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Competition
A -/- interaction in which organisms negatively affect one another through shared use of limiting resources
Intraspecific competition
Competition between individuals of the same species
Interspecific competition
Competition between individuals of different species
Direct or interference competition
Organisms directly prevent others from accessing resources through aggression or physical interactions
Indirect or exploitative competition
Organisms reduce resource availability for competitors by consuming or using shared resources
Fundamental niche
The full range of environmental conditions and resources a species could use without competition
Realized niche
The portion of a species' fundamental niche it actually occupies when competition and other biological interactions are present
Competitive exclusion principle
Two species with complete niche overlap cannot coexist indefinitely when competing for the same limiting resources
Competitive coexistence
Two competing species persist together when their ecological differences prevent one from eliminating the other
Resource partitioning
Species reduce competition by using different resources or different portions of a shared resource
Character displacement
Evolutionary divergence in traits between competing species that reduces resource overlap
Gause competition experiments
Paramecium aurelia outcompeted P. caudatum when grown together because it more effectively exploited their shared bacterial food resource
Recognizing the superior competitor in Gause experiments
The superior competitor maintains population growth while the inferior competitor declines toward extinction in mixed culture
Resource utilization curves
Graphs showing the frequency with which species use resources along a resource gradient
Competition and resource utilization curves
Competition is greatest where the resource utilization curves overlap most
Resource partitioning and utilization curves
Less overlap between curves indicates reduced competition and greater potential for coexistence
Lotka-Volterra competition model
A modification of logistic growth that incorporates the effects of both intra- and interspecific competition on population growth
Lotka-Volterra equation for species 1
dN1/dt = r1N1[1-(N1+αN2)/K1]
Lotka-Volterra equation for species 2
dN2/dt = r2N2[1-(N2+βN1)/K2]
Components of Lotka-Volterra equations
N = population size • r = intrinsic growth rate • K = carrying capacity • α and β = interspecific competition coefficients
Alpha α
The competitive effect of one species 2 individual on species 1 expressed in equivalent species 1 individuals
Beta β
The competitive effect of one species 1 individual on species 2 expressed in equivalent species 2 individuals
Alpha greater than 1
An individual of species 2 has a stronger competitive effect on species 1 than another species 1 individual
Alpha less than 1
An individual of species 2 has a weaker competitive effect on species 1 than another species 1 individual
Beta greater than 1
An individual of species 1 has a stronger competitive effect on species 2 than another species 2 individual
Beta less than 1
An individual of species 1 has a weaker competitive effect on species 2 than another species 2 individual
Alpha or beta equal to 1
Individuals of either species have equivalent competitive effects on the focal species
Lotka-Volterra isocline
A line representing combinations of the two population sizes where one species has zero population growth
Lotka-Volterra graph axes
Species 1 population size N1 is on the x-axis and species 2 population size N2 is on the y-axis
Species 1 isocline intercepts
Connect K1 on the x-axis with K1/α on the y-axis
Species 2 isocline intercepts
Connect K2/β on the x-axis with K2 on the y-axis
Growth on an isocline
The species represented by that isocline has zero population growth
Growth below the species 1 isocline
Species 1 increases because its total competitive pressure is below its carrying capacity
Growth above the species 1 isocline
Species 1 decreases because its total competitive pressure exceeds its carrying capacity
Growth below the species 2 isocline
Species 2 increases because its total competitive pressure is below its carrying capacity
Growth above the species 2 isocline
Species 2 decreases because its total competitive pressure exceeds its carrying capacity
Lotka-Volterra population vectors
Species 1 growth produces horizontal arrows and species 2 growth produces vertical arrows
Interpreting combined population vectors
Combine horizontal and vertical arrows to determine whether populations move toward coexistence or competitive exclusion
Lotka-Volterra outcome 1 species 1 wins
Species 1 excludes species 2 when K1 > K2/β and K1/α > K2
Lotka-Volterra outcome 2 species 2 wins
Species 2 excludes species 1 when K1 < K2/β and K1/α < K2
Lotka-Volterra outcome 3 stable coexistence
Isoclines intersect with K1/α > K2 and K2/β > K1 → populations move toward a stable equilibrium
Lotka-Volterra outcome 4 unstable coexistence
Isoclines intersect with K1/α < K2 and K2/β < K1 → initial population sizes determine which species wins
Priority effects
The species that establishes first or begins at sufficiently high abundance can prevent its competitor from establishing
Stable versus unstable Lotka-Volterra equilibrium
Stable equilibrium attracts population trajectories while unstable equilibrium causes trajectories to move toward exclusion of one species
When should Lotka-Volterra be used
Use it to predict competition outcomes from population sizes • carrying capacities • competition coefficients and zero-growth isoclines
Limitation of Lotka-Volterra
The model summarizes competition without explicitly tracking individual resources or their different requirements
Tilman resource-ratio hypothesis
A model predicting competition outcomes based on species-specific requirements for two limiting resources and their availability
Tilman versus Lotka-Volterra graphs
Tilman graphs resource concentrations while Lotka-Volterra graphs population abundances
Zero net growth isocline ZNGI
A line representing resource combinations where population births and deaths balance and net growth equals zero
R* value
The minimum concentration of a limiting resource required for a species to maintain zero net population growth
Interpreting R* values
The species with the lower R* for a resource is the better competitor for that resource because it can persist at lower concentrations
Tilman population growth relative to ZNGI
Above and to the right of both resource thresholds growth is positive • on the ZNGI growth is zero • below either essential threshold growth is negative
Resource supply point S
The concentrations of the two resources available in the environment without consumers
Consumption vector
A directional arrow showing how a species reduces resource concentrations through consumption and the relative amounts of each resource it uses
Renewal vector
A directional arrow showing how environmental resource replenishment increases resource concentrations
Tilman outcome with nonintersecting ZNGIs
The species with lower R* requirements for both resources excludes the other when resource supply supports its persistence
Tilman outcome with intersecting ZNGIs
Each species is the better competitor for a different resource → outcomes depend on resource supply ratios and consumption vectors
Tilman outcome when resource X is limiting
The species with the lower R* for X wins when X is scarce relative to Y and resource consumption leads toward exclusion
Tilman outcome when resource Y is limiting
The species with the lower R* for Y wins when Y is scarce relative to X and resource consumption leads toward exclusion
Tilman stable coexistence
With intermediate resource supply ratios each species consumes more of the resource that limits itself → populations stabilize together
Tilman priority effects
When consumption patterns create an unstable equilibrium the initially established species can exclude the other
Tilman outcome when both resources are insufficient
Neither species can maintain positive population growth when essential resource concentrations remain below its requirements
Interpreting Tilman consumption vector slopes
A steeper consumption vector indicates greater Y consumption relative to X while a shallower vector indicates greater X consumption relative to Y
Predicting Tilman outcomes from a graph
Compare species R* values → locate the resource supply point → follow consumption vectors → determine whether exclusion or stable coexistence occurs
Three long-term outcomes of competition
Extinction • exclusion from part of a habitat or niche • evolutionary change that reduces competition
Extinction as a competition outcome
A competitively inferior species may disappear entirely when it cannot maintain a viable population
Exclusion as a competition outcome
A species may persist elsewhere but be excluded from habitats or resources occupied by a superior competitor
Evolution as a competition outcome
Competition can favor heritable changes in resource use that reduce overlap between species
Ghosts of competition past
Present-day species distributions and niche differences may reflect historical competition even when little competition is currently observabl