Naiman. 2001. Effects of salmon-derived nitrogen on
In addition to fertilizing trees, salmon carcasses provide food for riparian forest growth and implications for stream productivity. Ecology
insects and other scavengers. Birds and other predators consume 82(9):2403–09.
WHO LIVES WHERE, AND WHY? relative to demand is the critical determinant in the distribution of
that species. Ecologist Victor Shelford later expanded this princi-
“Why” questions often are the stimulus for scientific research, but ple of limiting factors by stating that each environmental factor has
the research itself centers on “how” questions. Why, we wonder, both minimum and maximum levels, called tolerance limits,
does a particular species live where it does? More to the point, how beyond which a particular species cannot survive (fig. 4.1) or is
is it able to live there? How does it deal with the physical resources unable to reproduce. The single factor closest to these survival lim-
of its environment and are some of its techniques unique? How its, he postulated, is the critical limiting factor that determines
does it interact with the other species present? And what gives one where a particular organism can live.
species an edge over another species in a particular habitat? At one time, ecologists accepted this concept so completely
In this section we will examine some specific ways organisms that they called it Liebig’s or Shelford’s law and tried to identify
are limited by the physical aspects of their environment. We then unique factors limiting the growth of every population of plants
will discuss how members of a biological community interact, and animals. For many species, however, we find that the inter-
pointing out a few of the difficulties ecologists encounter when action of several factors working together, rather than a single lim-
they attempt to discern patterns and make generalizations about iting factor, determines biogeographical distribution. If you have
community interactions and organization. ever explored the rocky coasts of New England or the Pacific
Northwest, for instance, you probably have noticed that mussels
and barnacles endure extremely harsh conditions but generally are
Critical Factors and Tolerance Limits sharply limited to an intertidal zone where they grow so thickly
Every living organism has limits to the environmental conditions that they often completely cover the substrate. No single factor
it can endure. Temperatures, moisture levels, nutrient supply, soil determines this distribution. Instead, a combination of tempera-
and water chemistry, living space, and other environmental factors ture extremes, drying time between tides, salt concentrations, com-
must be within appropriate levels for life to persist. In 1840, Jus- petitors, and food availability limits the number and location of
tus von Liebig proposed that the single factor in shortest supply these animals.
72 PART ONE Principles for Understanding Our Environment

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Zone of Zone of Optimal range Zone of Zone of

intolerance physiological physiological intolerance
stress Species abundant stress
nt Sp
ecie
que s in
nfre frequ
ies i
t Spec ent
Species ab
Species absen sent
Too low: Too high:

Optimum
lower limit of tolerance upper limit of tolerance
Environmental gradient
FIGURE 4.1 The principle of tolerance limits states that for every environmental factor, an organism has both maximum and minimum levels
beyond which it cannot survive. The greatest abundance of any species along an environmental gradient is around the optimum level of the critical fac-
tor most important for that species. Near the tolerance limits, abundance decreases because fewer individuals are able to survive the stresses imposed by
limiting factors.
For other organisms, there may be a specific critical factor that, effects of human activities. Lichens and eastern white pine are less
more than any other, determines the abundance and distribution of restricted in habitat than locoweeds, but are indicators of air pol-
that species in a given area. A striking example of cold intolerance lution because they are extremely sensitive to sulfur dioxide and
as a critical factor is found in the giant saguaro cactus (Carnegiea acid precipitation. Bull thistle is a weed that grows on disturbed
gigantea), which grows in the dry, hot Sonoran desert of southern soil but is not eaten by cattle; therefore, an abundant population
Arizona and northern Mexico (fig. 4.2). Saguaros are extremely sen- of bull thistle in a pasture is a good indicator of overgrazing.
sitive to low temperatures. A single exceptionally cold winter night Similarly, anglers know that trout species require clean, well-
with temperatures below freezing for 12 hours or more will kill oxygenated water, so the presence or absence of trout can be an
growing tips on the branches. Young saguaros are more susceptible indicator of water quality.
to frost damage than adults, but seedlings typically become estab-
lished under the canopy of small desert trees such as mesquite that
shield the young cacti from the cold night sky. Unfortunately, the
popularity of grilling with mesquite wood has caused extensive har-
vesting of the nurse trees that once sheltered small saguaros,
adversely affecting reproduction of this charismatic species.
Animal species, too, exhibit tolerance limits that often are
more critical for the young than for the adults. The desert pupfish
(Cyprinodon), for instance, occurs in small isolated populations in
warm springs in the northern Sonoran desert. Adult pupfish can
survive temperatures between 0°C and 42°C (a remarkably high
temperature for a fish) and are tolerant to an equally wide range of
salt concentrations. Eggs and juvenile fish, however, can only live
between 20°C and 36°C and are killed by high salt levels. Repro-
duction, therefore, is limited to a small part of the range of adult
fish, which is often restricted anyway by the size of the small
springs and desert seeps in which the species lives.
Sometimes the requirements and tolerances of species are use-
ful indicators of specific environmental characteristics. The presence
or absence of such species can tell us something about the commu-
nity and the ecosystem as a whole. Locoweeds, for example, are
small legumes that grow where soil concentrations of selenium are FIGURE 4.2 Saguaro cacti, symbolic of the Sonoran desert, are
an excellent example of distribution controlled by a critical environmen-
high. Because selenium is often found with uranium deposits, tal factor. Extremely sensitive to low temperatures, saguaros are found
locoweeds have an applied economic value as environmental only where minimum temperatures never dip below freezing for more than
indicators. Such indicator species also may demonstrate the a few hours at a time. © William P. Cunningham.
www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 73

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Natural Selection, Adaptation,

and Evolution
How is it that mussels have developed the ability to endure pound-
ing waves, daily exposure to drying sun and wind, and seasonal
threats of extreme cold or hot temperatures? What enables desert
pupfish to tolerate hot, mineral-laden springs? How does the
saguaro survive in the harsh temperatures and extreme dryness of
the desert? We commonly say that each of these species is
“adapted” to its special set of conditions, but what does that mean?
In this section, we will examine one of the most important con-
cepts in biology: how species acquire traits that allow them to live
in unique ways in particular environments.
In common use, to adapt means to modify slightly, usually
temporarily. We use the term adapt in two ways. One is a limited
range of physiological modifications (called acclimation) available
to individual organisms. If you keep house plants inside all winter, FIGURE 4.3 Giraffes don’t have long necks because they stretch
for example, and then put them out in full sunlight in the spring, to reach tree-top leaves, but those giraffes that happened to have longer
necks got more food and had more offspring, so the trait became fixed in
they get sunburned. If the damage isn’t too severe, your plants will the population. © Corbis/Volume 6.
probably grow new leaves with a thicker cuticle and denser pig-
ments that protect them from the sun. But this change isn’t per-
manent. Another winter inside will make them just as sensitive to their prey. One of the most unusual species is the woodpecker
the sun as before. Furthermore, the changes they acquire are not finch, which pecks at tree bark for hidden insects. Lacking the
passed on to their offspring. woodpecker’s long tongue, however, the finch uses a cactus spine
In biological terms, adaptation refers specifically to inherited as a tool to extract bugs.
traits that gradually change a population or a species, not an indi- The amazing variety of colors, shapes, and sizes of dogs, cats,
vidual. These inherited traits allow a species to live in a particular rabbits, fish, flowers, vegetables, and other domestic species is evi-
environment. This process is explained by the theory of evolution, dence of deliberate selective breeding. The various characteristics
developed by Charles Darwin and Alfred Wallace. According to of these organisms arose through mutations. We simply kept the
this theory, species change gradually through competition for ones we liked. Note that sexual reproduction helps to redistribute
scarce resources and natural selection, a process in which those genetic material in new and novel combinations that greatly
members of a population that are best suited for a particular set of increase the variation and diversity we see in both wild and domes-
environmental conditions will survive and produce offspring more tic species. Organisms that reproduce asexually can evolve, but
successfully than their ill-suited competitors. often do so very slowly.
Natural selection acts on preexisting genetic diversity created What environmental factors cause selective pressure and influ-
by a series of small, random mutations (changes in genetic mater- ence fertility or survivorship in nature? They include (1) physio-
ial) that occur spontaneously in every population. These mutations logical stress due to inappropriate levels of some critical
produce a variety of traits, some of which are more advantageous environmental factor, such as moisture, light, temperature, pH, or
than others in a given situation. Where resources are limited or specific nutrients; (2) predation, including parasitism and disease;
environmental conditions place some selective pressure on a pop- (3) competition; and (4) chance. In some cases the organisms that
ulation, individuals with those advantageous traits become more survive environmental catastrophes or find their way to a new habi-
abundant in the population, and the species gradually evolves or tat where they start a new population may simply be lucky rather
becomes better suited to that particular environment. Although each than more fit or better suited to subsequent environmental condi-
change may be very slight, many mutations over a very long time tions than their less fortunate contemporaries.
have produced the incredible variety of different life-forms that we Be sure you understand that while selection affects individu-
observe in nature (fig. 4.3). als, evolution and adaptation work at the population level. Indi-
The variety of finches observed by Charles Darwin on the viduals don’t evolve; species do. Each individual is locked in by
Galápagos Islands is a classic example of speciation driven by genetics to a particular way of life. Most plants, animals, or
availability of different environmental opportunities (fig. 4.4). microbes have relatively limited ability to modify their physical
Originally derived from a single seed-eating species that somehow makeup or behavior to better suit a particular environment. Over
crossed the thousands of kilometers from the mainland, the finches time, however, random genetic changes and natural selection can
have evolved into a dozen or more distinct species that differ change an entire population.
markedly in appearance, food preferences, and habitats they Given enough geographical isolation or selective pressure, the
occupy. Fruit eaters have thick parrot-like bills; seed eaters have members of a population become so different from their ancestors
heavy, crushing bills; insect eaters have thin probing beaks to catch that they may be considered an entirely new species that has

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Warbler finch Cactus ground finch
Woodpecker Sharp-beaked
finch ground finch
er
Warbl
Small finch Small ground
insectivorous finch
ing bills
tree finch Prob Gr
Cactus ou
eaters
nd
es
f in
ch
Medium
ch
Cr
ills
fin
es
us
ground
gb
hin
e
finch
n
Tre
Large
spi
Insect eaters
g
insectivorous
bills
G ra
tree finch Parrot-like
Seed eaters
Large
bill
Fruit eaters ground

finch
Vegetarian
tree finch
FIGURE 4.4 Some species of Galápagos Island finches. Although all are descendents of a common ancestor, they now differ markedly in appear-
ance, habitat, and feeding behavior. Ground finches (lower right) eat cactus leaves; warbler finches (upper left) eat insects; others eat seeds or have mixed
diets. The woodpecker finch (upper left) pecks tree bark as do woodpeckers, but lacks a long tongue. Instead, it uses cactus spines as tools to extract insects.
Source: From Peter H. Raven and George B. Johnson, Biology, 4th edition. Copyright © 1996 McGraw Hill Company, Inc. All rights reserved. Reprinted by permission.
replaced the original one. Alternatively, isolation of population sub- needs to swim in order to eat; it has webbed feet because some
sets by geographical or behavioral factors that prevent exchange of ancestor happened to have a gene for webbed feet that gave it some
genetic material can result in branching off of new species that advantage over other ducks in its particular pond and because those
coexist with their parental line. Suppose that two populations of genes were passed on successfully to its offspring. A variety of dif-
the same species become separated by a body of water, a desert, or ferent genetic types are always present in any population, and nat-
a mountain range that they cannot cross. Over a very long time— ural selection simply favors those best suited for particular
often millions of years—random mutations and different environ- conditions. Whether there is a purpose or direction to this process
mental pressures may cause the populations to evolve along such is a theological question rather than a scientific one and is beyond
dissimilar paths that they can no longer interbreed successfully the scope of this book.
even if the opportunity to do so arises. They have now become sep-
arate species as in the case of the Galápagos finches. The barriers
that divide subpopulations are not always physical. In some cases,
The Ecological Niche
behaviors such as when and where members of a population feed, Habitat describes the place or set of environmental conditions in
sleep, or mate—or how they communicate—may separate them which a particular organism lives. A more functional term, the
sufficiently for divergent evolution and speciation to occur even ecological niche, is a description of either the role played by a
though they occupy the same territory. species in a biological community or the total set of environmental
Natural selection and adaptation can cause organisms with a factors that determine species distribution. Niches as community
similar origin to become very different in appearance and develop roles—describing how a species obtains food, what relationships it
different habits over time, but they can also result in unrelated has with other species, and the services it provides its community,
organisms coming to look and act very much alike. We call this for example—were first described by the British ecologist, Charles
latter process convergent evolution. The cactus-eating Galápagos Elton in 1927. Thirty years later, the American limnologist G. E.
finches (fig. 4.4), for example, look and act very much like par- Hutchinson proposed a more biophysical definition of this concept.
rots even though they are genetically very dissimilar. The features Every species, he pointed out, has a range of physical and chemi-
that enable parrots to eat fruit successfully work well for these cal conditions (temperature, light levels, acidity, humidity, salinity,
finches also. etc.) as well as biological interactions (predators and prey present,
A common mistake is to believe that organisms develop cer- defenses, nutritional resources available, etc.) within which it can
tain characteristics because they want or need them. This is incor- exist. Figure 4.1, for example, shows the abundance of a hypothet-
rect. A duck doesn’t have webbed feet because it wants to swim or ical species along a single factor gradient. If it were possible to

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Abundance
Niche
breadth
Competition
(a) Resource gradient
Divergence
Abundance
Less
niche
breadth
FIGURE 4.5 The giant panda feeds exclusively on bamboo.

Although its teeth and digestive system are those of a carnivore, it is not (b) Resource gradient
a good hunter, and has adapted to a vegetarian diet. In the 1970s, huge
acreages of bamboo flowered and died, and many pandas starved.
© William P. Cunningham. FIGURE 4.6 Resource partitioning and niche specialization
caused by competition. Where niches of two species overlap along a
resource gradient, competition occurs (shaded area in (a)). Individuals
occupying this part of the niche are less successful in reproduction so that
graph simultaneously all of the factors that affect a particular characteristics of the population diverge to produce more specialization,
species, a multidimensional space would result that describes the narrower niche breadth, and less competition between species (b).
ecological niche available to that species.
The idea of niches can be further defined in terms of funda-
mental niche and realized niche. A species’ fundamental niche is either migrate to a new area, become extinct, or change its behav-
the full range of resources or habitat it could exploit if there were ior or physiology in ways that minimize competition. We call this
no competition with other species. A species’ realized niche, the latter process of niche evolution resource partitioning (fig. 4.6).
resources or habitat it actually uses, may be much less than its fun- It can produce high levels of specialization that allow several
damental niche. species to utilize different parts of the same resource and coexist
Some species, like raccoons or coyotes, are generalists that eat within a single habitat (fig. 4.7).
a wide variety of food and live in a broad range of habitats (includ- Niche specialization also can create behavioral separation that
ing urban areas). Others, such as the panda (fig. 4.5), are special- allows subpopulations of a single species to diverge into separate
ists that occupy a very narrow niche. Specialists often tend to be species. Why doesn’t this process continue until there is an infi-
rarer than generalists and less resilient to disturbance or change. nite number of species? The answer is that a given resource can be
A few species such as elephants, chimpanzees, and baboons partitioned only so far. Populations must be maintained at a mini-
learn how to behave from their social group and can invent new ways mum size to avoid genetic problems and to survive bad times. This
of doing things when presented with new opportunities or challenges. puts an upper limit on the number of different niches—and there-
Most organisms, however, are limited by genetically determined fore the number of species—that a given community can support.
physical structure and instinctive behavior to established niches. Perhaps you haven’t thought of time as an ecological factor,
Over time, though, niches can evolve, just as physical char- but niche specialization in a community is a 24-hour phenomenon.
acteristics do. The law of competitive exclusion states that no two Swallows and insectivorous bats both catch insects, but some insect
species will occupy the same niche and compete for exactly the species are active during the day and others at night, providing
same resources in the same habitat for very long. Eventually, one noncompetitive feeding opportunities for day-active swallows and
group will gain a larger share of resources while the other will night-active bats.

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Cape May
warbler 60 ft
Blackburnian
warbler
50 ft
Black-throated
green warbler
40 ft
30 ft
Bay-breasted
warbler 20 ft
FIGURE 4.7 Resource partitioning

and the concept of the ecological niche are
10 ft demonstrated by several species of wood war-
blers that use different strata of the same forest.
This is a classic example of the principle of
competitive exclusion.
Yellow-rumped Ground Source: Original observation by R. H. MacArthur.
warbler
SPECIES INTERACTIONS
Predation and competition for scarce resources are major factors in
evolution and adaptation. Not all biological interactions are com-
petitive, however. Organisms also cooperate with, or at least tol-
erate, members of their own species as well as individuals of other
species in order to survive and reproduce. In this section, we will
look more closely at the different interactions within and between
species that shape biological communities.
Exploitation Predation and Parasitism

All organisms need food to live. Producers make their own food,
and consumers eat organic matter created by other organisms. In
most communities, as we saw in chapter 3, photosynthetic organ-
isms are the producers. Consumers include herbivores, carnivores, FIGURE 4.8 Insect herbivores are predators as much as are lions
omnivores, scavengers, and decomposers. With which of these cat- and tigers. In fact, insects consume the vast majority of biomass in the world.
egories do you associate the term predator? Ecologically, the term Complex patterns of predation and defense have often evolved between
has a much broader meaning than you might expect. A predator insect predators and their plant prey. © Ray Coleman/Photo Researchers, Inc.
in an ecological sense, is an organism that feeds directly upon
another living organism, whether or not it kills the prey to do so build hard shells that protect them from fish but can be crushed by
(fig. 4.8). By this definition herbivores, carnivores, and omnivores limpets and other mollusks. Predators also may change their feed-
that feed on live prey are predators, but scavengers, detritivores, ing targets. Adult frogs, for instance, are carnivores, but the tad-
and decomposers that feed on dead things are not. poles of most species are grazing herbivores. Sorting out the
Predatory relationships can be complex, as in the case of trophic levels in these communities can be very difficult.
marine shellfish. Many crustaceans, mollusks, and worms release Predation is an important factor in evolution. Predators prey
eggs directly into the water, and the eggs and free-living larval and most successfully on the slowest, weakest, least fit members of their
juvenile stages are part of the floating community, or plankton target population, thus reducing competition, preventing excess pop-
(fig. 4.9). Planktonic animals feed upon each other and are food ulation growth, allowing successful traits to become dominant in
for successively larger carnivores, including small fish. As prey the prey population, and making the prey population stronger and
species mature, their predators change. Barnacle larvae are plank- healthier. As the poet Robinson Jeffers said, “What but the wolf’s
tonic and are eaten by fish. Adult barnacles, on the other hand, tooth whittled so fine/The fleet limbs of the antelope?”.

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FIGURE 4.9 Microscopic plants and animals form the basic

levels of many aquatic food chains and account for a large percentage of
total world biomass. Many oceanic plankton are larval forms that have
habitats and feeding relationships very different from their adult forms.
© D. P. Wilson/Photo Researchers, Inc.
Prey species have evolved many protective or defensive adap-

tations to avoid predation. In plants, for instance, this often takes
the form of thick bark, spines, thorns, or chemical defenses. Ani-
mal prey may become very adept at hiding, fleeing, or fighting back
against predators. Predators, in turn, evolve mechanisms to over-
come the defenses of their prey. This process in which species exert
selective pressure on each other is called coevolution.
Parasites are organisms that feed on a host, or take resources
from it, without killing the host. Some parasites do little damage:
a mosquito takes blood but usually causes little damage. Others
cause significant harm and may eventually kill a host. Pathogens
(disease-causing organisms) are often considered parasites. Your
immune system is an evolved defense against pathogens in our
environment.
Keystone Species
A keystone species is a species or group of species whose feeding
activity has an inordinate influence on the structure of its community.
Originally, keystone species were thought to be top predators, such
as wolves, whose presence limits the abundance of herbivores and
FIGURE 4.10 Giant kelp is a massive alga that forms dense
“forests” off the Pacific coast of California. It is a keystone species in that
thereby reduces their grazing or browsing on plants. Recently, it has it provides food, shelter, and structure essential for a whole community.
been recognized that less conspicuous species also play essential Removal of sea otters allows sea urchin populations to explode. When the
community roles. Certain tropical figs, for example, bear during sea- urchins destroy the kelp, many other species suffer as well. © Randy Morse/Tom
sons when no other fruit is available for frugivores (fruit-eating ani- Stack & Associates.
mals). If these figs were removed, many animals would starve to

death during periods of fruit scarcity. With those animals gone, many essential key. A classic example is in the Pacific kelp forests, where
other plant species that depend on them at other times of the year for towering columns of kelp (algae) shelter myriad fish, shellfish, and
pollination and seed-dispersal would disappear as well. mammals (fig. 4.10). The sheltering kelp could be regarded as the
Even microorganisms can play vital roles. In some forest eco- key to community structure. Sea urchins, however, feed on the kelp
systems, mycorrhizae (fungi associated with tree roots) are essen- and determine their number and distribution while sea otters reg-
tial for mineral mobilization and absorption. If the fungi die, so do ulate urchins and kelp provides a resting place for dozing otters.
the trees and many other species that depend on a healthy forest Which of these species is the most important? Each depends on
community. Rather than being a single species, mycorrhizae are and affects the others. Perhaps we should think in terms of a “key-
actually a group of species that together fulfill a keystone function. stone set” of organisms in some ecosystems. (See “Oreas, Otters,
Often a number of species are intricately interconnected in Urchins, and Kelp: Disrupting a Marine Food Web,” on the Online
biological communities so that it is difficult to tell which is the Learning Center in the chapter 4 Case Studies.)

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Competition Intraspecific competition can be especially intense because

members of the same species have the same space and nutritional
Competition is another kind of antagonistic relationship within a requirements; therefore, they compete directly for these environ-
community. For what do organisms compete? To answer this ques- mental resources. How do plants cope with intraspecific competi-
tion, think again about what all organisms need to survive: energy tion? The inability of seedlings to germinate in the shady
and matter in usable forms, space, and specific sites for life activ- conditions created by parent plants acts to limit intraspecific com-
ities (What Do You Think? p. 80). Plants compete for growing petition by favoring the mature, reproductive plants.
space for root and shoot systems so they can absorb and process Symbiotic relationships often enhance the survival of one or
sunlight, water, and nutrients. Animals compete for living, nesting, both partners. Symbiotic relationships often entail some degree of
and feeding sites, as well as for food, water, and mates. Competi- coadaptation or coevolution of the partners, shaping—at least in
tion among members of the same species is called intraspecific part—their structural and behavioral characteristics. An interest-
competition, whereas competition between members of different ing case of mutualistic coadaptation is seen in Central and South
species is called interspecific competition. American swollen thorn acacias and their symbiotic ants. Acacia
You can observe interspecific competition if you look closely ant colonies live within the swollen thorns on the acacia tree
at a patch of weeds growing on good soil early in the summer. First branches and feed on two kinds of food provided by the trees: nec-
of all, many weedy species attempt to crowd out their rivals by tar produced in glands at the leaf bases and special protein-rich
producing prodigious numbers of seeds. After the seeds germinate, structures produced on leaflet tips. The acacias thus provide shel-
the plants race to grow the tallest, cover the most ground, and get ter and food for the ants. Although they spend energy to provide
the most sun. You may observe several strategies to do this. For these services, the trees are not physically harmed by ant feeding.
example, vines don’t build heavy stems of their own; they simply Animals also have developed adaptive responses to intraspe-
climb up over their neighbors to get to the light. cific competition. Two major examples are varied life cycles and
Species also race to new territory. Plants with highly mobile territoriality. The life cycles of many invertebrate species have
seeds can reach and colonize open ground ahead of other species juvenile stages that are very different from the adults in habitat and
(fig. 4.11). Some plants secrete substances that inhibit the growth feeding. Compare a leaf-munching caterpillar to a nectar-sipping
of seedlings near them, including their own and those of other adult butterfly or a planktonic crab larva to its bottom-crawling
species. This strategy is particularly significant in deserts where adult form. In these examples, the adults and juveniles of each
water is a limiting factor. species do not compete because they occupy different ecological
We often think of competition among animals as a bloody bat- niches.
tle for resources. A famous Victorian description of the struggle You may have observed robins chasing other robins during the
for survival was “nature red in tooth and claw.” In fact, a better mating and nesting season. Robins and many other vertebrate
metaphor is a race. Have you ever noticed that birds always eat species demonstrate territoriality, an intense form of intraspecific
fruits and berries just before they are ripe enough for us to pick? competition in which organisms define an area surrounding their
Having a tolerance for bitter, unripe fruit gives them an advantage home site or nesting site and defend it, primarily against other
in the race for these food resources. Many animals tend to avoid members of their own species. Territoriality helps to allocate the
fighting if possible. It’s not worth getting injured. Most confronta- resources of an area by spacing out the members of a population.
tions are more noise and show than actual fighting. It also promotes dispersal into adjacent areas by pushing grown
offspring outward from the parental territory.
Territory size depends on the size of the species and the
resources available. A pair of robins might make do with a subur-
ban yard, but a large carnivore like a tiger may need thousands of
square kilometers.
Symbiosis
In contrast to predation and competition, symbiotic interactions
between organisms can be nonantagonistic. Symbiosis is the inti-
mate living together of members of two or more species. Com-
mensalism is a type of symbiosis in which one member clearly
benefits and the other apparently is neither benefited nor harmed.
Cattle often are accompanied by cattle egrets, small white shore
birds who catch insects kicked up as the cattle graze through a field.
The birds benefit while the cattle seem indifferent. Many of the
mosses, bromeliads, and other plants growing on trees in the moist
FIGURE 4.11 Dandelions and other opportunistic species gen- tropics are also considered to be commensals (fig. 4.12). These epi-
erally produce many highly mobile offspring. © William P. Cunningham. phytes get water from rain and nutrients from leaf litter and dust

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What do you think?
Understanding Competition 1. Things need to be organized in such a

way that the outcome can clearly be
Ecology is a relatively young science. Con- linked to a particular cause. In other
sequently, many ecological processes are words, differences in insect survival rates
incompletely understood. How a community need to be clearly attributable to com-
comes to have its particular organization is petition and not to other factors. Karban
one area of uncertainty. Some ecologists feel accomplished this by making his plant/
that physical factors are the most important insect groups as uniform as possible,
determinants in community organization, except for the presence or absence of
while others feel that interspecific competi- competitors. He eliminated genetic dif-
tion is most important. ferences between plants by using plants
How can we find out which view is from the same clone. He was careful to
correct? Ecologists employ the scientific put the same numbers of insects on
method, as described in chapter 2, to bet- each plant to eliminate animal density as
ter understand community dynamics. This a factor, and so on.
process is mostly refined common sense
and its basic elements can be useful in 2. The data collected must be a reliable
everyday life. representation of the larger situation and
Once ecologists have decided on the not simply the result of chance. This is
concept to be investigated, they look for a usually accomplished by replicating the
specific situation that can either be observed procedure many times. Instead of setting
or manipulated to provide relevant informa- up just a few plants with one or both
tion. For example, ecologist Richard Karban insects present, Karban set up 30 plants
was interested in how competition affected with each treatment. The procedure was Spittlebugs produce mounds of foam under
a community. He learned that larvae of two repeated a second year. This gave him a which they hide from predators while feeding on
insect species, the meadow spittlebug and cumulative total of 60 plants that had just host plants. © Milton Tierney/Visuals Unlimited.
the calendula plume moth, both feed and spittlebugs, 60 plants having just moths,
develop on the seaside daisy, a common and 60 plants each having both or nei- petition can play an important role in influ-
beach plant on the American west coast. ther spittlebugs and moths. With such a encing densities of plant-feeding insects.
The specific question to be investigated large number of replications it was highly Notice the caution expressed in these
was: Does competition affect these two likely that differences in survival rates words. He did not claim to have proven any-
insect species, therefore impacting commu- were, in fact, the result of competition thing. Instead, his study “supports the con-
nity organization? and not simply chance occurrences. tention.” Second, he states competition
Competition might reduce survival rate, 3. Finally, conclusions must be justified by “can play an important role,” instead of using
larval growth, or both. Karban’s procedure the data. Karban’s statistical analysis stronger language. And finally, he restricts
involved setting up four groups of plants at revealed that spittlebug persistence was these conclusions to plant-feeding insects.
Bodega Bay, CA: one got both spittlebugs nearly 40 percent higher when the plume Karban carefully avoids drawing conclusions
and moths, another got only spittlebugs, moths were absent. Plume moth persis- beyond the realm supported by his data.
another only moths, and a fourth had nei- tence was not significantly affected by Based on a healthy skepticism, clarity of
ther. He compared survival rates of spittle- spittlebug presence, however. language, critical evaluation of relationships
bugs and moths when competitors were and information, and caution in coming to
His overall conclusion was:
present and absent. judgment, critical thinking in science has
There are three important general consid- Evidence from this and other studies sup- been a very successful tool in enhancing
erations in designing scientific investigations: ports the contention that interspecific com- understanding.
fall, and often neither help nor hurt the trees on which they grow. mutualistic relationships may be more important in evolution than
In a way, the robins and sparrows that inhabit suburban yards are we have commonly thought. Aggressive interactions often are dan-
commensals with humans. gerous and destructive, while cooperation and compromise may have
Lichens are a combination of a fungus and a photosynthetic part- advantages that we tend to overlook. Survival of the fittest often may
ner, either an alga or a cyanobacterium. Their association is a type of mean survival of those organisms that can live best with one another.
symbiosis called mutualism, in which both members of the partner- What do the acacias get in return and how does the relation-
ship benefit (fig. 4.13). Some ecologists believe that cooperative, ship relate to community dynamics? Ants tend to be aggressive

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defenders of their home areas, and acacia ants are no exception. Defensive Mechanisms
They drive off herbivorous insects that attempt to feed on their
home acacia, thus reducing predation. They also trim away vegeta- Many species of plants and animals have toxic chemicals, body
tion that grows around their home tree, thereby reducing competi- armor, and other ingenious defensive adaptations to protect them-
tion. This is a fascinating example of how a symbiotic relationship selves from competitors or predators. Arthropods, amphibians,
fits into community interactions. It is also an example of coevolu- snakes, and some mammals, for instance, produce noxious odors
tion based on mutualism rather than competition or predation. or poisonous secretions to induce other species to leave them alone.
Plants also produce a variety of chemical compounds that make
them unpalatable or dangerous to disturb. Perhaps you have brushed
up against poison ivy or stinging nettles in the woods or you have
encountered venomous insects or snakes and appreciate the wisdom
of leaving them alone. Often, species possessing these chemical
defenses will evolve distinctive colors or patterns to warn potential
enemies (fig. 4.14).
FIGURE 4.13 Lichens, such as the various species growing on

this log, are a combination of algae and fungi in a classic example of mutu-
alistic symbiosis. © William P. Cunningham.
FIGURE 4.12 Plants compete for light and growing space in

this Indonesian rainforest. Epiphytes, such as the ferns and bromeliads
shown here, find a place to grow in the forest canopy by perching on the FIGURE 4.14 Poison arrow frogs of the family Dendrobatidae
limbs of large trees. This may be a commensal relationship if the epiphytes use brilliant colors to warn potential predators of the extremely toxic secre-
don’t hurt their hosts. Sometimes, however, the weight of epiphytes breaks tions from their skin. Native people in Latin America use the toxin on
off branches and even topples whole trees. © William P. Cunningham. blowgun darts. © Michael Fogden/Animals Animals/Earth Scenes.

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FIGURE 4.15 An example of Batesian mimicry. The dangerous wasp (left) has bold yellow and black bands to warn predators away. The much
rarer longhorn beetle (right) has no poisonous stinger, but looks and acts like a wasp and thus avoids predators as well. © Edward S. Ross.
dead leaves or twigs to hide from predators. Predators also use cam-
ouflage to hide as they lie in wait for their prey. The scorpion fish
(fig. 4.16) blends in remarkably well with its surroundings as it waits
for smaller fish to come within striking distance. Not all cases of mim-
icry are to avoid or carry out predation, however. Some tropical
orchids have evolved flower structures that look exactly like female
flies. Males attempting to mate unwittingly carry away pollen.
COMMUNITY PROPERTIES
The processes and principles that we have studied thus far in this
chapter—tolerance limits, species interactions, resource partition-
ing, evolution, and adaptation—play important roles in determining
the characteristics of populations and species. In this section we will
look at some fundamental properties of biological communities and
ecosystems—productivity, diversity, complexity, resilience, stabil-
ity, and structure—to learn how they are affected by these factors.
FIGURE 4.16 This highly camouflaged scorpion fish lies in wait
for its unsuspecting prey. Natural selection and evolution have created the
elaborate disguise seen here. © Brian Parker/Tom Stack & Associates. Productivity
A community’s primary productivity is the rate of biomass pro-
Sometimes species that actually are harmless will evolve colors, duction, an indication of the rate of solar energy conversion to
patterns, or body shapes that mimic species that are unpalatable or chemical energy. The energy left after respiration is net primary
poisonous. This is called Batesian mimicry after the English natu- production. Photosynthetic rates are regulated by light levels, tem-
ralist H. W. Bates, who described it in 1857. Wasps, for example, perature, moisture, and nutrient availability. Figure 4.17 shows
often have bold patterns of black and yellow stripes to warn off approximate productivity levels for some major ecosystems. As
potential predators. The rarer longhorn beetle (fig. 4.15), although it you can see, tropical forests, coral reefs, and estuaries (bays or
has no stinger, looks and acts much like wasps and thus avoids preda- inundated river valleys where rivers meet the ocean) have high lev-
tors as well. Another form of mimicry, called Müllerian mimicry, els of productivity because they have abundant supplies of all these
named for the German biologist Fritz Müller, who described it in resources. In deserts, lack of water limits photosynthesis. On the
1878, involves two species, both of which are unpalatable or dan- arctic tundra or in high mountains, low temperatures inhibit plant
gerous and have evolved to look alike. When predators learn to avoid growth. In the open ocean, a lack of nutrients reduces the ability
either species, both benefit. of algae to make use of plentiful sunshine and water.
Species also evolve amazing abilities to avoid being discovered. Some agricultural crops such as corn (maize) and sugar cane
You very likely have seen examples of insects that look exactly like grown under ideal conditions in the tropics approach the produc-

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tivity levels of tropical forests. Because shallow water ecosystems water on a warm, dry, sunny day while it makes only a few kilo-
such as coral reefs, salt marshes, tidal mud flats, and other highly grams of sugars and other energy-rich organic compounds.
productive aquatic communities are relatively rare compared to the
vast extent of open oceans—which are effectively biological
deserts—marine ecosystems are much less productive on average
Abundance and Diversity
than terrestrial ecosystems. Abundance is an expression of the total number of organisms in
Even in the most photosynthetically active ecosystems, only a a biological community, while diversity is a measure of the num-
small percentage of the available sunlight is captured and used to ber of different species, ecological niches, or genetic variation pre-
make energy-rich compounds. Between one-quarter and three- sent. The abundance of a particular species often is inversely
quarters of the light reaching plants is reflected by leaf surfaces. related to the total diversity of the community. That is, communi-
Most of the light absorbed by leaves is converted to heat that is ties with a very large number of species often have only a few
either radiated away or dissipated by evaporation of water. Only 0.1 members of any given species in a particular area. As a general
to 0.2 percent of the absorbed energy is used by chloroplasts to rule, diversity decreases but abundance within species increases as
synthesize carbohydrates. we go from the equator toward the poles. The arctic has vast num-
In a temperate-climate oak forest, only about half the incident bers of insects such as mosquitoes, for example, but only a few
light available on a midsummer day is absorbed by the leaves. species. The tropics, on the other hand, have vast numbers of
Ninety-nine percent of this energy is used to evaporate water. species—some of which have incredibly bizarre forms and
A large oak tree can transpire (evaporate) several thousand liters of habits—but often only a few individuals of any particular species
in a given area.
Consider bird populations. Greenland is home to 56 species
of breeding birds, while Colombia, which is only one-fifth the size
Desert of Greenland, has 1,395. Why are there so many species in Colom-
bia and so few in Greenland?
Climate and history are important factors. Greenland has such
Tundra a harsh climate that the need to survive through the winter or
escape to milder climates becomes the single most important crit-
ical factor that overwhelms all other considerations and severely
Grassland,
shrubland limits the ability of species to specialize or differentiate into new
forms. Furthermore, because Greenland was covered by glaciers
until about 10,000 years ago, there has been little time for new
Coniferous
forest species to develop.
Temperate
deciduous
forest
Intensive
agriculture
Tropical
rainforest
Estuaries,
coral reefs
Coastal
zone
Open ocean
0 2 4 6 8 10 12 14 16 18 20
1,000 kcal/m2/year
FIGURE 4.17 Relative biomass accumulation of major world ecosystems. Only plants and some bacteria capture solar energy. Animals con-
sume biomass to build their own bodies.

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Resilience and Stability

Key Concepts Many biological communities tend to remain relatively stable and
constant over time. An oak forest tends to remain an oak forest, for
• Natural selection explains species change and adaptation: members example, because the species that make it up have self-perpetuating
of a population that are most suited to survive environmental condi- mechanisms. We can identify three kinds of stability or resiliency in
tions are most likely to survive and reproduce. ecosystems: constancy (lack of fluctuations in composition or func-
• Predation can lead to evolution as species develop evasive character- tions), inertia (resistance to perturbations), and renewal (ability to
istics. Symbiotic relationships can enhance changes through compet- repair damage after disturbance).
itive advantages of both species. In 1955, Robert MacArthur, who was then a graduate student
• A species’ niche is its ecological role or its environmental conditions. at Yale, proposed that the more complex and interconnected a com-
• Ecological communities can be compared by quantitative measures munity is, the more stable and resilient it will be in the face of dis-
such as primary productivity, complexity, and diversity. turbance. If many different species occupy each trophic level, some
• Ecological succession refers to the gradual change from one set of can fill in if others are stressed or eliminated by external forces,
species to another in a location. making the whole community resistant to perturbations and able to
recover relatively easily from disruptions. This theory has been
controversial, however. Some studies support it, while others do
Many areas in the tropics, by contrast, have relatively abun- not. For example, Minnesota ecologist David Tilman, in studies of
dant rainfall and warm temperatures year-round so that ecosystems native prairie and recovering farm fields, found that plots with high
there are highly productive. The year-round dependability of food, diversity were better able to withstand and recover from drought
moisture, and warmth supports a great exuberance of life and than those with only a few species.
allows a high degree of specialization in physical shape and behav- On the other hand, in a diverse and highly specialized ecosys-
ior. Coral reefs are similarly stable, productive, and conducive to tem, removal of a few keystone members can eliminate many other
proliferation of diverse and amazing life-forms. The enormous associated species. Eliminating a major tree species from a tropi-
abundance of brightly colored and fantastically shaped fish, corals, cal forest, for example, may destroy pollinators and fruit distribu-
sponges, and arthropods in the reef community is one of the best tors as well. We might replant the trees, but could we replace the
examples we have of community diversity. whole web of relationships on which they depend? In this case,
Productivity is related to abundance and diversity, both of which diversity has made the forest less resilient rather than more.
are dependent on the total resource availability in an ecosystem as Diversity is widely considered important and has received a
well as the reliability of resources, the adaptations of the member great deal of attention. In particular, human impacts on diversity are
species, and the interactions between species. You shouldn’t assume a primary concern of many ecologists (Case Study, p. 87).
that all communities are perfectly adapted to their environment.
A relatively new community that hasn’t had time for niche special-
ization, or a disturbed one where roles such as top predators are miss-
Edges and Boundaries
ing, may not achieve maximum efficiency of resource use or reach An important aspect of community structure is the boundary
its maximum level of either abundance or diversity. between one habitat and its neighbors. We call these relationships
edge effects. Sometimes, the edge of a patch of habitat is rela-
tively sharp and distinct. In moving from a woodland patch into
Complexity and Connectedness a grassland or cultivated field, you sense a dramatic change from
Community complexity and connectedness generally are related to the cool, dark, quiet forest interior to the windy, sunny, warmer,
diversity and are important because they help us visualize and open space of the field or pasture (fig. 4.19). In other cases, one
understand community functions. Complexity in ecological terms habitat type intergrades very gradually into another, so there is no
refers to the number of species at each trophic level and the num- distinct border.
ber of trophic levels in a community. A diverse community may Ecologists call the boundaries between adjacent communi-
not be very complex if all its species are clustered in only a few ties ecotones. A community that is sharply divided from its
trophic levels and form a relatively simple food chain. neighbors is called a closed community. In contrast, communi-
By contrast, a complex, highly interconnected community (fig. ties with gradual or indistinct boundaries over which many
4.18) might have many trophic levels, some of which can be com- species cross are called open communities. Often this distinc-
partmentalized into subdivisions. In tropical rainforests, for tion is a matter of degree or perception. As we saw earlier in this
instance, the herbivores can be grouped into “guilds” based on the chapter, birds might feed in fields or grasslands but nest in the
specialized ways they feed on plants. There may be fruit eaters, forest. As they fly back and forth, the birds interconnect the
leaf nibblers, root borers, seed gnawers, and sap suckers, each com- ecosystems by moving energy and material from one to the other,
posed of species of very different size, shape, and even biological making both systems relatively open. Furthermore, the forest
kingdom, but that feed in related ways. A highly interconnected edge, while clearly different from the open field, may be sunnier
community such as this can form a very elaborate food web. and warmer than the forest interior, and may have a different

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Sperm whale
Minke whale Killer whale
Humpback whale
Squid Leopard seal
Fin whale
Open-water community
King Weddell seal

penguin
Blue whale Zooplankton Other penguins

Krill
Elephant seal
Sei whale Phytoplankton Pelagic fish

and bacteria
Protozoa
Ross seal
Light, circulation, Detritus Albatross

temperature, and
nutrients
Skua
Bottom community
Bacteria Echinoderm
and protozoa Bottom fish
Crustacean and octopod
Alga Mollusk
FIGURE 4.18 A complex and highly interconnected community can have many species at each trophic
level and many relationships, as illustrated by this Antarctic marine food web.
combination of plant and animal species than either field or for- Many popular game animals, such as white-tailed deer and
est “core.” pheasants that are adapted to human disturbance, often are most
Depending on how far edge effects extend from the boundary, plentiful in boundary zones between different types of habitat.
differently shaped habitat patches may have very dissimilar Game managers once were urged to develop as much edge as pos-
amounts of interior area (fig. 4.20). In Douglas fir forests of the sible to promote large game populations. Today, however, most
Pacific Northwest, for example, increased rates of blowdown, wildlife conservationists recognize that the edge effects associated
decreased humidity, absence of shade-requiring ground cover, and with habitat fragmentation are generally detrimental to biodiversity.
other edge effects can extend as much as 200 m into a forest. A Preserving large habitat blocks and linking smaller blocks with
40-acre block (about 400 meters square) surrounded by clear-cut migration corridors may be the best ways to protect rare and endan-
would have essentially no true core habitat at all. gered species (see chapter 13).

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Ecological Succession
Biological communities have a history in a given landscape. The
process by which organisms occupy a site and gradually change
environmental conditions by creating soil, shade, shelter, or
increasing humidity is called ecological succession or develop-
ment. Primary succession occurs when a community begins to
develop on a site previously unoccupied by living organisms, such
as an island, a sand or silt bed, a body of water, or a new volcanic
flow (fig. 4.21). Secondary succession occurs when an existing
community is disrupted and a new one subsequently develops at
the site. The disruption may be caused by some natural catastrophe,
such as fire or flooding, or by a human activity, such as deforesta-
tion, plowing, or mining. Both forms of succession usually follow
an orderly sequence of stages as organisms modify the environ-
ment in ways that allow one species to replace another.
In primary succession on a terrestrial site, the new site first is
FIGURE 4.19 Ecological edges are known as ecotones. Tem- colonized by a few hardy pioneer species, often microbes, mosses,
perature, wind, and humidity differ at the edges in a landscape. Edge con-
ditions do extend into patches of habitat. Small or linear fragments may
and lichens that can withstand harsh conditions and lack of
be mostly edge. © Corbis/Volume 262. resources. Their bodies create patches of organic matter in which
protists and small animals can live (fig. 4.22). Organic debris accu-
mulates in pockets and crevices, providing soil in which seeds can
become lodged and grow. We call this process of environmental
modification by organisms ecological development or facilita-
tion. The community of organisms often becomes more diverse
and increasingly competitive as development continues and new
niche opportunities appear. The pioneer species gradually disap-
pear as the environment changes and new species combinations
replace the preceding community. In a global sense, the gradual
changes brought about by living organisms have created many of
Core area: 0 ha the conditions that make life on earth possible. You could con-
Total area: 39 ha sider evolution to be a very slow, planetwide successional and
developmental process.
Examples of secondary succession are easy to find. Observe an
abandoned farm field or clear-cut forest (fig. 4.23) in a temperate
climate. The bare soil first is colonized by rapidly growing annual
plants (those that grow, flower, and die the same year) that have
light, wind-blown seeds and can tolerate full sunlight and exposed
soil. They are followed and replaced by perennial plants (those that
live for several to many years), including grasses, various non-
woody flowering plants, shrubs, and trees. As in primary succes-
sion, plant species progressively change the environmental
Total area: 47 ha Core area: 20 ha conditions. Biomass accumulates and the site becomes richer, bet-
ter able to capture and store moisture, more sheltered from wind
FIGURE 4.20 Shape can be as important as size in small pre- and climate change, and biologically more complex. Species that
serves. While these areas are close to the same size, no place in the top fig-
ure is far enough from the edge to have characteristics of core habitat,
cannot survive in a bare, dry, sunny, open area find shelter and food
while the bottom patch has a significant core. as the field turns to prairie or forest.
Eventually, in either primary or secondary succession, a com-
munity often develops that resists further change. Ecologists call
COMMUNITIES IN TRANSITION this a climax community because it appears to be the culmination
of the successional process. An analogy is often made between com-
So far our view of communities has focused on the day-to-day munity succession and organism maturation. Beginning with a
interactions of organisms with their environments, set in a context primitive or juvenile state and going through a complex develop-
of survival and selection. In this section, we’ll step back to look at mental process, each progresses until a complex, stable, and mature
some transitional aspects of communities, including where com- form is reached. It’s dangerous to carry this analogy too far, how-
munities meet and how communities change over time. ever, because no mechanism is known to regulate communities in

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Where Have All the Songbirds Gone?
ures in the United States and Canada may

E very June, some 2,200 amateur ornithol-
ogists and bird watchers across the
United States and Canada join in an annual
be just as big a problem for woodland song-
birds. Many of the most threatened species
bird count called the Breeding Bird Survey. are adapted to deep woods and need an
Organized in 1966 by the U.S. Fish and area of 10 hectares (25 acres) or more per
Wildlife Service to follow bird population pair to breed and raise their young. As our
changes, this survey has discovered some woodlands are broken up by roads, hous-
shocking trends. While birds such as robins, ing developments, and shopping centers, it
starlings, and blackbirds that prosper becomes more and more difficult for these
around humans have increased their num- highly specialized birds to find enough con-
ber and distribution over the past 30 years, tiguous woods to nest successfully.
many of our most colorful forest birds have Predation and nest parasitism also pre-
declined severely. The greatest decreases sent a growing threat to many bird species. This thrush has been equipped with a lightweight
have been among the true songbirds such In human-dominated landscapes, raccoons, radio transmitter and antenna so that its move-
as thrushes, orioles, tanagers, catbirds, opossums, crows, bluejays, squirrels, and ments can be followed by researchers. Courtesy
vireos, buntings, and warblers. These long- house cats thrive. They are protected from Dr. David Mech.
distance migrants nest in northern forests larger predators like wolves or owls and find
but spend the winters in South or Central abundant supplies of food and places to hide. invaded by cowbirds. Another study in the
America or in the Caribbean Islands. Scien- Cats are a particular problem. By some esti- Shawnee National Forest in southern Illinois
tists call them neotropical migrants. mates, there are 100 million feral cats in the found that 80 percent of the scarlet tanager
In many areas of the eastern United United States, and 73 million pet cats. A com- nests contained cowbird eggs and that 90
States and Canada, three-quarters or more of parison of predation rates in the Great Smoky percent of the wood thrush nests were
the neotropical migrants have declined signif- Mountain National Park and in small rural and taken over by these parasites. The sobering
icantly since the survey was started. Some suburban woodlands shows how devastating conclusion of this latter study is that there
that once were common have become locally predators can be. In a 1,000-hectare study probably is no longer any place in Illinois
extinct. Rock Creek Park in Washington, D.C., area of mature, unbroken forest in the national where scarlet tanagers and wood thrushes
for instance, lost 75 percent of its songbird park, only one songbird nest in fifty was raided can breed successfully.
population and 90 percent of its long-distance by predators. By contrast, in plots of 10 What can we do about this situation?
migrant species in just 20 years. Nationwide, hectares or less near cities, up to 90 percent Elsewhere in this book, we discuss sustain-
cerulean warblers, American redstarts, and of the nests were raided. able forestry and economic development pro-
ovenbirds declined about 50 percent in the Nest parasitism by brown-headed cow- jects that could preserve forests at home and
single decade of the 1970s. Studies of radar birds is one of the worst threats for woodland abroad. Preserving corridors that tie together
images from National Weather Service sta- songbirds. Rather than raise their young important areas also will help. In areas where
tions in Texas and Louisiana suggest that only themselves, cowbirds lay their eggs in the people already live, clustering of houses pro-
about half as many birds fly across the Gulf of nests of other species. The larger and more tects remaining woods. Discouraging the
Mexico each spring now compared to the aggressive cowbird young either kick their fos- clearing of underbrush and trees from yards
1960s. This could mean a loss of about half a ter siblings out of the nest, or claim so much and parks leaves shelter for the birds.
billion birds in total. food that the others starve. Well adapted to Could we reduce the number of preda-
What causes these devastating losses? live around humans, there are now about 150 tors or limit their access to critical breeding
Destruction of critical winter habitat is clearly million cowbirds in the United States. areas? Would you accept fencing or trap-
a major issue. Birds often are much more A study in southern Wisconsin found ping of small predators in wildlife preserves?
densely crowded in the limited areas avail- that 80 percent of the nests of woodland How would you feel about a campaign to
able to them during the winter than they are species were raided by predators and that keep house cats inside during the breeding
on their summer range. Unfortunately, forests three-quarters of those that survived were season?
throughout Latin America are being felled
at an appalling rate. Central America, for Ethical Considerations
instance, is losing about 1.4 million hectares
Some wildlife managers are already trapping cowbirds. The Kirtland’s warbler is one of the
(2 percent of its forests or an area about the rarest songbirds in the United States. It nests only in young, fire-maintained jackpine forests
size of Yellowstone National Park) each year. in Michigan. Controlled burning to maintain habitat for this endangered species was started
If this trend continues, there will be essen- in the 1960s, but the population continued to decline. Studies showed that 90 percent of the
tially no intact forest left in much of the region nests were being parasitized by cowbirds. Since 1972, refuge managers have trapped and
in 50 years. killed some 7,000 cowbirds each year to protect the warblers. In the past two decades, the
But loss of tropical forests is not the only number of breeding pairs of warblers has risen from about 150 to nearly 400. Would it be pos-
sible to do something similar on a nationwide scale? Could we trap and kill 150 million cow-
threat. Recent studies show that fragmen-
birds? How much should we reduce one species to save another? What do you think?
tation of breeding habitat and nesting fail-
cun39564_ch04.qxd 12/5/03 4:38 PM Page 88
White spruce
Balsam fir
Paper birch
Aspen
Black spruce
Jack pine
Grasses
Herbs
Shrubs
Tree seedlings
Lichens
Exposed rocks Mosses
Pioneer community Climax community

Time
Time
FIGURE 4.21 One example of primary succession, shown in five stages (left to right). Here,
bare rocks are colonized by lichens and mosses, which trap moisture and build soil for grasses, shrubs,
and eventually trees.
H. A. Gleason, who saw community history as a much more indi-

vidualistic and random process driven by many environmental fac-
tors. He argued that temporary associations are formed according
to the conditions prevailing at a particular time and the species
available to colonize a given area. You might think of the Glea-
sonian model as a time-lapse movie of a busy railroad station. Pas-
sengers come and go; groups form and then dissipate. Patterns and
assemblages that seem significant to us may not mean much in
the long run.
The process of succession may not be as deterministic as we
once thought, yet mature or highly developed ecological commu-
nities may tend to be resilient and stable over long periods of time
because they can resist or recover from external disturbances.
Many are characterized by high species diversity, narrow niche
specialization, well-organized community structure, good nutrient
conservation and recycling, and a large amount of total organic
matter. Community functions, such as productivity and nutrient
cycling, tend to be self-stabilizing or self-perpetuating. What once
FIGURE 4.22 Primary succession occurs where there had been were regarded as “final” climax communities, however, may still
no living things, as on this lava in Hawaii. Fungi, algae, and bacteria grew
here first, providing rooting material for these ferns. © William P. Cunningham. be changing. It’s probably more accurate to say that the rate of suc-
cession is so slow in a climax community that, from the perspec-
tive of a single human lifetime, it appears to be stable.
the same way that genetics and physiology regulate development Some landscapes never reach a stable climax in the traditional
of the body. sense because they are characterized by, and adapted to, periodic dis-
The concept of succession to a climax community was first ruption. They are called equilibrium communities or disclimax
championed by the pioneer biogeographer F. E. Clements. He communities. Grasslands, the chaparral shrubland of California, and
viewed this process as being like a parade or relay, in which some kinds of coniferous forests, for instance, are shaped and main-
species replace each other in predictable groups and in a fixed, tained by periodic fires that have long been a part of their history.
regular order, and as being driven almost entirely by climate. This They are, therefore, often referred to as fire-climax communities
community-unit theory was opposed by Clements’s contemporary, (fig. 4.24). Plants in these communities are adapted to resist fires,

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What can
you do?
Working Locally for Ecological Diversity
You might think that diversity and complexity of ecological systems are
too large or too abstract for you to have any influence. But you can
contribute to a complex, resilient, and interesting ecosystem, whether
you live in the inner city, a suburb, or a rural area.
• Keep your cat indoors. As discussed in the Case Study (p. 00),
our lovable domestic cats are also very successful predators.
Migratory birds, especially those nesting on the ground, have not
evolved defenses against these predators.
• Plant a butterfly garden. Use native plants that support a diverse
insect population. Native trees with berries or fruit also support
birds. (Be sure to avoid non-native invasive species: see chapter
11). Allow structural diversity (open areas, shrubs, and trees) to
support a range of species.
• Join a local environmental organization. Often, the best way to
FIGURE 4.23 This area was once a cool, shady black spruce be effective is to concentrate your efforts close to home. City
stand. The forest floor was covered by a deep, moist layer of sphagnum parks and neighborhoods support ecological communities, as do
moss. Clear-cutting and burning have turned it into a dry, sunny, barren
farming and rural areas. Join an organization working to maintain
ground on which few of the former residents can survive. Secondary suc-
cession will probably restore previous conditions if the climate doesn’t ecosystem health: start by looking for environmental clubs at your
change and further disturbance is prevented. © William P. Cunningham. school, parks organizations, a local Audubon chapter, or a local
Nature Conservancy branch.
• Take walks. The best way to learn about ecological systems in
your area is to take walks and practice observing your environ-
ment. Go with friends and try to identify some of the species and
trophic relationships in your area.
• Live in town. Suburban sprawl consumes wildlife habitat and
reduces ecosystem complexity by removing many specialized
plants and animals. Replacing forests and grasslands with lawns
and streets is the surest way to simplify, or eliminate, ecosystems.
reseed quickly after fires, or both. In fact, many of the plant species
we recognize as dominants in these communities require fire to elim-
inate competition, to prepare seedbeds for germination of seedlings,
or to open cones or thick seed coats. Without fire, community struc-
ture may be quite different.
Introduced Species and Community Change

Succession requires the continual introduction of new community
members and the disappearance of previously existing species.
New species move in as conditions become suitable; others die or
move out as the community changes. New species also can be
introduced after a stable community already has become estab-
lished. Some cannot compete with existing species and fail to
become established. Others are able to fit into and become part of
the community, defining new ecological niches. If, however, an
FIGURE 4.24 This lodgepole pine forest in Yellowstone
introduced species preys upon or competes more successfully with
National Park was once thought to be a climax forest, but we now know
that this forest must be constantly renewed by periodic fire. It is an exam- one or more populations that are native to the community, the entire
ple of an equilibrium, or disclimax, community. © William P. Cunningham. nature of the community can be altered.

cun39564_ch04.qxd 12/5/03 12:58 PM Page 90
Human introductions of Eurasian plants and animals to non-

Eurasian communities often have been disastrous to native species
because of competition or overpredation. Oceanic islands offer
classic examples of devastation caused by rats, goats, cats, and
pigs liberated from sailing ships. All these animals are prolific,
quickly developing large populations. Goats are efficient, non-
specific herbivores; they eat nearly everything vegetational, from
grasses and herbs to seedlings and shrubs. In addition, their sharp
hooves are hard on plants rooted in thin island soils. Rats and pigs
are opportunistic omnivores, eating the eggs and nestlings of
seabirds that tend to nest in large, densely packed colonies, and
digging up sea turtle eggs. Cats prey upon nestlings of both
ground- and tree-nesting birds. Native island species are particu-
larly vulnerable because they have not evolved under circum-
stances that required them to have defensive adaptations to these
predators (What Can You Do? p. 89).
Sometimes we introduce new species in an attempt to solve FIGURE 4.25 Mongooses were released in Hawaii in an effort
problems created by previous introductions but end up making the to control rats. The mongooses are active during the day, however, while
situation worse. In Hawaii and on several Caribbean Islands, for the rats are night creatures, so they ignored each other. Instead, the mon-
gooses attacked defenseless native birds and became as great a problem
instance, mongooses were imported to help control rats that had as the rats. © Gerard Lacz/Peter Arnold, Inc.
escaped from ships and were destroying indigenous birds and dev-
astating plantations (fig. 4.25). Since the mongooses were diurnal
(active in the day), however, and rats are nocturnal, they tended to cally engineered organisms are based on concerns that they are
ignore each other. Instead, the mongooses also killed native birds novel organisms, and we might not be able to predict how they will
and further threatened endangered species. Our lessons from this interact with other species in natural ecosystems—let alone how
and similar introductions have a new technological twist. Some of they might respond to natural selective forces. It is argued that we
the ethical questions currently surrounding the release of geneti- can’t predict either their behavior or their evolution.
Summary
• Organisms are adapted to live within certain ranges of envi- • Resource partitioning occurs when species adapt to use a sin-
ronmental conditions. Tolerance limits are the maximum or gle resource differently.
minimum conditions, such as temperature or moisture, that an • Species interact in many ways. Some general classes of inter-
organism can survive. Since many environmental factors affect action include predation, parasitism, symbiosis, and competi-
survival, it is useful to consider critical factors that limit a tion. All of these interactions can exert selective pressure, as
species’ growth or expansion. organisms develop defenses against predators or parasites, as
• Evolution is gradual change of organisms by natural selec- they develop traits that improve competitiveness, or as they
tion. Natural selection refers to a higher rate of survival and develop mutually beneficial interactions. Both interspecific
reproduction among individuals that happen to have advanta- (between species) and intraspecific (within a species) compe-
geous traits. Environmental conditions can exert selective pres- tition can lead to changes in traits or behavior.
sure by making some traits more advantageous than others. • Defensive mechanisms can include Batesian mimicry, in
• An ecological niche is usually described as its ecological role which a harmless species looks like a dangerous one, and Mül-
in a community; a niche can also be the place or set of envi- lerian mimicry, in which two dangerous species look like each
ronmental conditions in which an organism lives. Generalist other, and thus both discourage predation.
species can occupy a range of habitats and ecological roles or • Primary productivity, or the rate of biomass accumulation, is
environmental conditions. Highly specialized species occupy a basic characteristic of communities. Abundance and species
narrower niches. diversity are also important characteristics.
90
cun39564_ch04.qxd 12/5/03 12:58 PM Page 91
• Complexity refers to the number of species at each trophic when an existing community is disrupted and a new, different
level and the number of trophic levels in a community. Many community develops.
ecologists believe that complexity contributes to stability in • The idea of a climax community is a stable community that
an ecosystem, or resilience to abrupt change such as fire, flood, appears to be the culmination of successional processes.
or drought. Others believe that complex communities can be A contrasting idea is that species occur individualistically,
less resilient than simple ones. each according to its ability to colonize an area.
• Edges, where contrasting conditions meet, are important fea- • Introduced species are one of the greatest modern threats to
tures in biological communities. Ecotones, or zones of transi- biological diversity and ecosystem complexity. When intro-
tion, have great diversity. Edges also reduce habitat quality for duced species are free of predators, they can become abundant
interior species. and cause significant damage to ecosystems.
• Primary succession occurs when pioneer species occupy areas
previously lacking living things. Secondary succession occurs
9. Which world ecosystems are most productive in terms of bio-

Questions for Review mass (fig. 4.17)? Which are least productive? What units are
1. Explain how tolerance limits (fig. 4.2) to environmental fac- used in this figure to quantify biomass accumulation?
tors determine distribution of a highly specialized species such 10. Discuss the dangers posed to existing community members
as the saguaro cactus. Compare this to the distribution of a when new species are introduced into ecosystems. What type
generalist species such as cowbirds or starlings. What would of organism would be most likely to survive and cause prob-
the curve in fig. 4.1 look like for one of these species? lems in a new habitat?
2. Productivity, diversity, complexity, resilience, and structure
are exhibited to some extent by all communities and ecosys- Questions for Critical Thinking
tems. Describe how these characteristics apply to the ecosys-
tem in which you live. 1. Ecologists debate whether biological communities have self-
sustaining, self-regulating characteristics or are highly vari-
3. Resource partitioning (figs. 4.6, 4.7) is an important adaptive
able, accidental assemblages of individually acting species.
strategy. Explain resource partitioning, and think of an exam-
What outlook or worldview might lead scientists to favor one
ple in your local area.
or the other of these theories?
4. Define keystone species and explain their importance in com-
2. The concepts of natural selection and evolution are central to
munity structure and function.
how most biologists understand and interpret the world, and
5. All organisms within a biological community interact with yet the theory of evolution is contrary to the beliefs of many
each other. The most intense interactions often occur between religious groups. Why do you think this theory is so important
individuals of the same species. What concept discussed in to science and so strongly opposed by others? What evidence
this chapter can be used to explain this phenomenon? would be required to convince opponents of evolution?
6. Relationships between predators and prey play an important 3. What is the difference between saying that a duck has webbed
role in the energy transfers that occur in ecosystems. They also feet because it needs them to swim and saying that a duck is
influence the process of natural selection. Explain how preda- able to swim because it has webbed feet?
tors affect the adaptations of their prey. This relationship also
4. The concept of keystone species is controversial among ecol-
works in reverse. How do prey species affect the adaptations
ogists because most organisms are highly interdependent. If
of their predators?
each of the trophic levels is dependent on all the others, how
7. Competition for a limited quantity of resources occurs in all can we say one is most important? Choose an ecosystem with
ecosystems. This competition can be interspecific or intra- which you are familiar and decide whether it has a keystone
specific. Explain some of the ways an organism might deal species or keystone set.
with these different types of competition.
5. Some scientists look at the boundary between two biological
8. Each year fires burn large tracts of forestland. Describe the communities and see a sharp dividing line. Others looking at
process of succession that occurs after a forest fire destroys an the same boundary see a gradual transition with much inter-
existing biological community. Is the composition of the final mixing of species and many interactions between communi-
successional community likely to be the same as that which ties. Why are there such different interpretations of the same
existed before the fire? What factors might alter the final out- landscape?
come of the successional process? Why may periodic fire be
beneficial to a community?
91
cun39564_ch04.qxd 12/5/03 12:59 PM Page 92
6. The absence of certain lichens is used as an indicator of air pathogens 78 resource partitioning 76
pollution in remote areas such as national parks. How can we pioneer species 86 secondary succession 86
be sure that air pollution is really responsible? What evidence plankton 77 symbiosis 79
would be convincing? predator 77 territoriality 79
7. We tend to regard generalists or “weedy” species as less inter- primary productivity 82 tolerance limits 72
esting and less valuable than rare and highly specialized primary succession 86
endemic species. What values or assumptions underlie this
attitude? Further Readings
8. What part of this chapter do you think is most likely to be
Botkin, Daniel B. 1989. Discordant Harmonies: A New Ecology
challenged or modified in the future by new evidence or new
for the Twenty-First Century. Oxford University Press.
interpretations?
Ehrlich, Paul R., and Peter H. Raven. 1967. Butterflies and plants:
A study in coevolution. Evolution 18:586–608.
Key Terms Gleason, Henry A. 1926. The individualistic concept of the plant
abundance 83 environmental indicators 73 association. Bulletin of the Torrey Botanical Club 53:7–26.
Batesian mimicry 82 equilibrium communities 88 MacArthur, R. H. 1958. Population ecology of some warblers of
climax community 86 evolution 74 Northeastern coniferous forests. Ecology 39:599–619.
coevolution 78 fire-climax communities 88 Paracer, S., and V. Ahmadjian. 2000. Symbiosis: An Introduction to
commensalism 79 habitat 75 Biological Associations. Oxford University Press.
complexity 84 interspecific competition 79 Simberloff, D. 1997. Flagships, umbrellas, and keystones: Is single-
disclimax communities 88 intraspecific competition 79 species management passé in the landscape era? Biological
diversity 83 keystone species 78 Conservation 83:247–57.
ecological development 86 Müllerian mimicry 82
Tilman, David, et al. 1997. The influence of functional diversity
ecological niche 75 mutualism 80
and composition on ecosystem processes. Science 277(5330):
ecotones 84 natural selection 74
1300–02.
edge effects 84 parasites 78
WEB EXERCISES
Project FeederWatch
The FeederWatch Program coordinated by the Cornell Laboratory of 1. Explain the three graphs. Why does an increase in game fish (pisci-
Ornithology is an excellent example of citizen science. Thousands of vol- vores) cause a decrease in phytoplankton (algae) in a lake?
unteers collect data on bird frequency and distribution from backyard feed- 2. If you were designing a test of this hypothesis, how would you regu-
ers throughout winter months. The data are displayed on innovative late piscivore biomass experimentally?
animated maps that allow you to view dynamic information about a given
species in a particular region or state over time. Go to: http://birds. 3. What would you use as a control in your study?
cornell.edu/PFWMaproom/pfwmaproom.html to find a species and loca- 4. What do the authors mean by top down and bottom up controls?
tion that interests you; then consider the following questions: 5. Why do they call this a trophic cascade?
1. Does it surprise you that this species does or doesn’t occur in your
area? Alien Invaders: When Weeds Do Good and Bad Things
2. How would you account for the patterns you see on the map? Is it On the same regional perspectives page, look at the first case study in the
possible that the results show a bias in data collection rather than a real Southwest Region. You can also find an interesting international case study
variation in distribution of the species? about water hyacinth on the USGS Eros site at http://edcsnw3.cr.usgs.gov/
ip/hyacinth/hyacinth.html. Look at the Winam Gulf study for some impres-
3. Some species display seasonal movements. Can you detect a pattern
sive images of how this plant can clog lakes and waterways.
in changing distribution of the species you’ve chosen during the time
shown? How would you account for the pattern (or the lack of a pat- 1. When and why was water hyacinth introduced into the United States?
tern) you observe? 2. Where did it come from?
Trophic Cascades in Aquatic Food Webs 3. How fast does it spread?
Ecological relationships can affect physical qualities in our environment. To 4. Why is it a problem?
understand how this occurs, go to http://www.mhhe.com/environmental 5. What possible benefits does it convey?
science. Click on the title of your textbook to take you to the Online Learning 6. How is it controlled?
Center, and then click on the student edition. Click on “Regional Case Stud-
ies” on the left-hand navigational menu. Scroll down to the North region to 7. Drawing on what you’ve learned about community interactions in this
find a case study titled “Food Web Control of Primary Production in Lakes.” chapter, why is this plant so aggressive and so successful in its new
Read the text and study the graphics to answer the following questions. home?

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qxd 12/5/03 12:55 PM Page 71

Biological Communities and Species Interactions

Why Trees Need Salmon

72 PART ONE Principles for Understanding Our Environment

Zone of Zone of Optimal range Zone of Zone of

Too low: Too high:

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 73

Natural Selection, Adaptation,

74 PART ONE Principles for Understanding Our Environment

Warbler finch Cactus ground finch

Fruit eaters ground

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 75

(a) Resource gradient

FIGURE 4.5 The giant panda feeds exclusively on bamboo.

76 PART ONE Principles for Understanding Our Environment

FIGURE 4.7 Resource partitioning

Exploitation Predation and Parasitism

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 77

FIGURE 4.9 Microscopic plants and animals form the basic

Prey species have evolved many protective or defensive adap-

mals). If these figs were removed, many animals would starve to

78 PART ONE Principles for Understanding Our Environment

Competition Intraspecific competition can be especially intense because

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 79

What do you think?

Understanding Competition 1. Things need to be organized in such a

80 PART ONE Principles for Understanding Our Environment

FIGURE 4.13 Lichens, such as the various species growing on

FIGURE 4.12 Plants compete for light and growing space in

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 81

82 PART ONE Principles for Understanding Our Environment

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 83

Resilience and Stability

84 PART ONE Principles for Understanding Our Environment

Minke whale Killer whale

Squid Leopard seal

King Weddell seal

Blue whale Zooplankton Other penguins

Sei whale Phytoplankton Pelagic fish

Light, circulation, Detritus Albatross

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 85

86 PART ONE Principles for Understanding Our Environment

Where Have All the Songbirds Gone?

ures in the United States and Canada may

Pioneer community Climax community

H. A. Gleason, who saw community history as a much more indi-

88 PART ONE Principles for Understanding Our Environment

Introduced Species and Community Change

www.mhhe.com/environmentalscience/ CHAPTER 4 Biological Communities and Species Interactions 89

Human introductions of Eurasian plants and animals to non-

9. Which world ecosystems are most productive in terms of bio-

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