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Chapter 53

Community Ecology

PowerPoint Lectures for


Biology, Seventh Edition
Neil Campbell and Jane Reece

Lectures by Chris Romero


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• Overview: What Is a Community?

• A biological community
– Is an assemblage of populations of various
species living close enough for potential
interaction

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• The various animals and plants surrounding
this watering hole
– Are all members of a savanna community in
southern Africa

Figure 53.1
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• Concept 53.1: A community’s interactions
include competition, predation, herbivory,
symbiosis, and disease
• Populations are linked by interspecific
interactions
– That affect the survival and reproduction of the
species engaged in the interaction

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• Interspecific interactions
– Can have differing effects on the populations
involved

Table 53.1
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Competition
• Interspecific competition
– Occurs when species compete for a particular
resource that is in short supply

• Strong competition can lead to competitive


exclusion
– The local elimination of one of the two
competing species

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The Competitive Exclusion Principle
• The competitive exclusion principle
– States that two species competing for the
same limiting resources cannot coexist in the
same place

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Ecological Niches
• The ecological niche
– Is the total of an organism’s use of the biotic
and abiotic resources in its environment

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• The niche concept allows restatement of the
competitive exclusion principle
– Two species cannot coexist in a community if
their niches are identical

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• However, ecologically similar species can
coexist in a community
– If there are one or more significant difference
in their niches
EXPERIMENT Ecologist Joseph Connell studied two barnacle RESULTS When Connell removed Balanus from the lower
speciesBalanus balanoides and Chthamalus stellatus that have a strata, the Chthamalus population spread into that area.
stratified distribution on rocks along the coast of Scotland.

High tide High tide


Chthamalus
Chthamalus
Balanus realized niche

Chthamalus
fundamental niche
Balanus
realized niche

Ocean Ocean
Low tide Low tide

In nature, Balanus fails to survive high on the rocks because it is


unable to resist desiccation (drying out) during low tides. Its realized
niche is therefore similar to its fundamental niche. In contrast,
Chthamalus is usually concentrated on the upper strata of rocks. To CONCLUSION The spread of Chthamalus when Balanus was
determine the fundamental of niche of Chthamalus, Connell removed removed indicates that competitive exclusion makes the realized
Balanus from the lower strata. niche of Chthamalus much smaller than its fundamental niche.
Figure 53.2

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• As a result of competition
– A species’ fundamental niche may be different
from its realized niche

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Resource Partitioning
• Resource partitioning is the differentiation of
niches
– That enables similar species to coexist in a
community A. insolitus
usually perches
on shady branches.

A. ricordii

A. insolitus
A. distichus perches
on fence posts and A. alinigar A. christophei
other sunny
A. distichus
surfaces.
A. cybotes
A. etheridgei

Figure 53.3
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Character Displacement
• In character displacement
– There is a tendency for characteristics to be more
divergent in sympatric populations of two species
than in allopatric populations of the same two G. fortis

species G. fuliginosa

Beak
depth

Santa María, San Cristóbal


Percentages of individuals in each size class
40 Sympatric
20 populations

Los Hermanos

40
G. fuliginosa,
20
allopatric
0

Daphne

40

20 G. fortis, allopatric
0
8 10 12 14 16
Beak depth (mm)
Figure 53.4
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Predation
• Predation refers to an interaction
– Where one species, the predator, kills and eats
the other, the prey

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• Feeding adaptations of predators include
– Claws, teeth, fangs, stingers, and poison

• Animals also display


– A great variety of defensive adaptations

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• Cryptic coloration, or camouflage
– Makes prey difficult to spot

Figure 53.5
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• Aposematic coloration
– Warns predators to stay away from prey

Figure 53.6
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• In some cases, one prey species
– May gain significant protection by mimicking
the appearance of another

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• In Batesian mimicry
– A palatable or harmless species mimics an
unpalatable or harmful model

(b) Green parrot snake

Figure 53.7a, b (a) Hawkmoth larva


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• In Müllerian mimicry
– Two or more unpalatable species resemble
each other

(a) Cuckoo bee

Figure 53.8a, b
(b) Yellow jacket
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Herbivory
• Herbivory, the process in which an herbivore
eats parts of a plant
– Has led to the evolution of plant mechanical
and chemical defenses and consequent
adaptations by herbivores

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Parasitism
• In parasitism, one organism, the parasite
– Derives its nourishment from another
organism, its host, which is harmed in the
process

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• Parasitism exerts substantial influence on
populations
– And the structure of communities

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Disease
• The effects of disease on populations and
communities
– Is similar to that of parasites

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• Pathogens, disease-causing agents
– Are typically bacteria, viruses, or protists

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Mutualism
• Mutualistic symbiosis, or mutualism
– Is an interspecific interaction that benefits both
species

Figure 53.9
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Commensalism
• In commensalism
– One species benefits and the other is not
affected

Figure 53.10
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• Commensal interactions have been difficult to
document in nature
– Because any close association between
species likely affects both species

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Interspecific Interactions and Adaptation
• Evidence for coevolution
– Which involves reciprocal genetic change by
interacting populations, is scarce

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• However, generalized adaptation of organisms
to other organisms in their environment
– Is a fundamental feature of life

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• Concept 53.2: Dominant and keystone species
exert strong controls on community structure
• In general, a small number of species in a
community
– Exert strong control on that community’s
structure

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Species Diversity
• The species diversity of a community
– Is the variety of different kinds of organisms
that make up the community
– Has two components

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• Species richness
– Is the total number of different species in the
community

• Relative abundance
– Is the proportion each species represents of
the total individuals in the community

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• Two different communities
– Can have the same species richness, but a
different relative abundance
A
B

Community 1
A: 25% B: 25% C: 25% D: 25%

Community 2
Figure 53.11 A: 80% B: 5% C: 5% D: 10%

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• A community with an even species abundance
– Is more diverse than one in which one or two
species are abundant and the remainder rare

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Trophic Structure
• Trophic structure
– Is the feeding relationships between organisms
in a community
– Is a key factor in community dynamics

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• Food chains
Quaternary
consumers

– Link the trophic


levels from Carnivore Carnivore

producers to top
Tertiary
consumers

carnivores Carnivore Carnivore

Secondary
consumers

Carnivore Carnivore

Primary
consumers

Herbivore Zooplankton

Primary
producers

Plant Phytoplankton
Figure 53.12 A terrestrial food chain A marine food chain

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Food Webs
• A food web Humans

– Is a branching
food chain with Baleen
whales
Smaller toothed
whales
Sperm
whales

complex
trophic Crab-eater seals
Leopard
seals
Elephant
seals

interactions
Birds Fishes Squids

Carnivorous
plankton

Euphausids Copepods
(krill)

Phyto-
plankton

Figure 53.13
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• Food webs can be simplified
– By isolating a portion of a community that
interacts very little with the rest of the
community

Sea nettle Juvenile striped bass

Fish larvae

Figure 53.14 Fish eggs Zooplankton

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Limits on Food Chain Length
• Each food chain in a food web
– Is usually only a few links long

• There are two hypotheses


– That attempt to explain food chain length

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• The energetic hypothesis suggests that the
length of a food chain
– Is limited by the inefficiency of energy transfer
along the chain

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• The dynamic stability hypothesis
– Proposes that long food chains are less stable
than short ones

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• Most of the available data
– Support the energetic hypothesis

6 6
No. of species
5 5

Number of trophic links


No. of trophic
Number of species

4 links 4

3 3

2 2

1 1

0 0
High Medium Low
(control)
Productivity
Figure 53.15
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Species with a Large Impact
• Certain species have an especially large
impact on the structure of entire communities
– Either because they are highly abundant or
because they play a pivotal role in community
dynamics

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Dominant Species
• Dominant species
– Are those species in a community that are
most abundant or have the highest biomass
– Exert powerful control over the occurrence and
distribution of other species

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• One hypothesis suggests that dominant
species
– Are most competitive in exploiting limited
resources

• Another hypothesis for dominant species


success
– Is that they are most successful at avoiding
predators

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Keystone Species
• Keystone species
– Are not necessarily abundant in a community

– Exert strong control on a community by their


ecological roles, or niches

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• Field studies of sea stars
– Exhibit their role as a keystone species in intertidal
communities

With Pisaster (control)


20

Number of species
15

present
10 Without Pisaster (experimental)

0
1963 ´64 ´65 ´66 ´67 ´68 ´69 ´70 ´71 ´72 ´73

(a) The sea star Pisaster ochraceous feeds (b) When Pisaster was removed from an intertidal zone,
preferentially on mussels but will mussels eventually took over the rock face and eliminated
consume other invertebrates. most other invertebrates and algae. In a control area from
which Pisaster was not removed, there was little change
Figure 53.16a,b in species diversity.

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• Observation of sea otter populations and their
predation
– Shows the
100

80

(% max. count)
Otter number
effect the
60

40

otters have
20

0
(a) Sea otter abundance

on ocean 400

communities 300

Grams per
0.25 m2
200
100
0
(b) Sea urchin biomass

10
8
Number per

6
0.25 m2

4
2
0
1972 1985 1989 1993 1997
Year
Food chain before (c) Total kelp density Food chain after killer
Figure 53.17 killer whale involve- whales started preying
ment in chain on otters

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Ecosystem “Engineers” (Foundation Species)
• Some organisms exert their influence
– By causing physical changes in the
environment that affect community structure

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• Beaver dams
– Can transform landscapes on a very large
scale

Figure 53.18
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• Some foundation species act as facilitators
– That have positive effects on the survival and
reproduction of some of the other species in the
community
8

Number of plant species


6

0
With Without
Juncus Juncus
Salt marsh with Juncus
(foreground) Conditions
Figure 53.19
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Bottom-Up and Top-Down Controls
• The bottom-up model of community
organization
– Proposes a unidirectional influence from lower
to higher trophic levels

• In this case, the presence or absence of abiotic


nutrients
– Determines community structure, including the
abundance of primary producers

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• The top-down model of community
organization
– Proposes that control comes from the trophic
level above

• In this case, predators control herbivores


– Which in turn control primary producers

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• Long-term experiment studies have shown
– That communities can shift periodically from
bottom-up to top-down
100

75
herbaceous plant cover
Percentage of

50

25

0
0 100 200 300 400

Figure 53.20 Rainfall (mm)

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• Pollution
– Can affect community dynamics

• But through biomanipulation


– Polluted communities can be restored
Polluted State Restored State

Fish Abundant Rare

Zooplankton Rare Abundant

Algae Abundant Rare

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• Concept 53.3: Disturbance influences species
diversity and composition
• Decades ago, most ecologists favored the
traditional view
– That communities are in a state of equilibrium

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• However, a recent emphasis on change has
led to a nonequilibrium model
– Which describes communities as constantly
changing after being buffeted by disturbances

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What Is Disturbance?
• A disturbance
– Is an event that changes a community

– Removes organisms from a community

– Alters resource availability

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• Fire
– Is a significant disturbance in most terrestrial
ecosystems
– Is often a necessity in some communities

Figure 53.21a–c (a) Before a controlled burn. (b) During the burn. The detritus (c) After the burn. Approximately one
A prairie that has not burned for serves as fuel for fires. month after the controlled burn,
several years has a high propor- virtually all of the biomass in this
tion of detritus (dead grass). prairie is living.

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• The intermediate disturbance hypothesis
– Suggests that moderate levels of disturbance
can foster higher species diversity than low
levels of disturbance

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• The large-scale fire in Yellowstone National
Park in 1988
– Demonstrated that communities can often
respond very rapidly to a massive disturbance

(a) Soon after fire. As this photo taken soon after the fire shows, (b) One year after fire. This photo of the same general area taken the
the burn left a patchy landscape. Note the unburned trees in the following year indicates how rapidly the community began to
distance. recover. A variety of herbaceous plants, different from those in the
former forest, cover the ground.
Figure 53.22a, b
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Human Disturbance
• Humans
– Are the most widespread agents of
disturbance

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• Human disturbance to communities
– Usually reduces species diversity

• Humans also prevent some naturally occurring


disturbances
– Which can be important to community
structure

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Ecological Succession
• Ecological succession
– Is the sequence of community and ecosystem
changes after a disturbance

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• Primary succession
– Occurs where no soil exists when succession
begins

• Secondary succession
– Begins in an area where soil remains after a
disturbance

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• Early-arriving species
– May facilitate the appearance of later species
by making the environment more favorable
– May inhibit establishment of later species

– May tolerate later species but have no impact


on their establishment

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• Retreating glaciers
– Provide a valuable field-research opportunity on
succession
Canada
Grand

Ri

l.
gg
Alaska

G
Pacific Gl. 0 5 10

sG

ide
ui
r

l.

Br
G

Mc
l.
1940 1912 1948 Miles

Pl

l.
a
1941

tG
1899

te
au

en
G

em
1931

l.
1907 1879 1948 1911

s
Ca
1900
1879 1879 1892 1913
1935 1949

1860
Reid Gl.
1879
Johns Hopkins
Gl. Glacier
Bay

1830
1780

1760
Pleasant Is.

Figure 53.23 McBride glacier retreating

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• Succession on the moraines in Glacier Bay, Alaska
– Follows a predictable pattern of change in vegetation
and soil characteristics (a) Pioneer stage, with fireweed dominant

(b) Dryas stage

60

50
Soil nitrogen (g/m2)

40

30

20

10

0
Pioneer Dryas Alder Spruce
Successional stage
(c) Spruce stage

Figure 53.24a–d (d) Nitrogen fixation by Dryas and alder


increases the soil nitrogen content.

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• Concept 53.4: Biogeographic factors affect
community diversity
• Two key factors correlated with a community’s
species diversity
– Are its geographic location and its size

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Equatorial-Polar Gradients
• The two key factors in equatorial-polar
gradients of species richness
– Are probably evolutionary history and climate

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• Species richness generally declines along an
equatorial-polar gradient
– And is especially great in the tropics

• The greater age of tropical environments


– May account for the greater species richness

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• Climate
– Is likely the primary cause of the latitudinal
gradient in biodiversity

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• The two main climatic factors correlated with
biodiversity
– Are solar energy input and water availability
180

160 200
Tree species richness

140

Vertebrate species
100
120

(log scale)
richness
100
50
80

60
40

20
10
0 1
500 1,000 1,500 2,000
100 300 500 700 900 1,100
Actual evapotranspiration (mm/yr) Potential evapotranspiration (mm/yr)
(a) Trees (b) Vertebrates

Figure 53.25a, b

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Area Effects
• The species-area curve quantifies the idea that
– All other factors being equal, the larger the
geographic area of a community, the greater
the number of species

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• A species-area curve of North American
breeding birds
– Supports this idea
1,000
Number of species (log scale)

100

10

1
1 10 100 103 104 105 106 107 108 109 1010
Area (acres)
Figure 53.26
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Island Equilibrium Model
• Species richness on islands
– Depends on island size, distance from the
mainland, immigration, and extinction

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• The equilibrium model of island biogeography
maintains that
– Species richness on an ecological island
levels off at some dynamic equilibrium point

d)
n
Im

sla tion
io

Im
an
Im
m

(n
ct
Rate of immigration or extinction

Rate of immigration or extinction


Rate of immigration or extinction

)
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ig
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Equilibrium number Small island Large island Far island Near island

Number of species on island Number of species on island Number of species on island

(a) Immigration and extinction rates. The (b) Effect of island size. Large islands may (c) Effect of distance from mainland.
equilibrium number of species on an ultimately have a larger equilibrium num- Near islands tend to have larger
island represents a balance between the ber of species than small islands because equilibrium numbers of species than
immigration of new species and the immigration rates tend to be higher and far islands because immigration rates
extinction of species already there. extinction rates lower on large islands. to near islands are higher and extinction
rates lower.
Figure 53.27a–c
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• Studies of species richness on the Galápagos Islands

– Support the prediction that species richness


increases with island size
FIELD STUDY Ecologists Robert MacArthur and E. O. Wilson studied the
number of plant species on the Galápagos Islands, which vary greatly in size, in
relation to the area of each island.

RESULTS

400

200
Number of plant species (log scale)

100

50

25

10

0
0.1 1 10 100 1,000
Area of island (mi2)
(log scale)

CONCLUSION The results of the study showed that plant species


Figure 53.28 richness increased with island size, supporting the species-area theory.

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• Concept 53.5: Contrasting views of community
structure are the subject of continuing debate
• Two different views on community structure
– Emerged among ecologists in the 1920s and
1930s

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Integrated and Individualistic Hypotheses
• The integrated hypothesis of community
structure
– Describes a community as an assemblage of
closely linked species, locked into association
by mandatory biotic interactions

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• The individualistic hypothesis of community
structure
– Proposes that communities are loosely
organized associations of independently
distributed species with the same abiotic
requirements

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• The integrated hypothesis
– Predicts that the presence or absence of
particular species depends on the presence or
absence of other species
densities of
Population

individual
species

Environmental gradient
(such as temperature or moisture)

(a) Integrated hypothesis. Communities are discrete groupings


of particular species that are closely interdependent and nearly
Figure 53.29a always occur together.

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• The individualistic hypothesis
– Predicts that each species is distributed
according to its tolerance ranges for abiotic
factors
densities of
Population

individual
species

Environmental gradient
(such as temperature or moisture)
(b) Individualistic hypothesis. Species are independently
distributed along gradients and a community is simply the
assemblage of species that occupy the same area because of
Figure 53.29b similar abiotic needs.

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• In most actual cases the composition of
communities
– Seems to change continuously, with each
species more or less independently distributed

600
per hectare
Number of
plants

400

200

0
Wet Moisture gradient Dry

(c) Trees in the Santa Catalina Mountains. The distribution of tree species at one
elevation in the Santa Catalina Mountains of Arizona supports the individualistic
hypothesis. Each tree species has an independent distribution along the gradient,
apparently conforming to its tolerance for moisture, and the species that live
together at any point along the gradient have similar physical requirements.
Because the vegetation changes continuously along the gradient, it is impossible to
Figure 53.29c delimit sharp boundaries for the communities.

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Rivet and Redundancy Models
• The rivet model of communities
– Suggests that all species in a community are
linked together in a tight web of interactions
– Also states that the loss of even a single
species has strong repercussions for the
community

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• The redundancy model of communities
– Proposes that if a species is lost from a
community, other species will fill the gap

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• It is important to keep in mind that community
hypotheses and models
– Represent extremes, and that most
communities probably lie somewhere in the
middle

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