Food Webs – 3 BI2020 M Pinard

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 Previously
• Predation
– Process of Predation and Factors that Influence
– Functional Response Curves
– Predation Hypothesis
– Optimal Foraging Theory and Predator Preferences
– Predator-Prey Dynamics through L-V

• Food Webs
– Types and attributes
– Energy Limitation, Ecosystem Size, Productive Space Hypotheses
– Top-down and Bottom-up Regulation
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Top down effects

Predators can influence the

abundances of species in lower
trophic levels

Direct and indirect effects

Bottom-up effects

Basal resource can influence the

abundance of species at higher
trophic levels

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 See Mittlebach, Chapter 11
Trophic cascade

Islands without otters, sea

urchins boomed, kelp forest
communities degraded

Predator reduces prey abundance . . . cascades down such that the

prey’s resources increase in abundance 4
In 1987, bass were reintroduced to a lake. Bass
population increased rapidly following the
introduction (A).

As bass numbers increased, they decimated the

population of planktivorous minnows (B).

Loss of minnows allowed for the return of large-

bodied herbivorous zooplankton (C).

The return of Daphnia resulted in an increase in

water clarity (D).
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If you have a trophic cascade in 4 level system, top carnivore and herbivore numbers correlated

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If you have a trophic cascade in 4 level system, top carnivore and herbivore numbers correlated
. . . . Not always the case
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What do complex networks of species interactions tell
us about what controls the abundances of plants,
herbivores and carnivores?

Examine hypotheses about how top-down and

bottom-up forces combine to determine species
abundances at different trophic levels

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The world is green

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 “World is Green” Hypothesis (or HSS)

Carnivores competition

Herbivores predation

Producers competition
Hairston, Smith, Slobodkin (1960) Community structure, population control, and 10
competition. American Naturalist 94.: 421-425.
• Fretwell (1977) extended HSS (The World Is Green)
– Alternating importance of regulation by competition or predation
depending on the number of trophic levels

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• Exploitation Ecosystems Hypothesis Oksanen & Oksanen 2000 ext HSS
– Predicts that depending on productivity, the biomass at each
trophic level will shift to achieve balance or equilibrium

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See figure 11.2
• Exploitation Ecosystems Hypothesis Oksanen & Oksanen 2000 ext HSS
– Predicts that depending on productivity, the biomass at each
trophic level will shift to achieve balance or equilibrium

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See figure 11.2
• Exploitation Ecosystems Hypothesis (Oksanen & Oksanen 2000 ext HSS)
– Predicts that depending on productivity, the biomass at each
trophic level will shift to achieve balance or equilibrium

Figure from original publication

Much research has been conducted exploring

the hypothesis

Allowed research to focus on components of a

system, how systems compare

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Equilibrium biomass Oksanen et al. simple model

TL=trophic levels

Potential primary productivity

Energy limitation hypothesis?

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 Serratia marcescens, gram negative
bacterium, basal level
Cultured alone (filled) or with
protest (open)

Colpidium striatum, ciliated protest

Filled squares – protest response to
productivity gradient

Experimental demonstration of this stair-step response, where one and two trophic
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level food chains respond differently to an increase in resources.
 In these examples, all
trophic levels increase
with potential
productivity

Why don’t these

examples support the
predictions?

A – North American lakes (Ginzburg & Akcakaya 1992)

B – North American grasslands (Chase et al. 2000)

From Mittlebach, page 232

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 Shift from palatable to non-palatable species
may affect pattern of response

The figure above illustrates the relationship between algal biomass in lakes and
phosphorous availability. In the figures to the right, the same relationship is plotted, but the
top panel includes algal species that are small enough to be consumed by herbivores, and
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the bottom panel includes large, grazer resistant algae. Taken from Mittelbach, p 233
Mittlebach, page 234 (figure 11.9)
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 Kangaroo vs Dingo - National Geographic

Dingo: Wild Dog at War

BBC Wildlife Video of dingo and walleroo at waterhole
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How does variation in primary productivity
influence the effects of dingo culling on kangaroo
and pasture biomass?

Kinchega National Park, New South Wales

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Choquenot & Forsyth (2013) Oikos 122: 1292-1306.
In arid areas, kangaroos have a negative impact on pasture biomass
and ecosystem function.
What controls kangaroo abundance?
Could kangaroo numbers be controlled by dingoes?

How important are top

down controls this
ecosystem?

Kinchega National Park, New South Wales

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Choquenot & Forsyth (2013) Oikos 122: 1292-1306.
Simulations
- stochastic rainfall events
- Culls of 25-100% of densities

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Internal social regulation of dingo densities
uncouples system from expected trophic interactions
(predation no longer regulates kangaroo abundance
with increased primary productivity)

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Example of a simulation built around EEH that provides insights into role of dingo culling on
trophic cascades, starting point for hypotheses that can be tested in the field
Strength of cull (% original dingo density) was
important for dingo density

Culling increased kangaroo densities and

decreased pasture biomass (i.e., example of
trophic cascade)

A 10% cull had an effect on kangaroos that was

similar to higher % culls

Effects of culling were stronger at lower levels

of productivity (results not presented here)

Prey switching to reptiles allowed dingo

densities to reach max at lower productivity
levels (results not presented here)
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 Main messages
Top-down or bottom-up regulation is false dichotomy
– Relative strengths of predator limitation and resource
limitation vary with trophic level and ecosystem productivity

Most systems show an increase in abundance of all trophic levels

with increased productivity,
– Heterogeneity in species composition within a trophic level
– Trade-offs between competitive ability and vulnerability to
predation
– Species replacements along productivity gradients

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 Main messages
Relatively simple models are useful despite being unrealistic

Drive research on components of systems, comparisons

between systems

Generates insights into process, mechanism, implications

Applications for conservation, agriculture, fisheries, building

resilience for global change . . . .

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 Summary of Models
• HSS Why the world is green Hairton, Smith and Stobodkin 1960

– World is green despite herbivores eating plants because

carnivores control herbivores

• Fretwell Extension of HSS (1977)

– Alternating importance of regulation by competition or
predation depending on the number of trophic levels

• Hypothesis of Exploitation Ecosystems (EEH)

– Predicts that biomass at each trophic level shifts
depending on productivity (Oksanen & Oksanen 2000) 29