Lecture 10

COMMUNITY STRUCTURE
Community Structure
Paine ‘80 - Keystone Species

Keystone species
-a relatively low biomass species that is a
major factor in determining community structure
Community Structure
Paine ‘80 - Keystone Species
Introduced concept of ‘links’ in community

e.g. Pisaster and Mytilus

Pisaster Tonicella Katherina

Mytilus Lithophyllum Hedophyllum

Community Structure
Paine ‘80 - Keystone Species
Introduced concept of ‘links’ in community

e.g. Pisaster and Mytilus

strong link weak links

Pisaster Tonicella Katherina

Mytilus Lithophyllum Hedophyllum

Community Structure
Paine ‘80 - Keystone Species
Introduced concept of ‘links’ in community

e.g. Pisaster and Mytilus

strong link weak links

Pisaster Tonicella Katherina

Mytilus Lithophyllum Hedophyllum

Module 1 Module 2
Keystone species

Enhydra lutris

Strongylocentrotus purpuratus

Macrocystis pyrifera
But in reality - much more complex
But in reality - VERY much more complex
 Another take on this - Menge and Sutherland ‘87

Rat Gulls
Food Web

Fish

Crabs Sea stars

Whelk Limpet Snails

Urchin

Barnacles Red algae

Crusts
Polychaete Emphemerals
Bivalves
 Another take on this - Menge and Sutherland ‘87

Interaction web
Rat Gulls
strong links
Fish
weak links

Crabs Sea stars

Whelk Limpet Snails

Urchin

Barnacles Red algae

Crusts
Polychaete Emphemerals
Bivalves
Interactions between species in an intertidal community
Can be looked at in one of two ways

a) Hierarchy
A

A A C
B C
A
B B
C

b) Network

A A B C
C
C
B
B C A A
B

Time
Community Structure
Multispecies interactions - Underwood

Tesseropora - barnacle Patelloida -acamaeid limpet

Cellana - patellid limpet Morula - predatory whelk

Community Structure
Multispecies interactions - Underwood

Exclusion cages
Community Structure
Multispecies interactions - Underwood

Experiment 1 - Effect of Cellana on survival of Tesseropa

Morula - excluded

Tesseropora

Cellana
-grazes open rock
Community Structure
Multispecies interactions - Underwood

Experiment 1 - Effect of Cellana on survival of Tesseropa

Higher survivorship at
intermediate density
% Tesseropora
surviving

0 2 4 6 8 10
# Cellana/enclosure
Community Structure
Multispecies interactions - Underwood

Experiment 2 - Effect of Tesseropa on growth of Cellana

Cellana growth
rate

Tesseropa density
Community Structure
Multispecies interactions - Underwood

Experiment 2 - Effect of Tesseropa on growth of Cellana

Experiment 2 - Effect of Tesseropa on growth of Patelloida

Cellana growth
rate

Patelloida
growth rate

Tesseropa density
Community Structure
Multispecies interactions - Underwood

General conclusion

- major interactions among all components of system

- no keystone species - no links are ‘weak’

 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
Habitat
amelioration b) high and low recruitment

Relative
importance

high low

Stress

HIGH RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
Habitat
amelioration b) high and low recruitment

Relative
Competition
importance

high low

Stress

HIGH RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
Habitat
amelioration b) high and low recruitment

Predation
Relative
Competition
importance

high low

Stress

HIGH RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
Habitat
amelioration b) high and low recruitment

Predation
Relative
Competition
importance

Associational
defenses

high low

Stress

HIGH RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
b) high and low recruitment

Relative
Competition
importance

high low

Stress

LOW RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
b) high and low recruitment

Predation

Relative
Competition
importance

high low

Stress

LOW RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
b) high and low recruitment

Predation

Relative
Competition
importance

Associational
defenses

high low

Stress

LOW RECRUITMENT
 Community Structure
How do the forces shaping communities interact?
(Menge and Sutherland, Amer. Nat. 110:351)

-consider situations of
a) high and low stress
b) high and low recruitment

Habitat Predation
amelioration
Relative
Competition
importance

Associational
defenses

high low

Stress

LOW RECRUITMENT
Community Structure
Relationship to diversity

Diversity

high low

Stress
Community Structure
Relationship to diversity
Negative effects of
competition and
Negative effects of
predation
stress

Diversity

high low

Stress
Community Structure
Disturbance and Diversity

Connell - Intermediate Disturbance Hypothesis

Diversity

Low High
Disturbance
(frequency, intensity, extent)
Community Structure
Disturbance and Diversity

Connell - Intermediate Disturbance Hypothesis

Competitive
dominants fill
space

Diversity

Low High
Disturbance
(frequency, intensity, extent)
Community Structure
Disturbance and Diversity

Connell - Intermediate Disturbance Hypothesis

Competitive Few species can

dominants fill invade or persist
space

Diversity

Low High
Disturbance
(frequency, intensity, extent)
Models of succession

1. Inhibition
- initial colonizers inhibit future colonizers.

2. Tolerance
- initial colonizers do not affect later colonizers. Organisms best suited to
local conditions will persist (e.g. Chthalamus/Semibalanus)

3. Facilitation
- initial colonizers facilitate success of later colonizers (e.g. protection of
barnacles growth of algae
Models of succession
-probably work in different areas

Habitat
Recruitment
stress

Inhibition low high

Tolerance midrange low

Facilitation high high or low

 Community Structure:

Top-Down vs. Bottom-Up Control

Hairston, Smith and Slobodkin (1960)

Community Dynamics

Carnivores Detritivores
control

Resource limited

Frees plants
Herbivores Plants
from herbivore
control
Critiques

Too Simple

1) Species differences matter

2) Plant dominance could be explained by good defences

Other inferences of Hairston et al, 1960

1) Exceptions not important ??

2) All communities have 3 trophic levels X

3) Omnivory not important X

4) External abiotic factors - not controllers X

 Robles et al, 1995

Mean
Density
(seastars/m2)

Recruitment index of Mytilus

Menge and Sutherland, 1976

Effects of predation by whelks.

Predation is weak

High wave energy - effects of predation -weak

Moderate wave energy

- effects of predation - strong

Menge Sutherland
 Ecological Relationships in Kelp Forests

Kelp Orca

Sea Otter
Urchins
Bottom Up Control

Fretwell, 1977, 1987

- availability of plant material governs structure of food chains

- Low productivity - 1 link (plants)

- Higher productivity - add links

Transplant mussels and barnacles (filter feeders) to urchin-dominated and
kelp-dominated substrates
Transplant mussels and barnacles (filter feeders) to urchin-dominated and
kelp-dominated substrates

Expected (top down)

Urchin

Kelp
Transplant mussels and barnacles (filter feeders) to urchin-dominated and
kelp-dominated substrates

Expected (top down) Observed (bottom up)

Urchin Urchin Kelp

Kelp
Clearly - can be a complex interaction

Increased nutrient

Increased algae

Increased benthic filter feeders

Increased consumers (predation)

control
Interaction of Systems High flow

Leonard et al, 1998 Low flow

Interaction of Systems

• increased seaweed growth

• increased filter feeder growth
• increased consumer pressure
• increased larval settlement
• increased sedimentation
• low consumer efficiency
• lower densities of
• higher densities of organisms with planktonic
organisms with planktonic larvae
• increased consumer mortality
•larvae
more spatial competition
• less spatial competition
 Leonard et al, 1998

Hydrodynamics

Flow rate

Time
 Leonard et al, 1998
Community structure
High flow Low flow
T
i
d
e

h
e
i
g
h
t
Percent cover Percent cover

barnacles Fucus

mussels Bare space

 Leonard et al, 1998

High flow
Recruitment rates
Low flow

Barnacles Mussels Snails

Density
(#/100 cm2)
Leonard et al, 1998

Crab predation

High flow

Low flow

On Littorina, Nucella, Mytilus

Predation Intensity
(% mortality)
 Leonard et al, 1998

crabs

grazers

mussels barnacles
diatoms

Nutrients Larvae Plankton

 Leonard et al, 1998

crabs crabs

whelks
grazers grazers

mussels barnacles mussels barnacles

diatoms diatoms

Nutrients Larvae Plankton Nutrients Larvae Plankton

Interference competition,
exploitative competition
for resources other than
Depletion of more
food (-) nutritious,
Predators palatable or
(-) accessible prey
(-)

Induced morphological or - (+)

chemical defenses,
hiding, retreat to refuges

Consumers
+ (-)

Cover from (for) predators

Stimulation of area-
specific primary
productivity

Plants
(+)
Powers. ‘92. Ecology 73: 733