23

Chapter

Terrestrial Ecosystems

Spectacular fall color is a hallmark of the eastern deciduous mixed hardwood forest.

Chapter Guide
23.1 Terrestrial Ecosystems Reflect Adaptations of the Dominant Plant
Life-Forms
23.2 Tropical Forests Characterize the Equatorial Zone
23.3 Tropical Savannas Are Characteristic of Semiarid Regions with Seasonal
Rainfall
23.4 Grassland Ecosystems of the Temperate Zone Vary with Climate
and Geography
23.5 Deserts Represent a Diverse Group of Ecosystems
23.6 Mediterranean Climates Support Temperate Shrublands
23.7 Forest Ecosystems Dominate the Wetter Regions of the Temperate Zone
23.8 Conifer Forests Dominate the Cool Temperate and Boreal Zones
23.9 Low Precipitation and Cold Temperatures Define the Arctic Tundra
Ecological Issues & Applications Forest Management

526

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  C h ap t er 23   •  Terrestrial Ecosystems    527

I
n 1939, ecologists F. E. Clements (Carnegie Insti­ 450
tution of Washington) and V. E. Shelford (University of
400
Illinois) introduced an approach for combining the broad-scale
distribution of plants and associated animals into a single classifi­

Mean annual precipitation (cm)

350
cation system. In their book Bio-ecology, Clements and Shelford Tropical
referred to these biotic units as biomes. Biomes are classified 300 rain forest

according to the predominant plant types. There are at least eight Temperate
250
major terrestrial biome types, but there may be more, depending rain forest

on how finely the biomes are classified. These include tropical 200
Tropical
seasonal
forest, temperate forest, conifer forest (taiga or boreal forest), forest
Temperate forest
tropical savanna, temperate grasslands, chaparral (shrublands), 150
tundra, and desert (Figure 23.1). These broad categories reflect
100 Thorn forest
the relative contribution of three general plant life-forms: trees, (Savanna) Woodland
Taiga
shrubs, and grasses. A closed canopy of trees characterizes forest 50 Thorn scrub (Grassland)
Shrubland
ecosystems. Woodland and savanna ecosystems are character­ Desert Tundra
ized by the codominance of grasses and trees (or shrubs). As the 30 25 20 15 10 5 0 –5 –10 –15
names imply, shrubs are the dominant plant form in shrublands, Mean annual temperature (°C)
and grasses dominate in grasslands. Desert is a general category Tropical—Subtropical—Warm temperate—Cold temperate—Arctic–Alpine
used to refer to areas with scarce plant cover.
When the plant ecologist Robert Whittaker of Cornell Figure 23.2  The pattern of terrestrial biomes in relation to
temperature and moisture. Where the climate varies, soil can
University plotted these biome types on gradients of mean
shift the balance between types. The dashed line encloses
annual temperature and mean annual precipitation, he found
environments in which either grassland or one of the types
they formed a distinctive climatic pattern, as graphed in dominated by woody plants may prevail.
Figure  23.2. As the graph indicates, boundaries between bi­ (Adapted from Whittaker 1970.)
omes are broad and often indistinct as they blend into one an­
other. Besides climate, other factors such as topography, soils,

Paleartic Figure 23.1  Major

biomes and biogeographical
realms of the world.
(Adapted from Olson et al. 2001.)

Neartic

Oceania

Oceania Indo-
Malay

Afrotropic
Neotropic

Australasia

Antartic

Tropical and subtropical moist broadleaf forests Temperate grasslands, savannas, and shrublands
Tropical and subtropical dry broadleaf forests Flooded grasslands and savannas
Tropical and subtropical coniferous forests Montane grasslands and shrublands
Temperate broadleaf and mixed forests Tundra
Temperate coniferous forests Mediterranean forests, woodlands, and scrub
Boreal forests/taiga Deserts and xeric shrublands
Tropical and subtropical grasslands, savannas, Mangroves
and shrublands

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 528    Part se ve n   •  Eco l o g ical B io g e og r a p h y

and exposure to disturbances such as fire can influence which resources to stems than do trees. The production of woody tis­
of several biome types occupy an area. sue gives the advantage of height and access to light, but it also
If we plot the relationship between mean annual tempera­ has the associated cost of maintenance and respiration. If this
ture and precipitation for locations on the land surface, another cost cannot be offset by carbon gain through photosynthesis,
general pattern emerges from Whittaker’s analysis of the rela­ the plant is unable to maintain a positive carbon balance and
tionship between biomes and climate. The range of observed dies (see Chapter 6). As a result, as environmental conditions
values for mean annual precipitation declines with decreasing become adverse for photosynthesis (dry, low nutrient concen­
mean annual temperature (note that the range of biomes de­ trations, or short growing season and cold temperatures), trees
fined by precipitation along the y-axis decreases with declin­ decline in both stature and density until they can no longer
ing temperatures along the x-axis). In geographic terms, this persist as part of the plant community.
relationship indicates a decrease in the range of environmental Within the broad classes of forest and woodland ecosys­
conditions defined by moisture availability as one moves from tems in which trees are dominant or codominant, leaf form
the tropics to the temperate and arctic regions (see label on is another plant characteristic that ecologists use to classify
x-axis of Figure 23.2 and Figure 2.17). This relationship be­ ecosystems. Leaves can be classified into two broad categories
tween climate and geography reflects the systematic latitudinal based on their longevity. Leaves that live for only a single year
pattern of environmental conditions discussed in Chapter 2 that or growing season are classified as deciduous, whereas those
result directly from seasonal variations in the influx of solar that live beyond a year are called evergreen. The deciduous
radiation to Earth’s surface. Mean annual temperature decreases leaf is characteristic of environments with a distinct growing
from the equator to the poles, whereas seasonal variation in tem­ season. Leaves are typically shed at the end of the growing sea­
peratures (and day length) increases (see Figures 2.5 and 2.8). son and then regrown at the beginning of the next. Deciduous
The result is a decline in the growing season (period over which leaf type is further divided into two categories based on dor­
photosynthesis and plant growth can be maintained). mancy period. Winter-deciduous leaves are characteristic of
The systematic variation in climate with latitude is not temperate regions, where the period of dormancy corresponds
limited to temperature. Average annual precipitation decreases to low (below freezing) temperatures (Figures 23.3a and
with increasing latitude as a result of the interaction of humid­ 23.3b; also see discussion of plant adaptations in Chapter 6).
ity and temperature (see Section 2.6 and Figure 2.17). With de­ Drought-deciduous leaves are characteristic of environments
clining temperatures, the amount of moisture that can be held with seasonal rainfall, especially in the subtropical and tropi­
in the air declines, reducing the overall amount of precipitation cal regions, where leaves are shed during the dry period
(see Figure 2.15). As we shall see in this chapter, these sys­ (Figures  23.3c and 23.3d). The advantage of the deciduous
tematic patterns of climate across the globe dictate the general habit is that the plant does not incur the additional cost of
distribution of terrestrial biomes on Earth’s surface. maintenance and respiration during the period of the year when
environmental conditions restrict photosynthesis.
Evergreen leaves can likewise be classified into two broad
23.1  Terrestrial Ecosystems categories. The broadleaf evergreen leaf type (Figure 23.4a)
Reflect Adaptations of the is characteristic of environments with no distinct growing
season where photosynthesis and growth continue year-round,
Dominant Plant Life-Forms such as tropical rain forests. The needle-leaf evergreen form
Given that the broad classification of terrestrial biomes pre­ (Figure  23.4b) is characteristic of environments where the
sented in Figures 23.1 and 23.2 (forest, woodland/savanna, growing season is very short (northern latitudes) or nutrient
shrubland, and grassland) reflects the relative contribution of availability severely constrains photosynthesis and plant growth.
three general plant life-forms (trees, shrubs, and grasses), the A simple economic model has been proposed to explain
question of what controls the distribution of biomes relative the adaptation of this leaf form (see discussion of leaf longev­
to climate becomes: Why are there consistent patterns in the ity in Chapter 6, Section 6.11). The production of a leaf has a
distribution and abundance of these three dominant plant life- “cost” to the plant that can be defined in terms of the carbon
forms that relate to climate and the physical environment? The and other nutrients required to construct the leaf. The time
answer to this question lies in the adaptations that these three required to “pay back” the cost of production (carbon) will
very different plant life-forms possess, as well as the advan­ be a function of the rate of net photosynthesis (carbon gain).
tages and constraints arising from these adaptations under dif­ If environmental conditions result in low rates of net photo­
ferent environmental conditions. synthesis, the period of time required to pay back the cost of
Although the broad categories of grasses, shrubs, and trees production will be longer. If the rate of photosynthesis is low
each represent a diverse range of species and characteristics, enough, it may not be possible to pay back the cost over the
they have fundamentally different patterns of carbon allocation period of a single growing season. A plant adapted to such en­
and morphology (see Chapter 6). Grasses allocate less carbon vironmental conditions cannot “afford” a deciduous leaf form,
to the production of supportive tissues (stems) than do woody which requires producing new leaves every year. The leaves of
plants (shrubs and trees), enabling grasses to maintain a higher evergreens, however, may survive for several years. So under
proportion of their biomass in photosynthetic tissues (leaves). this model, we can view the needle-leaf evergreen as a plant
For woody plants, shrubs allocate a lower percentage of their adapted for survival in an environment with a distinct growing

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  C h ap t er 23   •  Terrestrial Ecosystems    529

season, in which conditions limit the plant’s ability to produce

enough carbon through photosynthesis during the growing sea­
son to pay for the cost of producing the leaves.
On combining the simple classification of plant life-forms
and leaf type with the large-scale patterns of climate presented
previously, we can begin to understand the distribution of bi­
(a)
ome types relative to the axes of temperature and precipitation
shown in Figure 23.2. Ecosystems characteristic of warm, wet
climates with no distinct seasonality are dominated by broadleaf
evergreen trees and are called tropical (and subtropical) rain for­
est. As conditions become drier, with a distinct dry season, the
broadleaf evergreen habit gives way to drought-deciduous trees
that characterize the seasonal tropical forests. As precipitation
declines further, the stature and density of these trees declines,
giving rise to the woodlands and savannas that are characterized
by the coexistence of trees (shrubs) and grasses. As precipita­
tion further declines, trees can no longer be supported, giving
rise to the arid shrublands (thorn scrub) and desert.
The temperature axis represents the latitudinal gradient
from the equator to the poles (see geographical labels on x-axis
of Figure 23.2). Moving from the broadleaf evergreen forests
(b) of the wet tropics into the cooler, seasonal environments of the
temperate regions, the dominant trees are winter-deciduous.
These are the regions of temperate deciduous forest. In areas
of the temperate region where precipitation is insufficient to
support trees, grasses dominate and give rise to the prairies

Figure 23.4  Examples of evergreen trees. (a) Broadleaf

evergreen trees dominate the canopy of this tropical rain
forest in Queensland, Australia. (b) Needle-leaf evergreen
trees (foxtail pine) inhabit the high-altitude zone of the
Sierra Nevada in western North America.
(a)

(c)

(b)

(d)

Figure 23.3  Examples of winter- and drought-deciduous trees.

Temperate deciduous forest in central Virginia during (a) summer
and (b) winter seasons. Semiarid savanna/woodland in Zimbabwe,
Africa, during (c) rainy and (d) dry seasons.

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 530    Part se ve n   •  Eco l o g ical B io g e og r a p h y

Iquitos Gatun Ipoh Madang Figure 23.5  Geographic

104 m 29 m 39 m 6m
°C 26.4 2845 mm °C 27.1 3149 mm 0° °C 26.4 2551 mm °C 27.3 3509 mm distribution of Earth’s tropical
40 500 40 500 40 500 40 500
400 400 400 400 forest ecosystems and associated
30 30 30 30
20
300
20
300
20
300
20
300 climate diagrams showing
200 200 200 200 long-term patterns of monthly
10 100 10 100 10 100 10 100
0 0 0 0 0 0 0 0 temperature and precipitation for
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
selected locations. Note the lack
of seasonality in mean monthly
temperatures, which are above
40° 40° 20°C. Although the rainfall in some
regions is seasonal, note that
20° 20° minimum monthly precipitation
is typically above 60 mm, and
0° 0° total annual precipitation above
2000 mm.
20° 20° (Adapted from Archibold 1995.)

40° 40°

Uaupes Manaus Yabassi Bogor

84 m 47 m 40 m 265 m
°C 25.2 2915 mm °C 26.7 2102 mm °C 27.5 2540 mm °C 25.0 4230 mm
40 500 40 500 40 500 40 500
30 400 30 400 30 400 30 400
300 300 300 300
20 20 20 20
200 200 200 200
10 100 10 100 10 100 10 100
0 0 0 0 0 0 0 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

of North America, the steppes of Eurasia, and the pampas of emphasize the unique physical and biological characteristics de­
Argentina. Moving poleward, the temperate-deciduous forests fining these broad categories of terrestrial ecosystems (biomes).
give way to the needle-leaf–dominated forests of the boreal
region (conifer forest or taiga). As temperatures become more
extreme and the growing season shorter, trees can no longer be 23.2  Tropical Forests Characterize
supported, and the short-stature shrubs and sedges (grasslike
plants of the family Cyperaceae) characteristic of the tundra the Equatorial Zone
dominate the landscape ecosystems of the arctic region. The tropical rain forests are restricted primarily to the equato­
In the following sections, we will examine the eight major rial zone between latitudes 10° N and 10° S (Figure 23.5),
categories of terrestrial biomes outlined in Figure 23.2. We begin where the temperatures are warm throughout the year and
each section by relating their geographic distribution to the broad- rainfall occurs almost daily. The largest and most continuous
scale constraints of regional climate, as outlined in Figure 23.2, region of rain forest in the world is in the Amazon basin of
as well as to associated patterns of seasonality in temperature South America (Figure 23.6). The second largest is located in
and precipitation (see Quantifying Ecology 23.1) that func­ Southeast Asia, and the third largest is in West Africa around
tion as constraints on the dominant plant life-forms and patterns the Gulf of Guinea and in the Congo basin. Smaller rain forests
of primary and secondary productivity. In our discussion, we occur along the northeastern coast of Australia, the windward

Figure 23.6  Tropical rain forests in (a) Amazon Basin (South America) and (b) Malaysia
(Southeast Asia). Despite being taxonomically distinct, these two tropical rain forest regions are
dominated by broadleaf evergreen trees and support vigorous plant growth year-round. Tropical
rain forests represent the most diverse and productive terrestrial ecosystems on our planet.
(a) (b)

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  C h ap t er 23   •  Terrestrial Ecosystems    531

Q u a n t if y in g Ec ol og y 23. 1   Climate Diagrams

A s illustrated in Figure 23.2, the distribution of terrestrial

biomes is closely related to climate. The measures of
regional climate used in Whittaker’s graph (Figure 23.2) are
around the world where a particular biome type is found. See
Figure 1 for a representative climate diagram, which we have
labeled to help you interpret the information it presents. As
mean annual temperature and precipitation. Yet, as we will see you study the diagram, take particular note of the patterns of
in the discussion of the various biome types, the distribution of seasonality.
terrestrial ecosystems is influenced by other aspects of climate
as well, namely seasonality of both temperature and precipita- 1. In Figure 23.12, what is the distinctive feature of the cli-
tion. Topographic features such as mountains and valleys also mate diagrams for these tropical savanna ecosystems?
influence the climate of a region. How do the patterns differ between sites in the Northern
To help understand the relationship between regional and Southern Hemispheres? What feature of Earth’s cli-
climate and the distribution of terrestrial ecosystems, for mate system discussed in Chapter 2 is responsible for
each biome discussed in this chapter (tropical forest, sa- these distinctive patterns?
vanna, etc.), we present a map showing its global distribution.
Accompanying the map is a series of climate diagrams. The 2. In Figure 23.23, what feature of the climate is common to
diagrams describe the local climate at representative locations all mediterranean ecosystems?

Location of site: Elevation above sea

Bulawayo, Zimbabwe level in meters

Mean annual Bulawayo Mean annual

temperature (°C) 1343 m
precipitation (mm)
°C 19.2 581 mm
300
40
Figure 1  Climate diagram for Bulawayo, 30 200
Zimbabwe. This city is in the Southern Hemisphere, Mean monthly 20 Precipitation (mm)
100
temperature (°C) 10
where the cooler winter season occurs during the
0 0
period of May–August. Note the distinct dry season J FMAMJ J ASOND

during the winter months, with the rainy season Mean monthly
Temperature (°C)
beginning in October (spring) and lasting through precipitation (mm)
Month
the summer months.

side of the Hawaiian Islands, the South Pacific Islands, the east above 60 millimeters (mm; see climate diagrams for representa­
coast of Madagascar, northern South America, and southern tive tropical rain forest sites in Figure 23.5). Within the lowland
Central America. forest zone, mean annual temperatures typically exceed 25°C
The climate of tropical rain forest regions varies geograph­ with an annual range less than 5°C.
ically but is typically characterized by a mean temperature of Tropical rain forests have a high diversity of plant and ani­
all months exceeding 18°C and minimum monthly precipitation mal life. Covering only 6 percent of the land surface, tropical

(a) (b) Figure 23.7  Examples of primate

species that inhabit the tropical
rain forests of the world: (a) the
chimpanzee (Pan troglodytes) inhabits
the tropical rain forests of Central
Africa, and (b) the orangutan (Pongo
pygmaeus) inhabits the tropical rain
forests of Borneo (Southeast Asia).

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 532    Part se ve n   •  Eco l o g ical B io g e og r a p h y

40
Emergent canopy
(trees widely spaced)
35

30
Height aboveground (m)

Upper canopy 25
(medium-spaced
crowns)
20

15

Lower canopy 10

5
Understory
(shrubs and saplings)
Ground cover 0
(herbs and ferns)

Figure 23.8  Vertical stratification of a tropical rain forest.

rain forests account for more than 50 percent of all known The continually warm, moist conditions in rain forests
plant and animal species. Tree species number in the thousands. promote strong chemical weathering and rapid leaching of
A 10-km2 area of tropical rain forest may contain 1500 species soluble materials. The characteristic soils are oxisols, which
of flowering plants and up to 750 species of trees. The rich­ are deeply weathered with no distinct horizons (see Chapter 4
est area is the lowland tropical forest of peninsular Malaysia, for discussion and classification of soils). Ultisols may develop
which contains some 7900 species. in areas with more seasonal precipitation regimes and are typi­
Nearly 90 percent of all nonhuman primate specie live in the cally associated with forested regions that exhibit seasonal soil
tropical rain forests of the world (Figure 23.7). Sixty-four spe­ moisture deficits. Areas of volcanic activity in parts of Central
cies of New World primates—small mammals with prehensile and Southeast Asia, where recent ash deposits quickly weather,
tails—live in the trees. The Indo-Malaysian forests are inhabited are characterized by andosols (see Figure 4.12).
by a number of primates, many with a limited distribution within The warmer, wetter conditions of the tropical rain forest
the region. The orangutan, an arboreal ape, is confined to the result in high rates of net primary productivity and subsequent
island of Borneo. Peninsular Malaysia has seven species of pri­ high annual rates of litter input to the forest floor. Little litter
mates, including three gibbons, two langurs, and two macaques. accumulates, however, because decomposers consume the dead
The long-tailed macaque is common in disturbed or secondary
forests, and the pig-tailed macaque is a terrestrial species adapt­
Figure 23.9  Plank-like buttresses help to support tall rain
able to human settlements. The tropical rain forest of Africa is
forest trees.
home to mountain gorillas and chimpanzees. The diminished
rain forest of Madagascar holds 39 species of lemurs (see
Chapter 9, Ecological Issues & Applications and Figure 9.20).
Tropical rain forests may be divided into five vertical lay­
ers (Figure 23.8): emergent trees, upper canopy, lower canopy,
shrub understory, and a ground layer of herbs and ferns.
Conspicuous in the rain forest are lianas—climbing vines—
growing upward into the canopy, epiphytes growing on the
trunks and branches, and strangler figs (Ficus spp.) that grow
downward from the canopy to the ground. Many large trees
develop plank-like outgrowths called buttresses (Figure 23.9).
They function as prop roots to support trees rooted in shallow
soil that offers poor anchorage. The floor of a tropical rain for­
est is thickly laced with roots, both large and small, forming a
dense mat on the ground.

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  C h ap t er 23   •  Terrestrial Ecosystems    533

dry tropical forest. Much of the original forest, especially in

Central America and India, has been converted to agricultural
and grazing land.

23.3  Tropical Savannas Are

Characteristic of Semiarid Regions
with Seasonal Rainfall
(a) The term savanna was originally used to describe the treeless
areas of South America. Now it is generally applied to a range
of vegetation types in the drier tropics and subtropics char­
acterized by a ground cover of grasses with scattered shrubs
or trees. Savanna includes an array of vegetation types repre­
senting a continuum of increasing cover of woody vegetation,
from open grassland to widely spaced shrubs or trees and to
woodland (Figure 23.11). In South America, the more densely
wooded areas are referred to as cerrado. The campos and llano
are characterized by a more open appearance (lower density of
trees), and thorn scrub is the dominant cover of the caatinga.
In Africa, the miombo, mopane, and Acacia woodlands can
be distinguished from the more open and park-like bushveld.
Scattered individuals of Acacia and Eucalyptus dominate the
mulga and brigalow of Australia.
The physiognomic diversity of the savanna vegetation re­
flects the different climate conditions occurring throughout this
(b)

Figure 23.10  A tropical dry forest in Costa Rica during the

(a) rainy and (b) dry season. Most of the tropical dry forests Figure 23.11  Savanna ecosystems, such as the (a) cerrano of
in Central America have disappeared from land clearing for South America and (b) mulga woodlands of central Australia are
agriculture. characterized by a ground cover of grasses with scattered shrubs
or trees.
(a)
organic matter almost as rapidly as it falls to the forest floor.
Most of the nutrients available for uptake by plants are a result
of the rapidly decomposed organic matter that is continuously
falling to the soil surface. Growing plants, however, rapidly
absorb these nutrients. The average time for leaf litter to de­
compose is 24 weeks.
Moving from the equatorial zone to the regions of the
tropics that are characterized by greater seasonality in precipi­
tation, the broadleaf evergreen forests are replaced by the dry
tropical forests (Figure 23.10). Dry tropical forests undergo a
dry season whose length is based on latitude. The more distant
the f­ orest is from the equator, the longer is the dry season—in
some areas, up to eight months. During the dry season, the
drought-deciduous trees and shrubs drop their leaves. Before
the start of the rainy season, which may be much wetter than (b)
the wettest time in the rain forest, the trees begin to leaf. During
the wet season, the landscape becomes uniformly green.
The largest proportion of tropical dry forest is found in
Africa and South America, to the south of the zones dominated
by rain forest. These regions are influenced by the seasonal mi­
gration of the Intertropical Convergence Zone (see Section 2.6,
Figure 2.18). In addition, areas of Central America, northern
Australia, India, and Southeast Asia are also classified as

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 534    Part se ve n   •  Eco l o g ical B io g e og r a p h y

San Fernando Kano Nyala Gulbarga Figure 23.12  Geographic

73 m 472 m 675 m 458 m
°C 27.4 1432 mm °C 26.2 866 mm 0° °C 26.9 495 mm °C 27.2 753 mm distribution of Earth’s tropical
300 300 300 300
40 40 40 40 savanna ecosystems and associated
30 200 30 200 30 200 30 200 climate diagrams showing long-term
20 20 20 20
100 100 100 100 patterns of monthly temperature
10 10 10 10
0 0 0 0 0 0 0 0 and precipitation for selected
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
locations. Seasonal patterns of
temperature are similar to those of
the tropical forest sites presented in
40° 40° Figure 24.5, reflecting the tropical
and subtropical climates at these
20° 20° sites. Note, however, the distinct
seasonality of precipitation that
0° 0° characterizes these ecosystems
(also note the shift in timing of rainy
20° 20° season for Northern and Southern
Hemisphere locations, reflecting the
seasonal shift of the Intertropical
40° 40°
Convergence Zone).
(Adapted from Archibold 1995.)
Barra Ndola Bulawayo Daly Waters
408 m 1269 m 1343 m 210 m
°C 26.3 684 mm °C 19.7 1168 mm °C 19.2 581 mm °C 26.9 628 mm
300 300 300 300
40 40 40 40
30 200 30 200 30 200 30 200
20 20 20 20
100 100 100 100
10 10 10 10
0 0 0 0 0 0 0 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

widely distributed ecosystem (Figure 23.12). Moisture appears (year to year) variation in total precipitation (see climate
to control the density of woody vegetation, a function of both diagrams for representative savanna sites in Figure 23.12).
rainfall (amount and distribution) and soil—its texture, structure, Mean monthly temperatures typically do not fall below 18°C,
and water-holding capacity (Figure 23.13; also see Chapter 4). although during the coldest months in highland areas, tem­
Savannas are associated with a warm continental climate peratures can be considerably lower. There is seasonality in
with distinct seasonality in precipitation and a large interannual temperatures, and maximum temperatures occur at the end of

Figure 23.13  Diagram

showing the interaction between
annual precipitation and soil
texture in defining the transition
from woodland to savanna and
grassland in southern Africa.
Plants have more limited access
to soil moisture on the heavily
textured soils (clays) than on
the coarser sands, so annually
more precipitation is needed to
support the woody plants.

Clay

Grassland
Soil texture

Savanna

Woodland
to forest
Sand
200 600 1000
Rainfall (mm)

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  C h ap t er 23   •  Terrestrial Ecosystems    535

the wet season. The nature of the vegetation cover, however, diverse ungulate fauna of at least 60 species that share the vegeta­
is more closely determined by the amount and seasonality of tive resources. Some species, such as the wildebeest and zebra,
precipitation than by temperature. are migratory during the dry season (see Figure 7.10 for example).
Savannas, despite their differences in vegetation, exhibit a Savanna vegetation supports an incredible number of
certain set of characteristics. Savannas occur on land surfaces insects: flies, grasshoppers, locusts, crickets, carabid beetles,
of little relief—often on old plateaus, interrupted by escarp­ ants, and detritus-feeding dung beetles and termites. Mound-
ments and dissected by rivers. Continuous weathering in these building termites excavate and move tons of soil, mixing min­
regions has produced nutrient-poor oxisols, which are particu­ eral soil with organic matter. Some species construct extensive
larly deficient in phosphorus. Alfisols are common in the drier subterranean galleries and others accumulate organic matter.
savannas, whereas entisols are associated with the driest savan­ Preying on the ungulate fauna is an array of carnivores
nas (see Figure 4.12). Subject to recurrent fires, the dominant including the lion, leopard, cheetah, hyena, and wild dog.
vegetation is fire adapted. Grass cover with or without woody Scavengers, including vultures and jackals, subsist on the re­
vegetation is always present, and the woody component is mains of prey killed by carnivores.
short-lived—individuals seldom survive for more than several
decades. Savannas are characterized by a two-layer vertical
structure because of the ground cover of grasses and the pres­ 23.4  Grassland Ecosystems of the
ence of shrubs or trees (see Figure 16.12b). Temperate Zone Vary with Climate
The yearly cycle of plant activity and subsequent produc­
tivity in tropical savannas is largely controlled by the markedly and Geography
seasonal precipitation and corresponding changes in available Natural grasslands occupy regions where rainfall is between
soil moisture. Most leaf litter is decomposed during the wet 25 and 80 centimeters (cm) a year, but they are not exclusively
season, and most woody debris is consumed by termites during climatic. Many exist through the intervention of fire and hu­
the dry season. man activity. Conversions of forests into agricultural lands and
The microenvironments associated with tree canopies can the planting of hay and pasturelands extended grasslands into
influence species distribution, productivity, and soil character­ once forested regions. Formerly covering about 42 percent of
istics. Stem flow and associated litter accumulation result in the land surface of Earth, natural grasslands have shrunk to less
higher soil nutrients and moisture under tree canopies, often than 12 percent of their original size because of conversion to
encouraging increased productivity and the establishment of cropland and grazing lands.
species adapted to the more shaded environments. The natural grasslands of the world occur in the midlati­
Savannas can support a large and varied assemblage of tudes in midcontinental regions, where annual precipitation de­
­herbivores—invertebrate and vertebrate, grazing and browsing. clines as air masses move inward from the coastal environments
The African savanna, visually at least, is dominated by a large and (Figure 23.14; see Section 2.7 for discussion of continental

Saskatoon Lincoln Mariupol Semipalatinsk Figure 23.14  Geographic

476 m 375 m 73 m 206 m
°C 2.0 352 mm °C 11.6 698 mm 0° °C 8.5 445 mm °C 3.2 265 mm distribution of Earth’s temperate
30 120 30 120 30 120 30 120
20 100 20 100 20 100 20 100 grassland ecosystems and associated
10 80 10 80 10 80 10 80 climate diagrams showing long-term
60 60 60 60
0 40 0 40 0 40 0 40 patterns of monthly temperature and
-10 20 -10 20 -10 20 -10 20
-20 0 -20 0 -20 0 -20 0 precipitation for selected locations.
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
Most of the major grassland regions
are mid-continental with a distinct
seasonality in temperature. Mean
40° 40° annual precipitation is typically well
below 1000 mm, comparable in
20° 20° range to that observed for tropical
and subtropical savannas but less
0° 0° than that of temperate deciduous
forests (compare with Figure 24.27).
20° 20° (Adapted from Archibold 1995.)

40° 40°

Macachin Kroonstad Changan Ang’angxi

140 m 1348 m 395 m 149 m
°C 15.3 602 mm °C 16.4 606 mm °C 15.8 510 mm °C 2.7 363 mm
30 120 30 120 30 120 30 120
20 100 20 100 20 100 20 100
10 80 10 80 10 80 10 80
60 60 60 60
0 40 0 40 0 40 0 40
-10 20 -10 20 -10 20 -10 20
-20 0 -20 0 -20 0 -20 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

M23_SMIT7406_09_GE_C23.indd 535 30/01/15 1:48 PM

 536    Part se ve n   •  Eco l o g ical B io g e og r a p h y

patterns of precipitation). In the Northern Hemisphere, these

regions include the prairies of North America and the steppes
of central Eurasia. In the Southern Hemisphere, grasslands are
represented by the pampas of Argentina and the veld of the
high plateaus of southern Africa. Smaller areas occur in south­
eastern Australia and the drier parts of New Zealand.
The temperate grassland climate is one of recurring
drought, and much of the diversity of vegetation cover reflects
differences in the amount and reliability of precipitation.
Grasslands do the least well where precipitation is lowest and
the temperatures are high. They are tallest in stature and the
most productive where mean annual precipitation is greater
than 800 mm and mean annual temperature is above 15°C.
Thus, native grasslands of North America, influenced by de­
clining precipitation from east to west, consist of three main
types distinguished by the height of the dominant species:
tallgrass, mixed-grass, and shortgrass prairie (Figure  23.15).
Tallgrass prairie (Figure  23.16a) is dominated by big (a)
bluestem (Andropogon gerardi), growing 1  meter (m) tall
with flowering stalks 1 to 3.5 m tall. Mixed-grass prairie
(Figure  23.16b), typical of the Great Plains, is composed
largely of needlegrass–grama grass (Bouteloua–Stipa). South
and west of the mixed prairie and grading into the desert re­
gions is the shortgrass prairie (Figure 23.16c), dominated by
sod-forming blue grama (Bouteloua gracilis) and buffalo grass
(Buchloe dactyloides), which has remained somewhat intact,
and desert grasslands. From southeastern Texas to southern
Arizona and south into Mexico lies the desert grassland,

(b)

Shortgrass prairie
Mixed-grass prairie
Tallgrass prairie
Flint Hills (c)

Figure 23.15  Map showing the original extent of shortgrass, Figure 23.16  North American grasslands. (a) A remnant
mixed-grass, and tallgrass prairies in North America before the tallgrass prairie in Iowa; (b) the mixed-grass prairie has been
arrival of Europeans. called “daisyland” for the diversity of its wildflowers; (c) shortgrass
(After Reichman 1987.) steppe in western Wyoming.

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  C h ap t er 23   •  Terrestrial Ecosystems    537

similar in many respects to the shortgrass plains, except The great ungulate herds have been destroyed and replaced
that three-awn grass (Aristida spp.) replaces buffalo grass. with sheep, cattle, and horses.
Confined largely to the Central Valley of California is annual Many forms of Australian marsupial mammals evolved
grassland. It is associated with a mediterranean climate (see that are the ecological equivalents of placental grassland mam­
Section 23.6) characterized by rainy winters and hot, dry sum­ mals. The dominant grazing animals are several kangaroo
mers. Growth occurs during early spring, and most plants are species, especially the red kangaroo (Macropus rufus) and the
dormant in summer, turning the hills a dry tan color accented gray kangaroo (Macropus giganteus).
by the deep green foliage of scattered California oaks. Grasslands evolved under the selective pressure of graz­
At one time, the great grasslands of the Eurasian conti­ ing. Thus, up to a point, grazing stimulates primary production.
nent extended from eastern Europe to western Siberia south Although the most conspicuous grazers are large herbivores,
to Kazakhstan. These steppes, treeless, except for ribbons and the major consumers in grassland ecosystems are invertebrates.
patches of forest, are divided into four belts of latitude, from The heaviest consumption takes place belowground, where the
the mesic meadow steppes in the north to semiarid grasslands dominant herbivores are nematodes.
in the south. The most visible feature of grassland is the tall, green,
In the Southern Hemisphere, the major grasslands exist in ephemeral herbaceous growth that develops in spring and dies
southern Africa and southern South America. Known as pam- back in autumn. One of the three strata in the grassland, it
pas, the South American grasslands extend westward in a large arises from the crowns, nodes, and rosettes of plants hugging
semicircle from Buenos Aires and cover about 15 percent of the soil. The ground layer and the belowground root layer are
Argentina. These pampas have been modified by the introduc­ the other two major strata of grasslands. The highly developed
tion of European forage grasses and alfalfa (Medicago sativa), root layer can make up more than half the total plant biomass
and the eastern tallgrass pampas have been converted to wheat and typically extends fairly deep into the soil.
and corn. In Patagonia, where annual rainfall averages about Depending on their history of fire and degree of grazing
25 cm, the pampas change to open steppe. and mowing, grasslands accumulate a layer of mulch that
The velds of southern Africa (not to be confused with retains moisture and, with continuous turnover of fine roots,
­savanna) occupy the eastern part of a high plateau 1500 to 2000 m adds organic matter to the mineral soil. Dominant soils of the
above sea level in the Transvaal and the Orange Free State. grasslands are mollisols with a relatively thick, dark-brown
Australia has four types of grasslands: arid tussock grass­ to black surface horizon that is rich in organic matter (see
land in the northern part of the continent, where the rainfall Figure  4.12). Soils typically become thinner and paler in the
averages between 20 and 50 cm, mostly in the summer; arid drier regions because less organic material is incorporated into
hummock grasslands in areas with less than 20 cm rainfall; the surface horizon.
coastal grasslands in the tropical summer rainfall region; and
subhumid grasslands along coastal areas where annual rainfall Figure 23.17  North American grasslands were once
is between 50 and 100 cm. However, the introduction of fer­ dominated by (a) large grazing ungulates such as bison and (b)
tilizers, nonnative grasses, legumes, and sheep grazing have burrowing mammals such as the prairie dog.
changed most of these grasslands.
(a)
Grasslands support a diversity of animal life dominated
by herbivorous species, both invertebrate and vertebrate. Large
grazing ungulates and burrowing mammals are the most con­
spicuous vertebrates (Figure 23.17). The North American
grasslands were once dominated by huge migratory herds of
millions of bison (Bison bison) and the forb-consuming prong­
horn antelope (Antilocarpa americana). The most common
burrowing rodent was the prairie dog (Cynomys spp.), which
along with gophers (Thomomys and Geomys spp.) and the
mound-building harvester ants (Pogonomyrex spp.), appeared
to be instrumental in developing and maintaining the ecologi­
cal structure of the shortgrass prairie.
The Eurasian steppes and the Argentine pampas lack
herds of large ungulates. On the pampas, the two major large (b)
herbivores are the pampas deer (Ozotoceros bezoarticus), and,
farther south, the guanaco (Lama guanicoe), a small relative of
the camel. These species, however, are greatly reduced in num­
ber compared with the past.
The African grassveld once supported great migratory
herds of wildebeest (Connochaetes taurinus) and zebra (Equus
spp.) along with the associated carnivores, the lion (Panthera
leo), leopard (Panthera pardus), and hyena (Crocuta crocuta).

M23_SMIT7406_09_GE_C23.indd 537 30/01/15 1:48 PM

 538    Part se ve n   •  Eco l o g ical B io g e og r a p h y

1500 along the Intertropical Convergence Zone subsides to form the

semipermanent high-pressure cells that dominate the climate of
Aboveground NPP

1000 tropical deserts (see Figure 2.17). The warming of the air as it
(g/m2/yr)

descends in addition to cloudless skies result in intense radia­

tion heat during the summer months.
500 Temperate deserts lie in the rain shadow of mountain bar­
riers or are located far inland, where moist maritime air rarely
penetrates. Here, temperatures are high during the summer but
0 500 1000 1500 2000
can drop to below freezing during the winter months. Thus, the
Mean annual precipitation (mm)
lack of precipitation, rather than continually high temperature,
Figure 23.18  Relationship between aboveground net primary is the distinctive characteristic of all deserts.
production (NPP) and mean annual precipitation for 52 grassland Most of the arid environments are found in the Northern
sites around the world. Each point represents a different grassland Hemisphere. The Sahara, the world’s largest desert, covers
site. North American grasslands are indicated by dark-green dots. approximately 9 million km2 of North Africa. It extends the
(Adapted from Lauenroth 1979.) breadth of the African continent to the deserts of the Arabian
Peninsula, continuing eastward to Afghanistan and Pakistan
and finally terminating in the Thar Desert of northwest India.
The productivity of temperate grassland ecosystems is
The temperate deserts of Central Asia lie to the north. The
primarily related to annual precipitation (Figure 23.18), yet
most westerly of these is the Kara Kum desert region of
temperature can complicate this relationship. Increasing tem­
Turkmenistan. Eastward lie the high-elevation deserts of west­
peratures have a positive effect on photosynthesis but can actu­
ern China and the high plateau of the Gobi Desert.
ally reduce productivity by increasing the demand for water.
A similar transition to temperate desert occurs in western
North America. Here, the Sierra Nevada effectively blocks
23.5  Deserts Represent a Diverse the passage of moist air into the interior of the Southwest.
Mountain ranges run parallel to the Sierras throughout the
Group of Ecosystems northern part of this region, and desert basins occur on the east­
The arid regions of the world occupy from 25 to 35 percent of ern sides of these ranges.
Earth’s landmass (Figure 23.19). The wide range reflects the Apart from the drier parts of southern Argentina, the des­
various approaches used to define desert ecosystems based on erts of the Southern Hemisphere all lie within the subtropical
climate conditions and vegetation types. Much of this land lies high-pressure belt that mirrors that of the Northern Hemisphere
between 15° and 30° latitude, where the air that is carried aloft (see preceding discussion). Cold ocean currents also contribute

Barstow Riyadh Termez Jacobabad Figure 23.19  Geographic

653 m 624 m 302 m 56 m
°C 17.8 108 mm °C 26.3 105 mm 0° °C 17.4 133 mm °C 27.1 88 mm distribution of Earth’s desert (arid)
40 40 40 40 40 40 40 40
ecosystems and associated climate
30 30 30 30 30 30 30 30
diagrams showing long-term
20 20 20 20 20 20 20 20
10 10 10 10 10 10 10 10
patterns of monthly temperature
0 0 0 0 0 0 0 0
and precipitation for selected
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
locations. Seasonal variations in
temperature differ among these
representative sites; but at all sites,
40° 40° annual precipitation is well below
the potential evaporative demand,
20° 20° resulting in a low level of soil
moisture available to provide for
0° 0° primary productivity.
(Adapted from Archibold 1995.)
20° 20°

40° 40°

Antofagasta Alexander Bay Bilma William Creek

849 m 21 m 359 m 76 m
°C 16.6 4 mm °C 18.5 52 mm °C 26.8 23 mm °C 20.3 127 mm
40 40 40 40 40 40 40 40
30 30 30 30 30 30 30 30
20 20 20 20 20 20 20 20
10 10 10 10 10 10 10 10
0 0 0 0 0 0 0 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

M23_SMIT7406_09_GE_C23.indd 538 30/01/15 1:48 PM

  C h ap t er 23   •  Terrestrial Ecosystems    539

Cool deserts—including the Great Basin of North

America, the Gobi, Takla Makan, and Turkestan deserts of
Asia—and high elevations of hot deserts are dominated by
Artemisia and chenopod shrubs (Figure 23.20). They may be
considered shrub steppes or desert scrub. In the Great Basin
of North America, the northern, cool, arid region lying west
of the Rocky Mountains is the northern desert scrub. The cli­
mate is continental, with warm summers and prolonged cold
winters. The vegetation falls into two main associations: one
is sagebrush, dominated by Artemisia tridentata, which often
(a) forms pure stands; the other is shadscale (Atriplex confertifo-
lia), a C4 species, and other chenopods (halophytes—tolerant
of saline soils).
A similar type of desert scrub exists in the semiarid inland
of southwestern Australia. Many chenopod species, particu­
larly the saltbushes of the genera Atriplex and Maireana, form
extensive low shrublands on low riverine plains.
The hot deserts range from those lacking vegetation to ones
with some combination of chenopods, dwarf shrubs, and suc­
culents (Figure 23.21). Creosote bush (Larrea divaricata) and
bur sage (Franseria spp.) dominate the deserts of southwestern
North America—the Mojave, the Sonoran, and the Chihuahuan.
Areas of favorable moisture support tall growths of Acacia spp.,
saguaro (Cereus giganteus), palo verde (Cercidium spp.), ocoti­
llo (Fouquieria spp.), yucca (Yucca spp.), and ephemeral plants.

Figure 23.21  Two examples of hot deserts. (a) The Chihuahuan

(b) Desert in Nuevo Leon, Mexico. The substrate of this desert is sand-
sized particles of gypsum. (b) Dunes in the Saudi Arabian desert
Figure 23.20  Two examples of desert scrub. (a) Northern near Riyadh. Note the extreme sparseness of vegetation.
desert shrubland in Wyoming is dominated by sagebrush
(a)
(Artemisia). Although classified as a cool desert plant, sagebrush
forms one of the most important shrub types in North America.
(b) Saltbrush shrubland in Victoria, Australia, is dominated by
Atriplex and is an ecological equivalent of the Great Basin
shrublands in North America.

to the development of arid coastal regions (see Section 2.4).

Drought conditions are severe along a narrow strip of the coast
that includes Chile and Peru. The drier parts of Argentina lie in
the rain shadow of the Andes.
The deserts of southern Africa include three regions. The
Namib Desert occupies a narrow strip of land that runs along
the west coast of Africa from southern Angola to the border of
the cape region of South Africa. This region continues south
and east across South Africa as the Karoo, which merges with
the Kalahari Desert to the north in Botswana. The most exten­ (b)
sive region of arid land in the Southern Hemisphere is found in
Australia, where more than 40 percent of the land is classified
as desert.
Deserts are not the same everywhere. Differences in mois­
ture, temperature, soil drainage, topography, alkalinity, and
salinity create variations in vegetation cover, dominant plants,
and groups of associated species. There are hot deserts and cold
deserts, extreme deserts and semideserts, ones with enough
moisture to verge on being grasslands or shrublands, and grada­
tions between those extremes within continental deserts.

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 540    Part se ve n   •  Eco l o g ical B io g e og r a p h y

The infrequent rainfall coupled with high rates of evapora­

tion limit the availability of water to plants, so primary produc­
tivity is low. Most desert soils are poorly developed aridisols
and entisols, and the sparse cover of arid lands limits the abil­
ity of vegetation to heavily modify the soil environment (see
Figure 4.12). Underneath established plants, however, “islands
of fertility” can develop because of higher litter input and the
enrichment by wastes from animals that seek shade, particu­
larly under shrubs.

23.6  Mediterranean Climates

Support Temperate Shrublands
Figure 23.22  A spadefoot toad, named for the black, sharp- Shrublands—plant communities where the shrub growth form is
edged “spades” on its hind feet, emerges from its desert burrow either dominant or codominant—are difficult types of ecosystems
to breed when the rains come. to categorize, largely because of the difficulty in characterizing
the term shrub itself. In general, a shrub is a plant with multiple
Both plants and animals adapt to the scarcity of water by woody, persistent stems but no central trunk and a height from
either drought evasion or drought resistance. Drought-evading 4.5 to 8 m. However, under severe environmental conditions,
plants flower only when moisture is present. They persist as even many trees do not exceed that size. Some trees—particularly
seeds during drought periods, ready to sprout, flower, and individuals that coppice (resprout from the stump) after destruc­
produce seeds when moisture and temperature are favorable. tion of the aboveground tissues by fire, browsing, or cutting—are
If no rains come, these ephemeral species do not germinate multistemmed, and some shrubs can have large, single stems. In
and grow. addition, the shrub growth form can be a dominant component
Drought-evading animals, like their plant counterparts, of a variety of tropical and temperate ecosystems, including the
adopt an annual cycle of activities or go into estivation or some tropical savannas and scrub desert communities (see Section 23.3,
other dormant stage during the dry season. For example, the respectively). However, in five widely disjunct regions along the
spadefoot toad (Scaphiopus; Figure 23.22) remains under­ western margins of the continents, between 30° and 40° latitude,
ground in a gel-lined underground cell, making brief reproduc­ are found the mediterranean ecosystems dominated by evergreen
tive appearances during periods of winter and summer rains. shrubs and sclerophyllous trees that have adapted to the distinc­
If extreme drought develops during the breeding season, many tive climate of summer drought and cool, moist winters.
animals such as lizards and birds do not reproduce. The five regions of mediterranean ecosystems include
Desert plants may be deep-rooted woody shrubs, such as the semiarid regions of western North America, the regions
mesquite (Prosopis spp.) and Tamarix, whose taproots reach bordering the Mediterranean Sea, central Chile, the cape re­
the water table, rendering them independent of water supplied gion of South Africa, and southwestern and southern Australia
by rainfall. Some plants, such as Larrea and Atriplex, are deep- (Figure 23.23). The mediterranean climate has hot, dry sum­
rooted perennials with superficial laterals that extend as far as mers, with at least one month of protracted drought, and
15 to 30 m from the stems. Other perennials, such as the vari­ cool, moist winters (see representative climate diagrams in
ous species of cactus, have shallow roots that often extend no Figure  23.23). About 65 percent of the annual precipitation
more than a few centimeters below the surface. falls during the winter months. Winter temperatures typically
Despite their aridity, desert ecosystems support a surpris­ average 10–12°C with a risk of frost. The hot, dry summer
ing diversity of animal life, including a wide assortment of climates of the mediterranean regions arise from the seasonal
beetles, ants, locusts, lizards, snakes, birds, and mammals. change in the semipermanent high-pressure zones that are
The mammals are mostly herbivorous species. Grazing herbi­ centered over the tropical deserts at about 20° N and 20° S (see
vores of the desert tend to be generalists and opportunists in discussion in Section 23.5). The persistent flow of dry air out
their mode of feeding. They consume a wide range of species, of these regions during the summer brings several months of
plant types, and parts. Desert rodents—particularly the family hot, dry weather. Fire is a frequent hazard during these periods.
Heteromyidae—and ants feed largely on seeds and are impor­ All five regions support similar-looking communities of
tant in the dynamics of desert ecosystems. Seed-eating her­ xeric broadleaf evergreen shrubs and dwarf trees known as
bivores can eat up to 90 percent of the available seeds. That sclerophyllous (scleros, “hard”; phyll, “leaf”) vegetation with
consumption can distinctly affect plant composition and plant a herbaceous understory. Sclerophyllous vegetation possesses
populations. Desert carnivores, such as foxes and coyotes, small leaves, thickened cuticles, glandular hairs, and sunken
have mixed diets that include leaves and fruits; even insec­ stomata—all characteristics that function to reduce water loss
tivorous birds and rodents eat some plant material. Omnivory, during the hot, dry summer period (Figure 23.24). Vegetation
rather than carnivory and complex food webs, seems to be the in each of the mediterranean systems also shares adaptations to
rule in desert ecosystems. fire and to low nutrient levels in the soil.

M23_SMIT7406_09_GE_C23.indd 540 30/01/15 1:48 PM

  C h ap t er 23   •  Terrestrial Ecosystems    541

Rabat Sevilla Athens Adana Figure 23.23  Geographic

68 m 30 m 107 m 21 m
°C 17.3 523 mm °C 18.9 573 mm 0° °C 17.8 402 mm °C 18.8 619 mm distribution of Earth’s mediterranean
30 200 30 200 30 200 30 200
ecosystems and associated climate
150 150 150 150
20 20 20 20 diagrams showing long-term
100 100 100 100
10
50
10
50
10
50
10
50
patterns of monthly temperature and
0 0 0 0 0 0 0 0 precipitation for selected locations.
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
Note that these ecosystems are
characterized by a winter rainy
season and dry summers.
40° 40°
(Adapted from Archibold 1995.)

20° 20°

0° 0°

20° 20°

40° 40°

Oakland Talca Cape Town Esperance

134 m 122 m 12 m 4m
°C 13.9 802 mm °C 14.8 700 mm °C 17.4 627 mm °C 16.3 679 mm
30 200 30 200 30 200 30 200
150 150 150 150
20 20 20 20
100 100 100 100
10 10 10 10
50 50 50 50
0 0 0 0 0 0 0 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

The largest area of mediterranean ecosystem forms a discon­ as mallee is dominated by low-growing Eucalyptus, 5 to 8 m in
tinuous belt around the Mediterranean Sea in southern Europe height, with broad sclerophyllous leaves.
and North Africa. Much of the area is currently or was once dom­ In North America, the sclerophyllous shrub community is
inated by mixed evergreen woodland supporting species such as known as chaparral, a word of Spanish origin meaning a thicket
holm oak (Quercus ilex) and cork oak (Quercus suber). Often, of shrubby evergreen oaks (Figure 23.26). California chaparral,
these two species grow in mixed stands in association with dominated by scrub oak (Quercus berberidifolia) and chamise
strawberry tree (Arbutus unedo) and various species of shrubs. (Adenostoma fasciculatum), is evergreen, winter-active, and
The easternmost limit of these ecosystems is in the coastal areas summer-dormant. Another shrub type, also designated as chap­
of Syria, Lebanon, and Israel, where they grade into the arid arral, is found in the Rocky Mountain foothills. Dominated by
lands of the Middle East. Here, deciduous oak species are more Gambel oak (Quercus gambelii), it is winter-deciduous.
abundant. Desert vegetation extends across North Africa as far The matorral shrub communities of central Chile occur
as Tunisia, with mediterranean shrub and woodland extending in the coastal lowlands and on the west-facing slopes of the
through the northern coastal areas of Algeria and Morocco. Andes. Most of the matorral species are evergreen shrubs
The mediterranean zone in southern Africa is restricted to 1 to 3 m in height with small sclerophyllous leaves, although
the mountainous region of the Cape Province, where the vegeta­ drought-deciduous shrubs are also found.
tion is known as fynbos. The vegetation is composed primar­ For the most part, mediterranean shrublands lack an un­
ily of broadleaf proteoid (Proteaceae) and ericoid (Ericaceae) derstory and ground litter and are highly flammable. Many spe­
shrubs that grow to a height of 1.5 to 2.5 m (Figure 23.25). In cies have seeds that require the heat and scarring action of fire
southwest Australia, the mediterranean shrub community known to induce germination. Without fire, chaparral grows taller and

Figure 23.24 
Sclerophyllous leaves
of some tree and shrub
species inhabiting
mediterranean shrublands
(chaparral) of California:
(a) chamise (Adenostoma
fasciculatum), (b) scrub oak
(Quercus dumosa), and
(c) chinquapin (Chrysolepis
(a) (b) (c) sempervirens).

M23_SMIT7406_09_GE_C23.indd 541 30/01/15 1:48 PM

 542    Part se ve n   •  Eco l o g ical B io g e og r a p h y

Figure 23.25  Mediterranean vegetation (fynbos) of the

Western Cape region of South Africa.

denser, building up large fuel loads of leaves and twigs on the

ground. In the dry season the shrubs, even though alive, nearly
explode when ignited.
After fire, the land returns either to lush green sprouts
coming up from buried root crowns or to grass if a seed source
Figure 23.26  Chaparral is the dominant mediterranean shrub
is nearby. As the regrowth matures, the chaparral vegetation
vegetation of southern California.
once again becomes dense, the canopy closes, the litter accu­
mulates, and the stage is set for another fire.
Shrub communities have a complex of animal life that var­
ies with the region. In the mediterranean shrublands, similarity the Southern Hemisphere, temperate evergreen forests become
in habitat structure has resulted in pronounced parallel and con­ predominant. Deciduous forest once covered large areas of
vergent evolution among bird species and some lizard species, Europe and China, parts of North and South America, and
especially between the Chilean mattoral and the California the highlands of Central America. The deciduous forests of
chaparral. In North America, chaparral and sagebrush com­ Europe and Asia, however, have largely disappeared, cleared
munities support mule deer (Odocoileus hemionus), coyotes over the centuries for agriculture. In eastern North America,
(Canis latrans), a variety of rodents, jackrabbits (Lepus spp.), the deciduous forest consists of several forest types or as­
and sage grouse (Centrocercus urophasianus). The Australian sociations (Figure 23.28), including the mixed mesophytic
mallee is rich in birds, including the endemic mallee fowl forest of the unglaciated Appalachian plateau, the beech–maple
(Leipoa ocellata), which incubates its eggs in a large mound. and northern hardwood forests (with pine and hemlock) in
Among the mammalian life are the gray kangaroo (Macropus northern regions that eventually grade into the boreal forest
giganteus) and various species of wallaby (Macropodidae). (see Section 24.8), the maple–basswood forests of the Great
The diverse topography and geology of the mediterranean Lakes states, the oak–chestnut (now oak since the die-off of the
environments give rise to a diversity of soil conditions, but American chestnut) or central hardwood forests, which cover
soils are typically classified as alfisols (see Figure 4.12). The most of the Appalachian Mountains, the magnolia–oak forests
soils of the regions are generally deficient in nutrients, and of the Gulf Coast states, and the oak–hickory forests of the
litter decomposition is limited by low temperatures during the Ozarks. In North America, temperate deciduous forests reach
winter and low soil moisture during the summer months. These their greatest development in the mesic forests of the central
ecosystems vary in productivity depending on the annual pre­ Appalachians, where the number of tree species is unsurpassed
cipitation and the severity of summer drought. by any other temperate area in the world.
The Asiatic broadleaf forest, found in eastern China,
Japan, Taiwan, and Korea, is similar to the North American
23.7  Forest Ecosystems Dominate deciduous forest and contains several plant species of the same
the Wetter Regions of the genera as those found in North America and western Europe.
However, broadleaf evergreen species become increasingly
Temperate Zone present in Japan, South Korea, and southern China and in
Climatic conditions in the humid midlatitude regions give rise the wet foothills of the Himalayas. In southern Europe, their
to the development of forests dominated by broadleaf decidu­ presence reflects the transition into the mediterranean region.
ous trees (Figure 23.27). But in the mild, moist climates of Evergreen oaks and pines are also widely distributed in the

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  C h ap t er 23   •  Terrestrial Ecosystems    543

Madison Harrisburg Oxford Ryongyang Figure 23.27  Geographic

286 m 102 m 63 m 34 m
°C 7.9 795 mm °C 11.9 1028 mm 0° °C 10.1 652 mm °C 9.4 925 mm distribution of Earth’s temperate
30 120 30 120 30 120 80 250
20 100 20 20 100 200
forest ecosystems and associated
60
10 80 10 80 10 80 150 climate diagrams showing long-term
40
60 60
0 40 0 40 0 40 20 100 patterns of monthly temperature and
−10 20 −10 −10 20 0 50
−20 0 −20 0 −20 0 −20 0
precipitation for selected locations.
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
These ecosystems are characterized
by distinct seasonality in temperature
with adequate precipitation during
40° 40° the growing season to support a
closed canopy of trees.
20° 20° (Adapted from Archibold 1995.)

0° 0°

20° 20°

40° 40°

Punta Areñas Kazan Kaifeng Hokitika

5m 64 m 100 m 4m
°C 5.7 448 mm °C 3.1 435 mm °C 14.4 566 mm °C 11.1 2764 mm
30 120 30 120 60 200 30 300
20 100 20 100
40 150
10 80 10 80 20 200
60 60 20 100
0 40 0 40 10 100
−10 −10 0 50
20 20
−20 0 −20 0 −20 0 0 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

southeastern United States, where they are usually associated forests are also found in New Zealand, Tasmania, and parts of
with poorly developed sandy or swampy soils. southeastern Australia where the winter temperatures are mod­
In the Southern Hemisphere, temperate deciduous forests erated by the coastal environment. Climate regions in these
are found only in the drier parts of the southern Andes. In areas are similar to those of the Pacific Northwest of North
southern Chile, broadleaf evergreen rain forests have developed America, but here the predominant species are conifers.
in an oceanic climate that is virtually frost-free. Evergreen In the broadleaf deciduous forests of the temperate region,
the end of the growing season is marked by the autumn col­
ors of foliage shortly before the trees enter into their leafless
winter period (Figure 23.29). The trees resume growth in the
spring in response to increasing temperatures and longer day
lengths. Many herbaceous species flower at this time before the
developing canopy casts a heavy shade on the forest floor.
Highly developed, unevenly aged deciduous forests usu­
Mesophytic ally have four vertical layers or strata (see Figure 16.12a). The
Appalachian oak section upper canopy consists of the dominant tree species, below
Oak–hickory which is the lower tree canopy, or understory. Next is the shrub
layer, followed finally by the ground layer of herbs, ferns,
Southern mixed
and mosses. The diversity of animal life is associated with
Oak–pine section this vertical stratification and the growth forms of plants (see
Mississippi alluvial plain Figure  16.13). Some animals, particularly forest arthropods,
Subtropical evergreen spend most of their lives in a single stratum; others range over
Beach–maple–basswood
two or more strata. The greatest concentration and variety of
0 500 1000
km life in the forest occurs on and just below the ground layer.
Northern hardwoods–red pine
Many animals—the soil and litter invertebrates in ­particular—
Northern hardwoods–hemlock remain in the subterranean stratum. Others, such as mice,
Figure 23.28  Large-scale distribution of temperate forest shrews, ground squirrels, and forest salamanders, burrow into
communities in the eastern United States, derived from the soil or litter for shelter and food. Larger mammals live on
contemporary data. Compare with Figure 17.11, depicting the the ground layer and feed on herbs, shrubs, and low trees. Birds
original forest pattern. move rather freely among several strata but typically favor one
(Adapted from Dyer 2006.) layer over another (see Figure 16.13).

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 544    Part se ve n   •  Eco l o g ical B io g e og r a p h y

Figure 23.29 
A temperate forest
of the Appalachian
region: (a) the canopy
during autumn,
and (b) interior of
the forest during
spring. The forest
is dominated by
oaks (Quercus spp.)
and yellow poplar
(Liriodendron
tulipifera), with an
understory of redbud
(Cercis canadensis)
in bloom.

(a) (b)

Differences in climate, bedrock, and drainage are reflected

in the variety of soil conditions present. Alfisols, inceptisols,
23.8  Conifer Forests Dominate the
and ultisols are the dominant soil types with alfisols typically Cool Temperate and Boreal Zones
associated with glacial materials in more northern regions (see Conifer forests, dominated by needle-leaf evergreen trees,
Figure 4.12). Primary productivity varies geographically and is are found primarily in a broad circumpolar belt across the
influenced largely by temperatures and the length of the grow­ Northern Hemisphere and on mountain ranges, where low
ing season (see Section 20.3). Leaf fall in deciduous forests temperatures limit the growing season to a few months each
occurs over a short period in autumn, and the availability of year (Figure 23.30). The variable composition and structure
nutrients is related to rates of decomposition and mineraliza­ of these forests reflect the wide range of climatic conditions
tion (see Chapter 21). in which they grow. In central Europe, extensive coniferous

Fairbanks Kapuskasing Härnosänd Yakutsk Figure 23.30  Geographic

133 m 215 m 8m 100 m
°C −3.4 287 mm °C 0.8 858 mm °C 4.4 697 mm °C −10.2 213 mm distribution of Earth’s conifer forest
20 100 20 100 20 100 20 100
80 80 80 80
ecosystems and associated climate
0 0 0 0
60 60 60 60 diagrams showing long-term
−20 40 −20 40 0° −20 40 −20 40 patterns of monthly temperature
−40 20 −40 20 −40 20 −40 20
0 0 0 0
and precipitation for selected
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
locations. Regions supporting
conifer forest ecosystems have a
60° 60° lower mean annual temperature and
shorter growing season than areas
that support temperate deciduous
forest (see Figure 23.27).
40° 40°
(Adapted from Archibold 1995.)

20° 20°

0° 0°

20° 20°

Prince Rupert Aspen Turukhansk Onor

16 m 2412 m 45 m 180 m
°C 7.6 2399 mm °C 4.8 471 mm °C −7.6 496 mm °C −1.0 570 mm
20 20 100 20 100 20 100
300 80 80 80
0 0 0 0
200 60 60 60
−20 −20 40 −20 40 −20 40
100
−40 −40 20 −40 20 −40 20
0 0 0 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

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  C h ap t er 23   •  Terrestrial Ecosystems    545

(a) (b)

Figure 23.31  Two coniferous forest types. (a) A Norway spruce in the Tarvisio region of
Italy. (b) A montane coniferous forest in the Rocky Mountains. The dry, lower slopes support
ponderosa pine; the upper slopes are cloaked with Douglas fir.

forests, dominated by Norway spruce (Picea abies), cover the productive coastal forest extending along the coastal strip
slopes up to the subalpine zone in the Carpathian Mountains from Alaska to northern California.
and the Alps (Figure  23.31a). In North America, sev­ The largest expanse of conifer forest—in fact, the larg­
eral coniferous forests blanket the Rocky, Wasatch, Sierra est vegetation formation on Earth—is the boreal forest, or
Nevada, and Cascade mountains. At high elevations in the taiga (Russian for “land of little sticks”). This belt of conifer­
Rocky Mountains grows a subalpine forest dominated by ous forest, encompassing the high latitudes of the Northern
Engelmann spruce (Picea engelmannii) and subalpine fir Hemisphere, covers about 11 percent of Earth’s terrestrial
(Abies lasiocarpa). Middle elevations have stands of Douglas surface (see Figure  23.30). In North America, the boreal for­
fir, and lower elevations are dominated by open stands est covers much of Alaska and Canada and spills into northern
of ponderosa pine (Pinus ponderosa; Figure  23.31b) and New England, with fingers extending down the western moun­
dense stands of the early successional conifer, lodgepole pine tain ranges and into the Appalachians. In Eurasia, the boreal
(Pinus contorta). The largest tree of all, the giant sequoia forest begins in Scotland and Scandinavia and extends across
(Sequoiadendron giganteum), grows in scattered groves on the continent, covering much of Siberia, to northern Japan.
the western slopes of the California Sierra. In addition, the Three major vegetation zones make up the taiga
mild, moist climate of the Pacific Northwest supports a highly (Figure 23.32): (1) the forest–tundra ecotone with open stands

Figure 23.32  Major

subdivisions of the North
American boreal forest.
(Adapted from Payette et al. 2001.)

Forest-tundra
Atlantic
Lichen woodland Ocean
Closed-crown forest
Hudson
Bay
Pacific
Ocean

Canada
0 500 1000 U.S.A.
km

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 546    Part se ve n   •  Eco l o g ical B io g e og r a p h y

The boreal forest has a unique animal community. Caribou

(Rangifer tarandus), wide-ranging and feeding on grasses,
sedges, and especially lichens, inhabit open spruce–lichen
woodlands. Joining the caribou is the moose (Alces alces), called
elk in Eurasia, the largest of all deer. It is a lowland mammal
feeding on aquatic and emergent vegetation as well as alder
and willow. Competing with moose for browse is the cyclic
snowshoe hare (Lepus americanus). The arboreal red squirrel
(Sciurus hudsonicus) inhabits the conifers and feeds on young
pollen-bearing cones and seeds of spruce and fir, and the quill-
bearing porcupine (Erethizon dorsatum) feeds on leaves, twigs,
and the inner bark of trees. Preying on these is an assortment of
predators including the wolf, lynx (Lynx canadensis and L. lynx),
pine martin (Martes americana), and owls. The taiga is also the
nesting ground of migratory neotropical birds and the habitat of
northern seed-eating birds such as crossbills (Loxia spp.), gros­
Figure 23.33  Black spruce is a dominant conifer in the North beaks (Coccothraustes spp.), and siskins (Carduelis spp.).
American taiga. Of great ecological and economic importance are major
herbivorous insects such as the spruce budworm (Choristoneura
fumiferana). Although they are major food items for insectivo­
of stunted spruce, lichens, and moss; (2) the open lichen wood­ rous summer birds, these insects experience periodic outbreaks
land with stands of lichens and black spruce; and (3) the main during which they defoliate and kill large expanses of forest.
boreal forest (Figure 23.33) with continuous stands of spruce Compared to more temperate forests, boreal forests have
and pine broken by poplar and birch on disturbed areas. This generally low net primary productivity; they are limited by low
boreal–mixed forest grades into the temperate forest of south­ nutrients, cooler temperatures, and the short growing season.
ern Canada and the northern United States. Primarily occupy­ Likewise, inputs of plant litter are low compared to the forests
ing formerly glaciated land, the taiga is also a region of cold of the warmer temperate zone. However, rates of decomposi­
lakes, bogs, rivers, and alder thickets. tion are slow under the cold, wet conditions, resulting in the
A cold continental climate with strong seasonal varia­ accumulation of organic matter. Soils are primarily spodosols
tion dominates the taiga. The summers are short, cool, and characterized by a thick organic layer (see Figure 4.12). The
moist; the winters are long, harsh, and dry, with a prolonged mineral soils beneath mature coniferous forests are compara­
period of snowfall. The driest winters and the greatest sea­ tively infertile, and growth is often limited by the rate at which
sonal fluctuations are in interior Alaska and central Siberia, mineral nutrients are recycled through the ecosystem.
which experience seasonal temperature extremes (differences
between ­minimum and maximum annual temperatures) of as
much as 100°C. 23.9  Low Precipitation and Cold
Much of the taiga is under the controlling influence of Temperatures Define the Arctic
permafrost, which impedes infiltration and maintains high soil
moisture. Permafrost is the perennially frozen subsurface that Tundra
may be hundreds of meters deep. It develops where the ground Encircling the top of the Northern Hemisphere is a frozen
temperatures remain below 0°C for extended periods of time. plain, clothed in sedges, heaths, and willows, dotted with
Its upper layers may thaw in summer and refreeze in winter. lakes, and crossed by streams (Figure 23.34). Called tundra,
Because the permafrost is impervious to water, it forces all wa­ its name comes from the Finnish tunturi, meaning “a tree­
ter to remain and move above it. Thus, the ground stays soggy less plain.” The arctic tundra falls into two broad types: tun­
even though precipitation is low, enabling plants to exist in the dra with up to 100 percent plant cover and wet to moist soil
driest parts of the Arctic. (Figure 23.35), and polar desert with less than 5 percent plant
Fires are recurring events in the taiga. During periods of cover and dry soil.
drought, fires can sweep over hundreds of thousands of hectares. Conditions unique to the Arctic tundra are a product of at
All of the boreal species, both broadleaf trees and conifers, are least three interacting forces: (1) the permanently frozen deep
well adapted to fire. Unless too severe, fire provides a seedbed layer of permafrost; (2) the overlying active layer of organic
for regeneration of trees. Light surface burns favor early succes­ matter and mineral soil that thaws each summer and freezes the
sional hardwood species. More severe fires eliminate hardwood following winter; and (3) vegetation that reduces warming and
competition and favor spruce and jack pine regeneration. retards thawing in summer. Permafrost chills the soil, retarding
Because of the global demand for timber and pulp, vast ar­ the general growth of plant parts both above- and belowground,
eas of the boreal forest across North America and Siberia are be­ limiting the activity of soil microorganisms, and diminishing
ing clear-cut with little concern for their future, see this chapter, the aeration and nutrient content of the soil.
Ecological Issues & Applications. This exploitation can alter the Alternate freezing and thawing of the upper layer of
nature and threaten the survival of the boreal forest. soil creates the unique, symmetrically patterned landforms

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  C h ap t er 23   •  Terrestrial Ecosystems    547

Nome Baker Lake Godthaab Kazachye Figure 23.34  Geographic

4m 4m 27 m 22 m
°C −3.6 463 mm °C −11.9 150 mm °C −0.7 736 mm °C −13.9 199 mm distribution of Earth’s tundra
20 120 20 120 20 120 20 120
10 100 10 100 10 100 10 100 ecosystems and associated
0 80 0 80 0 80 0 80 climate diagrams showing
−10 60 −10 60 −10 60 −10 60
−20 40 −20 40 −20 40 −20 40 long-term patterns of monthly
−30 20 −30 20 −30 20 −30 20
−40 0 −40 0 0° −40 0 −40 0 temperature and precipitation
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND
for selected locations. Tundra
ecosystems are characterized
by lower mean temperatures, a
shorter growing season, and lower
60° 60°
annual precipitation than regions
supporting conifer forest (see
Figure 24.30 for comparison).
40° 40° (Adapted from Archibold 1995.)

20° 20°

0° 0°

20° 20°

Oruro Murmansk Lhasa Khatanga

3707 m 46 m 3600 m 24 m
°C 10.7 288 mm °C 0.1 386 mm °C 8.8 199 mm °C −13.8 237 mm
20 120 20 120 20 120 20 120
10 100 10 100 10 100 10 100
0 80 0 80 0 80 0 80
−10 60 −10 60 −10 60 −10 60
−20 40 −20 40 −20 40 −20 40
−30 20 −30 20 −30 20 −30 20
−40 0 −40 0 −40 0 −40 0
J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND J FMAMJ J ASOND

typical of the tundra (Figure 23.36 ). The frost pushes stones buffeting by the wind, and abrasion from wind-carried particles
and other material upward and outward from the mass to of soil and ice can survive. Low ground is covered with a com­
form a patterned surface of frost hummocks, frost boils, earth plex of cotton grasses, sedges, and Sphagnum. Well-drained
stripes, and stone polygons. On sloping ground, creep, frost sites support heath shrubs, dwarf willows and birches, herbs,
thrusting, and downward flow of supersaturated soil over the mosses, and lichens. The driest and most exposed sites support
permafrost form solifluction terraces, or “flowing soil.” This scattered heaths and crustose and foliose lichens growing on
gradual downward creep of soils and rocks eventually rounds the rock. Arctic plants propagate themselves almost entirely
off ridges and other irregularities in topography. Such mold­ by vegetative means, although viable seeds many hundreds of
ing of the landscape by frost action, called cryoplanation, is years old exist in the soil.
far more important than erosion in wearing down the Arctic Plants are photosynthetically active on the Arctic tundra
landscape. about three months out of the year. As snow cover disappears,
Structurally, the vegetation of the tundra is simple. The plants commence photosynthetic activity. They maximize use
number of species tends to be low, and growth is slow. Only of the growing season and light by photosynthesizing during
those species able to withstand constant disturbance of the soil, the 24-hour daylight period, even at midnight when light is

Figure 23.35  (a) The plant cover that characterizes the wide expanse of the Arctic tundra
in the Northwest Territories of Canada presents a stark contrast to (b) the polar desert that is
characterized by dry soils and sparse plant cover.
(a) (b)

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 548    Part se ve n   •  Eco l o g ical B io g e og r a p h y

the surface, where there are abundant populations of segmented

whiteworms (Enchytraeidae), collembolas, and flies (Diptera),
chiefly crane flies. Summer in the Arctic tundra brings hordes
of black flies (Simulium spp.), deer flies (Chrysops spp.), and
mosquitoes.
Dominant vertebrates on the Arctic tundra are herbivores,
including lemmings, Arctic hare, caribou, and musk ox (Ovibos
moschatus). Although caribou have the greatest herbivore bio­
mass, lemmings, which breed throughout the year, may reach
densities as great as 125 to 250 per hectare; they consume
three to six times as much forage as caribou do. Arctic hares
(Lepus arcticus) that feed on willows disperse over the range
in winter and congregate in more restricted areas in summer.
Caribou are extensive grazers, spreading out over the tundra
in summer to feed on sedges. Musk oxen are more intensive
(a)
grazers, restricted to more localized areas where they feed on
sedges, grasses, and dwarf willow. Herbivorous birds are few,
dominated by ptarmigan and migratory geese.
The major Arctic carnivore is the wolf (Canus lupus),
which preys on musk ox, caribou, and, when they are abun­
dant, lemmings. Medium-sized to small predators include
the Arctic fox (Alopex lagopus), which preys on Arctic hare,
and several species of weasel, which prey on lemmings. Also
feeding on lemmings are snowy owls (Nyctea scandiaca) and
the hawk-like jaegers (Stercorarius spp.). Sandpipers (Tringa
spp.), plovers (Pluvialis spp.), longspurs (Calcarius spp.), and
waterfowl, which nest on the wide expanse of ponds and boggy
ground, feed heavily on insects.
At lower latitudes, alpine tundra occurs in the higher
mountains of the world. The alpine tundra is a severe envi­
ronment of rock-strewn slopes, bogs, meadows, and shrubby
thickets (Figure  23.37). It is a land of strong winds, snow,
(b)
cold, and widely fluctuating temperatures. During summer, the
temperature on the surface of the soil ranges from 40 to 0°C.
Figure 23.36  Patterned landforms typical of the tundra The atmosphere is thin so light intensity, especially ultraviolet,
region: (a) frost hummocks, and (b) polygons. Alternate freezing is high on clear days. Alpine tundras have little permafrost,
and thawing of the upper layer of soil creates the symmetrically and it is confined mostly to very high elevations. Lacking per­
patterned landforms. mafrost, soils are drier. Only in alpine wet meadows and bogs
do soil moisture conditions compare with those of the Arctic.
one-tenth that of noon. The nearly erect leaves of some Arctic Precipitation, especially snowfall and humidity, is higher in the
plants permit almost complete interception of the low angle of alpine regions than in the Arctic tundra, but steep topography
the Arctic sun. induces a rapid runoff of water.
Much of the photosynthate goes into the production of
new growth, but about one month before the growing season
ends, plants cease to allocate photosynthate to aboveground Figure 23.37  Rocky Mountains alpine tundra.
biomass. They withdraw nutrients from the leaves and move
them to roots and belowground biomass, sequestering 10 times
the amount stored by temperate grasslands.
Structurally, most of the tundra vegetation is underground.
Root-to-shoot ratios of vascular plants range from 3:1 to 10:1.
Roots are concentrated in the upper soil that thaws during
the summer, and aboveground parts seldom grow taller than
30  cm. It is not surprising, then, that the belowground net
­annual production is typically three times that of the aboveg­
round productivity.
The tundra hosts fascinating animal life, even though the
diversity of species is low. Invertebrates are concentrated near

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  C h ap t er 23   •  Terrestrial Ecosystems    549

Eco l o g i c a l The Extraction of Resources from Forest

Issues & Applications Ecosystems Involves an Array of Management
Practices

Fruits and nuts Cordwood: Pulpwood Global distribution of original and remaining forests
Fuel
Foliage: Oils Charcoal
Extracts Tannin
Decorations Dyes
Wood alcohol
Bark: Tannin Poles
Dyes
Piles
Drugs
Oils Posts
Logs: Lumber
Sap: Sugar and syrup Veneer
Plywood
Gums: Chewing gum Pressboard
Ointments Stumps: Veneer
Perfumes Turpentine
Flavorings Tropical current Temperate and boreal current
Charcoal Tropical original Temperate and boreal original
Resins Wood tar
Adhesives Pine oil
Drugs Figure 23.39  Globally, about half of the forest that was present
under modern (post-Pleistocene) climatic conditions, and before the
Roots: Tea and oil
Smoking pipes spread of human influence, has disappeared—largely because of
human activities.
Figure 23.38  A variety of products derived from forests. (Adapted from United Nations, FAO.)

Forest ecosystems cover approximately 35 percent of Earth’s production of forest resources requires achieving a balance
surface and provide a wealth of resources, including fuel, between net growth and harvest. To achieve this end, foresters
building materials, and food (Figure 23.38). Although planta­ have an array of silvicultural and harvesting techniques from
tions provide a growing percentage of forest resources, more clear-cutting to selection cutting.
than 90 percent of global forest resources are still harvested Clear-cutting involves removing the forest and reverting
from native forests. it to an early stage of succession (Figure 23.40a). The area
Globally, about half of the forest that was present un­ harvested can range from thousands of hectares to small patch
der modern (post-Pleistocene) climatic conditions—and be­ cuts of a few hectares designed to create habitat for wildlife spe­
fore the spread of human influence—has disappeared largely cies that require an opening within the forest (see Section 19.4).
through the impact of human activities (Figure  23.39). Postharvest management varies widely for clear-cut areas. When
The spread of agriculture and animal husbandry, the harvest­ natural forest stands are clear-cut, there is generally no follow-
ing of forests for timber and fuel, and the expansion of popu­ up management. Stands are left to regenerate naturally from
lated areas have all taken their toll on forests. The causes and existing seed and sprouts on the site and the input of seeds from
timing of forest loss differ among regions and forest types, as adjacent forest stands. With no follow-up management, clear-cut
do the current trends in change in forest cover. In the face of areas can be badly disturbed by erosion that affects subsequent
increasing demand and declining forest cover, the sustainable recovery of the site as well as adjacent aquatic communities.

Figure 23.40  Examples of (a) clear-cut and (b) shelterwood (seed-tree) forest harvest.
(a) (b)

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 550    Part se ve n   •  Eco l o g ical B io g e og r a p h y

Harvest by clear-cutting is the typical practice on forest 300

Harvestable timber volume (m3)

plantations, but here intensive site management follows clear­ Operable window
ing. Plant materials that are not harvested (branches, leaves, 250
and needles) are typically burned to clear the site for planting.
200
After clearing, seedlings are planted and fertilizer applied to
encourage plant growth. Herbicides are often used to discour­ 150
age the growth of weedy plants that would compete with the
seedlings for resources. 100
The seed-tree, or shelterwood, system is a method of 50
regenerating a new stand by removing all trees from an area ex­
cept for a small number of seed-bearing trees (Figure 23.40b). 0
25 50 75 100
The uncut trees are intended to be the main source of seed for
Stand age
establishing natural regeneration after harvest. Seed trees can (a)
be uniformly scattered or left in small clumps, and they may or
may not be harvested later.
In many ways, the shelterwood system is similar to a 14 Stems per m3
clear-cut because generally not enough trees are left standing decrease as the
12 average tree
to affect the microclimate of the harvested area. The advantage

Stems per m3
size increases
of the shelterwood approach is that the seed source for natural
regeneration is not limited to adjacent stands. This can result 10
in improved distribution (or stocking) of seedlings as well as a
8
more desirable mix of species.
Like any silviculture system, shelterwood harvesting re­
6
quires careful planning to be effective. Trees left on the site
must be strong enough to withstand winds and capable of pro­ 0
25 50 75 100
ducing adequate seed, seedbed conditions must be conducive Stand age
to seedling establishment (this may require a preparatory treat­ (b)
ment during or after harvest), and follow-up management may Figure 23.41  Two criteria are used to determine when a
be required to fully establish the regeneration. stand of trees is suitable for harvest (referred to as the operable
In selection cutting, mature single trees or groups of trees window): (1) salable volume of wood per hectare (m3/ha), and (2)
scattered through the forest are removed. Selection cutting pro­ average tree size as measured by the stems per cubic meter (the
duces only small openings or gaps in the forest canopy. Although number of trees required to make a cubic meter of wood volume).
this form of timber harvest can minimize the scale of disturbance In the example shown, dashed horizontal lines represent the
within the forest caused by direct removal of trees, the network criteria of (a) minimum salable wood volume of 100 m3/ha, and (b)
of trails and roads necessary to provide access can be a major minimum average tree size of 9 stems/m3. Dashed vertical lines
indicate the earliest stage at which both criteria are met.
source of disturbance (to both plants and soils). Selective cutting
(Adapted from Oriens et al. 1986.)
also can cause changes in species composition and diversity be­
cause only certain species are selectively removed. Interpreting Ecological Data
Regardless of the differences in approach, some general
Q1. What does graph (a) imply about the change in average tree
principles apply if the harvesting of resources is to be sus­ size (diameter or height) with stand age?
tainable. Forest trees function in the manner discussed for
Q2. Assume that a decision is made to harvest the stand when
competition in plant populations (Chapter 11, Section 11.3). it reaches the minimum salable wood volume (100 m3/ha). How
Whether a forest is planted as seedlings or grown by natural re­ many stems (trees) per m3 would be harvested?
generation, its establishment begins with a population of small
individuals (seedlings) that grow and compete for the essential
resources of light, water, and nutrients. As biomass in the forest (planting density) can be controlled to influence the timing of
increases, the density of trees decreases and the average tree the stand’s availability for harvest (Figure 23.42).
size increases as a result of self-thinning (Figure 23.41; also After trees are harvested, a sufficient time must pass for
see Section 11.5, Figure 11.9). For a stand to be considered the forest to regenerate. For sustained yield, the time between
economically available for harvest (referred to as being in an harvests must be sufficient for the forest to regain the level
operative state), minimum thresholds must be satisfied for of biomass it had reached at the time of the previous harvest.
the harvestable volume of timber per hectare and average tree Rotation time depends on a variety of factors related to the tree
size (see Figure 23.41); these thresholds vary depending on species, site conditions, type of management, and intended use
the species. In plantation forestry, for a given set of thresholds of the trees being harvested. Wood for paper products (pulp­
(timber volume and average tree size), the initial stand density wood), fence posts, and poles are harvested from fast-growing

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  C h ap t er 23   •  Terrestrial Ecosystems    551

300
600 Uncut watershed

Nitrate (kg/ha)
250 Initial Clear-cut watershed
Salable volume (m3/ha)

trees/ha 500
200 400
4000
150 0

1966–1967

1972–1973

1975–1976
1963–1964

1969–1970
100 2000

50 500

0
25 50 75 100
Figure 23.43  Temporal changes in the nitrate concentration
Stand age (years) of stream water for two forested watersheds in Hubbard Brook,
(a)
New Hampshire. The forest on one watershed was clear-cut (noted
by arrow), and the other forest was undisturbed. Note the large
18
increase in concentrations of nitrate in the stream on the clear-cut
16 watershed. This increase is a result of increased decomposition
14 and nitrogen mineralization after clear-cutting. The nitrogen then
leached into the surface water and groundwater.
Stems per m3

12
Initial (Adapted from Likens and Borman 1995.)
10 trees/ha
8
4000
6 As with agricultural crops, a significant amount of nu­
500 2000 trients are lost from the forest when trees are harvested and
4
removed (see Chapter 22, Ecological Issues & Applications).
0
25 50 75 100 The loss of nutrients in plant biomass is often compounded
(b)
Stand age (years) by further losses from soil erosion and various postharvest
management practices—particularly the use of fire. The re­
Figure 23.42  Effects of initial stand density on timing of stand duction of nutrients reduces plant growth, requiring a longer
availability for harvest (operable window). As in Figure  23.41, rotation period for subsequent harvests or causing reduced
dashed horizontal lines represent criteria of (a) minimum forest yield if the rotation period is maintained. Forest manag­
marketable wood volume of 100 m3/ha and (b) a minimum ers often counter the loss of nutrients by using chemical fertil­
average tree size of 9 stems/m3. Dashed vertical lines indicate izers, which create other environmental problems for adjacent
the earliest stage at which both criteria are met. Note that the
aquatic ecosystems (see Chapter 24, Ecological Issues &
intermediate planting density of 2000 trees per hectare provides
Applications).
the earliest operable window.
(Adapted from Oriens et al. 1983.)
In addition to the nutrients lost directly through biomass
removal, logging can also result in the transport of nutrients
Interpreting Ecological Data? from the ecosystem by altering processes involved in inter­
Q1. The analysis includes three initial planting densities: 500,
nal cycling. The removal of trees in clear-cutting and other
2000, and 4000 trees/ha. At which stand age does each of the forest management practices increases the amount of radia­
three initial planting densities achieve the minimum constraint for tion (including direct sunlight) reaching the soil surface. The
average tree size? resulting increase in soil temperatures promotes decomposi­
Q2. Given the requirements of minimum wood volume and tion of remaining soil organic matter and causes an increase
average tree size as defined, which of the initial planting densities in net mineralization rates (see Sections 21.4 and 21.5). This
meets these requirements at the earliest stand age (earliest
increase in nutrient availability in the soil occurs at the same
operable window)?
time that demand for nutrients is low because plants have been
removed and net primary productivity is low. As a result, there
species, allowing a short rotation period (15–40 years). These is a dramatic increase in the leaching of nutrients from the soil
species are often grown in highly managed plantations where into ground and surface waters (Figure 23.43). This export of
trees can be spaced to reduce competition and fertilized to nutrients from the ecosystem results from decoupling the two
maximize growth rates. Trees harvested for timber (saw logs) processes of nutrient release in decomposition and nutrient up­
require a much longer rotation period. Hardwood species used take in net primary productivity.
for furniture and cabinetry are typically slower growing and Sustained yield is a key concept in forestry and is prac­
may have a rotation time of 80 to 120 years. Sustained forestry ticed to some degree by large timber companies and federal
of these species works best in extensive areas where blocks of and state forestry agencies. But all too often, industrial for­
land can be maintained in different age classes. estry’s approach to sustained yield is to grow trees as a crop

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 552    Part se ve n   •  Eco l o g ical B io g e og r a p h y

rather than maintaining a forest ecosystem. Their manage­

ment approach represents a form of agriculture in which trees
are grown as crops: trees are clear-cut, the site is sprayed
with herbicides, planted or seeded with one species, then
clear-cut and planted again. Clear-cutting practices in some
national forests, especially in the Pacific Northwest and the
Tongass National Forest in Alaska, hardly qualify as sus­
tained-yield management. Even more extensive clear-cutting
of forests is taking place in the northern forests of Canada,
especially in British Columbia (Figure 23.44), and in large
areas of Siberia.
The problem of sustained-yield forestry is its economic
focus on the resource with little concern for the forest as a
biological community. A carefully managed stand of trees,
often reduced to one or two species, is not a forest in an eco­
logical sense. Rarely will a naturally regenerated forest, and
certainly not a planted one, support the diversity of life found
in old-growth forests. By the time the trees reach economic or
Figure 23.44  Large-scale clear-cut in British Columbia, financial maturity—based on the type of rotation—they are
Canada. cut again.

S u mm a r y
Ecosystem Distribution and Plant Adaptations  23.1 Tropical Savannas  23.3
Terrestrial ecosystems can be grouped into broad categories Savannas are characterized by a codominance of grasses and
called biomes. Biomes are classified according to the predomi­ woody plants. Such vegetation is characteristic of regions with
nant plant types. There are at least eight major terrestrial biome alternating wet and dry seasons. Savannas range from grass
types: tropical forest, temperate forest, conifer forest (taiga or with occasional trees to shrubs to communities where trees
boreal forest), tropical savanna, temperate grasslands, chapar­ form an almost continuous canopy as a function of precipitation
ral (shrublands), tundra, and desert. These broad categories and soil texture. Productivity and decomposition in savanna
reflect the relative contribution of three general plant life-
­ ecosystems are closely tied to the seasonality of precipitation.
forms: trees, shrubs, and grasses. Interaction between moisture Savannas support a large and varied assemblage of both
and temperature is the primary factor limiting the nature and invertebrate and vertebrate herbivores. The African savanna is
geographic distribution of terrestrial ecosystems. dominated by a large, diverse population of ungulate fauna and
associated carnivores.
Tropical Forests  23.2
Seasonality of rainfall determines the types of tropical forests. Temperate Grasslands  23.4
Rain forests, associated with high seasonal rainfall, are domi­ Natural grasslands occupy regions where rainfall is between
nated by broadleaf evergreen trees. They are noted for their enor­ 250 and 800 mm a year. Once covering extensive areas of the
mous diversity of plant and animal life. The vertical structure of globe, natural grasslands have shrunk to a fraction of their orig­
the forest is divided into five general layers: emergent trees, high inal size because of conversion to cropland and grazing lands.
upper canopy, low tree stratum, shrub understory, and a ground Grasslands vary with climate and geography. Native grass­
layer of herbs and ferns. Conspicuous in the rain forest are the lands of North America, influenced by declining precipitation
lianas or climbing vines, epiphytes growing up in the trees, and from east to west, consist of tallgrass prairie, mixed-grass
stranglers growing downward from the canopy to the ground. prairie, shortgrass prairie, and desert grasslands. Eurasia has
Many large trees develop buttresses for support. Nearly 90 per­ steppes; South America, the pampas; and southern Africa, the
cent of nonhuman primate species live in tropical rain forests. veld. Grassland consists of an ephemeral herbaceous layer that
Tropical rain forests support high levels of primary pro­ arises from crowns, nodes, and rosettes of plants hugging the
ductivity. The high rainfall and consistently warm temperatures ground. It also has a ground layer and a highly developed root
also result in high rates of decomposition and nutrient cycling. layer. Depending on the history of fire and degree of grazing
Dry tropical forests undergo varying lengths of dry season, and mowing, grasslands accumulate a layer of mulch.
during which trees and shrubs drop their leaves (drought- Grasslands support a diversity of animal life dominated
deciduous). New leaves are grown at the onset of the rainy by herbivorous species, both invertebrate and vertebrate.
season. Most dry tropical forests have been lost to agriculture Grasslands once supported herds of large grazing ungulates
and grazing and other disturbances. such as bison in North America, migratory herds of wildebeest

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  C h ap t er 23   •  Terrestrial Ecosystems    553

in Africa, and marsupial kangaroos in Australia. Grasslands Conifer Forests  23.8

evolved under the selective pressure of grazing. Although the Coniferous forests of temperate regions include the montane pine
most conspicuous grazers are large herbivores, the major con­ forests and lower-elevation pine forests of Eurasia and North
sumers are invertebrates. The heaviest consumption takes place America and the temperate rain forests of the Pacific Northwest.
belowground, where the dominant herbivores are nematodes. North of the temperate coniferous forest is the circum­
polar taiga, or boreal forest, the largest biome on Earth.
Deserts  23.5 Characterized by a cold continental climate, the taiga consists
Deserts occupy about one-seventh of Earth’s land surface and of four major zones: the forest ecotone, open boreal woodland,
are largely confined to two worldwide belts between 15° N and main boreal forest, and boreal–mixed forest ecotone.
30° S latitude. Deserts result from dry descending air masses Permafrost, the maintenance of which is influenced by tree
within these regions, the rain shadows of coastal mountain and ground cover, strongly influences the pattern of vegetation,
ranges, and remoteness from oceanic moisture. Two broad as do recurring fires. Spruces and pines dominate boreal forest
types of deserts exist: cool deserts, exemplified by the Great with successional communities of birch and poplar. Ground
Basin of North America, and hot deserts, like the Sahara. cover below spruce is mostly moss; in open spruce and pine
Deserts are structurally simple—scattered shrubs, ephemeral stands, the cover is mostly lichen.
plants, and open, stark topography. In this harsh environment, Major herbivores of the boreal region include caribou,
ways of circumventing aridity and high temperatures by either moose, and snowshoe hare. Predators include the wolf, lynx,
evading or resisting drought have evolved in plants and ani­ and pine martin.
mals. Despite their aridity, deserts support a diversity of animal
life, notably opportunistic herbivorous species and carnivores. Tundra  23.9
The Arctic tundra extends beyond the tree line at the far north of
Shrubland  23.6 the Northern Hemisphere. It is characterized by low temperature,
Shrubs have a densely branched, woody structure and low low precipitation, a short growing season, a perpetually frozen
height. Shrublands are difficult to classify because of the vari­ subsurface (the permafrost), and a frost-molded landscape. Plant
ety of climates in which shrubs can be a dominant or codomi­ species are few, growth forms are low, and growth rates are slow.
nant component of the plant community. But in five widely Over much of the Arctic, the dominant vegetation is cotton grass,
disjunct regions along the western margins of the continents sedge, and dwarf heaths. These plants exploit the long days of
between 30° and 40° latitude are found the mediterranean eco­ summer by photosynthesizing during the 24-hour daylight period.
systems. Dominated by evergreen shrubs and sclerophyll trees, Most plant growth occurs underground. The animal community is
these biomes have adapted to the distinctive climate of sum­ low in diversity but unique. Summer in the Arctic brings hordes
mer drought and cool, moist winters. These shrublands are fire of insects, providing a rich food source for shorebirds. Dominant
adapted and highly flammable. vertebrates are lemming, Arctic hare, caribou, and musk ox.
Major carnivores are the wolf, Arctic fox, and snowy owl.
Temperate Forests  23.7 Alpine tundras occur in the mountains of the world. They
Broadleaf deciduous forests are found in the wetter environ­ are characterized by widely fluctuating temperatures, strong
ments of the warm temperate region. They once covered large winds, snow, and a thin atmosphere.
areas of Europe and China, but their distribution has been re­
duced by human activity. In North America, deciduous forests Forest Management  Ecological Issues
are still widespread. They include various types such as beech– & Applications
maple and oak–hickory forest; the greatest development is in More than 90 percent of global forest resources, which in­
the mixed mesophytic forest of the unglaciated Appalachians. clude fuel, building materials, and food, are harvested from
Well-developed deciduous forests have four strata: upper can­ native forests. Sustainable production of forest resources re­
opy, lower canopy, shrub layer, and ground layer. Vertical struc­ quires achieving a balance between net growth and harvest.
ture influences the diversity and distribution of life in the forest. To achieve this end, foresters have an array of silvicutural and
Certain species are associated with each stratum. harvesting techniques.

Study Questions
1. How do trees, shrubs, and grasses differ in their patterns 4. What types of trees characterize tropical rain forest
of carbon allocation? (leaf type)?
2. What are tree buttresses? 5. Apart from species composition, what are the defining
3. How does the warm, wet environment of tropical rain characteristics of savanna vegetation?
forests influence rates of net primary productivity and 6. What major environmental factor controls annual cycles
decomposition? in savanna plant communities?

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 554    Part se ve n   •  Eco l o g ical B io g e og r a p h y

7. How does seasonality influence rates of net primary 13. What climate is characteristic of temperate grasslands?
­productivity and decomposition in savanna ecosystems? 14. How does annual precipitation influence the structure and
8. What features of regional climate lead to the formation of productivity of grassland ecosystems?
tropical rain forest ecosystems? 15. What type of trees characterizes boreal forest?
9. What climate characterizes mediterranean ecosystems? 16. What is permafrost, and how does it influence the
10. What type of leaves characterize mediterranean plants? ­structure and productivity of boreal forest ecosystems?
11. What types of leaves characterize the trees of temperate 17. What physical and biological features characterize Arctic
forest ecosystems? tundra?
12. How does seasonality of temperature influence the 18. How does alpine tundra differ from Arctic tundra?
­structure and productivity of temperate forest ecosystems?

Further Readings
Archibold, O. W. 1995. Ecology of world vegetation. London: Quinn, R. D., and S. C. Keeley. 2006. Introduction to
Chapman & Hall. California chaparral. Berkeley: University of California
An outstanding reference for those interested in the geography Press.
and ecology of terrestrial ecosystems. An overview of the chaparral ecosystem with an excellent
Bliss, L. C., O. H. Heal, and J. J. Moore, eds. 1981. Tundra ­discussion of the role and impact of fire.
ecosystems: A comparative analysis. New York: Reichle, D. E., ed. 1981. Dynamic properties of forest
Cambridge University Press. ­ecosystems. Cambridge, UK: Cambridge University
A major reference book on the geography, structure, and Press.
­function of these high-latitude ecosystems. The major reference source on the ecology and function of
Bonan, G. B., and H. H. Shugart. 1989. “Environmental forest ecosystems throughout the world.
­factors and ecological processes in boreal forests.” Reichman, O. J. 1987. Konza prairie: A tallgrass natural
Annual Review of Ecology and Systematics 20:1–18. ­history. Lawrence: University Press of Kansas.
An excellent review article providing a good introduction to An excellent introduction to the tallgrass prairie ecosystem;
­boreal forest ecosystems. emphasizes the interrelationships of plants, animals, and
Bruijnzeel, L. A., F. N. Scatena, and L. S. Hamilton 2011. Tropical landscapes.
montane cloud forests. New York: Cambridge University Press. Richards, P. W. 1996. The tropical rain forest: An ecological
Comprehensive overview of high elevation cloud forests, exam­ study, 2nd ed. New York: Cambridge University Press.
ining geographic distribution, climate, soils, hydrological pro­ A thoroughly revised edition of a classic book on the ecology of
cesses, and cloud forest conservation and management. tropical rain forests.
Evenardi, M., I. Noy-Meir, and D. Goodall, eds. 1986. Hot Sinclair, A. R. E., and P. Arcese, eds. 1995. Serengeti
deserts and arid shrublands of the world. Ecosystems of II: Dynamics, management, and conservation of an
the World 12A and 12B. Amsterdam: Elsevier Scientific. ­ecosystem. Chicago: University of Chicago Press.
A major reference work on the geography and ecology of the A masterful study of an ecosystem. A valuable reference on the
world’s deserts. ecology of tropical savanna ecosystems.
French, N., ed. 1979. Perspectives on grassland ecology. Sinclair, A., C. Packer, and S. Mduma, eds. 2008. Serengeti
New York: Springer-Verlag. III. Human Impacts on Ecosystem Dynamics. Chicago:
This text provides a good summary of grassland ecology. University Chicago Press.
Murphy, P. G., and A. E. Lugo. 1986. “Ecology of tropical dry This book is a follow up to Serengeti II. It updates information
­forests.” Annual Review of Ecology and Systematics 17:67–88. on ecosystem dynamics, conservation, and changes brought
This article provides an overview of the distribution and ­ecology about by the growing population of pastoralists and agricultural­
of tropical dry forests, one of the most endangered terrestrial ists surrounding the Serengeti.
ecosystems.

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