1 oe Chapter 4
anee of a desert. Desertification has been ongoing for t_housands of years in
the Mediterranean region and was known even to anc1ent Greek observ
Worldwide, 9 million km 2 of arid lands have been converted to ~s.
man-made deserts. These areas are not f unctional desert ecosystems ~ -
wastelands, lacking the flora and fauna cha~acteristic of natural de~t t
The process of desertification is most severe m the Sahel region of Afric s.
just south of the Sahara, where most of the native large mammal spec· a,
are threatened with extinction. The human dimension of the prob~~
illustrated by the fact that the Sahel region is estimated to_ have 2.5 tun;:
more people (100 million currently) than the la~d c~n susta1nably support.
Further desertification appears to be_ almost me~1table,_ especially given
the higher temperatures and lower ramfall associated w1th predictions of
future climate change (Verstraete et al. 2009). In such areas, the solution
will be programs involving the implementation of sustainable agricultura!
practices, the elimination of poverty, increased political stability, and con-
trol of the human population.
Habitat Fragmentation
In addition to being destroyed outright, habitats that forrnerly occupied wide,
unbroken areas are often divided into pieces by roads, fields, towns, and a
broad range of other human constructs. Habitat fragmentation is the process
whereby a large, continuous area of habitat is both reduced in area and di-
vided into two or more fragments (FIGURE 4.8). When habitat is destroyed,
a patchwork of habitat fragments may be left behind. These fragments are
often isolated from one another by a highly modified or degraded landscape,
and their edges experience altered conditions, referred to as edge effects,
such as increased wind, fire, species invasion, or predation (Murcia 1995;
Porensky and Young 2013). The fragments are often on the least desirable
land for human uses, such as steep slopes, poor soils, and inaccessible areas.
Fragmentation almost always occurs during a severe reduction in habi-
tat area, but it can also occur when habitat area is reduced to only a minor
degree by roads, railroads, canals, power lines, fences, oil pipelines, fire
lanes, or other barriers to the free movement of species. In many ways,
habitat fragments resemble islands of original habitat in an inhospitable,
human-dominated landscape. Habitat fragmentation is a serious threat
to biodiversity, as species are often unable to survive under the altered
conditions found in fragments.
Habitat fragments differ from the original habitat in three importan!
ways:
l. Fragments have a greater amount of edge per area of habitat (and íhus
a greater exposure to edge effects ).
2. The center of a habitat fragment is closer to an edge. .
3. When a formerly continuous habitat hosting a large populat~on
· ·
1 5 di-
vided into fragments, each fragment hosts a smaller population.
Threats to Biodiversity 107
l•I ha
FIGURE 4.8 The Biological Dynamics of Forest Fragmentation (BDFF) study
in Brazil is the largest and longest-running experiment of tropical forest frag-
mentation; forest fragments of different sizes were created 1980 when the land
was cleared for cattle ranches and farms . Shown in this aerial photograph are
forest fragments of 2 ha, 14 ha, and 87 ha. These fragments were surveyed for
species composition, microclimate, and other ecosystem characteristics befare
the forest was cleared, and have been regularly surveyed over the past 36 years.
In addition, there have been forest blocks of equivalent size inside intact forest
that have been surveyed as controls. In the photograph, note that the fragments
var¡ in their distance from the intact forest. An important factor in the experi-
ment is that land use around the fragments has changed over time, with many of
the cattle pastures being abandoned, and undergoing succession to secondary
forests. For example, the left side of the 14 ha fragment is secondary vegetation
while the right side has recently been cleared. See Figure 4.10 for sorne of the
key results. (© DigitalGlobe.)
Threats posed by habitat fragmentation
LIMIT~ T? DISPERSAL AND COLONIZATION Fragmentation may limit
ªspecies potential for dispersa! and colonization by creating barriers to
nor;al movements (Stouffer et al. 2011). In an undisturbed environment,
s, ~pores, and animals move passively or actively across the landscape.
sca~r hme, populations of a species may build up and go extinct on a local
biol~ ~s t;e species disperses from one suitable site to another and the
of pogic~ ~ommunity undergoes succession. Ata landscape level, a series
soinefu ahons exhibiting this pattern of extinction and recolonization is
\%:es ref~rre~ to as a metapopulation (see Figure 6.12).
is often n; habitat 1s fragmented, the potential for dispersa! and colonization
re uced. Many bird, mammal, and insect species of the forest interior
10I Cl\&ptW 4
wlll i,ot cross even short stretches of op~ ground (Laura~e et al, 2009), lt
thay do venture into the open, they may find predators waihng on the for@lt
cdge to catch and eat them. Agricultura! fields_100 m w íde may reprl!lent
an lmpassable barrier to the dispersa! of many mvertebrate speciet. 'Road
too, may be significant barriere to anin~.al movement. Ma~y speciet
ci·ossing roads, which represent an env1ronment totally different frorn t~
av~!
habitat they are leaving. Por animals that do attempt to cross roads, motor
vehicles are a major source of mortality (Beebee 2013), as 1:138 been observed
Lt, the endangered Florida panther (Kroll 2015). To deal wtth such problems
highway officials are building animal u~erpasses, overpasses, and ot~
improvements to minimize ani~al m~r~hty. . .
As species go extmct w1th1n md1v~ual fragments thtough
natural successional and metapopulatton processes, new spe.
des will be unable to arrive because of barriere to dispersa¡
The barriers that
colonization, and the number of species present in the habítat
fraoment a. [Link].t
fragment will decline over time. Extinction will be most rapíd
reduce the ability of and severe in small habitat fragments.
[Link] t o forage, find
m ates. disperse, [Link] RESTRICTED ACCESS TO FOOD AND MATES Many animal
oolonize new locations. species need to move freely across the landscape, either as indí-
Habitat fragmentation viduals or in social groups, to feed on widely scattered resources
often creates small (Becker et al. 2010). A given resource may be needed for only a
subpopulations that few weeks each year, or even only once in a few years, but when
are vulnerable to local a habitat is fragmented, species confined to a single habitat frag-
extinction. ment may be unable to migrate over their normal home range
------•----- in search of that scarce resource. Gibbons and other primates,
for example, typically remain in forests and forage wídely for
fruits. Finding scattered trees with abundant fruit crops may be
crucial during episodes of fruit scarcity. Clearings and roads that break up
the forest canopy may prevent these primates from reaching nearby fruit-
ing trees because the primates are unable or unwilling to descend to the
ground and cross the intervening open landscape. Fences may prev~t the
natural migration of large grazing animals such as wildebeest and b1son,
forcing them to overgraze unsuitable habitat, which eventually leads to
starvation and further degradation of the habitat (Gates et al. 2012). d
Barriers to dispersal can also restrict the ability of widely scattere
species to find mates, leading to a loss of reproductive potentíal for
animal species. Plants may have reduced seed production íf butte~ 1 ~!
and bees are less able to migrate among habitat fragments to pollina
flowers.
CREATION OF SMALLER POPULATIONS · nmaY
Habitat fragme~ta_tio ide-
precipitate population decline and extinction by dividing an ex~st inJ
~eas.
spread population into two or more subpopulations in r~ stnd~i:g de·
These smaller populations are then more vulnerable to mbree
Threats to Biodiversity 109
ression, genetic drift, and o~er problems associated with small popu-
p . n size (see Chapter 5). While a large area may support a single large
Iatioulation, it is possible that none of the smaller subpopulations in the
popments will be sufficiently large to persist, even if the total area is the
frage connecting the fragments with properly designed movement cor-
saJll · may be thk
ridors e ey to mam · tammg
· · popu1atlons.
·
rNTERSPECIES INTERACTIONS. Ha~itat fragment~tion increases the
vuinerability of the fragments to mvas1on by nonnatlve and native pest
·es (Aguirre-Acosta et al. 2014; Flory and Clay 2009). Road edges them-
5~5 may represent dispersa! routes for invasive species. The forest edge
:epresents a high-energy, high-nutrient, disturbed environment in which
manY pest specie~ of p!ants and animals can increase in number and then
disperse into the interior of the fragment.
omnivorous animals may increase in population size along forest
edges, where they can eat foods, including eggs and nestlings of birds,
from both undisturbed and disturbed habitats. In the coniferous forests
of Finland, predation on nests was found to be higher in edges of clear-cut
areas than in either the forest interior or forest corridors, and sometimes
higher than in the clear-cut area (Huhta and Jokimaki 2015). The invading
predators include not only native omnivores-such as raccoons, skunks,
and blue jays in North America-but also introduced domesticated cats.
In a study done in the southeastern United States, successful hunting cats
were found to capture an average of 2.4 prey items during 7 days of roam-
ing, with Carolina anoles (Anolis carolinensis) being the most common prey
species (Loyd et al. 2013). In rural Poland, an examination of both scat and
stomach contents and prey brought home by cats found mammals to be
the most common prey, followed by birds (Krauze et al. 2012).
Nest-parasitizing cowbirds, which live in fields and edge habitats, use
habitat edges as invasion points, flying up to 15 km into forest interiors,
where they lay their eggs in the nests of forest songbirds (Cox et al. 2012;
Lloyd et al. 2005). The combination of habitat fragmentation, increased
nest predation, and destruction of tropical wintering habitats is probably
r~sponsible for the dramatic decline of certain migratory songbird spe-
cies of North America, such as the cerulean warbler (Dendroica cerulea),
f art~cu~rly in the eastern half of the United States (Valiela and Martinetto
!,
00 With 2015). Populations of deer and other herbivores can also build
up In e~ge areas, where plant growth is lush, eventually overgrazing the
;ege_tation and selectively eliminating certain rare and endangered plant
pecies for severa! kilometers into the forest interior.
pr ~b_itat fragmentation also puts wild populations of animals in closer
to:
th oxuruty to domesticated animals. Diseases of domesticated animals can
cíes
spread more readily to wild species, which often have no immunity
There_ is also the potential for diseases to spread from wild spe-
omeshcated plants, animals, and even people, once the level of
r
11 O Chapter 4
contact increases. Fragmented forest habitats characteristic of suburb
development often have high ~ensit~es of_white-foot_ed mice and blac~
legged ticks with high rates of mfe~tion ~1th Lyme ~1~a~, along With a
corresponding increase in Lyme d1sease m people hvmg m those areas
(Tran and Waller 2013).
Edge effects
Habitat fragmentation greatly increases the ~~unt of edge relative to the
amount of interior habitat A simple example will illustrate these charactens.
tics and the problems they can cause. Considera square conservation reserve
1000 m (1 km) on each side (FIGURE 4.9). The total area of the reserve is 11cm2
(100 ha). The perimeter (or edge) of the reserve totals 4000 m. A point in the
middle of the reserve is 500 m from the nearest perimeter. If the principal
edge effect for birds in the reserve is predation by domesticated cats and
introduced rats, which forage 100 m into the forest from the perimeter of the
reserve and prevent forest birds from successfully raising their young, then
only the reserve's interior-64 ha-is available to the birds for breeding. lf
the reserve is divided into four fragments by a road and a railroad, each of
which is 495 m in area, the nesting habitat is further reduced to 8.7 ha in each
fragment, for a total of 34.8 ha. Even though the road and railroad remove
only 2% of the reserve area, they reduce the habitat available to the birds
by about half. The implications of edge effects can be seen in the decreased
(A) (B)
1000m
(1 km)
Interior= 64 ha
••
Railroad ll:tt!:om:om:om:tttlj:jtmlmttmmlI!lllii
---[Link]~\
1-----·1ooom-------i~
•• Road
Interior = 8.7 ha x 4 = 34.S ha
FIGURE 4.9 A hypothetical example shows how habitat area is reduced bY
fragmentation and edge effects. (A) A 1 krn2 protected area. Assuming that -~;i:
effects (gray) ~enetrate 1?0 m into the reserve, approximately 64 ha are av~ and
as usable hab1tat far nesting birds. (B) The bisection of the reserve by 8 roa th t
a railroad, although [Link] up little actual area extends the edge effects so Jya
'
almost half the breeding habitat is destroyed. Edge · ate
effects are proportion
greater when forest fragments are irregular in shape, as is usually the case.
Threats to Biodiversity 111
ability of birds to live and breed in small forest fragments com-
pared with larger blocks of forest habitat.
Habitat fragmentation
The microenvironment ata forest fragment edge is different
increases edge effects-
from that in the forest interior. Sorne of the more important dif-
changes in light,
ferences are greater fluctuations in levels of light, temperature,
humidity, temperature,
humidity, and wind (Laurance et al. 2011). These edge effects
and wind that may be
are most evident up to 100 m inside the forest, although certain
less favorable for many
effects are detectable up to 400 m from the forest edge (FIGURE
species living there.
4.10). Because so many plant and animal species are precisely
adapted to certain levels of temperature, humidity, and light,
changes in those conditions eliminate many species from for-
est fragments. Shade-tolerant wildflower species of temperate
forests, late-successional tree species of tropical forests, and
Increased wind disturbance
Elevated tree mortality
Altered species composition of leaf-litter invertebrates
Altered abundance and diversity of leaf-litter invertebrates
Altered height of greatest foliage density
Reduced canopy height
Faster recruitment of disturbance-adapted trees
1Q¡
.-. -.
i
¡;¡¡ Lower canopy-foliage density Emergent
Increased air temperature
Reduced understory-bird abundance
Lower relative humidity
Higher understory-foliage density
Lower soil-moisture content
Reduced density of fungal fruiting bodies
100 200 300 400 500
Edge penetration distance (m)
FIGURE 4.10 Edge effects in the Amazon rain forestas measured in the ex-
periment shown in Figure 4.8. The bars indicate how far into the forest fragment
the specified effect occurs. Far example, trees growing wíthin 300 m of an edge
have a higher mortality rate, and the average height of trees in the forest canopy
(see drawíng) is reduced wíthin 100 m of the edge. (After Laurance et al. 2002.)
_J
112 Cbapter 4
hum1'd'1ty-sens1•ti·ve [Link] as amphibians are often rapidly elift..
~11-
nated by habitat fragmentat10n. . . . . .
When a forest is fragmented, mcr~ased w~d, lower hum1~1ty, anct
higher temperatures make fires more likely <1:1narte et ~1 2012). Pires Illay
5
read into habitat fragments from nearby _agncultural f1el~s that are bein
!urned regularly, as in sugarcc . ~arvesttn?' or from the irregular életivf
ties of farmers practicing slash-é d ~um agnculture. Forest frasments Il1ay
be particularly susceptible to t ! amage when wood has accumulated
at the edge of the forest where t. •e have died or ha~e ?een blown down
by the wind. In Borneo and the _l azilian Amazon, ~1lho~s of hectares of
tropical moist forest bumed dunr, ·unusually dry p~ods m 1997 and 199s.
A combination of factors contrib ·ted to these environmental disaster .
forest fragmentation due to road e nstruction, farming, th~ accumulatio~
of brush following selective loggir ~, and human-caused fires.
Environmental Degradation and Pollution
Even when a habitat is unaffected by overt destruction or fragmentation
the ecosystems and species in that habitat can be profou?dly affected b;
human activities. Ecosystems can be damaged and spec1es driven to ex-
tinction by externa! factors that do not obviously change the structure of
the dominant plants or other features in the community. Sometimes this
damage and its cause(s) are quite obvious (FIGURE 4.11). For example,
keeping too many cattle in a grassland community gradually damages
it, often eliminating many native species and favoring invasive species
that can tolerate grazing and trampling. On the other hand, out of sight
from the public, fishing trawlers drag across an estimated 15 million km2
of ocean floor each year, an area 150 times greater than the area of forest
cleared in the same period. Toe trawling destroys delicate creatures such
as anemones and sponges, reduces species diversity and biomass, and
alters community structure (Hinz et al. 2009; Maynou and Cartes 2012).
Other types of environmental damage are not visually apparent even
though they occur all around us, every day, in nearly every part of the
world. The most subtle and universal form of environmental degradation
is pollution, commonly caused by pesticides, herbicides, sewage, fertiliz~rs
from agricultura! fields, industrial chemicals and wastes, em~-
.. sions from factories and automobiles, and sediment deposits
from eroded hillsides. Toe general effects of pollution on water
Pollution of the air, quality, air quality, and even the global climate are cause ~or
water, and soil by great concern, not only because of their threats to biodive1:5ity,,
chemicals, wastes, and but also because of their effects on human health and ag ricuol··
1
the by-products of energy ture (Dearborn and Kark 2010). Although environmental P e
production destroys . .
1ut10n . . in the cas
1s sometimes highly visible and dramatlc, as . are
habitats in insidious of massive oil spills, the subtle, unseen forros of pollutione 50
ways. probably the most threatening-primarily because they ar
.. insidious.