PART NINE

Ecology
The Physical Environment
and Biogeography of Life
KEY CONCEPTS
52
52.1 Ecology Is the Study of the
Interrelationships among
Organisms and the Environment
52.2 Global Climate Is a Fundamental
Component of the Physical
Environment
52.3 Topography, Vegetation, and
Humans Modify the Physical
Environment
52.4 Biogeography Is the Study of
How Organisms Are Distributed
on Earth
52.5 Geographic Area and Humans
Affect Regional Species Diversity

© Morley Read/Alamy Stock Photo

Amazonian forests, such as this, contain half of all

the described species on Earth.

▶InvestigatingLIFE

The Largest Experiment on Earth

Most people know that there are many species on Earth, and How does habitat fragmentation affect the species living in
that new species are being discovered every day, but not ev- the Amazon Basin? Fragmentation makes habitats smaller, and
eryone knows that roughly half of all species worldwide reside it isolates populations. Deforestation forces species to sustain
in one place: the tropical rainforests of the Amazon Basin. The themselves in a smaller area or move to more suitable areas.
statistics are staggering: an estimated 390 billion trees repre- In the late 1970s, some ecologists asked a deceptively simple
senting 16,000 species grow in Amazonia, one in five bird spe- question about Amazon deforestation: What is the minimum
cies globally resides there, and one-fifth of all fresh water falls area needed to maintain species diversity within a rainforest
on its slopes and valleys. The Amazon Basin is by far the largest fragment? They wanted to know whether the species remain-
watershed on Earth, with thousands of tributaries leading to the ing in the forest fragments could maintain themselves there,
Amazon River and then out to sea. To date, 2,200 freshwater whether the sizes of the forest fragments mattered to this
fish species have been described, which is more than all the fish maintenance, and whether species would venture from one
species in the Atlantic Ocean. fragment to another. These ecologists conducted a large and
It is reasonable to assume, then, that when the Amazon Basin long-running experiment in the Amazon forests near Manaus,
is threatened by human activities, so, too, is global biodiversity. Brazil. As you will see in Key Concept 52.5, the size of the frag-
By far, the main destructive force is deforestation, which began ments and the ability of species to move across the deforested
about 50 years ago. As roads pushed their way into Amazonia’s landscape are key to maintaining the extraordinary diversity left
forests, more rainforest was logged and converted to agriculture in the Amazon Basin.
and settlements. It is estimated that nearly 20% of the Amazon
Basin has been clear-cut. Because soil fertility is generally poor
in tropical rainforests, the cleared land is often useful only for
a decade or less, resulting in abandonment and new cycles of
QA
and
How do geographic area and isolation
affect the biogeography of life on Earth?
(Find the answer on page 1194.)
deforestation and fragmentation.

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 CHAPTER 52 The Physical Environment and Biogeography of Life

pests and diseases safely and effectively, and to deal with natural
Ecology Is the Study of the disasters such as floods and fires. The greater our understanding
KEY CONCEPT Interrelationships among of ecological connections, the more likely it is that we can ac-
52.1 Organisms and the complish these things without causing a cascade of unanticipated
consequences for ourselves and the other life on Earth.
Environment
Ecology is studied at many levels of organization
Learning Objectives
Ecology is studied at many levels of organization, from molecules
52.1.1 Distinguish between ecology and environmentalism, to the entire globe. Some ecologists work at the level of genes to
and give examples of each. understand how individuals might adapt to a changing environ-
52.1.2 Explain the levels of organization in ecology, and be ment, while other researchers seek to understand how tempera-
able to distinguish them from one another based on ture might modify nutrient cycling at the ecosystem level. The
spatial scale.
work of ecologists focuses on many scales or levels of organization:
52.1.3 Describe the research methods used by ecologists, individuals, populations, communities, ecosystems, landscapes,
and identify how and why ecological studies might
differ from studies in other scientific fields. or the global biosphere (Figure 52.1).
As prior chapters have conveyed, a population is a group of
individuals of the same species within a given area that have the
Whether you notice ants collecting crumbs on a sidewalk or watch potential to interbreed and interact with one another. For example,
a video of polar bears hunting on sea ice, you are studying ecol- the individual sea stars in Figure 52.1 make up a population at this
ogy. From simple experiences such as these, most people know particular rocky shore location. The sea star population in turn
that organisms on Earth are interconnected with one another interacts with other species, including its prey, sessile mussels, and
and with their environment. Ecology is the exploration of these its competitors, whelks and other sea star species. These species
interconnections and is most simply defined as the study of the in- form a community, which is an assemblage of interacting species
terrelationships among organisms and the physical environment. living together at the same place and time.
Ecologists study these interactions at different scales and with Ecological studies often encompass both the living, or biotic,
different methods, but they all agree that ecology is a scientific and the nonliving, or abiotic, components of the environment.
endeavor, first and foremost. The biotic components of an organism’s environment are other
“Ecology” is sometimes equated with “environmentalism,” but organisms, so ecology includes the study of interactions within
the two terms are different. Ecology is a science that generates and among species. The abiotic components of an organism’s en-
knowledge about interactions in the natural world. Environmen- vironment are the myriad physical and chemical characteristics
talism is the use of ecological knowledge, along with economics, of the system. For example, in the rocky shore community, tides
ethics, and many other considerations, to inform both personal are a key abiotic factor because they limit where sea stars can feed
decisions and public policy related to stewardship of the natural on mussels and compete with whelks.
world. So while an ecologist might study the number of bluefin The term ecosystem is often used when describing a commu-
tuna in the world’s oceans, the environmentalist would focus on the nity of organisms in which their physical environment is explicitly
social, economic, and political capital needed to enforce a sustain- taken into account. Ecosystem ecology particularly focuses on
able harvest or, alternatively, to ban bluefin tuna fishing altogether. the movement of energy and nutrients through a group of organ-
Ecology is a relatively new branch of the biological sciences; in isms. The movement of energy and nutrients, as well as organisms,
fact, it did not have a formal name until 150 years ago when Ernst can affect higher levels of organization, including landscapes,
Haeckel, a German biologist, constructed the new word “ecology,” geographic areas that include multiple ecosystems (sometimes
from the Greek root oikos, meaning “household.” referred to as meta-ecosystems). Ultimately, meta-ecosystems
are linked to one another at larger geographic scales to form the
Modern ecology has a more “use-inspired” focus biosphere, consisting of all living organisms on Earth plus their
Although it was recognized in Darwin’s day that humans could respective environments.
affect the natural world, and that human populations were under-
going unprecedented growth, for the most part early ecologists Answering ecological questions requires
believed they were studying pristine systems largely untouched observations, experiments, and models
by human activities. Today most ecologists have a much greater Just as in other fields in biology, ecologists use a mixture of ob-
awareness of ways in which humans shape the natural world, be servations, experiments, and models to test ecological theories
it in the form of global climate change, the introduction of novel (see Key Concept 1.3). Ecologists are unique, though, in using
species to new locations, or the logging of Earth’s tropical for- a wide range of venues and scales in their experiments, includ-
ests. Because of the dominant role that humans play in almost ing short laboratory experiments, long-term field manipulations,
every ecological system on Earth, a “use-inspired” motivation and even decades-long ecosystem-wide experiments, such as the
often shapes ecologists’ research and teaching. They understand Biological Dynamics of Forest Fragments Project described in
that scientific knowledge of ecology greatly improves our ability Key Concept 52.5. However, at large spatial scales or over long
to grow food for ourselves reliably and sustainably, to manage periods of time, well-replicated and controlled experiments are

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 KEY CONCEPT 52.2 Global Climate Is a Fundamental Component of the Physical Environment

KEY CONCEPT
usually not possible. For example, when studying the ecological
effects of climate change on a particular species, ecologists might 52.1 Recap and Assess
have to combine observations of the temperature range of that Ecology is the scientific study of the interrelationships among
species with short-term experiments on the effects of warming organisms and the environment. Ecology is studied at multiple
on the species’ distribution and then use models to project how levels of organization, from individuals to the biosphere. Ecolo-
distributions could change with continued warming. Often this gists use a mixture of observations, experiments, and models to
type of research requires multiple techniques and collaborative test ecological theories.
efforts to be successful. 1. Compare how an ecologist versus an environmentalist might
approach an environmental problem.
2. Using observations, experiments, and models at different levels
of organization, describe how you might test the hypothesis that
The individual the widespread death of the ochre sea star (Pisaster ochraceus)
ochre sea star, (shown in Figure 52.1) is the result of a disease.
Pisaster ochraceus…
…is part of a population
of this species. The sea star
We will begin our study of ecology in the spirit of the discipline
population, in turn… in Haeckel’s time, focusing on variations in the distribution and
abundance of organisms worldwide and the factors that determine
these patterns. Our first step will be to examine attributes of the
Courtesy of Sally Hacker

physical environment, which is the most basic determinant for

where organisms can live. We’ll begin with climate.
Courtesy of Sally Hacker

KEY CONCEPT Global Climate Is a

Fundamental Component of
…is one of the interacting populations of
different species found in this community
52.2 the Physical Environment
or ecosystem (if we also consider the
physical environment). Learning Objectives
52.2.1 Compare weather and climate, and give examples
of both.
52.2.2 Given that Earth is a sphere, has an atmosphere, and
rotates around its axis, explain the latitudinal variation
© Konrad Wothe/Minden Pictures

in temperature, precipitation, and prevailing winds.

52.2.3 Relate Earth’s prevailing wind patterns to ocean
currents, and explain how ocean currents affect
The rocky intertidal ecosystem is
climate.
nested within a coastal landscape
composed of multiple ecosystems 52.2.4 Connect the tilt of Earth’s axis and its orbit to the
also known as a meta-ecosystem. existence of seasons on Earth, and explain why the
Northern and Southern Hemispheres have opposite
seasonal patterns.

The terms “weather” and “climate” both refer to atmospheric con-

© Jean Carter/AGE Fotostock

ditions—temperature, humidity, precipitation, and wind direction

and velocity—but they consider different time scales. Weather
is the short-term state of atmospheric conditions at a particular
place and time, whereas climate refers to the average atmospheric
conditions, and the extent of their variation, at a particular place
based on data from NASA and NOAA

over a longer time (years to millennia). Climate is what you expect

NASA image by Reto Stöckli,

The biosphere includes given past conditions; weather is what you get on any one day.
all of Earth’s ecosystems.
For example, a warming climate will cause glaciers to melt over
decades or centuries (see Figure 1.17), whereas a single large snow-
storm is unlikely to slow that retreat. In addition, the responses
of organisms to weather are usually short-term—seeking shelter
Figure 52.1 Scale Is Important to Ecological Systems These from a sudden rainstorm, for example, or shivering to keep warm
photos from the Oregon coast of the United States illustrate how eco- when the temperature drops. Climate, by contrast, acts as a sort of
logical systems can be studied at multiple scales, from the individual filter for organisms, ultimately determining their distribution and
to the biosphere. abundance. If organisms cannot tolerate the climate of a particular

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 CHAPTER 52 The Physical Environment and Biogeography of Life

Reflected solar radiation Outgoing radiation

Reflected by clouds Reflected

and atmospheric gases by surface
Incoming
solar Emitted by Emitted by clouds
radiation atmosphere
Greenhouse gases
Carbon dioxide (CO2)
Methane (CH4)
Nitrous oxide (N2O)

Back
Absorbed by radiation
atmosphere

Surface
Absorbed radiation
by surface Absorbed by surface

Figure 52.2 Radiant Energy Warms the Planet Solar energy input Lindsey, R. 2009. Climate and Earth’s energy budget. Earth Observa-
(yellow arrows) is absorbed by Earth’s atmosphere and surface. Much of tory, EOS Project Science Office, NASA Goddard Space Flight Center.
this energy is radiated from Earth’s surface in the form of heat (orange Greenbelt, MD.)
arrows). Of that absorbed at the surface, most is prevented from escap-
ing back into space by greenhouse gases in the atmosphere. The widths
Q: What effect do greenhouse gases have on Earth’s energy
balance?
of the arrows are roughly proportional to the sizes of the energy fluxes.
(After K. E. Trenberth et al. 2009. Bull Am Meteorol Soc 90: 311–323; View in Achieve
Animation 52.1 Radiant Energy Balance

place, they will not be found there. Here we consider the processes it has both land and water. As you will see, it is the combination of
that create climate patterns. these factors that determines the climate we experience on Earth.

Solar radiation drives global climate patterns Earth is a sphere, creating latitudinal variation in
The energy that drives global climate patterns ultimately originates global temperature and precipitation
from the Sun. Of the solar radiation that hits Earth, about 30% is Because Earth is a sphere, the intensity of solar radiation hitting
reflected back to space by clouds, atmospheric gases, and Earth’s its surface varies with latitude. This latitudinal variation in solar
surface (Figure 52.2). The rest is absorbed either by the atmosphere energy depends primarily on the angle of the Sun’s rays striking
(20%) or the surface of Earth (50%). The atmo-
sphere, a thin layer of gases surrounding Earth
North Pole
composed of nitrogen gas (N2, 78%), oxygen (O2, Toward the poles, more of the (90°N)
21%), argon (0.9%), carbon dioxide (CO2, 0.037%), Sun’s rays are absorbed
and traces of hydrogen, helium, ozone, methane because they must travel a
longer distance through the Direction
(CH4), and many other gases, moderates Earth’s atmosphere. of Earth’s
surface temperatures by trapping heat energy. rotation
If Earth had no atmosphere, its average surface
At and near the equator,
temperature would be about –18°C, rather than sunlight strikes Earth at a
its actual +17°C. Carbon dioxide, methane, ni- steep angle, delivering more °)
r (0
trous oxide (N2O), water vapor, and certain other heat and light per unit of area. ato
Equ
gases in the atmosphere are known as green-
house gases because they allow sunlight into Toward the poles, the Sun’s rays
Earth’s atmosphere but trap heat radiating back strike Earth at an oblique angle
and are spread over a larger area,
out toward space (see Figure 52.2). so that their energy is diffused.
South Pole
The amount of radiation received on Earth (90°S)
varies with latitude and with the seasons. Figure 52.3 Solar Energy Input Varies with Latitude The angle of incoming sunlight
These differences are driven by some well- affects the intensity of solar energy that reaches a given unit of Earth’s surface. (After C.
known characteristics of Earth: it is a sphere D. Ahrens. 2005. Essentials of Meteorology: An Invitation to the Atmosphere. Brooks/Cole,
that orbits the Sun, it spins on a tilted axis, and Cengage Learning: USA.)

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 KEY CONCEPT 52.2 Global Climate Is a Fundamental Component of the Physical Environment

(A) Average annual air temperature

Celsius) than the Southern Hemisphere (see
60°N Figure 52.4A). Although the reasons for this
difference are uncertain, one hypothesis is that
40°N because land heats up the air above it more than
does the ocean (which absorbs more heat), the
20°N greater proportion of land to ocean in the North-
Equator ern Hemisphere creates warmer air tempera-
0°

© Robert A. Rohde/Global Warming Art

tures than those in the Southern Hemisphere.
20°S This is particularly true in summer, when the
temperature difference is at its greatest. In light
40°S of global warming, it is worth noting that tem-
peratures are increasing faster in the Northern
60°S –40 –20 0 20 40 60 80 °F Hemisphere than in the Southern Hemisphere.
Besides controlling temperature, solar en-
–50 –40 –30 –20 –10 0 10 20 30 °C
Average annual air temperature ergy input also determines atmospheric circu-
lation and associated precipitation patterns. As
(B) Average annual precipitation you saw earlier, air in the region surrounding
the equator receives the greatest input of solar
energy. When a parcel of air is warmed, it ex-
60°N
pands, becomes less dense, and rises (see Fig-
ure 52.5A). As it rises, however, it starts to cool
40°N
at higher elevations. Cool air cannot hold as
20°N much water vapor as warm air, so the expand-
ing, cooling air releases moisture in the form of
Equator
0° precipitation. This is particularly evident in the
tropics, where there is abundant rainfall year-
20°S round (Figure 52.4B). Eventually the rising air
at the equator reaches the boundary between
40°S the troposphere, the atmospheric layer above
Earth’s surface, and the stratosphere, the next
60°S
layer up. The stratosphere is warmer than the
04 7 10 14 19 27 38 99 247
Average annual precipitation (cm) troposphere at this boundary, causing the tropi-
cal air to stop rising and eventually move either
Figure 52.4 Global Temperature and Precipitation Vary with Latitude Average annual north or south as newly warmed air rises to re-
air temperature (A) and precipitation (B) vary with Earth’s latitude but can be modified by place it (Figure 52.5A). Eventually this air starts
topography (i.e., Earth’s shape and surface features) (see Figure 52.9). (A, after Berkeley to cool as it exchanges heat with the atmosphere
Earth. 2016. Land and Ocean Baseline Comparison Map. http://www.berkeleyearth.org/; B, and meets colder air traveling from higher lati-
courtesy of the Nelson Institute Center for Sustainability and the Global Environment [SAGE], tudes. When this cool, dry air, which lost its
through their "Atlas of the Biosphere," https://nelson.wisc.edu/sage/data-and-models/atlas/.
moisture as it rose over the equator, reaches
University of Wisconsin-Madison.)
latitudes of about 30°N and 30°S, it begins to
sink under high atmospheric pressure, mak-
Earth. At high latitudes (i.e., areas toward the North and South ing the climate at these latitudes dry (Figure 52.5B). Earth’s great
Poles), incoming solar energy is distributed over a larger area (and deserts—including the Sahara of Africa, the Gobi of China, and
thus is less intense) than at the equator, where sunlight strikes the deserts of Australia and the North American Southwest—are
the surface perpendicularly (Figure 52.3). Moreover, when the located at these latitudes (see Figure 52.5B).
Sun’s radiation comes in at an angle, it must pass through more While some of the descending air flows back toward the equator,
of Earth’s atmosphere, resulting in more of its energy being ab- some of it flows toward the poles, setting up another atmospheric
sorbed or reflected before reaching Earth’s surface. Differences cell of air circulation at higher latitudes. At about 60° latitude, air
in solar energy input explain why the average global temperature rises again due to its warming at Earth’s surface (see Figure 52.5B).
varies with latitude, with warmer temperatures at the equator and As this air rises, it cools and releases precipitation, creating wet
colder
Life 12e temperatures at the poles (Figure 52.4A). The average air climate conditions at temperate latitudes. Eventually this air reaches
temperature overPress
Oxford University the course of a year decreases about 0.76 degrees the stratosphere and moves north or south. The cold, dry air trav-
Dragonfly
Celsius Media
for everyGroup
degree of latitude (~110 km) at sea level. eling at higher latitudes eventually reaches the cold polar region,
Life12e_52.00.ai Date 02-11-19
In addition to there being a latitudinal variation in temperature, where it descends. Despite the amount of snow and ice at the poles,
the Northern Hemisphere is slightly warmer (by ~2 degrees Cel- high latitudes at or near 90° actually receive little precipitation and
sius) and more variable in temperature (by a range of ~7 degrees can be thought of as “polar deserts” (see Figure 52.5B).

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 CHAPTER 52 The Physical Environment and Biogeography of Life

Figure 52.5 Air Circulation in Earth’s Atmosphere Drives Climate (A) Solar energy
drives patterns of atmospheric circulation. (B) The differential heating of Earth’s surface by
solar radiation gives rise to atmospheric circulation cells, which influence atmospheric pres- 90ºN
High pressure Polar
sure, air temperatures, and precipitation worldwide. (After C. D. Ahrens. 2005. Essentials of Sparse precipitation
in all seasons zone
Meteorology: An Invitation to the Atmosphere. Brooks/Cole, Cengage Learning: USA.)
Low pressure 60º
Ample precipitation
(A) Rising air (B) in all seasons Temperate
Descending zone
1 In the tropics, warm, moist air Winter wet, summer dry
air rises, expands and 30°N High pressure 30º
Dry in all seasons
cools, drops its moisture,
and flows poleward. Summer wet, winter dry

Tropics
Low pressure
Sunlight 0° Abundant precipitation in all seasons 0º
Tropics
Summer wet, winter dry
High pressure
2 The now cool, dry air sinks 30°S Dry in all seasons 30º
under high atmospheric Winter wet, summer dry
pressure, making the latitudes Temperate
of 30°N and 30°S dry. Low pressure zone
Ample precipitation
in all seasons 60º
High pressure Polar
Sparse precipitation zone
Earth spins on an axis, producing prevailing winds in all seasons
90ºS
and ocean currents
You have seen that Earth’s spherical shape creates predictable pat-
terns of temperature and precipitation, but Earth’s rotation, which
moves east to west, is responsible for generating global wind and
ocean currents. Because Earth is a sphere, the velocity of its rota- How, then, are the circulation patterns of ocean currents gen-
tion around its axis is fastest at the equator, where its diameter is erated? Ocean currents are driven by prevailing winds, which
greatest, and is slowest close to the poles. This difference creates move water by means of frictional drag. The trade winds, for
the Coriolis effect, which is the deflection of air or water as a result example, cause currents to converge at the equator and move
of differences in Earth’s rotational speed at different latitudes. For westward until they encounter a continent (Figure 52.7). At that
example, air traveling toward the equator (driven by the circula- point, the strong Equatorial Countercurrent brings some of the
tion patterns described above and in Figure 52.5) moves at a slower water back eastward. The remaining water divides, some mov-
speed than that of the planet beneath it and is thus deflected to ing northward and some southward along continental shores.
the west. Conversely, air traveling toward
either pole is moving faster than that of the
surface beneath it and is deflected to the 90°N As the winds move across
east. This interaction of Earth’s rotation Earth’s surface in response
and north–south air mass movement sets to the circulation of air
Easterlies masses, they are deflected
up a pattern of circulating surface air re- 60°
by the Coriolis effect, which
ferred to as prevailing winds (Figure 52.6). produces easterly (winds
Prevailing winds blow from east to west in Westerlies blow from east to west)
the tropics (the trade winds); from west to 30° trade winds in the tropics…
east in mid-latitudes (the westerlies); and
NE Trade Winds
from east to west again above 60°N or 60°S Direction
latitude (the easterlies). 0° of Earth’s
rotation
SE Trade
Winds
30°
Figure 52.6 Prevailing Winds Created
by Earth’s Rotation and Atmospheric Westerlies …and westerlies at
Circulation The deflection of air as a result temperate latitudes.
of differences in Earth’s rotational speed at 60°
Easterlies
different latitudes (also known as the Coriolis
effect) coupled with atmospheric circulation of
air masses combine to create a global pattern 90°S
of prevailing surface winds across the planet.

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 KEY CONCEPT 52.2 Global Climate Is a Fundamental Component of the Physical Environment

90°N
1 The trade winds push
water toward the equator… North
Atlantic
Drift
60°N 2 …where it moves 4 The Equatorial Counter-
westward until it current arises between
reaches a continent… Labrador
the hemispheric gyres. Current

30°N N. Equatorial Gulf

Kuroshio
Current Stream
Current
Latitude

0°

30°S Humboldt Benguela

South
Equatorial Current Current
Current

60°S Antarctic Circumpolar Current

Antarctic Circumpolar Current
3 …then moves north or south
along the coast, forming great
circular currents called gyres.
90°S

Figure 52.7 Oceanic Circulation Created by Winds and Continents Red arrows indicate warm surface water, and blue arrows indicate cold
The surface currents of the ocean are driven by the prevailing winds surface water.
(see Figure 52.6), Earth’s rotation, and the placement of the continents.

These patterns of water movement set up rotating circulation The tilt of Earth’s axis and its orbit
patterns called gyres (Greek gyros, “spiral”), which rotate clock- result in seasons
wise in the Northern Hemisphere and counterclockwise in the The tilt of Earth’s axis, coupled with Earth’s orbit around the
Southern Hemisphere. Sun, is responsible for seasonal changes in climate. Earth’s axis is
Because ocean currents transport heat, they have a tremen- tilted at an angle of 23.5 degrees, changing the amount of sunlight
dous effect on Earth’s climates. The poleward movement of a particular region receives over the course of a year as Earth or-
warm water from the tropics transfers large amounts of heat bits the Sun (Figure 52.8). This tilt causes seasonal variation in
to high latitudes. The Gulf Stream, for example, carries warm
water from the tropical Atlantic Ocean (including the Gulf
of Mexico) north across the Atlantic to northern Equinox
Europe, making Europe’s climate consider- September 22
ably milder than that of some corre-
sponding latitudes in North America.
Earth’s orbit
Similarly, currents flowing toward
the equator from high latitudes
bring cool, wet winters to some
western coastal regions that When the Southern When the Northern
are otherwise warm and dry. Hemisphere is tilted Hemisphere is tilted
toward the Sun, it is toward the Sun, it is
summer there and summer there and
winter in the Northern winter in the Southern
Hemisphere. Hemisphere.
23.5°
Figure 52.8 Seasonal Change
Is the Result of the Tilt of Earth’s Solstice Solstice
North Pole
Axis and Its Orbit Because Earth’s December 21 June 21
axis of rotation is tilted, orientation
relative to the Sun changes over the
course of a year as the planet orbits the
Sun. The resulting variation in solar radiation
creates seasonal climate variation. (After C. D.
Ahrens. 2005. Essentials of Meteorology: An Invitation Equinox
to the Atmosphere. Brooks/Cole, Cengage Learning: USA.) March 20

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 CHAPTER 52 The Physical Environment and Biogeography of Life

temperature and day length. Higher latitudes experience greater Earth’s topography affects the local and regional
seasonal variation than lower latitudes do. Around the equator, physical environment
day length and seasonal temperatures change only slightly over Earth’s topography, meaning its shape and surface features, has
the course of the year, although there are seasonal shifts in pre- been formed over geological time and provides the basis for much
cipitation patterns. of the variation in local and regional physical conditions. Here we
KEY CONCEPT
consider how topography modifies the physical environment with
respect to land, oceans, rivers, and lakes.
52.2 Recap and Assess
Climate and weather both refer to atmospheric conditions, but LAND The topography of land plays a large role in the regional
climate represents conditions over years whereas weather repre- and local physical environment, including climate. Mountains
sents conditions over a span of days. Earth is a sphere, has an create elevational gradients in temperature, precipitation, and
atmosphere, and rotates around its axis, which creates latitudinal
variation in solar radiation, temperature and precipitation, and sunlight. For example, mountains show progressively colder tem-
prevailing winds and ocean currents. Earth’s tilt and orbit around peratures and greater precipitation at higher elevations, creating
the Sun result in the seasons. different environmental conditions over relatively short distances.
1. Does a month of extremely hot weather tell us that Earth’s When a mountain range occurs adjacent to an ocean, a rain shadow
climate is warming? Explain your answer. is often created, whereby one side of the mountain has a wet climate
2. Refer to Figure 52.3. Why does solar energy input into Earth’s and the other side has a dry climate (Figure 52.9A). Rain shadows
atmosphere vary with latitude? What are the consequences for occur when winds deliver moist air evaporated from the ocean
global climate?
to the windward side of the mountain, where the air rises, cools,
3. Refer to Figures 52.6 and 52.7. If you were sailing, using wind and releases rain or snow. On the leeward side of the mountain
power only, from the east coast of North America to Europe
across the Atlantic Ocean, describe the best route given winds (that is, opposite from the direction of the winds), the now-dry air
and currents. descends, warms, and produces arid conditions. The High Desert,
4. Explain why Earth’s Northern and Southern Hemispheres have on the leeward side of the Cascade Range in the Pacific Northwest
opposite seasons. of the United States, is one such area.
Other topographic features, such as canyons and valleys, also
We have considered large, global patterns in climate, but we modify local climate in significant ways. Valleys can experience
know that the physical environment can vary within certain re- temperature inversions, extremes in temperature that produce
gions or even locally. Just travel across a large mountain range morning fog and intense afternoon heat. Temperature inversions
and you can experience lush forests on one side and bone-dry occur when the valley concentrates heat from solar radiation dur-
deserts on the other. We will turn next to what controls smaller, ing the day. As that heat rises, it forms an inversion layer that traps
more regional and local variation in the physical environment, cold, dense, and moisture-laden air that descends into the valley
both on land and in water. overnight and produces fog (Figure 52.9B).

OCEANS Beneath the oceans, which make up 70% of Earth’s

surface, dramatic differences in topography influence the marine
KEY CONCEPT Topography, Vegetation, environment. If you could drain the oceans, you would notice ma-
and Humans Modify the jor differences in topography, ranging from the Hawaiian island
52.3 Physical Environment
Mauna Loa, the tallest “mountain” on Earth at 9,170 meters high
from base to peak, to the Mariana Trench, the deepest “valley” at
Learning Objectives 10,994 meters deep. Variations in water depth affect light penetra-
tion, water temperature, water pressure, and water movement (i.e.,
52.3.1 Explain and compare the causes and effects of rain waves and tides). These physical discontinuities create distinct
shadows and temperature inversions.
zones that are identified principally by their physical conditions
52.3.2 Describe how ocean-floor topography creates and the characteristic biota they contain, similar to biomes on land
variable marine environments.
(see Key Concept 52.4). The depth of an ocean basin varies from
52.3.3 Compare and contrast the effects of vegetation on the shoreline to the relatively shallow continental shelf and to the
temperature and precipitation at local and regional
deepest part of the ocean, sometimes known as the abyssal plain
scales.
(Figure 52.9C). Water depth affects how much light is available
52.3.4 Describe and contrast ways in which agriculture
to sustain the photosynthetic organisms that form the base of the
and urbanization may transform local and regional
climates. marine food chain. In both marine and freshwater environments,
roughly 90% of all aquatic life is found in the photic zone, the layer
of water reached by enough sunlight to support photosynthesis.
Three main factors can modify the pattern of global climate, de- Other zones that vary in environmental conditions include the
termining regional and local variation in climate and other aspects coastal zone, which extends from the shoreline to the edge of the
of the physical environment: (1) Earth’s topography, (2) vegetation, continental shelf, and the ocean zone, which extends beyond the
and more recently, (3) the effects of humans. coastal zone (see Figure 52.9C). The coastal zone is diverse and

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 KEY CONCEPT 52.3 Topography, Vegetation, and Humans Modify the Physical Environment

(A) Rain shadow (B) Temperature inversion

On the windward side, air Cool air
Winds pick up 7ºC
moisture over rises, cools, and releases
the ocean. moisture as rain or snow,
creating a wet climate. Inversion layer
On the leeward side, dry 10ºC
air descends and warms, 7ºC
resulting in little rain and Fog
arid conditions. 4ºC

Warm air (inversion layer)

traps cold and moisture-
laden air that descends
into the valley overnight,
producing fog.

(C) Aquatic zones (D) Ocean upwelling

Intertidal zone Coastal zone Ocean zone Photic zone
A prevailing wind blows The wind, along with the Coriolis
parallel to the coast. effect, creates a surface current
flowing away from the coast.

200
Pelagic zone Deep, cool water
Continental Wind
2,000 rises to replace
shelf the surface water.
Water depth (m)

4,000
Benthic
zone
6,000
14°
13°
8,000 12°
11°
Abyssal plain
10,000

Figure 52.9 Effects of Earth’s Topography on the Physical Agriculture Handbook 360. D after C. D. Ahrens. 2005. Essentials of
Environment Earth’s topography can affect both land and oceans. Meteorology: An Invitation to the Atmosphere. Brooks/Cole, Cengage
Mountains and valleys create rain shadows (A) and temperature inver- Learning: USA.)
sions (B). Ocean topography affects water depth, which creates aquatic
zones that vary in light, temperature, and water pressure (C), and along
Q: What kind of weather conditions would you expect to pro-
duce temperature inversions? How about ocean upwelling?
with wind, produces ocean upwelling (D). (B after M. Schroeder and
C. Buck. 1970. Fire Weather: A Guide for Application of Meterological View in Achieve
Information to Forest Fire Control Operations, p. 29. USDA Forest Service, Animation 52.2 Rain Shadow

highly productive, supporting high densities of pelagic, or floating, colder, and nutrient-rich bottom water to rise to the surface (Figure
plankton and fishes. On the benthos or bottom of the ocean, the 52.9D). Upwelling affects local and regional climate on the coast
coastal zone has a variety of living and nonliving habitats such by creating cooler and moister conditions. In addition, upwelled
as rocky reefs, soft sediment flats, coral reefs, and kelp beds. The waters support high rates of primary production by phytoplank-
portion of the benthic coastal zone lying between the high and ton, which in turn support dense consumer populations. Most of
low tide levels is the intertidal zone, where tidal movements create the world’s great fisheries are concentrated in upwelling zones.
conditions of highly variable light and temperature, alternately As we move away from the coastline, most of the ocean’s physical
exposing organisms to air and water. Intertidal zones can occur environment is made up of vast amounts of water that is, with the ex-
on the open coast as sandy or rocky beaches, or in more protected ception of the photic zone (which can extend to 200 meters in some
bays as estuaries. Estuaries form where rivers meet the sea, creat- regions), mostly dark, cold, and under tremendous pressure from the
ing variability in salinity, sediment, and light conditions. weight of the water above. Little photosynthesis can take place, and
In some coastal zones, where the continental shelf drops off many of the organisms inhabiting these regions subsist on decaying
steeply and prevailing winds are parallel to the coastline, upwell- organic matter that sinks down from the photic zone. Deep-ocean
ing can affect ocean and climate conditions. Upwelling is a process trenches and rift valleys support hydrothermal vent ecosystems sus-
in which offshore winds in combination with the Coriolis effect tained by chemoautotrophic prokaryotes that can metabolize the
push warmer surface waters away from the shore, allowing deeper, nutrients without the aid of sunlight (see Key Concept 24.3).

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 CHAPTER 52 The Physical Environment and Biogeography of Life

RIVERS AND LAKES In contrast to the vast oceans, freshwater convective heat loss by acting as a wind screen, this is more than
lakes and rivers make up less than 3% of Earth’s surface, but they balanced by the cooling effect produced by evapotranspiration:
are home to about 10% of all aquatic species. Earth’s topography the evaporative transfer of heat and water from the surfaces of
influences freshwater environments just as it does marine envi- plants into the atmosphere, which reduces air temperature and
ronments—through the depth of the water—but it also influences increases moisture. When forests are cut and replaced by vegeta-
the degree and direction of movement of fresh water. A river is tion such as grasses, the albedo effect and convective heat loss
water that flows downhill from its point of origin (the source) over intensify while evapotranspiration decreases, causing overall air
the land surface as a result of gravity until it reaches either the temperatures to rise and precipitation to decrease, creating more
ocean or a depression in Earth’s surface, where it forms a lake or arid conditions. The climate changes associated with deforesta-
pond. The headwaters (those close to the source) tend to be cool, tion have been seen in the Amazon Basin, where massive forest
fast-flowing, and well oxygenated. As a river flows downstream, clearing makes way for cattle ranching.
it widens, it slows down, it warms, and sediment accumulates, Oceans have their own “forests”—coastal areas dominated by
reducing light penetration and oxygen levels. seaweeds such as giant kelps or sea grasses that create calmer
The Amazon River, mentioned in the chapter opening, offers waters, promote sedimentation, and provide habitat for fishes and
a dramatic example of river dynamics. The Amazon is one of the invertebrates. In warmer coastal waters, corals generate complex
longest rivers on Earth, at approximately 7,000 km. Eventually reef structures that support ecosystems rivaling rainforests in di-
this source water reaches the Atlantic Ocean, where discharge is versity. In Chapter 56 you will learn more about organisms that
the largest of any river on Earth—the Amazon’s mouth is 240 km engineer their environment, but for now let’s turn to the final
wide and is referred to as “the River Sea.” topic in this key concept: the effects of humans on climate and
the physical environment.
Vegetation affects the local and regional physical
environment, especially climate Humans have transformed their physical
Earth’s climate would be very different without the influence environment, including urban climate
of organisms, including humans. Organisms can “engineer” It is estimated that humans have transformed more than 50% of
the environment in multiple ways that affect the surrounding Earth’s land surface, replacing it with infrastructure and agriculture
climate and physical conditions. The most obvious influence needed to support our burgeoning population. You have already
comes from vegetation, especially forests, which can have large seen how the conversion of forestland to pastureland can change
effects on temperature and precipitation at local and even re- regional climate, and in Chapter 56 you will learn more about how
gional scales. For example, measurements of energy exchange humans have changed global climate through the burning of fossil
in tropical forests that have been converted to pastureland show fuels. Structures such as dams, bridges, and nuclear power plants
that forests make the climate cooler and moister (Figure 52.10). can dramatically transform rivers and lakes. Urban areas, in par-
While forests absorb heat by reflecting less sunlight (reflection ticular, modify not only the immediate physical environment but
of solar radiation is known as the albedo effect) and decrease also the surrounding climate. A survey of 60 U.S. cities in 2014

Evapotranspiration at the leaf surface

of trees releases heat, thus cooling air
and increasing precipitation. Removing trees increases the albedo of the land The change in surface roughness
surface, lowering its absorption of solar radiation. increases heat loss by wind convection.

Solar Solar
Precipitation Precipitation
radiation radiation
These cooling effects are
more than offset by the
warming effects of lower
evapotranspiration from
the loss of trees. Reduced
evapotranspiration also
reduces the return of
Albedo Convective Evapotranspiration Albedo Convective Evapotranspiration moisture to the atmosphere
heat loss heat loss and thus precipitation rates.

Forested Deforested

Figure 52.10 Deforestation Can Affect Climate Evapotranspira- not enough to outweigh the warming effects of reduced evapotranspira-
tion by trees reduces air temperatures and increases precipitation. tion. The widths of the arrows are roughly proportional to the sizes of the
When land is deforested and the trees are gone, the cooling effects of fluxes. (After J. A. Foley, et al. 2003. Front Ecol Environ 1: 38–44.)
the albedo (reflection of solar radiation) and of convective heat loss are

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 KEY CONCEPT 52.4 Biogeography Is the Study of How Organisms Are Distributed on Earth

33 geographic and historical processes determine the global dis-

tribution of organisms on Earth.
Late afternoon temperature (°C)

32

31 KEY CONCEPT Biogeography Is the Study

Inversion layer
of How Organisms Are
30
52.4 Distributed on Earth
Warm
Learning Objectives
Cold Cold 52.4.1 Describe the principle characteristics of each
major biome, including climate, latitude and/
or continent(s), and representative plants and
Rural Suburban Commercial Downtown Urban Park Suburban Rural animals.
residential residential residential farmland 52.4.2 Use the observations and insights of Alfred
Russel Wallace to explain how biogeography
Figure 52.11 Cities Can Create Heat Islands and Country Breezes might influence the distribution of plants and
As this hypothetical urban heat island profile shows, cities and suburbs typi- animals.
cally have higher temperatures than rural, park, or agricultural areas. Country
breezes form when warm air from cities and suburbs is replaced by cool air 52.4.3 Compare and contrast the hypotheses posed
from rural areas. (After D. Reidel. 2004. In Climate Change Impacts and Adapta- for latitudinal patterns of species diversity,
tion: A Canadian Perspective, D. S. Lemmen and F. J. Warren (Eds.), p. 157. including species diversification rate, species
diversification time, and productivity.
Ottawa, ON: Climate Change Impacts and Adaptation Program, Natural
Resources Canada; and H. E. Landsberg. 1970. Science 170: 1265–1274.)
Biogeography is the scientific study of the distribution and
diversity of organisms on Earth. “Distribution” is simply where
showed that urban summer temperatures were on average 1.5 de- organisms are found. “Diversity,” however, has multiple facets,
grees Celsius hotter than rural summer temperatures, with the top which we will explore in more detail in Chapter 56, but for now
10 cities having an average urban–rural temperature differential of 5 you can think of it simply as the number and composition of
degrees Celsius. The cause of this so-called urban heat island effect species or higher taxa. For centuries, the processes that control
is simple: concrete, asphalt, and even the dark roofs of buildings these biogeographic patterns have puzzled naturalists. Why do
absorb heat from solar radiation and radiate that heat in the evening species vary in their distributions around Earth, and what con-
hours (Figure 52.11). Cities also produce more heat through the trols this variability?
burning of fossil fuels from cars, factories, and buildings. Parks can
serve to cool the surrounding air through the evapotranspiration of Patterns of biogeography are interconnected across
trees and other vegetation. In addition, the temperature differential a hierarchy of spatial and temporal scales
between cities and the surrounding countryside can create “country The patterns of biogeography play out over global, regional, land-
breezes,” air movement generated as rising warm air from the city scape, and local spatial scales (Figure 52.12). Spatial scales are
is replaced by cold surface air from surrounding rural areas. interconnected in a hierarchical way, with patterns at one scale
setting the conditions for patterns at other scales.
KEY CONCEPT
At the global scale, species are isolated from one another by
52.3 Recap and Assess continents or oceans, and for long periods of time. Thus the rates
Earth’s topography is responsible for rain shadows and tempera- of three processes—speciation, extinction, and dispersal—help
ture inversions, producing variation in temperature and precipita- determine differences in biogeography across the globe (see Fig-
tion. In the oceans, topographical variability produces differences ure 52.12A). At the regional scale, large-scale climate conditions
in water depth, which affects light penetration, water temperature, are relatively uniform and species are limited by their ability to
water pressure, and water movement. Vegetation, especially for-
ests, affects climate through the process of evapotranspiration. disperse, or move between locations (see Figure 52.12B). All the
Cities can create their own climate; cities are on average hotter species that are limited to a region are part of the regional spe-
than rural areas. cies pool (also called gamma diversity). At the landscape scale,
1. What are some of the major ways that Earth’s topography there are multiple community types—for example, lakes, rivers,
affects land and ocean physical conditions? forests, and prairies—arranged in a spatially heterogeneous way
2. Why are cities warmer than rural areas, and what are country across the region (see Figure 52.12C). The change in the number
breezes? How and why could the temperature in a city be of species from one community to another across the landscape
changed with the establishment of a large forested park?
is known as beta diversity. Finally, the local scale (also called
alpha diversity) typically encompasses a single community—a
With an understanding of how Earth’s physical environment, in- group of species assembled together as the result of the physi-
cluding climate, is determined at a variety of spatial scales, we will cal environment, biotic interactions with other species, and the
turn next to biogeography: how the physical environment and other regional species pool (see Figure 52.12D).

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 CHAPTER 52 The Physical Environment and Biogeography of Life

(A) Global scale Global patterns of species diversity and Figure 52.12 Biogeography Depends on Interconnected Spatial
composition are the result of variation in and Temporal Scales The arrows represent the relationship be-
speciation, extinction, and dispersal at tween, and processes important to, species diversity and composition
the spatial scale of continents and at at (A) global, (B) regional, (C) landscape, and (D) local scales.
evolutionary time scales.
Q: At what scales is the regional species pool important to
species diversity and composition?
Within regions, patterns of species
diversity and composition (gamma
diversity) are driven by dispersal In the remainder of this chapter we will consider the distribu-
(B) Regional
scale and extinction across the landscape. tional and diversity patterns of organisms at global and regional
biogeographic scales. Chapter 55 will consider the causes and
consequences of diversity at community and local scales.

Terrestrial biomes reflect global patterns of

temperature and precipitation
(C) Landscape Biomes are groupings of ecologically similar organisms shaped by the
scale environment in which they are found. The classification of biomes is
Local and regional scales most often and easily applied to terrestrial systems. Ecologists classify
are connected by turnover, which is
the difference in species diversity
terrestrial biomes principally by the growth forms of their dominant
and composition (beta diversity) plants, which reflect the evolution of those plants under annual pat-
when moving across the landscape terns of temperature and precipitation.
from one community type to another. The same classic types of biomes may be widely separated,
occurring on different continents, depending in large part on the
(D) Local scale Local patterns of species diversity presence of suitable climatic conditions (Figure 52.13). For exam-
and composition (alpha diversity)
are driven by physical conditions
ple, the desert biome occurs in such distant locations as Arizona in
and species interactions. the North American Southwest and the Namib Desert in Africa;

Equator

Polar ice Mountain zone Temperate shrubland and woodland Temperate grassland
Tundra Temperate deciduous forest Tropical seasonal forest Desert
Boreal forest Temperate evergreen forest Tropical rainforest

Figure 52.13 Global Distribution of Terrestrial Biomes Q: Does the Northern or Southern Hemisphere have a greater number
The distribution of terrestrial biomes is determined primarily by of biome types? What do you think might determine this difference?
annual patterns of temperature and precipitation.
View in Achieve Activity 52.1 Terrestial Biomes
View in Achieve Activity 52.2 Aquatic Biomes

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 KEY CONCEPT 52.4 Biogeography Is the Study of How Organisms Are Distributed on Earth

Tropical Rainforest

Equator

© Michael Doolittle/Alamy Stock Photo

© Morley Read/Alamy Stock Photo

There are over 200 bird species in this tropical rainforest at

Lambir Hills National Park, on the island of Borneo, Malaysia.
Note that the
Bromeliads commonly grow on trees in tropical
Yanganbi, D.R.C. scale changes
rainforests, such as this ceibo tree in Ecuador,
0° above 100 mm
but can also be found in desert biomes.
Average annual temperature 24.6°C of precipitation.
Total annual precipitation 1,828 mm
300

Peterson and R. S. Vose. 1997. Bull Am Meteorol Soc

Precipitation (mm) (
100
)

Data from NOAA GHCN-Monthly, version 2; T. C.

© Thomas Marent/Minden Pictures

80
Temperature (°C) (

30 60

20 40

10 20

78: 2837–2849
)
The mountain gorilla (Gorilla beringei beringei) is a 0 0
critically endangered species. This family lives in the J F M A M J J A S O N D
tropical rainforest of Volcanoes National Park, Rwanda. Month

Go to Achieve for a companion Data in Depth exercise.

both locations are extremely dry and dominated by succulent In the following pages we briefly describe seven terrestrial
plants such as cacti and by drought-tolerant shrubs and grasses. biomes (a subset of those shown in Figure 52.13). For each, a
Why use plants as a way of classifying biomes? Because plants plot of the seasonal patterns of temperature and precipitation—
are immobile, they must adapt to environmental conditions in known as a Walter climate diagram—is included. Devised by the
order to be successful over long periods of time. Thus the growth German biogeographers Heinrich Walter and Helmut Lieth, this
forms of plants strongly reflect their environment in ways that graphic technique plots temperature and precipitation data in
can be compared around the world. Furthermore, by providing a simple way that visualizes a “growing season”—those months
three-dimensional structure, by modifying physical conditions when average temperatures are above freezing (0°C) and when
near the ground, and by providing food, dominant plants strongly average precipitation is sufficient for plant growth. Walter cli-
influence the organisms living there. mate diagrams are predicated on the “rule of thumb” that plant
growth requires temperatures above 0°C and at least 2 mm of
Connect the Concepts Key Concept 37.1 describes how plants precipitation for each 1-degree-Celsius rise in temperature.
growing in a particular type of environment (e.g., the desert) may
They have two y-axis scales, one for temperature and one for
display similar growth forms (e.g., succulence).
precipitation; these axes align 0 mm of precipitation with 0°C

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 CHAPTER 52 The Physical Environment and Biogeography of Life

Desert

Equator

© Paula French/EyeEm/Getty Images

A desert biome in Namibia, Africa, is home to the
gemsbuck, a large antelope in the Oryx genus.

When the precipitation curve

falls below the temperature
Ouargla, Algeria
curve (yellow area), water 31°N
© Terry Donnelly/Alamy Stock Photo

availability limits plant growth. Average annual temperature 22.3°C

Total annual precipitation 39 mm
40 80

Peterson and R. S. Vose. 1997. Bull Am Meteorol Soc

Precipitation (mm) (
)

Data from NOAA GHCN-Monthly, version 2; T. C.

30 60
Temperature (°C) (

20 40
© ZSSD/Minden Pictures

Sonoran desert in bloom

with saguaro (Carnegiea 10 20
gigantea) and cholla

78: 2837–2849
(Cylindropuntia sp.) cacti
and poppies at Organ 0 0 )
Pipe Cactus National J F M A M J J A S O N D
Monument, Arizona, USA. Month

A desert tortoise (Gopherus agassizii ) in its native

habitat, the Mohave Desert, California, USA.

of temperature. The x axis shows 12 months, with the summer epiphytes—plants that grow on other plants, deriving their nutri-
solstice placed in the center of the axis. ents and moisture from air and water rather than soil.
Tropical rainforests provide humans with a range of products,
TROPICAL RAINFOREST The tropical rainforest biome is found including fruits, nuts, medicines, fuel, pulp, and furniture wood.
in equatorial regions where rainfall and temperatures are high Rainforests, however, are currently being cut down or converted
year-round. With no season unsuitable for growth, it is the most to agriculture at a rate of almost 1% per year. In some cases,
productive and species-rich of all biomes. In some areas, there rainforests are recovering, but the soils are often nutrient-poor,
are up to 500 species of trees per square kilometer. Although hindering regrowth.
historically these forests covered 12% of Earth’s surface, and
are now reduced through deforestation, they are home to more DESERT The desert biome is concentrated in two belts, centered
than half of all known species. These forests are home to many around 30°N and 30°S latitude (where warm, dry air sinks under

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 KEY CONCEPT 52.4 Biogeography Is the Study of How Organisms Are Distributed on Earth

Temperate Grassland

© Tom Bean/Alamy Stock Photo

Equator

The vast expanse of the Sand Hill grasslands at

Valentine National Wildlife Refuge, Nebraska, USA.

Denison, Nebraska, USA

The blue area between 41°N

© legna69/Getty Images
the two curves represents Average annual temperature 9.1°C
subfreezing average Total annual precipitation 727 mm
monthly temperatures. 120
Peterson and R. S. Vose. 1997. Bull Am Meteorol Soc
Precipitation (mm) (

100
)

Data from NOAA GHCN-Monthly, version 2; T. C.

Alamy Stock Photo

80 © Laura Romin & Larry Dalton/ A Przewalski’s horse (Equus ferus
przewalskii ), a rare and endangered
Temperature (°C) (

30 60 subspecies of the wild horse, is

native to the grassland steppe of
20 40 Mongolia.
10 20

0 0
78: 2837–2849
)

The wild black-footed ferret (Mustela

–10 nigripes) was reintroduced back into its
J F M A M J J A S O N D native habitat at Grasslands National Park,
Month Saskatchewan, Canada.

high atmospheric pressure; see Figures 52.4 and 52.5). The driest TEMPERATE GRASSLAND Temperate grasslands are found in
of these regions, where rains rarely fall, are far from the oceans, as many parts of the world, all of which are relatively dry for much
in the center of Australia and the middle of the Sahara in Africa. of the year. Most grasslands, such as the pampas of Argentina,
Desert plants have several structural and physiological adapta- the veldt of South Africa, and the Great Plains of North America,
tions that help them conserve water, as described in Key Concept have hot summers and relatively cold winters. In some grass-
37.1. Small desert animals are inactive during the hottest part of lands, most of the precipitation falls in winter (as in California
the day, remaining in underground burrows. Desert mammals grasslands); in others, the majority falls in summer (as in the
have physiological adaptations for conserving water, including a Great Plains and the Russian steppe).
reduced number of sweat glands and kidneys that produce highly Grassland vegetation is structurally simple but rich in species of
concentrated urine. Many desert animals require no water beyond perennial grasses and forbs (herbaceous plants). Grassland plants
what they can extract from the carbohydrates in their food. support herds of large grazing mammals and are adapted to graz-
Humans have used deserts for livestock grazing and agriculture ing and to fire. They store much of their energy underground and
for centuries. Deserts can be irrigated from deep wells or distant resprout quickly after being burned or grazed. There are com-
mountains, but such efforts typically fail as a result of saliniza- paratively few trees in temperate grasslands because trees cannot
tion, the buildup of salts from the evaporation of irrigation water. survive the periodic fires or the dry conditions.

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 CHAPTER 52 The Physical Environment and Biogeography of Life

Temperate Deciduous Forest

© Stuart Black/robertharding/Getty Images

Equator

Wellsboro, Pennsylvania, USA

41°N
Average annual temperature 7.6°C
Total annual precipitation 848 mm
100

Precipitation (mm) (
)
80

Temperature (°C) ( 30 60

20 40
Layers of autumn foliage in a
deciduous forest on Miyajima 10 20
Island, Japan.
0 0

)
–10
North American deciduous J F M A M J J A S O N D
forests are a common home for Month
the black bear (Ursus Data from NOAA GHCN-Monthly,
version 2; T. C. Peterson and R. S. Vose. 1997.
americanus). Bull Am Meteorol Soc 78: 2837–2849
David McIntyre
© Jack Milchanowski/AGE Fotostock

The deciduous forest biome is

home to the spotted salamander
(Ambystoma maculatum) which
breeds in vernal pools throughout
most of the eastern USA.

The topsoil of grasslands is usually rich and deep, and thus ex- Temperate deciduous forests have many more species than
ceptionally well suited to growing crops such as corn and wheat. As boreal forest ecosystems. Those with the highest species rich-
a consequence, most of the world’s temperate grasslands have been ness occur in the southern Appalachian Mountains of the United
turned over to agriculture and no longer exist in their natural state. States and eastern China and Japan—areas that were not cov-
ered by glaciers during the Pleistocene. Many plant genera are
TEMPERATE DECIDUOUS FOREST The temperate deciduous shared among the three geographically separate regions where
forest biome is found in eastern North America, eastern Asia, and this biome is found.
Europe. Temperatures in these regions fluctuate dramatically be- Although many animals are permanent residents of deciduous
tween summer and winter, although precipitation is fairly evenly forests, some (including many birds) migrate to find food resources
distributed throughout the year. The deciduous trees that domi- and escape the winter cold. Others that remain through the winter
nate these forests lose their leaves during the cold winters and hibernate (see Key Concept 38.5), often in underground burrows.
produce new leaves during the warm, moist summers. Many insects pass the winter in a state of diapause (suspended

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 KEY CONCEPT 52.4 Biogeography Is the Study of How Organisms Are Distributed on Earth

Boreal Forest and Temperate Evergreen Forest

Equator
Temperate
evergreen forests

© All Canada Photos/Alamy Stock Photo

are found along
the coasts.

A boreal forest scene in Gros Morne

National Park, Newfoundland, Canada.
The boreal forests of Alaska are
home to moose (Alces americanus),
the largest North American
terrestrial herbivore.
© Shin Yoshino/Minden Pictures

Fort Simpson, Northwest Territories, Canada

61°N
Average annual temperature –4.6°C
Total annual precipitation 333 mm
30 60
Peterson and R. S. Vose. 1997. Bull Am Meteorol Soc
Precipitation (mm) (
)

20 40
Data from NOAA GHCN-Monthly, version 2; T. C.

© Javier Fernández Sánchez/Getty Images

10 20
Temperature (°C) (

0 0

–10

–20
78: 2837–2849
)

–30
J F M A M J J A S O N D A Great Gray Owl (Strix nebulosa) is camouflaged
Month against a boreal forest tree in Finland.

development), the onset of which is triggered by the decreasing The dominant mammals of the boreal forest, such as moose
hours of daylight—a reliable predictor of winter. and hares, eat leaves, whereas the seeds in conifer cones support a
variety of rodents, birds, and insects. Many small mammals hiber-
BOREAL FOREST AND TEMPERATE EVERGREEN FOREST nate in winter, but voles, lemmings, and mice remain active under
The boreal forest biome (also known as taiga) occurs just above the snowpack, serving as food for predators such as foxes and owls.
50°N but below Arctic tundra, and at elevations below alpine tun- The temperate evergreen forest biome occurs along the coasts
dra on temperate-zone mountains. Winters in the boreal forest are of continents in both hemispheres at middle to high latitudes,
long and very cold; summers are short, although often relatively where winters are mild and wet and summers are cool and dry. In
warm. The boreal forests of the Northern Hemisphere are domi- the Northern Hemisphere, the dominant trees in temperate ever-
nated by coniferous trees such as spruces and firs. The short sum- green forests are conifers, some of which are the world’s most mas-
mers favor evergreen leaves, which are ready to photosynthesize sive tree species (including the giant sequoia and coast redwood).
as soon as temperatures warm. In winter, downward-drooping In the Southern Hemisphere, the dominant trees are southern
limbs allow the trees to shed snow easily. beeches (Nothofagus), some of which are evergreen.

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 CHAPTER 52 The Physical Environment and Biogeography of Life

Tundra

Equator
Arctic tundra
is found at
high latitudes.

© imageBROKER/Alamy Stock Photo

The muskox (Ovibos moschatus) was
introduced to the tundra biome of Dovrefjell
National Park, Norway, in 1932.
© Peter van der Veen/NIS/

A brilliant autumn tundra scene at Thorofare Pass in

Denali National Park and Preserve, Alaska, USA.
Minden Pictures

Olenek, Russia
73°N
Average annual temperature –14.3°C
Total annual precipitation 184 mm
20 40

Peterson and R. S. Vose. 1997. Bull Am Meteorol Soc

© David W. Hamilton/Getty Images

Precipitation (mm) (
20
)

10

Data from NOAA GHCN-Monthly, version 2; T. C.

0 0
Temperature (°C) (

–10

–20
The Arctic fox (Vulpes
lagopus) has a white coat –30

78: 2837–2849
that serves as camouflage

)
in the winter. –40
J F M A M J J A S O N D
Month

TUNDRA The tundra biome is found at high latitudes (above much of the year. Resident birds and mammals, such as the Wil-
65°) characterized by low temperatures and a short growing low Ptarmigan (Lagopus lagopus) and Arctic fox (Vulpes lagopus),
season. This biome is underlain by permafrost—soil permeated have thick feathers or fur that may change color with the seasons,
with permanently frozen water. The top few centimeters of the from brown in summer to white in winter.
soil thaw during the short summers, when the Sun may be above
the horizon 24 hours a day. Thus even though there is little pre- Biogeographic regions reflect evolutionary isolation
cipitation near the poles, the soil in lowland Arctic tundra is Climate interacts with local abiotic features to influence where
wet because water cannot drain through the permafrost. As a and how organisms live, but these are not the only factors deter-
result, trees cannot grow in the tundra biome because their roots mining where organisms are found. Evolutionary history—where
cannot penetrate the permafrost. Instead, the tundra biome is and when species originated and diverged—is key to determining
characterized by sedges, forbs, grasses, and low-growing shrubs their biogeography.
such as heaths, willows, and birches. Lichens and mosses are also Until European naturalists traveled the globe in the nineteenth
important vegetation. century, they had no way of knowing how organisms were distrib-
Tundra plants have several structural and physiological adapta- uted in other parts of the world. Alfred Russel Wallace, who along
tions that help them conserve heat, as described in Key Concept with Charles Darwin advanced the idea that natural selection
37.1. Most animals are either summer migrants or are dormant for could account for the evolution of life on Earth (see Key Concept

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 KEY CONCEPT 52.4 Biogeography Is the Study of How Organisms Are Distributed on Earth

19.1), was one of those global travelers. Wallace spent seven years discontinuity that first piqued his curiosity is known to this day
in the Malay Archipelago, where he noticed some remarkable pat- as Wallace’s Line (see Figure 52.14).
terns in the distributions of species. For example, he described the The biotas of different parts of the world differ enough to al-
dramatically different species that inhabited the adjacent islands low us to divide Earth into many continental-scale areas called
of Bali and Lombok. He pointed out that the differences could biogeographic regions (Figure 52.14), each containing character-
not be explained by the physical environment, because Bali and istic assemblages of species. The boundaries of the biogeographic
Lombok are only 24 kilometers apart. regions in Figure 52.14 were originally proposed by Wallace, and
Wallace saw that, based on the distributions of plant and ani- represent assemblages of species that change dramatically, often
mal species, he could draw a line that divided the Malay Archi- over short distances. A major process controlling the formation
pelago into two distinct halves. He correctly deduced that the of these biogeographic regions is continental drift. For example,
dramatic differences in flora and fauna were related to the depth we now know that over the course of the Triassic and Jurassic,
of the channel separating Bali and Lombok. This channel is so the supercontinent Pangaea separated into two great land masses,
deep that it would have remained full of water, and thus would Laurasia and Gondwana (see Figure 23.14), which subsequently
have been a barrier to the movement of terrestrial animals, even separated into the continents we know today. After the land mass-
during the glaciations of the Pleistocene, when sea level dropped es broke up, the descendants of the organisms widely disturbed
more than 100 meters and Bali and the islands to the west were across Pangaea evolved independently, forming new species and
connected to the Asian mainland. new species assemblages. The legacy of these continental move-
With these insights, Wallace established the conceptual foun- ments can be found in several existing taxonomic groups and in
dations of biogeography. In The Geographical Distribution of Ani- the fossil record. For example, the modern southern beeches—
mals, published in 1876, he detailed the factors known at the time trees of the genus Nothofagus—are found in both the Neotropical
that influence the distributions of animals, including past glacia- and the Australasian biogeographic regions. Evidence of fossilized
tion, land bridges, deep ocean channels, and mountain ranges. He Nothofagus pollen from 55 to 34 million years ago has also been
earned some measure of scientific immortality in that the Malay found in Australia, New Zealand, western Antarctica, and South

The Sahara and the Arabian The Himalayan mountain range

Desert separate the Palearctic separates the Oriental and
and Ethiopian regions. Palearctic regions.

70
49 PALEARCTIC

180
Wallace’s Line
NEARCTIC
180 separates the
17 45 ORIENTAL Oriental and
ETHIOPIAN Australasian
6 100 regions.
NEO- 100
The Mexican Plateau TROPICAL
separates the Nearctic
and Neotropical regions. AUSTRALASIAN
80
100–110

ANTARCTIC 45 ANTARCTIC

49
ANTARCTIC

Figure 52.14 Earth’s Biogeographic Regions Wallace identified six major tectonic plates. (After H. Kreft and W. Jetz. 2010. J Biogeogr
major biogeographic regions that are separated by climatic, topographic, 37: 2029–2053.)
or aquatic barriers to dispersal that cause their biotas to differ strikingly
from one another. The red arrows on the map show the time (in millions
Q: As land masses separated, would you expect speciation to
increase? Explain.
of years) since land masses came together. Black arrows show the time
since land masses separated. These areas roughly correspond to Earth’s View in Achieve
Activity 52.3 Major Biogeographic Regions

52_Life12e_Ch 52.indd 19 12/4/19 3:07 PM

 CHAPTER 52 The Physical Environment and Biogeography of Life

America, suggesting that beeches originated in Gondwana during Diversity varies with latitude and longitude
the Cretaceous and were geographically separated by the breakup About 200 years ago, the German explorer and naturalist Al-
of that land mass 100 million years ago (see Figure 52.14). exander von Humboldt spent 5 years traveling around Latin
America. He remarked in the account of his voyages that “the
Connect the Concepts Key Concepts 23.2 and 23.3 describe how nearer we approach the tropics, the greater the increase in the
Earth’s land masses have undergone continental drift over the millennia
variety of structure, grace of form, and mixture of colors, as also
as a result of plate tectonics, shaping major patterns of biogeography.
in perpetual youth and vigour of organic life.” Humboldt and
The evolutionary separation of species can be attributed to two other nineteenth-century scientific explorers such as Wallace
basic processes, vicariance and dispersal. Vicariance occurs when and Darwin became keenly aware of this pattern as they col-
a physical barrier prevents dispersal and divides a species into lected thousands of species in the tropics and compared them
two or more discontinuous populations. Dispersal occurs when with their more meager collections from Europe. As more obser-
the members of a species cross an existing barrier and establish a vations and collections have been made over the last 200 years,
new population elsewhere. the latitudinal gradient in species diversity has been established
Given that the processes of vicariance and dispersal both influ- for a wide variety of taxa, including some groups of birds, mam-
ence distribution patterns, how can biogeographers determine the mals, flowering plants, and insects such as swallowtail butter-
role of each process when reconstructing the evolutionary history flies (Figure 52.15A). Significant variation in species diversity
of a particular species? As you saw in Chapter 20, taxonomists is also seen with longitude, but the number of species depends
have developed powerful molecular methods of reconstructing largely on the longitudes chosen.
the phylogenetic relationships among organisms that can be used There are intriguing exceptions to the classic latitudinal pat-
to understand how organisms came to occupy their present-day terns of species diversity just described. Some taxa show a positive
distributions. Phylogenetic trees can be used to discover whether relationship between species number and latitude (Figure 52.15B).
the distribution of an ancestral species was influenced by a vicari- That is, among some taxa—for example, seabirds—species number
ant event, such as continental drift or a change in sea level, or was increases at higher latitudes. This pattern of seabird diversity cor-
simply the result of a dispersal event. relates with marine net primary productivity, which is substan-
Media Clip 52.1 Rafting to Madagascar tially higher in temperate and polar oceans than in the tropics.
Life12e.com/mc52.1

(A) Global swallowtail butterfly diversity (B) Global seabird diversity

150
Number of species

There are many more swallowtail

butterfly species in the tropics…
100
70ºN
3
60º 50
David McIntyre

9
50º 0
16 Northern Tropics Southern
40º Hemisphere Hemisphere
19 (polar/temperate) (polar/temperate)
Tropic of 30º
Cancer 30 Latitude
20º
65
Jerome Whittingham

10º
© iStock.com/

Equator 81
© Irina Tischenko/Shutterstock.com

0º Auks occur in
79 the Northern
10º Hemisphere.
Tropic of 69
Capricorn 20º
46 Pelicans
30º occur in
13 …than there are at the tropics.
40º temperate latitudes.
50ºS
10 20 30 40 50 60 70 80 90
Number of species

Figure 52.15 Latitudinal Gradients in Diversity (A) Among swallowtail butter-

© Jacqueline Abromeit/

flies (Papilionidae), species richness decreases with latitude both north and south of
the equator. (B) Seabirds, by contrast, show a positive relationship between species Penguins
Shutterstock.com

occur in the
diversity and latitude: the number of seabird species increases at higher latitudes.
Southern
(A after J. M. Scriber. 1973. Psyche 80: 355–373; B graph data from P. Harrison 1987. Hemisphere.
A Field Guide to Seabirds of the World. Penguin Random House: London.)

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 KEY CONCEPT 52.5 Geographic Area and Humans Affect Regional Species Diversity

Connect the Concepts Key Concept 56.2 describes net primary competition. The productivity hypothesis might explain the positive
production (NPP) as the amount of biomass incorporated into the relationship between seabird diversity and increasing latitude, given
tissues of primary producers and as the base of all food chains. that ocean productivity is generally higher at temperate and polar
latitudes. As you will see in Chapter 55, the role of productivity can
We consider productivity differences along with other expla- more easily be explored at smaller spatial scales, where experiments
nations of latitudinal gradients in diversity in the next section. can give us a better idea of its role in determining species diversity.
KEY CONCEPT
Geographic variation in diversity is explained
by multiple factors 52.4 Recap and Assess
Although most ecologists agree that latitudinal gradients in diver- Biogeographic patterns are interconnected across a hierarchy of
sity exist, there is much less consensus as to why they exist. Doz- scales ranging from global to local. Three strong biogeographic
ens of hypotheses have been proposed to explain these patterns, patterns can be seen: (1) biomes, or groupings of ecologically
but they are hard to evaluate—in part because there are multiple similar organisms, are shaped by the environment, (2) diversity
varies from continent to continent, forming biogeographic regions,
and confounding differences in such factors as geographic area, and (3) diversity varies with latitude.
climate, and productivity with latitude. More important, because 1. What are the differences and similarities between the
these factors operate at large spatial scales and over evolutionary tropical rainforest biome and the temperate evergreen
time, it is impossible to conduct experiments to isolate the factors forest biome?
and separate correlation from causation. 2. Refer to Figure 52.14. Why did Wallace’s Line help explain
Here we present three well-known hypotheses that have been what controls the formation of biogeographic regions?
advanced to account for latitudinal gradients in diversity. Cor- 3. Over Earth’s history, ice sheets extended into temperate
roborative evidence can be found for each of these hypotheses, latitudes. Does this fact better support the species
varying with taxon, locality, and scale, which suggests that none diversification rate or the species diversification time
hypothesis proposed to explain latitudinal variation in
of these hypotheses is mutually exclusive. Multiple factors, work- diversity? Explain.
ing at multiple spatial scales, are likely responsible for common
biogeographic patterns.
As you have seen, an important aspect of biogeography is the re-
SPECIES DIVERSIFICATION RATE This hypothesis proposes lationship between diversity and geographic area. We have con-
that in the tropics the rate of speciation is higher and the rate of sidered this relationship at global scales. We turn now to smaller
extinctions is lower, resulting in overall higher species diversity regional scales, to understand in more detail the importance of
than in temperate or polar regions. Two factors could account for geographic area to patterns of biogeography.
a higher speciation rate in the tropics: a larger geographic area
(this is the widest part of the planet, so it encompasses the great-
est amount of area) and a warm and stable climate. The rationale
KEY CONCEPT Geographic Area and
is that a larger geographic area with a consistent climate should
Humans Affect Regional
promote speciation because species will have larger geographic
ranges, giving them a greater chance of reproductive isolation (see
52.5 Species Diversity
Key Concept 22.2). In addition, these large and stable areas should
Learning Objectives
reduce extinctions. The net result should be an increase in the
number of species in the tropics. 52.5.1 Discuss the species–area relationship and use
the theory of island biogeography to explain
the pattern.
SPECIES DIVERSIFICATION TIME This hypothesis proposes
that the amount of time over which speciation has taken place 52.5.2 Describe how humans have affected
biogeographic patterns.
is greatest in the tropics—in other words, that the tropics have
a longer evolutionary history than temperate or polar regions. 52.5.3 Describe specific ways in which habitat
fragmentation negatively affects species diversity.
First championed by Wallace in 1878, this hypothesis proposes
that tropical regions, because they are more climatically stable
over time, have more time to diversify than temperate or polar Biogeographers have repeatedly documented what is termed the
regions, where severe climate conditions (such as ice ages) could species–area relationship, in which species diversity increases
have decreased speciation and increased extinction. Thus even if with increasing area. The relationship has most often been mea-
the rates of speciation and extinction are the same worldwide, the sured for islands or islandlike habitats—any isolated area sur-
tropics should have accumulated more species over time simply rounded by a “sea” of dissimilar habitat. In addition to showing
because of the lack of dramatic changes in climate. a pattern of increasing species diversity with increasing area,
islands also show a pattern of decreasing species number with
PRODUCTIVITY This hypothesis proposes that species diversi- distance from the source of those species, or the species pool
fication is promoted by higher productivity, which allows species (Figure 52.16). How do island area and distance act together to
more resources and thus decreased risk of species extinction due to produce species–area relationships?

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 CHAPTER 52 The Physical Environment and Biogeography of Life

The biologist Edward O. Wilson was

struck by the species–area relationship
from his research on ant biogeography and
was looking for explanations. He teamed
up with Robert MacArthur, a gifted math-
ematical ecologist, to develop the theory
Wake Hawaii of island biogeography to explain this
Island pattern. They based their theory on just
Marshall Among islands of a two processes: the immigration of new
Islands given size, those nearest
Palau species to an island and the extinction
to New Guinea have the
most bird species.
of species already present on that island
1,000 (Figure 52.17). The premise of island bio-
Near islands geography is that the number of species
Solomon Intermediate islands on an island represents a balance between
Islands
Number of bird species

Far islands
New the rate at which species immigrate to and
Guinea 100 colonize the island and the rate at which
Solomon
resident species become locally extinct.
Palau
Islands The rate of immigration is determined
Figure 52.16 The Species–Area Hawaii in part by the number of species in the re-
Relationship E. O. Wilson and Robert 10 gional species pool. In the case of oceanic
MacArthur plotted the number of species islands, the species pool comprises all the
of birds against the size and distance of Wake
Island species on the nearest mainland location.
islands from New Guinea. Larger and closer
Not all species that reach the island will
islands consistently contained more spe- 1 persist there, however. The more species
cies, a fact that these scientists incorpo- 26 260 2,600 26,000 260,000
rated in their theory of island biogeography. there are on an island, the greater the like-
Area (km2)
(After R. H. MacArthur and E. O. Wilson. lihood that some will go extinct, either due
1963. Evolution 17: 373–387.) to chance or the inability to coexist with
other species on the island. The number of
species that will be sustained on any par-
Mainland ticular island is determined by the balance
Immigration
of the immigration rate and the extinction
Extinction
rate for that island. Even though species
will come and go over time (i.e., immigrate
The point at which the
immigration and extinction and become extinct), that turnover will al-
curves intersect predicts Large, Small, ways result in some equilibrium number
the equilibrium number of near near of species, assuming a constant species
species on the island (S).
pool and natural extinction events.
Large,
Rate

far The actual number of species on an is-

Small Small, land will ultimately depend on two factors:
Near Large far
1. The size (area) of the island. The
Far smaller the island, the fewer resources
Small, isolated …than larger
islands are islands closer it provides, the greater the potential
predicted to to a source for competition, and the higher the
have fewer of colonists. extinction rate will be (see Figure
species…
52.17). Larger islands provide more
resources and can sustain larger
Ssmall, far Slarge, near populations (which tend to have
Slarge, far Ssmall, near lower extinction rates than small
populations).
Number of species present
2. Distance of the island from the
Figure 52.17 The Equilibrium Theory of Island Biogeography MacArthur and Wilson’s species pool. The farther the island is
theory emphasized the balance between species immigration rates and species extinction rates from the source of immigrants, the
for islands of different sizes and at different distances from a source of colonizing species. (After
lower the immigration rate—the rate
R. H. MacArthur and E. O. Wilson. 1963. Evolution 17: 373–387.)
at which new species arrive—will be
View in Achieve
(see Figure 52.17).
Simulation 52.1 Biogeography

52_Life12e_Ch 52.indd 22 12/4/19 3:07 PM

 KEY CONCEPT 52.5 Geographic Area and Humans Affect Regional Species Diversity

Between 1966 and 1969, Wilson and his student Daniel Simberl- becoming smaller and more fragmented by local- and regional-
off conducted an experiment to test the theory of island biogeog- scale forces such as agriculture, deforestation, and urbanization.
raphy, using arthropods living on mangrove islands in the Florida They are being transformed into islandlike habitats—isolated
Keys. You can view this experiment and the results online. patches of suitable habitat such as parks or forest fragments, sur-
View in Achieve rounded by extensive areas of unsuitable habitat similar to the cat-
Animation 52.3 Island Biogeography in the Florida Keys tle pastures of Amazonia described at the opening of this chapter.
The fragmentation of the Amazon rainforest led Thomas Love-
Humans exert a powerful influence on joy and his colleagues to initiate, in 1979, one of the largest and
biogeographic patterns longest-running ecological experiments ever conducted to inves-
The theory of island biogeography has provided surprising in- tigate the effects of tropical forest fragmentation on species diver-
sight into the role human’s play in regional biogeographic patterns. sity (Investigating Life: The Largest Experiment on Earth). The
As you saw in Key Concepts 52.3 and 52.4, Earth’s biomes are Biological Dynamics of Forest Fragments Project (BDFFP) took

▶InvestigatingLIFE The Largest Experiment on Earth

Experiment Experimental fragments …deforested
of different sizes were land.
Original Paper: G. Ferraz et al. 2003. Rates of species loss from
surrounded by…
Amazonian forest fragments. Proc Natl Acad Sci USA 100: 14069–

After R. Bierregaard et al. 2001. Lessons from Amazonia:

14073. © 2003 National Academy of Sciences U.S.A.

The Ecology and Conservation of a Fragmented Forest.

Thomas Lovejoy and his colleagues asked what was the minimum
area needed to maintain species diversity in the rainforest fragments
created by logging near Manaus, Brazil. They conducted an experi- 100
ment, starting in 1979, that took advantage of an existing Brazilian 1

Yale University Press: New Haven

law that required landowners who cut rainforest to leave half of it un-
touched. In 2003, Ferraz and colleagues reported on one particular
aspect of the experiment: the number of forest understory birds liv-

HYPOTHESIS▸ The number of understory bird species will

ing in different-sized forest fragments surrounded by deforested land. 10

decline faster after the isolation of the fragment, and in smaller

METHOD▸ RESULTS▸ The average initial number of species was 83 for the
rainforest fragments compared with larger rainforest fragments.

1. Starting in 1979, fragments of three sizes (1, 10, and 100 ha) 1-ha fragment, 92 for the 10-ha fragment, and 113 for the 100-ha
were established 80 km north of Manaus. The fragments were fragment. Despite finding a total of 164 understory bird species in
separated from continuous forest by at least 100 meters of the fragments over the course of 13 years, there was a significant
cleared land. drop in the number of species over time. The t50 was shortest in the
1-ha fragments and longest in the 100-ha fragments. Fragments of
2. Birds were captured in mist nests placed in the fragments prior
100 ha were predicted to lose half of their species in 12 years.
to deforestation and then for 12 years after fragment establish-
ment. The mist nets were placed in roughly the same locations 15
throughout the study.
3. Each newly captured bird was recorded for species, given an 10
t50 (years)

individually numbered band, and released back into the rainfor-

est from which it was caught.
5
4. The data gathered were used in species-loss curves for each
fragment, and a scaling factor (i.e., a number that scales, or
multiplies, by some quantity) was estimated to determine the 1
1 10 100
time it takes to lose half of the bird species (t50) in fragments of Size of fragment area (hectares)

CONCLUSION▸ The number of understory bird species declined

different sizes.
View in Achieve
Animation 52.4 Edge Effects after the fragmentation of tropical rainforest habitat by deforesta-
tion. Smaller rainforest fragments lost species faster than larger
forest fragments. Even the largest fragments were predicted to
lose half of their bird species.

InvestigatingLIFE Work with the Data follows on the next page.

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 CHAPTER 52 The Physical Environment and Biogeography of Life

Work with the Data

The data in the graph shown in the experiment can be used to 2. Suppose you are consulting for the Brazilian government on the
calculate a scaling factor for the time it takes to lose half of the conservation of rainforests near Manaus. Assuming that the t50
bird species within a fragment. The scaling factor shows that to for 1 ha is 5 years, what is the minimum size that a rainforest
increase the t50 (i.e., the time it takes fragments to lose half their fragment in the Manaus area needs to be to ensure that half of
species) by 10-fold, fragment area would need to increase 1,000- the bird species remain 50 years after the deforestation event
fold. Here you will use this scaling factor and other data provided that isolated them? How about a century after deforestation?
in the graph to determine the area needed for the conservation of 3. Suppose it takes 100 years for rainforests to fully recover after

QUESTIONS▸
understory birds in the Amazon forests outside Manaus, Brazil. a deforestation event in the Manaus region. Given that the aver-
age size of a rainforest fragment in this area is no bigger than
1. Graph the initial numbers of bird species by the size of the 1,000 ha, will the number of species decline by half before the
fragment area. Do these fragments follow the species–area recovery of rainforest occurs around the largest fragments?
relationship?

Go to Achieve for a companion Data in Depth exercise.

▶InvestigatingLIFE
advantage of an existing Brazilian law that required landown-
ers who cut rainforest to leave half of it untouched. Lovejoy was
able to survey the species diversity in different-sized fragments
(either 1, 10, or 100 hectares [ha] in size surrounded by deforested
land). Among several results, his group found that, after more
than 30 years of isolation, even the largest deforested fragments
QA
and
How do geographic area and isolation affect
the biogeography of life on Earth?
At the opening of the chapter, we asked how geographic area
surveyed (100 ha) had lost half of their bird species diversity. and isolation might affect the biogeography of life on Earth. You
The researchers were also surprised by two additional findings. have seen that this question is highly relevant at different spa-
First, even minimal distances of 80 meters between fragments tial scales—from whole continents to small islands—and also
resulted in strong avoidance of the clearings, and thus isolation, over different time scales—from millions of years to months.
by birds, insects, and tree-dwelling mammals that lived in the Because the spatial and temporal scales at which biogeo-
graphic patterns manifest themselves are so wide and inter-
fragments. Second, the fragmentation exposed species within
connected, it can be hard to test theories of biogeography with
a fragment to a variety of potential hazards, including extreme
experiments that are large and long enough to be meaningful.
heat, fires, hunting, predators, diseases, and invasive species. The Biological Dynamics of Forest Fragments Project (BDFFP),
These so-called edge effects not only reduce the immigration an experiment conducted in the most species-rich region of the
rate from one fragment to another, thus contributing to a smaller world, and at arguably the largest scale ever attempted, offers
species pool, but they effectively reduce the size of the fragment, support for theories proposed nearly 200 years ago by explor-
thus increasing the extinction rate, by making the edge of the ers such as Humboldt, Wallace, and Darwin, who hypothesized
fragment much less hospitable habitat. that the distribution of species on Earth is highly influenced by
the geographic area and isolation those species have experi-
KEY CONCEPT enced. The question remains, in the case of the Amazon Basin,
52.5 Recap and Assess whether species will ultimately adapt to the changing biogeo-
graphic landscape or become a casualty of an extinction event
Species–area relationships, in which species diversity increases that would be unrivaled in human history.
with increasing area, have been shown for islands and islandlike
habitats. Smaller islands have higher extinction rates and lower
immigration rates compared with larger islands. Earth’s biomes
Future directions
are becoming smaller and more fragmented by human activities, One of the major findings of the BDFFP is the importance of
leading to lower diversity in the remaining habitats. connectivity in maintaining species diversity. Management and
1. Why do island size and distance from a species pool affect permitting of forest clearing are focusing on creative ways to
island biogeography? connect forest fragments by taking advantage of existing land-
2. Why is habitat fragmentation by human activities threatening scape features to increase connectivity of disturbed habitat. For
species diversity? Besides ceasing the activity altogether, what example, in Amazonia strong laws exist to prohibit clearing for-
can be done to reduce this threat? est along rivers and steep slopes, providing strips of land that
can serve to connect fragments around newly cleared land.

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 VISUAL SUMMARY 52
© Morley Read/Alamy Stock Photo

You should be able to relate each summary to the adjacent figures. If you go to this visual summary in Achieve,
you can follow links to figures, animations, activities, and simulations that will help you consolidate the material.

KEY CONCEPT Ecology Is the Study of the Interrelationships among

52.1 Organisms and the Environment
■ Ecology differs from environmentalism, which involves Figure 52.1
the use of ecological knowledge to inform our stewardship

Courtesy of Sally Hacker

of natural resources.
■ Ecologists have a use-inspired focus to their research, moti-

Courtesy of Sally Hacker

vated by a desire to address Earth’s ecological problems.
■ Ecology is studied at multiple levels of organization, ranging
from individuals to the biosphere.
Questions
1. Define the difference between ecology and

© Konrad Wothe/Minden Pictures/

environmentalism.
2. What is meant by “use-inspired” ecological research?
3. How are individuals connected to communities?

Getty Images

KEY CONCEPT Global Climate Is a Fundamental Component of the

52.2 Physical Environment Go to Animation 52.1

■ Climate and weather both refer to atmospheric conditions, Figure 52.5

but climate represents conditions over years to High pressure
90ºN
Polar
millennia whereas weather represents conditions over Sparse precipitation
in all seasons zone

days to weeks. Low pressure 60º

■
Ample precipitation
Rising air in all seasons Temperate
Solar radiation drives global climate. Of the solar radiation Descending zone
air Winter wet, summer dry
that hits Earth, some is reflected back to space (30%) and 30°N High pressure 30º
Dry in all seasons
the rest is absorbed by the atmosphere (20%) or by Earth’s Summer wet, winter dry

surface (50%). Greenhouse gases trap heat radiating back Tropics

Sunlight Low pressure
0° 0º
out toward space. Abundant precipitation in all seasons

■
Tropics
Latitudinal differences in solar energy input drive patterns of Summer wet, winter dry

atmospheric circulation. 30°S

High pressure
Dry in all seasons 30º

■
Winter wet, summer dry
The interaction of Earth’s rotation and north–south air mass Low pressure
Temperate
zone
movements generate prevailing winds, which in turn drive Ample precipitation
in all seasons 60º
ocean currents. High pressure Polar

■
Sparse precipitation zone
in all seasons
The seasons are a consequence of Earth’s tilt and orbit 90ºS

around the Sun. Figure 52.6

90°N

Questions
Easterlies
60°
1. Does a week of extreme cold weather tell us that Earth’s
climate is not warming? Westerlies
30°
2. Why are the tropics warm and wet and the poles cold
and dry? NE Trade Winds
Direction
0° of Earth’s
3. What direction do ocean currents flow as a result of rotation

trade winds? SE Trade

Winds
30°
Westerlies

60°
Easterlies

90°S

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 VISUAL SUMMARY 52
KEY CONCEPT Topography, Vegetation, and Humans Modify
52.3 the Physical Environment Go to Animation 52.2
■ Earth’s land topography creates rain shadows and temperature Figure 52.9
inversions, producing variation in temperature and precipitation.
■ Ocean-floor topography affects water depth, which affects light
penetration and water temperature, pressure, and movement,
creating aquatic zones and ocean upwelling.
■ Vegetation, especially forests, affects climate through the process of
evapotranspiration.
■ Humans have transformed their environment, including urban climate.
Questions Wind
1. How does a rain shadow affect climate on either side of a
mountain range?
2. How do winds and ocean-floor topography create ocean upwelling?
3. Why would planting more trees in an urban area help with urban
14°
climate warming? 13°
12°
11°

KEY CONCEPT Biogeography Is the Study of How Organisms

52.4 Are Distributed on Earth Go to Activities 52.1–52.3
■ Patterns of biogeography are interconnected across a hierarchy Figure 52.12
of scales, ranging from global to regional to landscape to local.
■ Terrestrial biomes are groupings of ecologically similar dominant
plants shaped by patterns of temperature and precipitation.
■ Earth’s diversity varies from continent to continent, forming
biogeographic regions and reflecting evolutionary isolation as a
consequence of Earth’s geological history.
■ Earth’s diversity varies with latitude. Hypotheses explaining this
variation rely on multiple factors, including geographic area, time,
climate, and productivity.
Questions
1. How does the regional scale determine local community species
diversity and composition?
2. What criteria are used to determine Earth’s terrestrial biomes?
3. Earth’s diversity varies by continent and by latitude. How do
evolutionary processes contribute to these patterns?
Figure 52.14

70
49 PALEARCTIC

180
NEARCTIC
180
17 45 ORIENTAL
ETHIOPIAN
6 100
NEO- 100
TROPICAL

AUSTRALASIAN
80
100–110

ANTARCTIC 45 ANTARCTIC

49
ANTARCTIC

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 VISUAL SUMMARY 52
KEY CONCEPT Geographic Area and Humans Affect Regional Go to Animations 52.3 and 52.4
52.5 Species Diversity and Simulation 52.1
■ Species–area relationships, in which species diversity Figure 52.17
increases with area, have been shown for islands and
Mainland
area fragments. Immigration

■
Extinction
The theory of island biogeography states that the
number of species on an island is a balance between
immigration and extinction rates, which vary by island Large, Small,
size and distance from the species pool. near near

■ As humans fragment Earth’s biomes through agriculture, Large,

Rate
far
deforestation, and urbanization, remaining habitats be- Small Small,
come more isolated, or islandlike. Near Large far

Far
Questions
1. Describe why species diversity increases with increas-
ing island or fragment area.
2. Using Figure 52.17, explain why the small and far
islands have lower species numbers than the large and Number of species present
near islands.

Go to Achieve for the eBook, LearningCurve, animations, activities,

simulations, and additional resources and assignments.

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