Overview: What Is A Community? A Biological Community

• Overview: What Is a Community?
• A biological community
– Is an assemblage of populations of various
species living close enough for potential
interaction
Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Cummings
• 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
EXPERIMENTEcologist Joseph Connell studied two barnacle RESULTS
When Connell removed Balanus from the lower strata,
species Balanus balanoides andChthamalus stellatus that have a theChthamalus 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 CONCLUSIONThe spread of Chthamalus whenBalanus was removed
determine the fundamental of niche of Chthamalus, Connell 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
surfaces. A. distichus
A. cybotes
A. etheridgei
Figure 53.3
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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 Baleen
whales
Smaller toothed
whales
Sperm
whales
with 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
links 4
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
WithPisaster (control)
20
Number of species
15
present
10 WithoutPisaster (experimental)
0
1963 ´64 ´65 ´66 ´67 ´68 ´69 ´70 ´71 ´72 ´73
(a) The sea starPisaster ochraceous feeds (b) WhenPisaster 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
Pisaster was not removed, there was little change in
which
Figure 53.16a,b 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
on ocean
(a) Sea otter abundance
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
(c) Total kelp density Food chain after killer
Food chain before
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
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, the ( b) One year after fire. This photo of the same general area taken the
burn left a patchy landscape. Note the unburned trees in the following year indicates how rapidly the community began to recover.
distance. 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
.
g
Gl
Pacific Gl. Alaska 0 5 10
gs
M
de
ui
Gl
Bri
rG
Mc
l.
1940 1912 1948 Miles
Pl
l.
a
1941
tG
1 899
te
a
n
u
me
Gl
1907 1931 1911
se
1879 1948
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
Vertebrate species richness

Tree species richness
140
100
120
(log scale)
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
Im
sla t io n
d)
Im
io
Rate of immigration or extinction

Im g e
m
(n
an
ct
m
ig
(fa xt inc
)
ea
( la
isl
n
t in
nd
ig
ra
ig
io
ri
la o n
r
t io
ra
Ex
ra
ct
all
sla
E
i
t io
t io
t in
ct
n
Im
ri
m
nd
t in
nd
i sla
m
n
Ex
(s
( la E x
ig
)
(fa
is
nd
ri ra
e
)
sla t io n
rg
nd n t io
I
)
m
c
t in land
(s
)
m
m
Ex
ig
i s
all
ra
r
ea
t io
isl
(n
n
an
d)
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(mi 2)
(log scale)
CONCLUSION The results of the study showed that plant species richness
Figure 53.28 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
Number of
per hectare
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 delimit sharp
Figure 53.29c 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

Cummings

Overview: What Is A Community? A Biological Community

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• Overview: What Is a Community?

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

• Strong competition can lead to competitive

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

High tide High tide

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

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

• Animals also display

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(b) Green parrot snake

(a) Cuckoo bee

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

– Link the trophic

producers to top Tertiary

carnivores Carnivore Carnivore

Sea nettle Juvenile striped bass

Figure 53.14 Fish eggs Zooplankton

• There are two hypotheses

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Number of trophic links

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• Another hypothesis for dominant species

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– Can transform landscapes on a very large

Number of plant species

• In this case, the presence or absence of abiotic

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• In this case, predators control herbivores

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Figure 53.20 Rainfall (mm)

• But through biomanipulation

Fish Abundant Rare

Zooplankton Rare Abundant

Algae Abundant Rare

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Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

Copyright © 2005 Pearson Education, Inc. publishing as Benjamin

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