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community
group of species that occur at the same time and place and the interactions that bind them
pairwise interactions
interactions based on: harmed (-), benefited (+), unaffected (0)
types of pairwise interactions
predation, herbivory, parasitism, competition, amensalism, mutualsim, commensalism
predation
(+-) predators benefit, prey do not
herbivory
(+-) plant is harmed, organism feeding benefits
parasitism
(+-) host is harmed
(+-) of pairwise interactions just paired under?
predation
competition
(- -)sharing limiting resources, taking resource from another
amensalism (asymmetric competition)
(- 0) one species harmed, another is completely unaffected (elephants foraging in savannah, damaging plants, affecting elephants in no way)
mutualism
(++) plants and their pollinators
commensalism
(+0) algae in lakes, benefit on back of turtles, does not harm turtle, cattle forging on road
most important biotic processes? most every species involved
predation and competition
interactions affect the what of species?
distribution and abundance
competition
interaction that occurs when organisms of the same or different species use an essential resource that is in short supply (limiting resource)
hyenas chased away by lioness
interspecific, interference
characterizing competition
intraspecific/interspecific, exploitative/interference, symmetric (-,-) or asymmetric (-,0)
are fights to death common?
no, they are rare, effects of competition are subtle
competition impacts?
life history traits, delayed effects on population size (N) or growth (r)
life history traits
any trait related to lx, bx, e, i ,R0, r, k
body size
longevity
age at 1st reproduction
fecundity
vigor
sex ratio (producing females are more costly)
intraspecific competition
first considered as an ecological force by Thomas Malthus, people are going to run out of resources
Darwin’s “struggle for existence” emerges from ideas on resource limitation
affects demography (lx, bx, r) and abundances, and to a lesser extent distributions of species
was interspecific competition studied as well?
no, mostly ignored as an important ecological process until the classic study by Tansley
Tansley’s Bedstraw
studied two species, one likes the acidic soil, one with the basic high soil, G. saxatile,
Tansley Bedstraw findings
grown alone: bedstraws survive on both soil types
grown togtehre: each bedstraw species wins on its native soilW
What does this tell you about the distribution of bedstraws?
species presence/absence due to interspecific competition
environment affect outcome
basic soils: G. sylvestre wins
acidic soils: G. saxatile wins
present distribution may be due to competiton in past (ghost of competition past)
competition experiments
Gause
2 species of paramecium
liming resource: bacteria
grew them in test tubes with limiting resource of bacteria
grown alone:
low and high resources
P aurelia has highest k (more efficient at utilizing the same amount of resources)
being more efficient, can make you a better competitor
together: P aurelia has a reduction in K
causes P. caudatum to be extinct (compeition causes extinction to some species)
Competitive exclusion principle
species utilizing the same limiting resource cannot coexist together
one species will competitively exclude the other
aka Gause’s principle
corollary
species can coexist if they utilize limiting resources in a different way
Thomas Park beetles
interspecific competition in Tribolium flour beetles
grew in hot and humid conditions, red flour drove the other species extinct, in cold it was vice versa
Joe Connell barnacles
rocky intertidal; upper zone Balanus scarce and no withstand air, exclude Balanus from middle Chthamalus survives by dislodging them; Balanus is a stronger competitor, Chthamalus is only in upper zone bc it can withstand the air
competitive exclusion
Tansley’s bedstraw
Gause’s Paramecium
Park’s flour beetles
Connell’s barnacles
experimental conclusions
competitive exclusion, outcome depends on environment, distribution of species strongly influenced by interspecific competition
outcome depends on environment
Tansley: soil conditions
Park: temperature and humidity
Connell: tidal zones
compeititon coefficients
measures negative effects of one speicies on another
measures competitive ability
largest competition coefficient is a better competitor, larger conversion factor, stronger competitior bc you use more resources per individual
higher carrying capcity more efficient in using resources
a>1
organism has stronger negative effect on species 1 than it has on itself through intraspecific competition
interspecific competition > strength of intraspecific
b<1
species 1 has a stronger negative effect on itself than on species 2
intraspecific> interspecific competition
dN1/dt
realized population growth rate of species 1
r1N1
exponential growth
k1-N1/k1
intraspecific competition
aN2
interspecfic competition
Model predictions
predictions depend on a,B,k1,k2
r ha no effect on outcome of competition
four possible outcomes
1) species 1 wins 2) species 2 wins 3) stable coexistence 4) unstable coexistence
General Criteria for Coexistence
a and b are < 1
model assumptions
logistic growth
linear density dependence
a, b, k1, k2, r1, r2 constant
closed population
the supply of resources to the system is constant
no other mortality factors
model provides a reasonable description for 2 competing species
coexistence of competitors
desert plants; different root structures;occupy different niches
niches
role of an organism within the community
relationshiop of a species to all aspects of the abiotic environment
how did niches revise Gause’s corollary?
competing species can coexist by occupying different niches
niche conept
Hutchinson; based on activity (tolerance) range for an environmental variable; range of something an organism can survive and reproducee
each abiotic variable represents?
1 dimension (abiotic variable)
fundamental niche
the activity range over all possible (n) dimensions for abiotic factors
niche dimensions often have?
optimal dimensions
realized niche
portion of the fundamental niche that is occupied by a species in the presence of biotic factors (how predators, etc can change niche)
predators
negative effect, diminish the niche; cause the fundamental niche to compress (realized niche when predators are present, reduce the size of the fundamental niche; realized niche is generally smaller)
competitors
can prevent a species from occupying a space it normally could; reduce niche; negative effect, diminish the niche
mutualists
postive effect, can expand the niche, realized niche is bigger than the fundamental niche
generally even with the prescence of mutualists
the realized niche is smaller just due to the competitors and predators
problems with realized niche
only one organism to know for sure the realized niche (humans)
a complete quantification of the niche of an organism is nearly impossible
a species’ niche is not fixed, but can vary with
a species’ niche can vary with
age or life stage
time (ex: changing climate)
space
solutions on the provlems with the realized niche
measure only most important dimensions (ex: those affecting resource use)
overlap in realized niches
sparrows,
tells about
niche breadth (measure of range of the resources they use (the line in the picture)
if x axis represents a limiting resource,
competition should occur among individuals in area of overlap
greater overlap means
greater competition
competitive exlcusion in the field; intentional/accidental introductions
introduced to a new area; fire ants into the US
areas where fire ants were present, native species were 70% lower than when they were present
total number of fire ants is 90% lower in areas, excluding native ant species
competitive exclusion example
cuban brown anole; introduced brown with green; changed its niche, green anoles declined and shifted perches higher above ground
why do introduced species appear to be so damaging to native species?
NEED TO KNOW ANSWER FOR THE EXAM
if resources are limited,
species should modify their behavior to minimize overlap in resource use and reduce the negative effects of competition
alter their niche to reduce effects of competitionn
niche partitioning
the division of a limitng resource by 2 or more species that share it; reduce likelihood for competition
niche partioning
MacArthur’s Warbler birds
feed on insects in conifers
not eating the same part of the tree
avoiding overlap in resource use
species that use the same limiting resources can partition to all surivive and live in the same place
if eliminate one species, the other would be able to expand its niche
Warbler
where black throat was absent, the yellow would expand its forging up the tree, suggest that the species are niche partitioning
competitive release is going on
competitive release
when a species is released from competiton, expanding their niche to the other species’ niche
slimy salamanders
found them on different elevation, the lower salamander was able to expand its niche up the mountain
coexistence is due to?
niche partioning; little overlap in resource use, little competition
spiny mice
niche partioning; predators hunt during day; remove the one who is noctunral, the other one switches to forage at night time
the removal species is a dominant competitor forcing the other one to forage in the daytime
paradox of competitive exclusion
many closely related species co-occur together and have nearly identical niches
zooplankton
trees
species not competing
species well below k (following a disturbance)
resources abundant (as for new colonists or invasive species)
explanations of paradox of competitive exclusion
1) species not competing
2) fluctuating environments
dominant doesnt have neough time to extinct the other in the amount of time
3) evolve not to compete
example of competitive release
study with chipmunks
a=1.5, b=0.5 k1<k2, what is the most likely outcome of competitive interaction?
a is for species 2, b species 1, species 2 wins
fire ants example of ?
competitive exclusion
natural selection should favor either:
increased competitive ability (higher k or a/b)
reduced negative effects of competition
trait that can evolve
body size of the falcons; closely related to the size of prey they consume and size of offspring, small flacons eat small prey, big falcons hunt bigger prey; trait related to resource use that can evolve
fitness in the area of the overlap
natural selection should act against phenotypes in overlap; favor divergence in resource use in each species (directional natural selection)
evolutionary divergence IF genetic basis to the trait associated with resource use
feeding niche evolve (grow to have less overlap use)
adaptive evolution
character divergence (character displacement)
evolutionary change in a trait (=character) associated with rsource use tha treduces the effects of competition
adaptive evolution
significance of character divergence
reduces competition
explanation for the paradox of competitive exclusion
lack of competition today could be due to competition in the past (ghost of competition past)
coevolution occurs through the reciprocal evolution in two competing species
what needs to be proven in studies of character diveregence
1) species differ in resource use
2) resource use related to a particular trait (=character)
3) trait has genetic basis and can evolve
4) lack of overlap in resource use today is due to competition in the past
indirect approach in inferring competition in the past
comparison of:
niche breadth for a trait associated with resource use when each species occurs alone (allopatry)
sympatry vs allopatry
allopatry: overlap
sympatry: less overlap
in sympatry
evolve reduced niche overlap (in terms of the trait associated with resource use)
small niche breadth
in allopatry:
evolve large niche breadth
competitive release
Darwin’s finches
look similar except the bills
evolve different bill sizes to avoid competition
Rhinoceros Beetles
occur in allopatry and sympatry
body size related to food size
beetle species
allopatry, sijmilar size
sympatry, one species is bigger than the other (divererged in body size)
anolis lizards (character divergence)
sympatry: forge on ground vs higher, diverge in niches
allopatry: overlap foraging areas
which trait is evolving? skeletal structure of the lizards
three spined sticklebacks (character divergence)
body shape genetically based, where they feed
one smaller body faster to chase, other bigger body at the bottom of pond\
competitive release - allopatry
first experiment for character divergence
put divider in pond add the allpatric benthic species with wider niche
other side with competition
broad overlap when they are together in body size
3 generations later, benthic evolved, no overlap in resource use
significance of stickleback experiment
natural selection operates on competing species
allopatric benthic species evolved more benthic habits
divergence (and possible speciation) in progress
is character divergence common in nature?
carnivorous vertebrates most studied
differences larger in sympatry than allopatry
suggests character divergence is common
evolution & competition conclusions
competitors should evolve to reduce competition (coevoltuion)
evidence is mostly indirect, but strong
few direct experimental studies
competition affects species distribution, abundance and evolution
can help to explain paradox of competitive exclusion
prevalence of competition
only field studies
manipulate competitor density
results
intraspecific > interspecific
50-75% show competition
most are asymmetric (-,0)
describe an experiment where competition does or doesn’t occur among species? how would you be able to demonstrate that?
remove a competitor or add one
competition should have negative effect on fitness, remove a competitior fitness should improve (show if competition is occuring)
removing a competitor would also have competitive release
expand its niche when a competitor leaves
add a competitor, cause negative fitness on the already existing species (lower survivorshiop or reproduction), maybe niche partitioning
hypothetical example to test how competition is occurring in a species
NEED TO KNOW