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
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
Peace and
Grubb
(1982)
Plant
fertilization
and
Light
treatments
Failure of species to
coexist in laboratory
cultures led to the
competitive exclusion
principle
∆N
Exponential growth =r•N
∆t
Time
Population Dynamics
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
In other words….
Competition
Area within the frame represents carrying capacity (K) of either species
^
Species 2: N2 = K2 - βN1
^
Species 2: N 2 = [K2 - βK1] / [1 - α β]
E.g., the product αβ must be < 1 for N to be > 0 for both species (a
necessary condition for coexistence)
Mapping state-space
trajectories onto single
population trajectories
4 time steps
Mapping state-space
trajectories onto single
population trajectories
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
N2=K2-beta N1
Beta=alpha2,1
There Are Four Possible Outcomes of Interspecific
Competition
Connell (1983)
Reviewed 54 studies
45/54 (83%) were consistent with competition
Schoener (1983)
Reviewed 164 studies
148/164 (90%) were consistent with 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
Design:
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
Foraging location:
St. Maarten anoles: large overlap in perch ht.
St. Eustatius anoles: no overlap in perch ht.
Experiment:
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)
• 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:
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
Salamander
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)