BIOL 4620 - Animal Behaviour

Lecture 9
Foraging Behaviour and Optimal
Foraging

1
 Foraging
●Foraging - series of actions animals take to search for,
pursue, and handle food resources
●This behaviour is complex
●Involves making decisions based on:
○Size of food item
○Nutritional value (energetics)
○Ease of transport
○Distance to safe area to consume food

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 Obtaining Food
●How many different food gathering techniques/strategies
can you think of?

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 Carnivory
●Carnivory - consuming other animals as a primary food
source
Approaches:
●Pursuit predation - predator actively give chase to prey; costs
energy
●Ambush predation - various strategies can be used for surprise
attacks
○Stealth - camouflage and/or remain motionless
■Example: Smokescreening behaviour in Portia spiders
○Aggressive mimicry - resemble something harmless or
appealing to lure prey
■Example: Alligator snapping turtle
○Traps - construct or use behavioural traps to ensnare prey
■Example:
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 Sensory Specializations
●Pit vipers (Family Crotalidae) and
Family Boidae use prey body heat to help
them hunt
○Pit organ with receptors sensitive to
infrared radiation Pit organ location in pit vipers

●Star-nosed moles (Condylura cristata)

○Star-shaped nose with 22 fleshy
appendages
○Tactile receptors - sensitive to touch
Star-nosed mole

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 Optimality Theory
●Can use optimality theory to understand foraging behaviour
(as well as antipredator behaviour)
○Adaptations have greater benefits to costs ratio

• Note: sometimes difficult

to measure costs and
benefits in the same
fitness units
• If you could, you would be
able to predict the
characteristic with
greatest benefit
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 Optimal Foraging Theory
Using foraging for testing optimality theory:
●Foraging behaviour → series of decisions that can be
analyzed one by one
●Easy to identify common measure to compare decisions
○i.e., rate of energy intake
●Can identify constraints on foraging behaviour (gut
capacity, presence of predators)

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 Optimal Foraging - Diet Selection
●Decisions while foraging → what to eat, eat one type of food
or focus on a few types of foods, taste, ease of finding food,
etc.
●Start with a simple model - oversimplification of nature
but good starting point

●Scenario:
○Two kinds of food available to forager
○Forager only finds one food at a time
■Should it eat the food, or these are
strategies
■Should it ignore it and keep searching?
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 Optimal Foraging - Diet Selection
(II)
●Compare using common “currency” - i.e., energy gain
●Next identify constraints:
○Amount of time it takes to search for food 🡪 search time
○Amount of time it takes to process food 🡪 handling time

●If these can be described using variables then can get a

sense of an optimal strategy.

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 Northwestern Crows Example
●Northwestern crows (Corvus caurinus) forage for whelks
(molluscs)
○Foraging decisions: whelk choice; how high to fly

Northwestern crow Whelk

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 Northwestern Crows - Observations
●Reto Zach observed:
○Crows selected and dropped only large whelks (3.5 to 4.4
cm long)
○Crows flew up approximately 5 m to drop the whelks
○Several trials repeated by crows until the whelk broke
●Are these decisions optimal in terms of maximizing whelk
flesh available to eat per unit time spent foraging?

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 Northwestern Crows - Questions
●Reto Zach wondered:
○Were crows selective about which whelks they dropped?
○Were there differences in handling times of different
sized whelks?
○Do crows change behaviour if a whelk is hard to crack
open (i.e., adjust cost according to expected caloric gain
from meal)?
○How many times will it drop hard to crack whelks?

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 Northwestern Crows - Test
● Is the behaviour efficient in terms of generating broken
whelks?

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 Northwestern Crows - Results
●Reto Zach found:
○Crows selected mainly the largest and heaviest whelks
on beach → provide more energy so crows can select
most profitable prey
○Large whelks required fewer drops from 5 m distance
than medium or smaller size ones
○Chance that whelk would break was not affected by
number of previous unsuccessful drops
●Conclusions:
○Adaptive behaviour
○Gain more energy from foraging activities than they
expend in attempting to get food
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 Optimal Foraging Assumptions
●Key assumption in optimality models → does the behaviour
translate into maximum reproductive success?
○Has not been established for crows but evidence from red
crossbills
○Male red crossbills were placed on both
sides of a divided cage that contained either:
■A branch with pinecones with edible
seeds removed; or
■A branch with unaltered pinecones
○Female crossbills that observed associated
preferentially with males they had seen
securing many seeds

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 Refining Optimal Foraging Models
●It is important to consider other factors – can result in
different predictions
●Example: Eurasian oystercatchers and mussel selection
○Meire and Ervynck calculated profitability of different
sized prey based on calories vs. energy expended
opening
■Cost and benefit

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 Eurasian Oystercatchers Example
●Largest mussels need more
time to open, but they provide
more calories
●Model prediction:
○Oystercatchers should focus
on largest mussels
■BUT in reality they
don’t
■Preference for 35 – 40 mm
size
●Why?

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 Eurasian Oystercatchers -
Hypothesis 1 mussels
●Hypothesis 1 – largest
have lowest profitability because
some can’t be opened at all
●Initial analysis - only considered
prey that the birds already opened
(Model A)
●Some large prey selected are not
possible to open
●Wasted handling time
●Birds should focus on 50 mm size
mussels (Model B)
○But they don’t…

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 Eurasian Oystercatchers -
Hypothesis 2 large mussels not worth trying to open
●Hypothesis 2 – Many
because are covered in barnacles and impossible to open
●Observation: birds never touch the largest barnacle-
encrusted mussels; these should be eliminated from
consideration
●Factors: prey-opening time; time wasted trying but failing and
actual size range of realistic prey, best option is to focus on 30
– 45 mm mussels
●This is in fact what the birds do

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 Criticisms of Optimal Foraging
Theory
●Reality is that animals don’t always hunt for or select food as
efficiently as possible or eat the food that offers highest caloric
gains
●Not about perfection in evolution; use is for testing whether
correct variables have been identified which have shaped
evolution of the behaviour
●Important to consider many variables
●Other factors to consider:
○Predators may have shaped the evolution of an animal’s
foraging behaviour – predation risk should be considered
○Food availability/seasonality

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 Finding Food and Avoiding
Predators
●Dugongs - large marine mammals that feed on sea grasses
and are preyed upon by tiger sharks
●Variability in dugong foraging behaviour
○Cropping vs. excavation
●Excavation provides more energy but puts
them at greater risk (heads are buried)
●When tiger sharks are common, dugongs avoid
Excavation method and opt for cropping

Optimal foraging theory must consider not only energy gain but
also predation risk to accurately predict animal behaviour

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 Finding Food and Avoiding
Predators (II) have changed their foraging behaviour
●Elk living in Yellowstone
following the reintroduction of wolves into the area
○Spend more time hidden in woodlands rather than feeding
in exposed meadows → reduced production and survival of
calves

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 Game Theory and Foraging
Behaviour
●Just as game theory can be used to assess anti-predator
behaviour, so too can it be used to find the adaptive basis
of foraging behaviours
●But sometimes two different foraging
strategies can coexist in the
population via frequency-
dependent selection
○Example: African cichlid fish
■Two forms - one with the jaw
twisted to the right and the
other with the jaw twisted to the
left
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 Behavioural Variation and
Conditional Strategies
●Multiple foraging strategies not always due
to frequency-dependent selection
○Example: Ruddy turnstone sandpipers
●Uses many different foraging strategies
(dominants vs. subordinates)
●Variation not due to genetic differences but
due to conditional strategy
●Strategy in this case based on dominance
in flock

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 Summary
●Cost-benefit approaches are useful for assessing foraging
and adaptive value of the behaviour
●Game theory approach contends that the better strategy
takes into account competing strategies of other
individuals
●Optimality theory is based on the premise that optimal
traits have a better benefit to cost ratio than alternative
traits
●Alternative strategies also exist:
○Frequency-dependent selection
○Conditional strategy

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