Ecology Presantation 5

13
CHAPTER
Interspecific
Competition
© 2016 Pearson Education, Ltd.
Elements of ECOLOGY Lecture Presentation by

NINTH EDITION, GLOBAL EDITION
Carla Ann Hass
Penn State University
Thomas M. Smith • Robert Leo Smith
Chapter 13 Interspecific Competition
§ Interspecific competition is a cornerstone of

evolutionary ecology
§ How does competition affect the species involved?
What type of interaction is it?
§ How does competition lead to species divergence
and specialization?

Section 13.1 Interspecific Competition
Involves Two or More Species
§ Interspecific competition affects the populations of
two or more species adversely (- -)
§ Intraspecific and interspecific competition may occur
simultaneously
§ What are examples of this?

§ Eastern deciduous forests in North America
§ In some years, oak trees produce few acorns
§ Gray squirrels compete among themselves for acorns
§ They also compete with other species for acorns
§ white-tailed deer, wild turkeys, white-footed mice, blue
jays
§ Competition for this limited resource may lead to
some of these species foraging for other food that is
less in demand

§ There are two forms of interspecific competition, just
as for intraspecific competitive interactions
§ exploitation
§ interference
§ Review – What is the difference between these?

§ Alternatively, most types of interspecific interactions
can be classified as one of six types
§ consumption
§ preemption
§ overgrowth
§ chemical interaction
§ territorial
§ encounter
§ What are the characteristics of each of these
interactions?

§ Consumption competition
§ Individuals of one species inhibit individuals of
another by consuming a shared resource
§ acorn example
§ Preemption competition
§ Individuals of one species prevent occupation of an
area by individuals of another species
§ sessile organisms, such as barnacles

§ Overgrowth competition
§ Individuals of one species grow over individuals of
other species, inhibiting access to a resource
§ taller plants shading shorter plants
§ Chemical interaction competition
§ Individuals of one species release growth inhibitors or
toxins that inhibit or kill other species
§ Allelopathy in plants – secretion of chemicals that
inhibit germination of other species

§ Territorial competition
§ Behavior of one species that excludes another
species from a specific location that is defended as a
territory
§ a bird keeping other birds from nesting in its territory
§ Encounter competition
§ Nonterritorial encounters between individuals of
different species affect one or more of the species
involved
§ scavengers fighting over a dead animal carcass

Section 13.2 The Combined Dynamics of
Two Competing Populations Can Be
Examined Using the Lotka-Volterra Model
§ Two mathematicians, Alfred Lotka and Vittora
Volterra, independently developed mathematical
expressions to model consumption competition
§ How is the Lotka-Volterra model related to the
logistic equation for population growth?
§ What does this model tell us about the relationship
between the two competing species?
§ What does the zero-growth isocline represent?

§ The logistic growth equations for the two species
are:
dN1/dt = r1N1(1 - N1/K1)
dN2/dt = r2N2(1 - N2/K2)
§ How do you modify these equations to model the

competitive effect of one species on the population
growth of the other?

§ Add a term that includes the population size of the
other species (N1 or N2)
§ A competition coefficient quantifies the
§ per capita effect of species 2 on species 1 (a)
§ per capita effect of species 1 on species 2 (b)
§ The competition coefficients are factors that convert
an individual of the competing species into the
equivalent number of individuals of the species
whose growth is being examined based on their
shared use of resources that define K
§ For species 1, aN2 accounts for the competitive
effect of species 2 on species 1
§ This equation models the population growth of
species 1
dN1/dt = r1N1 (1 - (N1 + aN2)/K1)
§ For species 2, 𝛽N1 accounts for the competitive
effect of species 1 on species 2
§ This equation models the population growth of
species 2
dN2/dt = r2N2 (1 - (N2 + bN1)/K2)
§ Two species of grazing herbivores feeding on the
same plants
§ Individuals of species 2 have twice the body mass of
species 1 and eat at twice the rate
§ One species 2 individual = two species 1 individuals
§ The impact of an individual of species 2 on species
1 would be a = 2.0
§ The impact of an individual of species 1 on species
2 would be b = 0.5

§ If there is no interspecific competition
§ Either a or N2 = 0
§ Either b or N1 = 0
§ The population of each species grows logistically to
equilibrium at K
§ What happens when there is interspecific
competition?

§ For species 1, as N1 approaches K1, population
growth (dN1/dt) approaches zero
§ Species 2 is also competing for the limited resource
that determines K1
§ The total effect of species 2 on species 1 is aN2
§ As the combined population N1 + N2 approaches K1,
the growth rate of species 1 approaches zero
§ the greater the population size of species 2 (N2) or
the greater the competitive effect (a), the greater the
reduction in the growth rate of species 1
§ To determine the possible outcomes of competition
using the Lotka-Volterra model, define the zero-
growth isocline for each species
§ The combined values of the population size for
species 1 (N1) and species 2 (N2) at which dN1/dt = 0
and dN2/dt = 0
§ This happens when the combined population sizes
are equal to K1 for species 1 or K2 for species 2

§ Defining the zero-growth isocline for species 1 using
a graphical approach
§ N1 on the x-axis
§ N2 on the y-axis
§ Solve for combined values of N1 and N2 when dN/dt
for both species = 0
§ For species 1
§ When (1 - (N1 + aN2)/K1) = 0
§ K1 = N1 + aN2
§ Define the line by solving for the x- and y-intercepts
§ x-intercept is N2 = 0, so N1 = K1
§ y-intercept is N1 = 0, so N2 = K1/a

Figure 13.1a
K1/a
dN1/dt = 0
N2
dN1/dt < 0
dN1/dt > 0
0 K1
(a) N1
§ For combinations of (N1, N2)
§ Below the line, N1 + aN2 < K1
§ Species 1 growth rate will be positive
§ Above the line, N1 + aN2 > K1
§ Species 1 growth rate will be negative
§ Lines are parallel to the x-axis (species 1 axis)

Figure 13.1a
K1/a
dN1/dt = 0
N2
dN1/dt < 0
dN1/dt > 0
0 K1
(a) N1
§ Defining the zero-growth isocline for species 2 using
a graphical approach
§ N1 on the x-axis
§ N2 on the y-axis
§ Solve for combined values of N1 and N2 when dN/dt
for both species = 0
§ For species 2
§ When (1 - (N2 + bN1)/K1) = 0
§ K2 = N2 + bN1
§ Define the line by solving for the x- and y-intercepts
§ x-intercept is N2 = 0, so N1 = K2 /b
§ y-intercept is N1 = 0, so N2 = K2

Figure 13.1b
K2
dN2/dt = 0
N2
dN2/dt < 0
dN2/dt > 0
0 K2/b
(b) N1
§ For combinations of (N1, N2)
§ Below the line, N2 + βN1 < K2
§ Species 2 growth rate will be positive
§ Above the line N2 + βN1 > K1
§ Species 2 growth rate will be negative
§ Lines are parallel to the y-axis (species 2 axis)

Figure 13.1b
K2
dN2/dt = 0
N2
dN2/dt < 0
dN2/dt > 0
0 K2/b
(b) N1
Section 13.3 There Are Four Possible
Outcomes of Interspecific Competition
§ To interpret the combined dynamics of the two
competing species, the isoclines are drawn on the
same graph
§ What are the four possible outcomes of
competition?
§ How is each represented graphically?

§ Species 1 “wins” over species 2
§ The two isoclines are parallel, and the species 1
isocline is completely above that for species 2
§ This defines three areas in the graph
§ Point A – below the zero-growth isoclines for both
species
§ Both species have a positive growth rate
§ Point B – above the zero-growth isoclines for both
species
§ Both species have a negative growth rate

§ Point C – between the zero growth isoclines for
species 2 and species 1
§ Species 1 has a positive growth rate and species 2
has a negative growth rate
§ Eventually, species 1 will go to K1 and species 2 will
become extinct

Figure 13.2a
K1/a
B
K2
N2
A
0 K2/b K1
N1
(a)
§ Species 2 “wins” over species 1
§ The two isoclines are parallel, and the species 2
isocline is completely above that for species 1
§ This defines three areas in the graph
species
species

§ Species 1 has a negative growth rate and species 2
has a positive growth rate
§ Eventually, species 2 will go to K2 and species 1 will
become extinct

Figure 13.2b
K2
B
K1/a
N2
A
0 K1 K2/b
N1
(b)
§ Species 1 and species 2 coexist
§ The two isoclines cross at an equilibrium point
§ A combined value (N1, N2) at which the growth of both
species is zero
§ This defines four areas in the graph
species
species
§ Point D – between the zero growth isoclines for

§ The combined dynamics (black lines) for all four
areas point to the center where the isoclines meet
§ This point represents a stable equilibrium at which
the two species coexist

Figure 13.2c
K1/a
B
D
N2
K2
E
A C
0 K1 K2/b
N1
(c)
§ Either species 1 or species 2 will “win”
§ Which one depends on initial population sizes (N1, N2)
and growth rates of each population (r1, r2)
§ The two isoclines cross at an equilibrium point
§ a combined value (N1, N2) at which the growth of both
species is zero
§ This defines four areas in the graph
§ Point A – both species have a positive growth rate
§ Point B – both species have a negative growth rate

§ Point D – between the zero growth isoclines for

§ The combined dynamics (black lines) for points A
and B point to the center where the isoclines meet
§ The combined dynamics for points C and D point
away from the center, moving toward K1 and K2
§ This point represents an unstable equilibrium
§ One species will reach K and the other will go
extinct

Figure 13.2d
K2
B
N2
K1/a D
E
C
A
0 K2/b K1
N1
(d)
Section 13.4 Laboratory Experiments
Support the Lotka-Volterra Model
§ Why were laboratory experiments preferred over
field experiments for the first studies performed to
test the predictions of the Lotka-Volterra competition
model?

§ Laboratory experiments were preferred because it is
easier to determine the outcome of competition
under controlled conditions
§ Gause investigated competition between two
Paramecium species, P. aurelia and P. caudatum
§ P. aurelia has a higher rate of population growth and
can tolerate higher population density
§ Placed each species separately in a single tube with
a fixed amount of bacterial food
§ Both survived for at least 20 days

§ Placed both species together in a single tube with a
fixed amount of bacterial food
§ What were the results?

Figure 13.3
Population density measured by volume
Separately
200
P. aurelia
150
100
In mixed population
50
200
P. caudatum
150 Separately
100
In mixed population
50
0 2 4 6 8 10 12 14 16 18 20 22 24
Days

§ Placed both species together in a single tube with a
fixed amount of bacterial food
§ P. aurelia survived, while P. caudatum died out

Figure 13.3
Population density measured by volume
Separately
200
P. aurelia
150
100
In mixed population
50
200
P. caudatum
150 Separately
100
In mixed population
50
0 2 4 6 8 10 12 14 16 18 20 22 24
Days

§ Gause then performed similar experiments using P.
caudatum and a different Paramecium species, P.
bursaria
§ These species coexisted because of differences in
feeding behavior
§ P. caudatum fed on bacteria suspended in solution
§ P. bursaria fed on bacteria at the bottom of the tube

§ Clark investigated competition between two species
of Tribolium (flour beetles), T. castaneum and T.
confusum
§ The outcome of competition was variable and
depended on the environmental temperature and
humidity, as well as fluctuations in the total number
of eggs, larvae, pupae, and adults
§ It could take many generations to determine the
outcome

§ Tilman investigated competition between the
diatoms Asterionella formosa and Synedra ulna
§ Both must have silica to form cell walls
§ Population growth and decline were monitored, as
well as the level of silica in the water
§ When grown separately with silica continually added,
both species kept silica at a low level
§ When grown together, silica use by S. ulna reduced
the concentration below the level needed by A.
formosa for survival and reproduction

§ Asterionella formosa was driven to extinction

Figure 13.4
105
24°C 30
Af
104
Population density
20
Silicate (µm)
103
10
102
Si
101 0
0 10 20 30 40 50
(a) Time (days)
105 24°C
30
104
Population density
20
Silicate (µm)
Su
103
10
102
Si
101 0
0 10 20 30 40 50
(b) Time (days)
105 24°C
30
104
Su
Population density
20
Silicate (µm)
103
Af 10
102
Si
101 0
0 10 20 30 40 50
(c) Time (days)
Section 13.5 Studies Support the
Competitive Exclusion Principle
§ In the Lotka-Volterra model, three of the four
possible outcomes result in the extinction of one
species
§ What is the competitive exclusion principle?
§ What is meant by the term “complete competitors”?

§ The competitive exclusion principle states that
complete competitors cannot coexist
§ Complete competitors are two distinct species that
live in the same place and have exactly the same
ecological requirements
§ What assumptions does competitive exclusion
require?

§ Competitive exclusion requires that
§ The competitors require exactly the same resources
§ Environmental conditions remain constant
§ The conditions rarely exist for this to occur, but this
principle raises questions, including
§ How similar can two species be and still coexist?
§ What ecological conditions are necessary for species
that share a common resource base to coexist?

§ Research suggests that many factors affect the
outcome of interspecific competition
§ environmental factors that influence the survival,
growth, and reproduction of a species (but are not
resources that are consumed)
§ temperature, pH
§ spatial and temporal variation in resource availability
§ competition for multiple limiting resources
§ resource partitioning

Section 13.6 Competition Is Influenced by
Nonresource Factors
§ Environmental features that are not resources can
influence the outcome of competition between
species
§ How can nonresource factors influence competition?

Nonresource Factors
§ The effect of water temperature on competitive
ability of three species was examined in both field
and laboratory experiments
§ Three species from Rocky Mountain streams
§ brook trout – most abundant at high elevations
§ brown trout – most abundant at middle elevations
§ creek chub – most abundant at lower elevations
§ Interference competition is known to influence the
relative success of these species when they live
together

Nonresource Factors
§ Fish from each species were gradually acclimated to
seven different test temperatures ranging from 3 to
26°C
§ Fish from each species were matched by size (less
than 10 percent difference) and placed in an
experimental stream together
§ The species that consumed the highest number of
food items was competitively superior
§ How did water temperature affect competitive
ability?

Based on the results from the experiments shown in
this figure, the fish from the middle elevations are most
competitive at
A. the highest water temperature.

B. the moderate water temperatures.
C. the lowest water temperature.
D. all water temperatures.

Figure 13.5
12
Brook trout
10 Brown trout
Mean number of food items
Creek chub
0
0 4 8 12 16 20 24 28
Water temperature (°C)

Based on the results from the experiments shown in
this figure, the fish from the middle elevations are most
competitive at
A. the highest water temperature.

B. the moderate water temperatures.
C. the lowest water temperature.
D. all water temperatures.

Nonresource Factors
§ Another example of a nonresource factor influencing
competitive ability is seen in the tadpoles of two
species of tree frogs
§ Hyla gratiosa and Hyla femoralis have broadly
overlapping geographic distributions
§ They respond differently to water acidity

Nonresource Factors
§ At lower water pH (4.5), interspecific interactions
were minimal
§ At higher water pH (6.0), H. femoralis was a better
competitor, causing decreased survival and an
increased larval period for H. gratiosa
§ The H. gratiosa that did metamorphose did so at a
smaller size

Section 13.7 Temporal Variation in the
Environment Influences Competitive Interactions
§ How can variation in environmental conditions over
time affect competitive interactions?

§ A species that is more efficient at exploiting a
shared, limiting resource may exclude other species
§ But when environmental conditions are variable over
time, the competitive advantages of species may
change
§ If this is continuous, no one species may reach
sufficient density to displace its competitors
§ Environmental variation allows competitors to
coexist when one would exclude the other under
constant conditions

§ Study of grass species living in a savanna
community in southwest Zimbabwe, Africa
§ Over a 10-year period, the dominant grass species
shifted from Urochloa mosambicensis to
Heteropogon contortus
§ What caused this shift?

Figure 13.6a
80
Heteropogon
70
contortus
Percent occurrence
60
50
40
30
20 Urochloa
10 mosambicensis
1971/72 72/73 73/74 74/75 75/76 76/77 77/78 78/79 79/80 80/81
Year (rainy season)
(a)

§ This shift was a result of yearly variations in rainfall
§ The 1971–1972 and 1972–1973 rainy seasons had
much lower than average rainfall
§ U. mosambicensis can survive and grow better in
dry conditions than can H. contortus
§ When rainfall returned to average levels, H.
contortus became the dominant species

Figure 13.6b
1000
900
800
700
Rainfall (mm)
600
500
400
300
200
100
0
1971/72 72/73 73/74 74/75 75/76 76/77 77/78 78/79 79/80 80/81
Year (rainy season)
(b)

§ A similar pattern was observed at a prairie grassland
site at Hays, Kansas in the Great Plains
§ The role of climate variability on the relative
abundance of different prairie grasses was studied
from 1937 to 1968
§ Climate variation from one year to the next
correlated with variation in species performance
§ Changing competitive ability as a result of a
changing environment reduced the incidence of
competitive exclusion

§ Climate can also have a density-independent effect
on population growth
§ Periods of extremes (drought, flooding, extreme
temperatures) may reduce population size below
carrying capacity
§ If this happens frequently relative to the time needed
for recovery (approaching K) resources may be
abundant enough to reduce or eliminate competition

Section 13.8 Competition Occurs for
Multiple Resources
§ Competition between species often involves multiple
resources
§ Competition for one resource may affect an
organism’s ability to use other resources
§ Why is interspecific territoriality often an example of
multiple resource competition?

Multiple Resources
§ Many birds in temperate and tropical communities
exhibit interspecific territoriality
§ Some species defend territories against closely
related species
§ gray and dusky flycatchers in the western United
States
§ Some species defend territories against a broad
range of competitors
§ Acorn woodpeckers defend against other
woodpeckers, blue jays, and squirrels

Multiple Resources
§ In plants, there are many examples showing how
competition for one resource can influence the
ability of an individual to exploit other resources,
affecting both survival and growth
§ Both greenhouse and field experiments have been
performed
§ Greenhouse experiments – competition between
subterranean clover and skeletonweed

Multiple Resources
§ Plants were grown in the greenhouse in
§ monocultures – single populations
§ in pots so leaves and roots intermingled
§ mixtures – two populations
§ in the same pot; leaves and roots intermingled
§ in the same pot; roots intermingled, leaves separate
§ in separate pots; leaves intermingled, roots separate
§ The effects of competition were determined for
§ aboveground (light) resources
§ belowground (water and nutrients) resources

Figure 13.7
Shoots of the two species

are separated by a divider;
root systems are allowed Individuals of the two species
to access the same share both above- and below-
Individuals of the two species are
soil volume in the pot. ground resources.
grown in separate pots, eliminating
interspecific root competition, while
the shoots share access to the same
aboveground space and resources.
-35% -53% -69%
Skeletonweed Subterranean clover Root Shoot Root and shoot

Grown alone Grown alone competition competition competition

Multiple Resources
§ Results showed that clover was not significantly
affected by the presence of skeletonweed
§ Skeletonweed was adversely affected under all
three conditions when the plants were grown
together
§ What does this tell you about resource competition
between these two species?

Multiple Resources
§ The species are competing for both aboveground
and belowground resources
§ Compared to skeletonweed grown in monoculture,
when the two species were grown together, effects
on skeletonweed biomass were as follows:
§ roots intermingled – reduced by 35 percent
§ canopies intermingled – reduced by 53 percent
§ both intermingled – reduced by 69 percent
§ Clover plants were superior competitors for light,
water, and nutrients

Multiple Resources
§ Field studies in an old field grassland community in
Pennsylvania
§ Experimental design similar to greenhouse study
§ PVC root exclusion tubes were used to control root
interaction by varying the number of holes drilled in
the tube
§ tied back neighboring vegetation to vary light level
§ created 16 combinations of above- and belowground
competition
§ Control plants were completely isolated from
neighboring plants – no competition

Figure 11.20
15
Increasing root competition
Neighbor root biomass (g)
10
0
0 12.5 25 50 100
(a) Percent of tube open to neighbors
Decreasing plant growth

1.0
Target plant biomass (g)
0.5
0
0 12.5 25 50 100
(b) Percent of tube open to neighbors
Multiple Resources
§ Results showed a clear pattern of interaction
between aboveground and belowground competition
§ Increased competition for belowground resources
reduces growth rate and plant height
§ This reduction leads to a reduction in competitive
ability for light (aboveground resource)

Section 13.9 Relative Competitive Abilities
Change along Environmental Gradients
§ As environmental conditions change, the relative
competitive abilities of species change
§ What can cause these shifts in competitive ability?

§ Shifts in competitive ability can result from
§ changes in the carrying capacity for a species
because of a change in the resource base
§ changes in the physical environment that interact with
resource availability
§ These changes in competitive ability along
environmental gradients have been examined for
both plants and animals

§ For plants, the outcomes of competition across
gradients of resource availability have been
examined in both the field and the laboratory
§ Greenhouse studies in Australia have investigated
the changes in interspecific competition among plant
species across experimental gradients of nutrient
availability
§ One study examined six species of thistles
§ The gradient was established by applying a nutrient
solution

§ Plants were grown in
§ monoculture – single species
§ mixture – all six species grown together
§ Eleven nutrient treatments ranging from 1/64 to 16
times the recommended concentration of a standard
nutrient solution
§ After 14 weeks, plants were harvested and dry
weights obtained
§ Do you expect a difference between plants grown in
monoculture and mixed culture? Why?

§ Plants grown in mixed culture showed much
variation, and the response was different than the
response in monoculture
§ Interspecific competition affected growth for each
species
§ The competitive abilities changed along the nutrient
gradient

Figure 13.8b
10
Carthamus lanatus
9 Carduus pycnocephalus
Onopordum
Cirsium vulgare
8 Carduus nutans
Silybum marianum
Total biomass per plant (loge g)

7
2 Mixture
1
1/64 1/16 1/4 1 4 16
Relative nutrient concentration
(b)
§ Examine the relative response of species in the
mixed-species experiments
§ Divide the biomass value for a particular species at
a given nutrient level by the biomass value of the
species that has the highest biomass at that nutrient
level (the best competitor)
§ Range is 0 (worst competitor) to 1.0 (best
competitor)

Examining the relative responses of the three
dominant thistle species in the experiment along the
nutrient gradient, Silybum marianum is the best
competitor at the
A. lowest nutrient concentrations.

B. moderate nutrient concentrations.
C. highest nutrient concentrations.
D. There is no consistent pattern for competitive ability.

Figure 13.9
Silybum
1.0 marianum
Normalized ecological
0.8
performance
0.6 Carduus
pycnocephalus
0.4
0.2 Carthamus
lanatus
0
0 1/64 1/16 1/4 1 4 16
Relative nutrient concentration
Examining the relative responses of the three
dominant thistle species in the experiment along the
nutrient gradient, Silybum marianum is the best
competitor at the
A. lowest nutrient concentrations.

B. moderate nutrient concentrations.
C. highest nutrient concentrations.
D. There is no consistent pattern for competitive ability.

§ Field experiments explored the trade-offs in
competitive ability among five perennial species of
C4 grasses at different levels of soil fertility and
disturbance
§ Soil fertility was altered by applying different levels of
fertilizer
§ Disturbance resulting from herbivore grazing was
simulated by clipping at different levels
§ Grasses were grown in both monoculture and
mixtures at each treatment level

§ Competitive ability measured by response ratio
changed along soil fertility and disturbance gradients

Figure 13.10
Under conditions of high soil

phosphorus, the relative
Under conditions of low soil competitive abilities of the
phosphorus, Themeda is the two species reverse, and Aristida
superior competitor having a is the superior competitor.
greater overall biomass.
1.4 Themeda triandra

Aristida junciformis
1.2
In response ratio (ratio +1)
1.0
0.8
0.6
0.4
0.2
0.0
0 5 10 15 20 25
Soil phosphorus (mg/kg)
§ Often, multiple environmental factors interact to
affect the response of organisms
§ Field studies of cordgrass species in New England
salt marshes
§ Low marsh habitats are frequently flooded
§ High marsh habitats are less frequently flooded
§ Changes along this gradient include
§ nutrient availability
§ physical stress related to waterlogging, salinity, and
oxygen availability in soil/sediments

§ Plant zonation follows this difference
§ Spartina alterniflora is found in low marshes
§ Spartina patens is found in high marshes
§ Experiments involved
§ Transplanting S. patens to low marshes
§ Transplanting S. alterniflora to high marshes
§ Results
§ S. patens was stunted with or without competition
§ S. alterniflora grew well in the high marsh without
competition, but were excluded when S. patens was
present
§ S. patens is limited to the high marsh; it cannot
tolerate the environmental conditions in the low
marsh
§ S. alterniflora can live in both but is excluded from
the high marsh by S. patens

Figure 13.11
Upper boundary of Lower boundary of S. patens 20

S. alterniflora determined determined by low tolerance Control
by competition from S. patens to anoxic condition
Biomass (g/100 cm2)

With competition
15
Without competition
10
0
Low marsh High marsh
(b) Spartina patens control and transplants
Spartina alterniflora Spartina patens 20

zone zone
(a)
Biomass (g/100 cm2)

15
10
0
(c) Spartina alterniflora control and transplants

§ Four species of chipmunks live on the eastern slope
of the Sierra Nevada mountains
§ Four species with overlapping food requirements;
each lives in a different altitudinal zone
§ alpine chipmunk
§ lodgepole chipmunk
§ yellow-pine chipmunk
§ least chipmunk
§ What factors determine the distribution of these
species?

Figure 13.12
Alpine chipmunk
Alpine zone
Lodgepole chipmunk
Lodgepole pine zone
Yellow-pine chipmunk
Least chipmunk
Piñon pine, sagebrush zone
Sagebrush zone

§ The contact zones are determined, in part, by
aggression between species
§ The yellow-pine chipmunk is dominant over the least
chipmunk, which can live in all of the habitats on the
slope but is restricted to sagebrush
§ It tolerates heat stress better than the other species
§ If the yellow-pine chipmunk is removed, the least will
move into its habitat
§ If the least chipmunk is removed, the yellow-pine will
not move into its habitat

§ The lodgepole chipmunk is dominant over the
yellow-pine chipmunk, determining its upper range
§ The lodgepole chipmunk is susceptible to heat stress,
so it only lives in the lodgepole pine shaded forest
habitat
§ The lodgepole chipmunk is very aggressive and
likely determines the lower range of the alpine
chipmunk
§ The range of each species is determined by
aggressive exclusion and its ability to survive and
reproduce in the different environments

Quantifying Ecology 13.1 Competition
under Changing Environmental Conditions:
Application of the Lotka-Volterra Model
§ In the Lotka-Volterra model, two factors interact to
influence the outcome of competition
§ competition coefficients (a and b)
§ carrying capacities (K1 and K2)
§ How do changes in the environment affect carrying
capacity and influence competition?

§ The outcome of interspecific competition reflects the
relative abilities of the competing species to gain
access to and acquire resources essential for
survival, growth, and reproduction
§ If the environment changes, this can influence
resource availability, thus influencing the relative
carrying capacity for the competing species

§ Two species (1 and 2) use the same limiting food
resource – seeds
§ The two species have a broad, symmetrical overlap
in their diet

Quantifying Ecology Box Figure 1a
Proportion of diet
1 2
(a) Seed size (g)

§ If the rate of food intake (seeds eaten per unit time)
is the same, assume the same competition
coefficients, a = b = 0.5
§ Assume that the abundance and size distribution of
seeds vary with environmental conditions
§ Three environments – A, B, C
§ Seed size varies in each environment, increasing
from A to B to C

Quantifying Ecology Box Figure 1b
Abundance (number per m 2)

A B C
Seed size (g)

(b)

§ As the size distribution of seeds changes in each
environment, K changes for each species in each
environment
Environment
A B C
Species 1 (K) 225 150 75
Species 2 (K) 75 150 225

§ Examine the change in competition between the two
species in these three different environments
§ Use the Lotka-Volterra equations and zero-growth
isocline graphical analysis
Environment K1 K1/α K2 K2/β
A 225 450 75 150
B 150 300 150 300
C 75 250 225 450

§ The outcomes of competition are different in the
three different environments
§ What factors led to these different outcomes?

Quantifying Ecology Box Figure 2
500 500 500

K1/a
Environment A Environment C
400 400 Environment B 400
Species 1 wins Species 2 wins

300 K1/a 300 Species 1 & 2 300
coexist
K2
200 200 200
K2 K1/a
100 100 100
K2
0 0 0
100 200 300 400 500 100 200 300 400 500 100 200 300 400 500
(a) K2/b K1 (b) K1 K2/b (c) K1 K2/b

Influences the Niche of a Species
§ The ecological niche of a species is the range of
physical and chemical conditions under which it can
survive and reproduce
§ The fundamental niche is the ecological niche when
there are no interactions with other species
§ How can competition restrict the fundamental niche
of a species?

§ The realized niche is the part of the fundamental
niche that a species actually exploits as a result of
interactions with other species
§ Competition can lead a species to restrict its use of
resources such as space or food
§ Many studies have demonstrated that the
fundamental niche contracts when a competitor is
present

§ What role does competition play in the distribution of
Stipa neomexicana, a C3 perennial grass that is part
of semiarid grassland communities in southeastern
Arizona?
§ Stipa is found on dry ridge crests with low grass
cover
§ It does not occur in moist, low-lying areas with
greater grass cover
§ Experiment – neighboring plants were removed from
individual Stipa plants in three habitats
§ ridge-crest, midslope, lower-slope
§ Control – neighboring plants were not removed
§ Stipa has a higher growth rate, more flowers per
plant, and greater seedling survival on mid- and low-
slopes
§ It is usually not found in those habitats because it is
excluded by competition with other grasses
§ Its realized niche is only the suboptimal habitat

Figure 13.13
Removal Control
1.0 45
0.8 40
0.6
35
0.4 30
Ridge crest
25
0.2
Ridge crest
0.1 40
Midslope
Number of flowers per plant

1.0 35 Removal
Seedling survivorship, lx
70 Control
Mean basal area (cm2)

0.8 30
0.6 60
25 50
0.4 20
Midslope 40
30
0.2
20
55
10
0.1 50
Lower slope 0
1.0 45 Ridge Mid- Lower
0.8 crest slope slope
40
0.6 (c)
35
0.4 30
Lower slope 25
0.2 20
15
0.1
0 4 8 12 16 24 Jan May Aug Oct Jan May Aug
Age (months) 80 81
May Sep Jan May Sep May Date
80 81 82
(a) (b)
§ Competitive release – the expansion of a species
niche when a competitor is no longer present
§ This may happen when a species
§ invades a habitat that is free of potential competitors
§ is removed from a community, allowing the remaining
species to move into habitats that they had not been
able to occupy
§ This has been demonstrated through experiments

§ Competitive release in a lake ecosystem
§ Changes in the feeding niche of the three-spine
stickleback were looked for by manipulating the
presence or absence of two competitors
§ juvenile cut-throat trout – feed at the surface and in
the water column
§ prickly sculpin – benthic feeder
§ Sticklebacks feed in both habitats
§ Stomach content analysis shows the diets of these
three species overlap

§ Twenty experimental enclosures in Blackwater Lake,
Vancouver Island (five replicates of four enclosures)
all contained sticklebacks
§ competition – sculpin and trout
§ release from sculpin competition – trout only
§ release from trout competition – sculpin only
§ total release – sticklebacks only
§ The experiments ran undisturbed for 15 days
§ Then sticklebacks were removed and the prey in
their stomachs were counted and identified

§ Diversity of prey in stomach is an indicator of niche
breadth of the stickleback
§ release from sculpin – no change in niche breadth
§ release from trout – niche breadth expanded

Figure 13.14
1.8
Total population niche width 1.7
1.6
1.5
1.4
1.3
Trout No trout
Section 13.11 Coexistence of Species Often
Involves Partitioning Available Resources
§ All plants in grassland require light, water, and
essential nutrients
§ All large mammalian herbivores in a grassland
require plants to eat
§ How can these competitors coexist in the same
community?
§ How different must two species be in their resource
use to prevent competitive exclusion?

§ Observations of similar species living in the same
habitat suggest that they are able to coexist
because they partition the available resources
§ plants use nutrients in different proportions and
tolerate different levels of light
§ Each species uses a portion of the resource not
used by the other it lives with

§ Field studies provide many examples that represent
apparent resource partitioning
§ Three species of annual plants grow together on
prairie soil abandoned a year after plowing
§ These species vertically partition the environment,
resulting in different use of resources

§ Bristly foxtail – fibrous, shallow roots access a
variable water supply
§ recovers rapidly from drought, takes up water quickly
after rain, can photosynthesize when partly wilted
§ Indian mallow – sparse branched taproot accesses
medium soil depths
§ more water early in growing season, can
photosynthesize at low water availability
§ Smartweed – taproot moderately branches in upper
layer, penetrates deepest into soil
§ continuous supply of moisture from deep soil
Figure 13.15
10
20 Bristly foxtail
Setaria faberii
30
A horizon
40
B horizon
Depth (cm)
50
60 Indian mallow
Abutilon theophrasti
70
80
Smartweed
90
Polygonum pensylvanicum

§ Apparent resource partitioning is also seen in
animals living in the same habitat and using similar
resources
§ Small wild cats in the Middle East
§ Variation among species in the size of canine teeth
§ There is a general relationship between the size of
the canine teeth and the preferred prey species
§ There is also sexual dimorphism in this species, with
females’ canines generally larger than males’ (reflects
intraspecific competition)

§ Plotting the difference in tooth size gives a very
regular pattern with little overlap (x-axis), suggesting
that selection has reduced the overlap of tooth size
and thus the size of prey taken

Figure 13.16
F. silvestris
F. silvestris
F. chaus
F. caracal
F. chaus
F. caracal
4 5 6 7 8 9
Diameter of canine teeth (mm)

§ The morphological, physiological, and behavioral
changes are adaptations allowing resource
partitioning among species that coexist
§ These patterns of resource partitioning are
consistent with the hypothesis of phenotypic
divergence arising from coevolution between
competing species
§ This would happen over a long period of time
§ But difference among species may also reflect
adaptations that enable a species to use resources
of life in an environment independent of competition
§ How can you study competition that happened in
the past with little information about the competitors
and resource availability that influenced natural
selection?
§ An ecologist has referred to this hypothesis, resource
partitioning as a product of coeveolution between
competitors, as the “ghosts of competition past”
§ The role of past competition cannot be directly
observed
§ Is there a way to evaluate if this has happened in a
population?
§ Examine difference in characteristics of
subpopulations of a species that are in different
competitive environments
§ Peter and Rosemary Grant’s studies of Galápagos
ground finches
§ Two species
§ Medium ground finch – Geospiza fortis
§ Small ground finch – Geospiza fuliginosa
§ They feed on an overlapping array of seed sizes

§ These two species are found together on Santa
Cruz Island
§ On that island, the distribution of beak sizes in the
two species does not overlap

Figure 13.17a
Santa Cruz
Frequency
6 9 12 15
(a) Beak depth

§ These two species live separately on two islands
§ G. fortis on Daphne Major
§ G. fuliginosa on Los Hermanos
§ In those populations, the distribution of beak sizes in
the two species overlaps

Figure 13.17a
Santa Cruz
Frequency
6 9 12 15
(a) Beak depth

§ What does this suggest about natural selection and
beak size in the populations that live together versus
those that live separately?

Figure 13.17
Santa Cruz
Frequency
6 9 12 15
(a)
Daphne Major
Frequency
6 9 12 15 G. fortis
(b)
Los Hermanos
Frequency
G. fuliginosa
6 9 12 15
(c) Beak depth
§ On Santa Cruz, where the two species coexist, there
is competition for food (seeds)
§ Natural selection has favored
§ G. fortis individuals with larger beaks that can handle
and eat larger seeds
§ G. fuliginosa individuals with smaller beaks that can
handle and eat smaller seeds
§ The outcome of competition has been a shift in their
feeding niches through a change in beak size in
each species

§ Character displacement is this shift in a
morphological, behavioral, or physiological character
as a result of niche partitioning due to competition
§ This now has been directly observed in G. fortis on
Daphne Major
§ It was the only species of ground finch on that island
until 1982
§ That year, the large ground finch, Geospiza
magnirostris, emigrated from an adjacent island

§ The diet of these two species overlaps
§ G. magnirostris eats mainly the hard seeds of
Jamaican feverplant
§ Large-beaked G. fortis can also feed on these
§ The G. magnirostris population was small at first so
had little competitive effect
§ This changed in 2003
§ The population of G. magnirostris was larger
§ There was a drought in 2003 and 2004 that reduced
the seeds available

§ G. magnirostris depleted the supply of Jamaican
feverplant seeds
§ The G. fortis population had to depend on the
smaller seeds for food
§ This exerted strong directional selection, and the
average beak size in G. fortis declined
§ This is clear evidence that character displacement is
a coevolutionary process that can take place
through competitive interactions

Section 13.12 Competition Is a Complex
Interaction Involving Biotic and Abiotic Factors
§ Demonstrating competition in the field is more
challenging than demonstrating competition in the
lab
§ Why is this statement true?

§ Under natural conditions in the field, researchers
§ have little control over the environment
§ find it difficult to determine if a population is at or
below carrying capacity
§ lack complete knowledge of the life history
requirements of the species or subtle differences
between species
§ What types of effects can removal experiments
have?

§ Removal experiments seem to be straightforward
§ Remove a potential competitor; monitor results
§ Expect clear evidence of competition
§ But removing individuals of a species can have
direct and indirect environmental effects
§ Removing plants can
§ increase the light energy reaching the soil surface,
soil temperatures, and the evaporation rate
§ reduce soil moisture and increase decomposition rate
§ This will affect belowground resources

§ Competition is a complex interaction
§ that seldom involves two species interacting for a
single resource
§ that involves a variety of environmental factors that
vary in time and space
§ for which the outcome under one set of environmental
conditions can be different than the outcome under a
different set

Ecological Issues & Applications: Is Range
Expansion of Coyote a Result of
Competitive Release from Wolves?
§ How have human activities changed the ranges of
the gray wolf and the coyote?
§ Are there competitive interactions between wolves
and coyotes?
§ How did the loss of these competitive interactions
historically impact the range of the coyote?

§ Before European settlement, the gray wolf (Canis lupus)
ranged from the Atlantic to the Pacific coast, from Alaska
to northern Mexico
§ It occurred in all North American habitats
§ Wolves preyed on bison, deer, elk, and moose
§ As their prey declined, they preyed on sheep and
cattle
§ Bounties were paid to kills wolves starting in 1630
§ The last wolf in the Northeast was killed in 1897
§ Wolves were protected under the Endangered Species
Act (ESA) beginning in 1973
Figure 13.18
Gray wolf
Occupied wolf range
Historical wolf range
§ Before European settlement, the coyote (Canis
latrans) was found only in western North America
§ As settlers moved westward, coyotes were not seen
as a threat by famers and ranchers
§ Human habitat alteration (agriculture and logging)
changed the eastern habitat, and the coyote range
expanded eastward
§ Coyotes are now found from the Atlantic almost to
the Pacific, from Alaska to central Mexico

§ Are the wolf decline and the coyote expansion
linked?

Figure 13.19
Alaska
Canada
0 500
Pacific
Ocean Vancouver Miles
1900- 1950
1880- 1930 Chicago

New York
Historical Washington
range
Santa
Barbara 1940- 1990
Atlantic
Ocean
Gulf of
Mexico
Coyote range Mexico

Before 1700
Early 1900s
Today
Expansion route

§ Interference competition occurs between wolves and
coyotes
§ Wolves limit resource access by coyotes through
direct aggression
§ Coyotes are excluded from wolf territories
§ Wolves kill coyotes
§ Did eradication of gray wolves over most of their
historic range reduce the competitive pressures that
limited coyotes to their historic range?
§ Is coyote range expansion a result of competitive
release?
§ As a result of conservation efforts and the ESA
listing, gray wolf populations are increasing in the
United States
§ There are currently populations in Minnesota,
Wisconsin, and the Upper Peninsula of Michigan
§ Wolves have been reintroduced in Montana, Idaho,
and Wyoming
§ Some of these are areas with coyote populations,
allowing researchers to examine the effect of
competition on these two carnivores
§ Researchers collected data on wolf and coyote
populations in the Northern Rocky Mountains
§ data on coyote cause-specific mortality and survival
rates in wolf-free and wolf-abundant sites in Grant
Teton National Park (GTNP)
§ data on population densities for both species at three
areas across the Greater Yellowstone Ecosystem
(GYE)
§ If wolves are present, how does that affect coyote
populations?

§ Coyotes were numerically dominant across the GYE
§ However, density varied spatially and temporally
§ This variation was related to wolf abundance
§ At wolf-abundant sites in GTNP, coyote densities
were 33 percent lower
§ In Yellowstone National Park, after wolf
reintroduction, coyote densities declined 39 percent

§ These data support that competition with wolves
limits coyote populations

Figure 13.20
1.00
0.80
Density (coyotes per km2)
0.60
0.40
0.20
0
0 0.02 0.04 0.06 0.08
Density (wolves per km2)
§ What type of competitive interactions are seen
between the two species?
§ Overall, mortality of coyotes as a result of wolf
predation was low
§ lower for resident coyotes in packs defending
territories
§ higher for transient coyotes
§ Wolves responsible for 56 percent of deaths
§ Dispersal rates were also much higher for transient
coyotes in wolf-abundant sites

§ Field studies in Yellowstone National Park where
wolves have been reintroduced allow observation of
competitive interactions (interference) between the
two species
§ All wolves are radio-collared
§ Over a 12-year period, the researchers documented
337 wolf-coyote interactions
§ What types of interactions did they observe?

§ Majority of interactions (75 percent) were at wolf-
killed carcasses coyotes were trying to scavenge
§ Wolves initiated most encounters and dominated
most interactions
§ Wolves chased without physical contact (79 percent)
§ In 7 percent of encounters, a coyote was killed
§ All of the current data on wolf-coyote interactions
suggest that the decline of wolf population across
most of its historic range allowed for range
expansion of the coyote
§ Wolf and coyote at a wolf-killed elk carcass

Figure 13.21

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13

Elements of ECOLOGY Lecture Presentation by

§ Interspecific competition is a cornerstone of

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dN2/dt = r2N2(1 - N2/K2)

§ How do you modify these equations to model the

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Population density measured by volume

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Population density measured by volume

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A. the highest water temperature.

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A. the highest water temperature.

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