Community Ecology

Reading: Freeman, Chapter 50,

53

What is a community?
A community is an assemblage of
plant and animal populations that live in
a particular area or habitat.
Populations of the various species in a
community interact and form a system
with its own emergent properties.

Pattern vs. Process

Pattern is what we can easily observe
directly - vegetation zonation, species lists,
seasonal distribution of activity, and
association of certain species.
Process gives rise to the patternherbivory, competition, predation risk, nutrient
availability, patterns of disturbance, energy
flow, history, and evolution.

 Community ecology seeks to explain the

underlying mechanisms that create, maintain,
and determine the fate of biological communities.
Typically, patterns are documented by
observation, and used to generate hypotheses
about processes, which are tested.
Not all science is experimental. Hypotheses
tests can involve special observations, or
experiments.

Emergent Properties of a
Community
Scale
Spatial and Temporal Structure
Species Richness
Species Diversity
Trophic structure
Succession and Disturbance

 Scale is the size of a community.

Provided that the area or habitat is
well defined, a community can be a
system of almost any size, from a
drop of water, to a rotting log, to a
forest, to the surface of the Pacific
Ocean.

 Spatial Structure is the way species

are distributed relative to each other.
Some species provide a framework
that creates habitats for other
species. These species, in turn
create habitats for others, etc.

 Example: Trees in a rainforest are

stratified into several different levels,
including a canopy, several understories,
a ground level, and roots. Each level is
the habitat of a distinct collection of
species. Some places, such as the pools
of water that collect at the base of tree
branches, may harbor entire
communities of their own.

 Temporal structure is the timing of the

appearance and activity of species. Some
communities, i.e., arctic tundra and the decay of
a corpse, have pronounced temporal species,
other communities have less.
Example: Many desert plants and animals are dormant
most of the year. They emerge, or germinate, in
response to seasonal rains. Other plants stick around
year round, having evolved adaptations to resist
drought.

 Species Richness - is the

number of species in a community.
Clearly, the number of species we
can observe is function of the area
of the sample. It also is a function
of who is looking. Thus, species
richness is sensitive to sampling
procedure

 Diversity is the number of species in the

community, and their relative abundances.
Species are not equally abundant, some
species occur in large percentage of
samples, others are poorly represented.
Some communities, such as tropical
rainforests, are much more diverse than
others, such as the great basin desert.
Species Diversity is often expressed using
Simpsons diversity index: D=1-pi

Example Problem
A community contains the following species:

Number of Individuals
Species A
104
Species B
71
Species C
19
Species D
5
Species E
3
What is the Simpson index value for this
community?

Answer:
Total Individuals= (104+19+71+5+3)=202

PA=104/202=.51 PB=19/202=.09
PC=71/202=.35
PD=5/202=.03
PE=3/202=.02
D=1-{(.51)2+(.09)2+(.35)2+(.03)2+
(.02)2}
D=1-.40=.60

Clicker Question
In the example above, what was the
species richness?
A. .60
B. 202 individuals
C. 5 species
D. .40
E. None of the above

Succession, Disturbance
and Change
In terms of species and physical
structure, communities change
with time.
Ecological succession, the predictable change in
species over time, as each new set of species
modifies the environment to enable the
establishment of other species, is virtually
ubiquitous.

 Example; a sphagnum bog community may

persist for only a few decades before the
process of ecological succession changes
transform it into the surrounding Black Spruce
Forest.
A forest fire may destroy a large area of trees,
clearing the way for a meadow. Eventually, the
trees take over and the meadow is replaced.

 Disturbances are events

such as floods, fire, droughts,
overgrazing, and human
activity that damage
communities, remove
organisms from them, and
alter resource availability.

Some Agents of
Disturbance
Fire
Floods
Drought
Large Herbivores
Storms
Volcanoes
Human Activity

Disturbance, Invasion,

Succession

Disturbance creates opportunities for new

species to invade an area and establish
themselves.
These species modify the environment, and
create opportunities for other species to
invade. The new species eventually displace
the original ones. Eventually, they modify the
environment enough to allow a new series of
invaders, which ultimately replace them, etc.

 Invasion:
Disturbance creates an ecological vacuum that can be
filled from within, from outside, or both. For example,
forest fires clear away old brush and open up the canopy,
releasing nutrients into the soil at the same time. Seeds
that survive the fire germinate and rapidly grow to take
advantage of this opportunity. At the same time, windborne and animal-dispersed seeds germinate and seek to
do the same thing.
The best invaders have good dispersal powers and many
offspring, but they are often not the best competitors in
the long run.

Succession

Disturbance of a community is usually

followed by recovery, called ecological
succession.
The sequence of succession is driven by the
interactions among dispersal, ecological
tolerances, and competitive ability.
Primary succession-the sequence of species on newly
exposed landforms that have not previously been
influenced by a community, e.g., areas exposed by glacial
retreat.
Secondary succession occurs in cases which vegetation of an
area has been partially or completely removed, but where soil,
seeds, and spores remain.

 Early in succession, species are generally excellent

dispersers and good at tolerating harsh environments,
but not the best interspecific competitors.
As ecological succession progresses, they are replaced
with species which are superior competitors, (but not
as good at dispersing and more specialized to deal
with the microenvironments created by other species
likely to be present with them).
Early species modify their environment in such a way
as to make it possible for the next round of species.
These, in turn, make their own replacement by
superior competitors possible.

 A climax community is a more

or less permanent and final stage
of a particular succession, often
characteristic of a restricted area.
Climax communities are characterized by
slow rates of change, compared with more
dynamic, earlier stages.
They are dominated by species tolerant of
competition for resources.

An Influential ecologist named F.E. Clements argued that

communities work like an integrated machine. These
closed communities had a predictable composition.
According to Clements, there was only one true climax
in any given climatic region, which was the endpoint of
all successions.
Other influential ecologists, including Gleason,
hypothesized that random events determined the
composition of communities.
He recognized that a single climatic area could contain a
variety of specific climax types.

 Evidence suggests that for many habitats,

Gleason was right, many habitats never
return to their original state after being
disturbed beyond a certain point.
For example; very severe forest fires have
reduced spruce woodlands to a terrain of
rocks, shrubs and forbs.

 An incredibly rapid glacial retreat is occurring in Glacier

Bay, Alaska. In just 200 years, a glacier that once filled
the entire bay has retreated over 100km, exposing new
landforms to primary succession.
Clements would have predicted that succession today would
follow the sequence of ecological succession that has
occurred in the past for other parts of Alaska.
In fact, three different successional patterns seem to be
occurring at once, depending upon local conditions. Thus,
Clements view of succession is somewhat of an
oversimplification.

Are Climax Communities

Real?
Succession can take a long
time.
For example, old-field succession may
require 100-300 years to reach climax
community. But in this time frame, the
probability that a physical disturbance
(fire, hurricane, flood) will occur
becomes so high, the process of
succession may never reach completion.

 Increasing evidence suggests that some amount

of disturbance and nonequilibrium resulting from
disturbance is the norm for most communities.
One popular hypothesis is that communities are
usually in a state of recovery from disturbance.
An area of habitat may form a patchwork of
communities, each at different stages of ecological
succession. Thus, disturbance and recovery
potentially enable much greater biodiversity than is
possible without disturbance.

Are biological communities real

functional
units?
Do communities have a tightly prescribed organization
and composition, or are they merely a loose
assemblage of species?
This is an unsolved problem in ecology.
Clements argued that communities are stable,
functional units with a fixed composition-each
integrated part needs the others. Every area should
ultimately have the same species, given time.
Gleason argued that their composition is unstable and
variable-they are more like assemblages of everything
that can live together in one place

The Kiddie Pool

Experiment

Jenkins and Buikema conducted an experiment to

see whether artificial ponds would develop
predictable assemblages of freshwater
microorganisms.
-if this were the case, it would support the notion
that communities are real, integrated units.
-They set up 12 identical ponds and filled them
with sterile water. Came back in year to study the
composition of the resulting communities.

 Result-the ponds had very different

compositions of species.
Accidents of dispersal, and different dispersal
capabilities affected which species ended up
in each pond.
The early arrival of certain competitors, and
predators greatly affected the ability of later
species to colonize later.
-Gleasons view was supported. Composition
of communities is dictated largely by chance
and history.

 Trophic structure is the hierarchy of feeding.

It describes who eats whom
(a trophic interaction is a transfer of energy:
i.e., eating, decomposing, obtaining energy via
photosynthesis).
For every community, a diagram of trophic
interactions called a food web.
Energy flows from the bottom to the top.

A Simple Food Web

Killer Whales
Sharks
Harbor Seals
Yellowfin Tuna

Mackerel

Cod
Zooplankton

Halibut

Killer Whales
Harbor Seals
Mackerel
Zooplankton
Phytoplankton

One path
through a
food web is
a food
chain.

 The niche concept is very important

in community ecology.
A niche is an organisms habitat and
its way of making a living.
An organisms niche is reflected by
its place in a food web: i.e, what it
eats, what it competes with, what
eats it.
Each organism has the potential to
create niches for others.

 Keystone species are disproportionately important in

communities.
Generally, keystone species act to maintain species
diversity.
The extinction of a keystone species eliminates the
niches of many other species.
Frequently, a keystone species modifies the
environment in such a way that other organisms are
able to live, in other cases, the keystone species is a
predator that maintains diversity at a certain trophic
level.

Examples of Keystone
Species

California Sea Otters: This species preys upon sea

urchins, allowing kelp forests to become established.
Pisaster Starfish: Grazing by Pisaster prevents the
establishment of dense mussel beds, allowing other
species to colonize rocks on the pacific coast
Mangrove trees: Actually, many species of trees
are called mangrove trees. Their seeds disperse in
salt water. They take root and form a dense forest in
saltwater shallows, allowing other species to thrive

Trophic Cascades
Species at one trophic level influence species at
other levels; the addition or subtraction of species
affects the entire food web.
This causes positive effects for some species,
and negative effects for others. This is called a
trophic cascade. For instance, removing a
secondary consumer might positively affect the
primary consumers they feed upon, and
negatively affect the producers that are food for
primary consumers.

Top down vs. Bottom up

Most biological communities have both top-down and
bottom-up effects on their structure and
composition.
In a well known study of ponds by Matthew
Leibold, it was demonstrated that the biomass of
herbivores (zooplankton) was positively correlated
to the biomass of producers (algae), indicating a
top down effect.
He intentionally introduced fish to some ponds,
The result was a decrease in zooplankton and
increase in producers, indicating a top down effect.

Badly scanned from

Rose and Mueller (2006)

Types of Interspecific
Interactions
Effect on
Effect on

Neutralism
Competition
Commensalism
Amensalism
Mutualism
Predation,

Species 1

0
+
+
-

Parasitism, Herbivory

Species 2

0
0
0
+
+

Neutralism
Neutralism the most common type
of interspecific interaction. Neither
population affects the other. Any
interactions that do occur are
indirect or incidental.
Example: the tarantulas living in a
desert and the cacti living in a
desert

Competition
Competition occurs when organisms in the
same community seek the same limiting
resource. This resource may be prey, water,
light, nutrients, nest sites, etc.
Competition among members of the same
species is intraspecific.
Competition among individuals of different
species is interspecific.
Individuals experience both types of
competition, but the relative importance of the
two types of competition varies from population
to population and species to species

Styles of Competition
Exploitation competition occurs
when individuals use the same limiting
resource or resources, thus depleting
the amount available to others.
Interference competition occurs
when individuals interfere with the
foraging, survival, or reproduction of
others, or directly prevent their
physical establishment in a portion of a
habitat.

Some specific types of

competition
Consumptive competition
Preemptive competition
Overgrowth competition
Chemical composition
Territorial competition
Encounter competition

Example of Interference
Competition
The confused flour beetle, Triboleum

confusum, and the red flour beetle, Triboleum

castaneum cannibalize the eggs of their own
species as well as the other, thus interfering
with the survival of potential competitors.

In mixed species cultures, one species always

excludes the other. Which species prevails
depends upon environmental conditions,
chance, and the relative numbers of each
species at the start of the experiment.

Outcomes of Competition

Exploitation competition may cause the exclusion

of one species. For this to occur, one organism
must require less of the limiting resource to
survive. The dominant species must also reduce
the quantity of the resource below some critical
level where the other species is unable to replace
its numbers by reproduction.
Exploitation does not always cause the exclusion
of one species. They may coexist, with a
decrease in their potential for growth. For this to
occur, they must partition the resource.
Interference competition generally results in the
exclusion of one of the two competitors.

The Competitive Exclusion

Early in the twentieth century, two

Principle

mathematical biologists, A.J. Lotka and V.

Volterra developed a model of population
growth to predict the outcome of competition.
Their models suggest that two species cannot
compete for the same limiting resource for
long. Even a minute reproductive advantage
leads to the replacement of one species by
the other.
This is called the competitive exclusion
principal.

Evidence for Competitive

Exclusion.
A famous experiment by the Russian
ecologist, G.F. Gausse demonstrated that
Paramecium aurellia outcompetes and
displaces Paramecium caudatum in
mixed laboratory cultures, apparently
confirming the principle.
(Interestingly, this is not always the case.
Later studies suggest that the particular
strains involved affect the outcome of this
interaction).

Other experiments...
Subsequent laboratory studies on other
organisms, have generally resulted in
competitive exclusion, provided that
the environment was simple enough.
Example: Thomas Park showed that,
via interference competition, the
confused flour beetle and the red flower
beetle would not coexist. One species
always excluded the other.

Resource Partitioning
Species that share the same
habitat and have similar needs
frequently use resources in
somewhat different ways - so that
they do not come into direct
competition for at least part of the
limiting resource. This is called
resource partitioning.

 Resource partitioning obviates

competitive exclusion, allowing the
coexistence of several species using
the same limiting resource.
Resource partitioning could be an
evolutionary response to interspecific
competition, or it could simply be that
competitive exclusion eliminates all
situations where resource partitioning
does not occur.

 One of the best known cases of resource

partitioning occurs among Caribbean anoles.
As many as five different species of anoles may exist
in the same forest, but each stays restricted to a
particular space: some occupy tree canopies, some
occupy trunks, some forage close to the ground.
When the brown anole was introduced to Florida from
Cuba, it excluded the green anole from the trunks of
trees and areas near the ground: the green anole is
now restricted to the canopies of trees:the resource
(space, insects) has been partitioned among the two
species
(for now at least, this interaction may not be stable in the long
run because the species eat each others young).

Character Displacement

Sympatric populations of similar

species frequently have differences in
body structure relative to allopatric
populations of the same species.
This tendency is called character
displacement.
Character displacement is thought to
be an evolutionary response to
interspecific competition.

Example of Character

The best known case of character displacement occurs

Displacement

between the finches, Geospiza fuliginosa and Geospiza

fortis, on the Galapagos islands.
When the two species occur together, G. fuliginosa has a
much narrower beak that G fortis. Sympatric
populations of G fuliginosa eats smaller seeds than G
fortis: they partition the resource.
When found on separate islands, both species have
beaks of intermediate size, and exploit a wider variety of
seeds.
These inter-population differences might have evolved in
response to interspecific competition.

Competition and the Niche

An ecological niche can be thought of in terms
of competition.
The fundamental niche is the set of
resources and habitats an organism could
theoretically use under ideal conditions.
The realized niche is the set of resources and
habitats an organism actually used: it is
generally much more restricted due to
interspecific competition (or predation.)

Two organisms cannot occupy

exactly the same niche.
This is sometimes called Gausses
rule(although Gausse never put it
exactly that way).
-Experiments by Gausse (Paramecium), Peter
Frank (Daphnia), and Thomas Park (Triboleum)
have confirmed it for simple laboratory
scenarios.

-This creates a bit of a paradox, because so

many species exist in nature using the same
resources.

Amensalism
Amensalism is when one species
suffers and the other interacting
species experiences no effect.
Example: Redwood trees falling
into the ocean become floating
battering-rams during storms,
killing large numbers of mussels
and other inter-tidal organisms.

 Allelopathy involves the production and

release of chemical substances by one
species that inhibit the growth of another.
These secondary substances are
chemicals produced by plants that seen
to have no direct use in metabolism.
This same interaction can be seen as both
amensalism, and extremely one-sided
interference competition-in fact it is both.

Example: Allelopathy in the California

Chaparral
Black Walnut (Juglans nigra) trees excrete an
antibiotic called juglone. Juglone is known to
inhibit the growth of trees, shrubs, grasses, and
herbs found growing near black walnut trees.
Certain species of shrubs, notably Salvia
leucophylla (mint) and Artemisia californica
(sagebrush) are known to produce allelopathic
substances that accumulate in the soil during the
dry season. These substances inhibit the
germination and growth of grasses and herbs in an
area up to 1 to 2 meters from the secreting plants.

Commensalism

Commensalism is an interspecific interaction where one

species benefits and the other is unaffected.
Commensalisms are ubiquitous in nature: birds nesting in
trees are commensal.
Commensal organisms frequently live in the nests, or on
the bodies, of the other species.
Examples of Commensalism:
Ant colonies harbor rove beetles as commensals. These
beetles mimic the ants behavior, and pass as ants. They
eat detritus and dead ants.
Anemonefish live within the tentacles of anemones. They
have specialized mucus membranes that render them
immune to the anemones stings. They gain protection by
living in this way.

Mutualism
Mutualism in an interspecific interaction
between two species that benefits both
members.
Populations of each species grow,
survive and/or reproduce at a higher rate
in the presence of the other species.
Mutualisms are widespread in nature,
and occur among many different types
of organisms.

Examples of Mutualism

Most rooting plants have mutualistic

associations with fungal mychorrhizae.
Mychorrhizae increase the capability of plant
roots to absorb nutrients. In return, the host
provides support and a supply of
carbohydrates.
Many corals have endosymbiotic organisms
called zooxanthellae (usually a dinoflagellate).
These mutualists provide the corals with
carbohydrates via photosynthesis. In return,
they receive a relatively protected habitat
from the body of the coral.

Mutualistic Symbiosis
Mutualistic Symbiosis is a type of mutualism in
which individuals interact physically, or even live
within the body of the other mutualist.
Frequently, the relationship is essential for the
survival of at least one member.
Example: Lichens are a fungal-algal symbiosis (that
frequently includes a third member, a cyanobacterium.)
The mass of fungal hyphae provides a protected habitat
for the algae, and takes up water and nutrients for the
algae. In return, the algae (and cynaobacteria) provide
carbohydrates as a source of energy for the fungus.

Facultative vs. Obligate

Mutualisms
Facultative Mutualisms are not
essential for the survival of either
species. Individuals of each
species engage in mutualism when
the other species is present.
Obligate mutualisms are essential
for the survival of one or both
species.

Other Examples of
Mutualisms
Flowering plants and pollinators. (both
facultative and obligate)
Parasitoid wasps and polydna viruses.
(obligate)
Ants and aphids. (facultative)
Termites and endosymbiotic protozoa.
(obligate)
Humans and domestic animals. (mostly
facultative, some obligate)

Predation, Parasitism,
Herbivory
Predators, parasites, parasitoids, and
herbivores obtain food at the expense of
their hosts or prey.

 Predators tend to be larger

than their prey, and consume
many prey during their
lifetimes.

 Parasites and pathogens are

smaller than their host. Parasites
may have one or many hosts during
their lifetime. Pathogens are
parasitic microbes-many
generations may live within the
same host. Parasites consume their
host either from the inside
(endoparasites) or from the
outside (ectoparasites).

 Parasitoids hunt their prey like

predators, but lay their eggs within
the body of a host, where they
develop like parasites.

 Herbibores are animals that eat

plants. This interaction may
resemble predation, or parasitism.

Predator-Prey and ParasiteHost Coevolution

The relationships between
predator and prey, and parasites
and hosts, have coevolved over
long periods of time.

 About 50 years ago, an evolutionary biologist named J.B.S.

Haldane suggested that the interaction between parasite and
host (or predator and prey) should resemble an
evolutionary arms race:
First a parasite (or predator) evolves a trait that allows it to attack its
host (or prey).
Next, natural selection favors host individuals that are able to defend
themselves against the new trait.
As the frequency of resistant host individuals increases, there is natural
selection for parasites with novel traits to subvert the host defenses.
This process continues as long as both species survive.
Recent data on Plasmodium, the cause of malaria, support this model.

Example of Parasite-Host
Coevolution
The common milkweed, Asclepias syriaca has leaves
that contain cardiac glycosides: they are very
poisonous to most herbivores. This renders them
virtually immune to herbivory by most species.
Monarch butterfly larvae have evolved the ability to
tolerate these toxins, and sequester them within
their bodies. They are important specialist
hervivores of milkweeds.
These sequestered compounds serve the additional
purpose of making monarch larvae virtually inedible
to vertebrate predators.

Predator-Prey Population
Predation may be a density-dependent mortality
Dynamics
factor to the host population-and prey may
represent a limiting resource to predators.
The degree of prey mortality is a function of the
density of the predator population.
The density of the prey population, in turn, affects
the birth and death rates of the predator population.
i.e, when prey become particularly common,
predators increase in numbers until prey die back
due to increased predation, this, in turn, inhibits the
growth of prey.
Typically, there is a time lag effect.

 There is often a dynamic

balance between predators
and prey that is necessary for
the stability of both populations.
Feedback mechanisms may
control the densities of both
species.

Example of Regulation of Host Population

by a Herbivore

In the 19th century, prickly pear cactus, Opuntia sp.

was introduced into Australia from South America.
Because no Australian predator species existed to
control the population size of this cactus, it quickly
expanded throughout millions of acres of grazing land.
The presence of the prickly pear cactus excluded cattle
and sheep from grazing vegetation and caused a
substantial economic hardship to farmers.
A method of control of the prickly pear cactus was
initiated with the introduction of Cactoblastis cactorum,
a cactus eating moth from Argentina, in 1925. By 1930,
densities of the prickly pear cactus were significantly
reduced.

 Sometimes predator species can drive their

prey to localized extinction.
If there are no alternate prey, the predator then goes
extinct.
If the environment is coarse grained, this makes
the habitat available for recolonization by the prey
species.
Example: The parasitic wasp Dieratiella rapae is a
very efficient parasitoid. One female can oviposit
into several hundred aphids during its lifetime.
Frequently, aphids are driven locally extinct and the
adults must search for new patches when they
emerge. Once the aphid and the host are gone, the
host plants may become re-infested with aphids.

 In other cases, there are alternate

prey to support the predator and the
prey is permanently excluded.
Example: Freshwater fish such as
bluegills and yellow perch frequently
exclude small invertebrates such as
Daphnia pulex from ponds. The fish
then switch to other prey such as
insects larvae.

The time-lag effect may lead to

predator-prey
oscillations.

Most predators do not respond instantaneously to

the availability of prey and adjust their reproduction
accordingly.
If predator populations grow faster than prey
populations, they may overshoot the number of prey
that are able to support them
This leads to a rapid decline in the prey, followed by
a rapid decline in the predator.
Once the predator becomes rare, the prey
population may begin growing again.
This pattern is called a predator-prey oscillation.

Cycles in the population dynamics of the snowshoe

hare and its predator the Canadian lynx (redrawn
from MacLulich 1937). Note that percent mortality is
an elusive measure, it may, or may not, be useful
since mortality varies with environment and time.

In the 1920s, A. J. Lotka (1925) and V. Volterra (1926)

devised mathematical models representing host/prey
interaction.
The Lotka-Volterra curve assumes that prey
destruction is a function not only of natural enemy
numbers, but also of prey density, i.e., related to the
chance of encounter.
This model predicts the predator-prey oscillations
sometimes seen in nature. Populations of prey and
predator were predicted to flucuate in a regular
manner (Volterra termed this "the law of periodic
cycle").
Lotka-Volterra model is an oversimplification of
reality. In nature, many different factors affect the
densities of predators and their prey.