Principles of Ecology

Competition
 Modes of Competition
• Competition: use or defense of a resource by
one individual that reduces the availability of
the resource to other individuals
• Intraspecific:
– Competition with members of own species.
• Interspecific:
– Competition between individuals of different
species - reduces fitness of both.
Pioneering
experiment

A.G. Tansley
(1917)

British botanist

Two small
perennial
herbaceous
plant species
(Galium)

Two kinds of
soils

G. Saxatile grow
on acidic peaty
soils
G. Sylestre on
alkaline soils of
limestone hills
Competition results when resources are limited
• Intraspecific competition: regulate population growth in a
density-dependent manner.
• Evolution tends to favor the individuals with high resource use
efficiency and competition ability

• Interspecific competition: depress both populations. Under

intense interspecific competition, population of one species
may decline and die out.

• Outcome of interspecific competition:

depends on how efficiently individuals within each species
exploit share resources.
Supercompetitor can persist at
lower resource levels

As population
grow, resource
available for each
individual
decreases
Consumers compete for resources
• Resource: any substance or factor that is both
consumed by an organism and supports increased
population growth rates as its availability in the
environment increases
• Examples:
– food, water, nutrient,
– light, space
– Refuges, safe site
• No-consumeable physical and biological factors are
not resource: Temperature is not consumed, one
does not change T for another
 Space is an important resource for
sessile animals

Barnacles on the rocky coast of Maine. Above optimal range of

intertidal zone (small ones are larvae)
Competition between closely and distantly
related species
• Which one is more intense, closely related species or
distantly related species?

 On the Origin of Species:

 Competition should be most intense

between closely related species
 Structure, Habitat, food resources
Competition between distantly related
species is common

Example 1: barnacles, mussels,

alage, sponges, bryozoans,
tunicates in the intertidal zone
compete for spaces

Example 2: fish, squid, diving

birds, seals, and whales all eat
krills

Example 3: birds, lizards eat same

insects;
Ants, rodents, birds eat seeds in
the desert systems.
Renewable and nonrenewable resources

• Renewable: constantly renewed or regenerated

Natural resources outside ecosystem: such as light and
precipitation
Resource regenerated
– Birth of prey provide foods for predator
Consumers directly depress such resources
– Decomposition provide nutrients for plants
Indirectly linked to consumers through food chain or
abiotic factors.
• Non-renewable: space
– Once occupied, space becomes unavailable to others
 Limiting resources
• Consumers require many different resources, but not
all resources limit population growth

• Liebig’s law of minimum

Populations are limited by the single resource that is most
scarce relative to demand
Justus von Liebig (1840)

Limiting resources: may vary

David Tilman’s diatom study: both P and silicon
<0.2 mM of phosphate
or <0.6 mM silicate, diatom pop.growth stops.
Positive interaction and synergistic effect

Synergistic effect: Two resources together enhances

population growth more than the sum of both
individually

Peace and
Grubb
(1982)

Plant
fertilization
and
Light
treatments
Failure of species to
coexist in laboratory
cultures led to the
competitive exclusion
principle

G.F. Gause, Russian biologist

Protist: (bacteria here)

P. aurelia and P. caudatum

Same nutrient medium

Diatom experiment
David Tilman, University of
Minnesota

Asterinella formosa (Af) and

Synedra ulna(Su) compete for
silica for the formation of cell
walls.
Grow well alone
Insufficient silica, Su reduced the
silica to a low level and drove Af to
extinction
 Competitive exclusion principle
• Principle: Complete competitors can not coexist. One
species must go extinction
• Complete competitors: two species that live in the
same place and possess exactly the same ecological
requirements.
• Assumptions:
– Exactly the same resource requirement (no more,
no less)
– Environmental conditions remain constant
• In natural situations, two similar species can coexist,
why?
Population Dynamics
 Population Dynamics

∆N
Exponential growth =r•N
∆t

Occurs when growth

rate is proportional to
population size;
Requires unlimited
N
resources

Time
 Population Dynamics

Density-dependent per capita birth (b) & death (d) rates

Notice that per capita b

fitness increases with

decreases in
b r
population size or
from K d

Equilibrium
(= carrying
N capacity, K)
 Population Dynamics

∆N N
Logistic growth = r • N • (1 – )
∆t K

K = carrying capacity
∆N
=0
∆t

N
∆N
is maximized
∆t

∆N
=0
∆t

Time
The quantitative theory of competition was developed by
Vito Volterra and Alfred Lotka in 1925-6.
Once more there are separate differential equations - here for the growth of the two
competing species. Each is a logistic equation. Begin with equations for each species
growing alone.
The equation for species 1
dN1  K1  N 1 
 r1 N 1  
growing alone:

dt  K1 

The equation for species 2 dN 2  K2  N 2 

growing alone:
 r2 N 2  
dt  K2 
The growth curves for the two species, each growing alone…
The equations basically indicate that a portion of the resource that could have
contributed to the carrying capacity for each species growing alone is used up by the
competing species. Competition is incorporated by specifying constants, called the
competition coefficients, that represent the strength of the competitive interaction
There are two coefficients, each representing the effect of one species on the other.
We add them to the two logistic equations as follows:
 dN 1  K1  N 1  1,2 N 2 
 r1 N 1  
dt  K1 
and
dN 2  K2  N 2   2 ,1 N 1 
 r2 N 2  
dt  K2 

Note that these are standard logistic growth equations with a

new, additional term in each equation. The new terms are
1,2N2 and 2,1N1 .
Let’s explain this again

In other words….
 Competition

Lotka-Volterra Competition Equations:

In the logistic population growth model, the growth rate is

reduced by intraspecific competition:

Species 1: dN1/dt = r1N1[(K1-N1)/K1]

Species 2: dN2/dt = r2N2[(K2-N2)/K2]

Lotka & Volterra’s equations include functions to further reduce

growth rates as a consequence of interspecific competition:

Species 1: dN1/dt = r1N1[(K1-N1-f(N2))/K1]

Species 2: dN2/dt = r2N2[(K2-N2-f(N1))/K2]

 Competition

Lotka-Volterra Competition Equations:

The function (f) could take on many forms, e.g.:

Species 1: dN1/dt = r1N1[(K1-N1-αN2)/K1]

Species 2: dN2/dt = r2N2[(K2-N2-βN1)/K2]

The competition coefficients α & β measure the per capita effect of

one species on the population growth of the other, measured
relative to the effect of intraspecific competition

If α = 1, then per capita intraspecific effects = interspecific effects

If α < 1, then intraspecific effects are more deleterious

to Species 1 than interspecific effects

If α > 1, then interspecific effects are more deleterious

 2 2
1
2
1
1
2 1 1
1 1
1
1

Area within the frame represents carrying capacity (K) of either species

The size of each square is proportional to the resources an individual

consumes and makes unavailable to others (Sp. 1 = purple, Sp. 2 = green)

Individuals of Sp. 2 consume 4x resources consumed

by individuals of Sp. 1

For Species 1: dN1/dt = r1N1[(K1-N1-αN2)/K1] … where α = 4.

Redrawn from Gotelli (2001)
 2 2
1
2
1
1
2 1 1
1 1
1
1

Competition is occurring because both α & β > 0  α = 4 & β = ¼

In this case, adding an individual of Species 2 is more deleterious to

Species 1 than is adding an individual of Species 1…

but, adding an individual of Species 1 is less deleterious to

Species 2 than is adding an individual of Species 2

Redrawn from Gotelli (2001)

Populus Lab
Next…..

How important and widespread is

competition in ecological communities?
Lotka-Volterra Phenomenological Competition Model

Find equilibrium solutions to the equations, i.e., set dN/dt = 0:

^
Species 1: N 1 = K1 - αN2

^
Species 2: N2 = K2 - βN1

This makes intuitive sense: The equilibrium for N1 is the carrying

capacity for Species 1 (K1) reduced by some amount owing to the
presence of Species 2 (α N2)

However, each species’ equilibrium depends on the equilibrium of

the other species! So, by substitution…
^ ^
Species 1: N1 = K1 - α(K2 - βN1)
^ ^
Species 2: N2 = K2 - β(K1 - αN2)
Lotka-Volterra Phenomenological Competition Model

The equations for equilibrium solutions become:

^
Species 1: N 1 = [K1 - αK2] / [1 - α β]

^
Species 2: N 2 = [K2 - βK1] / [1 - α β]

These provide some insights into the conditions required for

coexistence under the assumptions of the model

E.g., the product αβ must be < 1 for N to be > 0 for both species (a
necessary condition for coexistence)

But they do not provide much insight into the dynamics of

competitive interactions, e.g., are the equilibrium points stable?
 4 time steps

State-space graphs help to

track population trajectories
(and assess stability)
predicted by models

Mapping state-space
trajectories onto single
population trajectories

From Gotelli (2001)

 4 time steps

State-space graphs help to

track population trajectories
(and assess stability)
predicted by models

4 time steps

Mapping state-space
trajectories onto single
population trajectories

From Gotelli (2001)

Remember that equilibrium Lotka-Volterra Model
solutions require dN/dt = 0
^
Species 1: N1 = K1 - αN2

Therefore:

When N2 = 0, N1 = K1
K1 / α
When N1 = 0, N2 = K1/α Isocline for Species 1
N2 dN1/dt = 0

K1

N1
Remember that equilibrium Lotka-Volterra Model
solutions require dN/dt = 0
^
Species 2: N2 = K2 - βN1

Therefore:

When N1 = 0, N2 = K2
K2
When N2 = 0, N1 = K2/β Isocline for Species 2
N2 dN2/dt = 0

K2 / β

N1
Plot the isoclines for 2 Lotka-Volterra Model
species together to
examine population
trajectories Competitive exclusion of
Species 2 by Species 1
K1/α > K2
K1 > K2/β

For species 1: K1 / α

K1 > K2α
(intrasp. > intersp.)
N2

K2
For species 2:
K1β > K2
(intersp. > intrasp.)

= stable equilibrium
K2 / β K1
N1
Plot the isoclines for 2 Lotka-Volterra Model
species together to
examine population
trajectories Competitive exclusion of
Species 1 by Species 2
K2 > K1/α
K2/β > K1

For species 1: K2

K2α > K1
(intersp. > intrasp.)
N2

K1/ α
For species 2:
K2 > K1β
(intrasp. > intersp.)

= stable equilibrium
K1 K2 / β

N1
Plot the isoclines for 2 Lotka-Volterra Model
species together to
examine population
trajectories Competitive exclusion in an
unstable equilibrium
K2 > K1/α K2
K1 > K2/β

For species 1:
K2α > K1
(intersp. > intrasp.) K1/ α
N2

For species 2:
K1β > K2
(intersp. > intrasp.)

= stable equilibrium
= unstable equilibrium K2 / β K1
N1
Plot the isoclines for 2 Lotka-Volterra Model
species together to
examine population
trajectories Coexistence in a stable equilibrium

K1/α > K2 K1 / α
K2/β > K1

For species 1:
K1 > K2α
(intrasp. > intersp.)
N2

K2
For species 2:
K2 > K1β
(intrasp. > intersp.)

= stable equilibrium
K1 K2 / β
N1
 (a) Species 1

N2=(K1-N1)/alpha

Alpha=alpha1,2

Diagonal line is zero growth isocline

(b) Species 2

N2=K2-beta N1

Beta=alpha2,1
There Are Four Possible Outcomes of Interspecific
Competition

• Possible outcomes of the Lotka–Volterra equations

– In two situations, one of the species is the superior
competitor and wins out over the other
– In one case, species 1 inhibits the population of species
2 while continuing to increase
– In one case, species 2 inhibits the population of species
1 while continuing to increase
(c) Species 1 inhibits growth of species 2 and latter
goes extinction
(d) Species 2 inhibits growth of species 1 and latter
goes extinction
There Are Four Possible Outcomes of Interspecific
Competition

• Possible outcomes of the Lotka–Volterra equations

– In a third situation, each species, when abundant, inhibits
the growth of the other (more than it inhibits its own
growth)
– Eventually one of the two species “wins”
– In a fourth situation, neither species eliminates the other
resulting in coexistence
– Each species inhibits its own population growth more
than that of the other species
 (e) Unstable situation, both inhibit in a density dependent
manner. Depending on initial density, either can make other
extinct
(f) Each species inhibits its own population growth
more than competitor. Neither can eliminate
competitor
 Coexistence on multiple resources

David Tilman: two diatom species, Cyclotella and Asterionella

Two Resouces: phosphorus (for DNA, phospholipids etc) and silicon
(for shell)
Ratio of Si/P: if Si/P is below this level, silicon limited, above,
phosphorus is limited
Cyclotella: limited at Si/P=6, low requirement for Si, high for P
Asterionella: limited at Si/P=90, high requirement for Si, low for P
• Examples in nature
 Competition

Connell (1983)
Reviewed 54 studies
45/54 (83%) were consistent with competition

Of 54 studies, 33 (61%) suggested asymmetric competition

Schoener (1983)
Reviewed 164 studies
148/164 (90%) were consistent with competition

Of 61 studies, 51 (85%) suggested asymmetric competition

Kelly, Tripler & Pacala (ms. 1993) [But apparently never published!]
Only 1/4 of plot-based studies were consistent with competition,
whereas 2/3 of plant-centered studies were consistent
with competition
 A classic competition study: MacArthur (1958)

Five sympatric warbler species with similar bill sizes and shapes broadly
overlap in arthropod diet, but they forage in different zones
within spruce crowns

Is this an example of the “ghost of competition past”

(sensu Connell [1980])?
Light and nutrient competition among rain forest tree seedlings (Lewis and
Tanner 2000)

Above-ground competition for light

is considered to be critical to seedling
growth and survival

Fewer studies exist of in situ below-

ground competition

Design:

Transplanted seedlings of two species (Aspidospermum - shade

tolerant; Dinizia - light demanding) into understory sites (1% light)
and small gaps (6% light) in nutrient-poor Amazonian forest

Reduced below-ground competition by “trenching” (digging a 50-cm

deep trench around each focal plant and lining it with plastic); this stops
neighboring trees from accessing nutrients and water
 2.0 2.0
gap
Relative height growth/day x 1000

Aspidospermum Dinizia
gap
1.5 1.5
understory

understory
1.0 1.0

0.5 0.5

0 0
No trench Trench No trench Trench No trench Trench No trench Trench

Results and Conclusion:

Trenching had as big an impact as increased light did on seedling

growth

Seedlings are apparently simultaneously limited by (and compete

for) nutrients and light

Could allelopathy also be involved?

 Effect of territorial honeyeaters on homopteran abundance
Loyn et al. (1983)

Flocks of Australian Bell Miners defend communal territories in eucalypt

forest, excluding other (sometimes much larger) species of birds

Up to 90% of miners’ diet is nymphs, secretions and lerps (shields) of

Homopterans (Psyllidae)

Experiment: Counted birds, counted lerps, removed miners

Results & conclusion: Invasion by a guild of 11 species of insectivorous

birds (competitive release), plus 3x increase in lerp removal rate, reduction
in lerp density, and 15% increase in foliage biomass
 Competition between seed-eating rodents
and ants in the Chihuahuan Desert
Brown & Davidson (1977)

Strong resource limitation – seeds are the primary

food of many taxa (rodents, birds, ants)

Almost complete overlap in the sizes of seeds

consumed by ants and rodents – demonstrates the
potential for strong competition
Design:
Long-term exclosure experiments – fences to exclude rodents, and
insecticide to remove ants; re-censuses of ant and rodent populations
through time

Results and Conclusion:

Excluded rodents and the number of ant colonies increased 70%
Excluded ants and rodent biomass increased 24%

Competition can apparently occur between distantly related taxa

 Competition between sexual and asexual species of gecko
Petren et al. (1993)

Humans have aided the dispersal of a sexual

species of gecko (Hemidactylus frenatus) to
several south Pacific islands and it is
apparently displacing asexual species

Experiment: Added H. frenatus and L.

lugubris alone and together to aircraft hanger
walls

Results and Conclusion: L. lugubris avoids H.

Lepidodactylus lugubris, frenatus at high concentrations of insects on
asexual native on lighted walls
south Pacific islands
Sometimes “obvious” hypothesized reasons
for competitive dominance are incorrect
Competition among Anolis lizards
(Pacala & Roughgarden 1982)

What is the relationship between the strength of

interspecific competition and degree of interspecific
resource partitioning?

2 pairs of abundant insectivorous diurnal Anolis

lizards on 2 Caribbean islands:

St. Maarten: A. gingivinus & A. wattsi pogus

St. Eustatius: A. bimaculatus & A. wattsi schwartzi

Competition among Anolis lizards
(Pacala & Roughgarden 1982)

Body size (strongly correlated with prey size):

St. Maarten anoles: large overlap in body size
St. Eustatius anoles: small overlap in body size

Foraging location:
St. Maarten anoles: large overlap in perch ht.
St. Eustatius anoles: no overlap in perch ht.

Experiment:

Replicated enclosures on both islands, stocked

with one (not A. wattsi) or both species
Competition among Anolis lizards
(Pacala & Roughgarden 1982)

Results and Conclusions:

St. Maarten (similar resource use)

Growth rate of A. gingivinus was halved
in the presence of A. wattsi

St. Eustatius (dissimilar resource use)

No effect of A. wattsi on growth or
perch height of A. bimaculatis

Strength of present-day competition in these

species pairs is inversely related to resource
partitioning
Competition among Anolis lizards
(Pacala & Roughgarden 1982)

Why do these pairs of anoles on nearby islands

(30 km) differ in degree of resource partitioning?

Hypothesis: Character displacement occurred on

St. Eustatius during long co-evolutionary history
(i.e., the ghost of competition past), whereas
colonization of St. Maarten occurred much more
recently, and in both cases colonization was by
similarly sized Anolis species

Character displacement:
Evolutionary divergence of traits in response to
competition, resulting in a reduction in the intensity
of competition
Competition among Anolis lizards
(Pacala & Roughgarden 1985)

Pacala & Roughgarden (1985) presented evidence

to suggest that both species pairs have a long
history of co-occurrence on their respective islands
and that different colonization histories resulted in
the observed patterns of resource partitioning

Both islands may have been colonized by Anolis

species differing in size, yet on St. Maarten the
larger Anolis colonized later and has subsequently
converged in body size on the smaller resident
 Character Displacement

Schluter & McPhail (1992) surveyed the literature on character

displacement and listed criteria necessary to exclude other potential
explanations for species that share similar traits in allopatry, but differ in
sympatry (similar to Connell’s [1980] requirements to demonstrate the
“ghost of competition past”):

1. Chance should be ruled out as an explanation for the pattern

(appropriate statistical tests, often involving null models)
2. Phenotypic differences should have a genetic basis
3. Enhanced differences between sympatric species should be the
outcome of evolutionary shifts, not simply the inability of similar-sized
species to coexist
4. Morphological differences should reflect differences in resource use
5. Sites of sympatry and allopatry should be similar in terms of physical
characteristics
6. Independent evidence should be obtained that similar phenotypes
actually compete for food
 • Why do species coexist?

• Mechanisms:
So far, we see examples of
– Lotka Volterra
the predictions of LV – Negative Feedback
competition model, in
nature, why do species – Niche niche partitioning
coexist?
– Character displacement
To do:
Read the paper asigned
and reply to the question.
13.3 Asymmetric competition can occur when
different factors limit the populations of
competitors

Connell et al
(1961)
 Chipmunks
Alpine
Cold tolerant
Lodgepole
Most
aggressive
Needs shade
Yellow Pine
aggressive
Least
Heat
tolerant

Sierra Nevada, CA
 Habitat productivity can influence
competition between plant species
Two hypotheses:

1.Plants compete more intensively when mineral nutrients are

less abundant in the soil (By Grubb and Tilman)
Plants compete more intensively when nutrients are less.
High nutrients are less likely to limit plant population;
thus the intraspecific competition is weak.

2.Competition is less intense when water and nutrients are less

abundant (Grime and Keddy)
Competition for light is more important than
competition for nutrients; limit in water and nutrients would
limit the population growth to a certain point that individual
plants are widely spread and do not compete for light.

Difference between these hypotheses lies in the relative

importance placed on belowground and aboveground
competition for resources --Light or nutrient. (Debate)
 Habitat productivity can influence
competition between plant species

Smooth cordgrass saltmeadow cordgrass, black grass, alder

 Habitat productivity can influence
competition between plant species

Saltmeadow vs Smooth Blackgrass vs saltmeadow

Fertilization alters the outcome of competition by removing

nutrient limitation on stress-tolerant plants, expand, away from
water.
 Competition may occur through direct
interference
 Exploitation: indirectly influencing each
other by consuming the same resources
(eat same grass by zebras , compete for
water uptake by trees, indirectly)

 Interference: direct influencing each other

by preventing others to occupy a habit or
access resources (birds, bees chase birds
and bees, animals release toxic
chemicals).
 Meadow vole (wet) and mountain vole

(dry). (Asymmetric competition also)

 Allelopathy (chemical competition)

Clumps of shrubby Salvia plants

Figure 16.14 Some plants (mint) are usually surrounded by
(eucalyptus) compete by chemical bare zones separating the sage from
means. neighboring grassy areas ( Figure
16.15)
Australian ironwood trees
Consumers can influence the outcome of competition

Keystone predator

Starfish prey on
mussels, barnacles,
limpets, and chitons

Remove starfish,
what would happen?

Species diversity
increase or decrease?

Why?
Grazing on
plant
diversity?
Predator can influence the
outcome of prey competition

Peter Morin, Rutgers

Salamander

Frog or toad tadpole

(300 each of 3 species)
 Apparent competition

Combined populations of
two prey species support a
larger predator population
neither can support alone.
As a result, two prey
populations reduced, gives
outward appearance of
interspecific competition.

Experimental supports:
Nettle aphid, grass aphid and ladybug beetle (Smith and Smith,
page 359)
Brought nettle aphid plants to grass aphid plants together
suppressed both population, as a results of larger ladybug beetle
population.
 Apparent competition mediated by
pathogens (microbes)

Corals can be indirectly harmed by the presence of algae

Antibiotics can reverse the negative effects of algae
on coral growth

Smith et al. 2006