Lecture 2: Functional Response Curves

and Predator Preferences M Pinard BI2020

 Predation
• Definitions, types, process and impacts (lecture 1)
– Effects of densities, probabilities of encounter, attack, consumption
– Attributes of predators and prey influence outcome
– Lethal and non-lethal impacts
• Functional responses
• Predator selectivity and prey responses
– Preferences, switching
– Optimal foraging theory
• Population dynamics (lecture 3)
– Tolerance or defence
– Population effects on prey, on predators
– Models of interactions and applications al . 2011
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Chap ers 5 an 5, Mori
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Chap ers 4 an
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 Functional response curves

• Relationship between an individual predator’s feeding rate

(=number of prey eaten), and the density of its prey

Feeding rate = a N
Prey eaten /
unit time Feeding rate = attack rate * prey density

Prey density

3
 Holling, 1959
Type 1 1. Consumption rate is a function of prey
Prey density alone, handling time is not
eaten / important
unit
time
Prey density
2. As prey density increases, handling
Type 2 time of the prey begins to limit
Prey consumption rate, so increases at a
eaten /
unit decreasing rate
time
Prey density
3. As prey density increases, predator
Type 3 searching efficiency increases; at a
Prey certain density handling time limits
eaten / consumption rate
unit
time Based on assumption that feeding rates are not dependent
Prey density on predator density 4
 Functional response curves
Type 3 – sigmoidal

Shrews (Sorex, Blarina) and deermouse

(Peromyscus) respond to sawfly cocoon
(Neodiprion) density

Bluebottle flies (Calliphora) respond to

sugar droplet density

Wasp (Aphelinus) respond to aphid density

5
 Type 1 Functional response curves
Prey
eaten / Predator feeding rate
unit time
Attack rate
Prey density
Handling time
Search time
Type 2
Prey
eaten /
How would an inducible defence change curve?
unit time
If multiple predators, how would curve change?
(expressed as prey eaten/time/predator)
Prey density
How would an increase in prey refuges change curve?
Type 3
Prey
eaten /
unit time

Prey density
6
 Summary Points
• Functional response curves capture how prey consumption rates depend on an
interaction between prey density and predator behaviour.

• Predators tend to be selective (both positively and negatively), with some showing
ranked preferences (e.g., carnivores) and others showing balanced preferences.
Variety of reasons why a predator may switch from one prey type to another

• Optimal foraging theory suggests predator should concentrate on most profitable

prey, and that they should switch when high value prey decrease in abundance.

• When handling time exceeds search time, expect specialist predation, when search
time exceeds handing time, expect generalist. When prey are rare and
environment is unproductive, expect generalist diet, when prey are abundant and
environment is productive, expect specialist.

• While little in nature works is an optimal way, theory based on optimisation

provides a framework for testing hypotheses. 7
 Predator preferences

“The food of every carnivorous animal lies therefore

between certain size limits, which depend partly on its own
size and partly on other factors. There is an optimum size
of food which is the one usually eaten and the limits
actually possible are not usually realized in practice.”
Charles Elton (1927), taken from p 103, Mittlebach
8
 Predator preferences
Proportion of prey in the diet compared w/ proportion
of the prey in the environment
Definition of predator preference

Positive or Negative Selectivity

Food preference
Ranked preference =most valuable of what is available
Balanced preferenceitems
= that provide a mixed and balanced diet

9
 Balanced preference (mixed diet)

Mixed diets are common

– To limit accumulation of toxins
– To avoid vitamin or mineral
deficiencies
– Take advantage of what you find

10
 Balanced preference (mixed diet)

Advantage of mixed diet for fitness

If food is low in carbohydrate or protein, advantage of the mixed diet is greater

Ricklefs & Relyea (2014) Ecology: The Economy of Nature. 7th Edition. W H Freeman and Company. Pp 107 11
Ranked preferences (most valuable)

Elner and Hughes 1978

Common for carnivores

Diet varies little in composition
Can classify food on a single scale
12
 Ranked preferences (most valuable)
15-Spined stickleback and prey

Optimum prey length (mm)

Predators (stickleback)
of different lengths

Optimum prey size based on Preferred prey length (mm)

prey mass/handling time
Kisaliogu & Gibson 1976 from Mittlebach pg 107 13
 Optimize time spent
Central place foraging Find more food if you travel further but . . .
………greater exposure to risk
……….…increased energy cost
…………………….increased time

Starlings feed on cranefly larvae in lawns

and pastures; hold larvae in base of bill so
becomes more difficult to grab the next
one so number of prey caught slows over
time

14
After Kacelnik A (1984) Central place foraging in starlings (Sturnus vulgaris). I. Patch residence time. J An Ecol 53: 283-299.
Ricklefs & Relyea (2014) Ecology: The Economy of Nature. 7th Edition. W H Freeman and Company. Pp 105
 Optimize time spent
Find more food if you travel further
Central place foraging but . . .
………greater exposure to risk
……….…increased energy cost
…………………….increased time

Starlings feed on cranefly larvae in lawns

and pastures; hold larvae in base of bill so
becomes more difficult to grab the next
one so number of prey caught slows over
time

Rate of food gathering diminishes over

time because amount of prey gathered
per unit time decreases

15
After Kacelnik A (1984) Central place foraging in starlings (Sturnus vulgaris). I. Patch residence time. J An Ecol 53: 283-299.
Ricklefs & Relyea (2014) Ecology: The Economy of Nature. 7th Edition. W H Freeman and Company. Pp 105
 Optimize time spent
To maximize food acquisition, need to balance travel time plus search time (time
spent finding the food once you arrive at destination)

Orange line = diminishing benefits line

16
After Kacelnik A (1984) Central place foraging in starlings (Stumus vulgaris). I. Patch residence time. J An Ecol 53: 283-299.
Ricklefs & Relyea (2014) Ecology: The Economy of Nature. 7th Edition. W H Freeman and Company. Pp 105
 Optimize time spent
How does the distance you live from the
grocery store affect how much you buy on a
given visit?

Tested a prediction that longer travel

times would case starlings to return to
nest with a larger number of mealworms

Experimental set-up
tables set distances, tubes to deliver mealworms,
released at progressively longer intervals

After Kacelnik A (1984) Central place foraging in starlings

(Stumus vulgaris). I. Patch residence time. J Animal Ecology
53: 283-299.

Ricklefs & Relyea (2014) Ecology: The Economy of Nature. 7th Edition. W H Freeman and Company. Pp 105 17
 Risk sensitive foraging
Foraging behaviour that is influenced by the
presence of predators

Tested the sensitivity of young creek chubs

(Semotilus atromaculatus) to food density
and predators

Trade-off between risk and energy gain

Behaviour consistent with rule to minimize

ratio of mortality/foraging rate (=number
of deaths per unit energy gained)

Gilliam & Fraser (1987) Ecology 68: 1856-1862. Taken from Ricklefs & Relyea pp. 106 18
 Predator preferences
Preferences – can be fixed
or can switch

Snails offered two species of mussel Guppies offered fruit fly larvae and tubificids
 Predator preferences
Fixed preferences
or switch Why switch?
Different prey occupy different
microhabitats, concentrate on most
profitable habitat

When prey is more common . . . .

develop search image for it
higher chance of encountering
higher chance of capturing it
increase efficiency in handling it
20
 Summary Points
• Functional response curves capture how prey consumption rates depend on an
interaction between prey density and predator behaviour.

• Predators tend to be selective (both positively and negatively), with some showing
ranked preferences (e.g., carnivores) and others showing balanced preferences.
Variety of reasons why a predator may switch from one prey type to another

• Optimal foraging theory suggests predator should concentrate on most profitable

prey, and that they should switch when high value prey decrease in abundance.

• When handling time exceeds search time, expect specialist predation, when search
time exceeds handing time, expect generalist. When prey are rare and environment
is unproductive, expect generalist diet, when prey are abundant and environment is
productive, expect specialist.

• While little in nature works is an optimal way, theory based on optimisation

provides a framework for testing hypotheses.
 Predator preferences
Optimal foraging theory
Optimal diet: 1. foragers should prefer most profitable prey

Energy gained during feeding period

fx Time searching + Time handling

22
 Predator preferences
Optimal foraging theory
Optimal diet: 2. switch prey

Should broaden diet to include low value prey when high

value prey decrease in abundance

23
 Predator preferences
Optimal foraging theory
Energy gained during feeding period fx Time searching + Time handling

Switching
An efficient forager should broaden its diet to include low value
prey when high value prey decrease in abundance

Generalist or specialist – which is better?

Specialists
– Monophagous (one prey type) [parasitoids, parasites]
– Oligophagous (few prey types) [parasites, true predators]
Generalists
– Polyphagous (many prey types) [grazers, true predators]

24
 Predator preferences
If interested in theory, read Begon et al. 20006 chapter 9

Theory
. . . . A predator will encounter a variety of food when searching
. . . . Specialists will keep searching until find preferred (most profitable)
. . . . Generalist will pursue a large proportion of what they encounter,
regardless of profitability of the prey.

Predictions
• When handling times are short, relative to search time, makes
sense to be a generalist [Insectivorous birds]
• When handling times are long, relative to search time, makes
sense to be a specialist [Lions]

• When environment is unproductive and prey is rare, predict a

more generalist diet, but when productive, predict specialisation
25
 Predator preferences
Optimal foraging theory
• Supported many experimental hypothesis tests
– relative importance of profitability (net energy gained / handling
time) versus encounter rate
– Interactions between profitability of different prey types relative
to prey abundance

• Review 2001 (35 studies, Sih & Christensen)

– 87% studies supported the predictions
– But all predators consumed some prey that were not predicted as
optimal
– Assumptions of model
• Perfect predator knowledge of prey quality and prey density
26
 Predator preferences

Describe the pattern here.

Is there evidence in support of the optimal foraging theory?

27
 So what?

• Optimal foraging theory has increased our

understanding of why some prey are selected
and some are ignored

• Exceptions to the rule are interesting . . .

. . . . . . .How does imperfect knowledge affect prey selection?

28
 Summary Points
• Functional response curves capture how prey consumption rates depend on an
interaction between prey density and predator behaviour.

• Predators tend to be selective (both positively and negatively), with some showing
ranked preferences (e.g., carnivores) and others showing balanced preferences.
Variety of reasons why a predator may switch from one prey type to another

• Optimal foraging theory suggests predator should concentrate on most profitable

prey, and that they should switch when high value prey decrease in abundance.

• When handling time exceeds search time, expect specialist predation, when search
time exceeds handing time, expect generalist. When prey are rare and
environment is unproductive, expect generalist diet, when prey are abundant and
environment is productive, expect specialist.

• While little in nature works is an optimal way, theory based on optimisation

provides a framework for testing hypotheses.
 Predation (3 lectures & 1 practical)
• Definitions, types, process and impacts (Lecture 1)
– Effects of densities, probabilities of encounter, attack, consumption
– Attributes of predators and prey influence outcome
– Lethal and non-lethal impacts
• Functional responses
• Predator selectivity and prey responses (Lecture 2)
– Preferences, switching, optimization al. 2011
et
– Optimal foraging theory , B e gon
a nd 10 r 2 012
9 o h
• Population dynamics (Lecture 3) Chap t ers
itt lebac
n d 6, M
– Tolerance or defence e rs5 a or 2 0 11
ap t i n
– Population effects on prey, on predators Ch 5, Mor
4 and
– Models of interactions and applications h ap ters
C

30
 References
• Begon et al. (2006) Chapter 10. The population dynamics of predation. In, Ecology.
From Individuals to Ecosystems. Pp. 297-
• Connell (1975) Some mechanisms producing structure in natural communities: a
model and evidence from field experiments. In: Cody & Diamond (Eds). Ecology
and Evolution of Communities, pp. 460-490. Harvard Univ Press, Cambridge, MA
• Hudson et al. (1998) Prevention of population cycles by parasite removal. Science
282: 2256-2258.
• Menge et al. (1986) A test of the Menge-Sutherland model of community
organization in a tropical rocky intertidal food web. Oecologia 71: 75-89.
• Scheffer, M. 1999. Searching explanations of nature in the mirror world of math.
Conservation Ecology 3(2): 11. [online] URL:
http://www.consecol.org/vol3/iss2/art11/
• Sih et al. (1985) Predation, competition and prey communities. A review of field
experiments. Ann Rev Ecol Syst 16: 269-311.
• Stenseth et al. (1997) Regulation in snowshoe hare and Canadian lynx: asymmetric
food web configurations between hare and lynx. PNAS 94 (10): 5147-5152.