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Define: Optimality models predict which decision an animal should make in order to maximize its inclusive fitness under a given set
of conditions hypothesized to drive the behaviour.


Parts of optimality Model 1. A set of decisions or Foraging bird may choose to eat a particular
strategies – an animal piece of food, or search for another one
performs an action out of a instead.
variety that are available (it
may not be conciously aware)

2. Currency – the rate of energy Many foraging models use the rate of energy
intake, time spent foraging intake as currency.

3. Constraints or limits on the Animal can optimize its behaviour only

animal – can be internal or within the range of its capability and needs,
intrinsic (nutritional needs, and constrains define this range.
ability to see), or external
(other organisms, physico-
chemical factors that allow for
effective foraging

•A lion sees a wilder beast – should it try to capture and feed on it or not?

•A bird sees a worm. Should eat it or keep on searching?


The Prey Model Most species of animals are Forager is searching for food and finds one prey
surrounded by all manner of things at a time. Considering 3 parts of the model:
that they might consider eating.
•Decision – eat prey or continue finding
•Currency – rate of energy intake/calorific
value/maximizing the rate
•Constraints – prey needs to be processed: nuts
need to be opened; live prey needs to be subdued
(handling time).
•Different prey requires different handling time.
Foragers cannot handle prey and search at the
same time

The predictions of the model Mathematical Expression of the Prey Model

1.The decision to eat prey 2 should •Assume 2 different types of prey = type 1 and
be based on search time of prey 1. type 2
if prey 1 is abundant then S1 will
be low so the inequality shifts to the •Each prey has its own energy (Ei) = E1 & E2
right [E2/h2 > E1/(S1 + h1)]
•Each has its own handling time (hi) = h1 & h2
2.The search time for prey 2 should
not be important. Prey type 2 could •Because we are interested in maximizing the
be everywhere around and predator rate of energy intake, we need to figure out what
will still ignore them as long as the is the rate would be for each prey item.
inequality is to the right side
3.The forager should instantly •Profitability – ratio of energy gained to
switch back to eating both kinds of handling time, (Pi = Ei/hi)
prey to eating only the higher
quality prey – zero-one rule (eat •Assume prey 1 more profitable than prey 2,
less profitable prey either none of E1/h1>E2/h2
the time (probability 0) or all the
time (probability 1) •Each prey can have its own search time, Si (the
amount of time taken to locate prey item).
The model can be expanded to
include more than 2 prey items •As density of prey item increases search time

An animal has found a prey: should it eat or

continue searching for a new one?
•Scenario 1 - If prey 1 is more profitable the
animal will eat it – first prediction of model –
eat the most profitable prey!

•Scenario 2 - If animal finds prey type 2 –

should it eat the less profitable prey (type 2) or
continue searching until the better prey is found
(type 1)?

•One has to compare the rate of energy intake of

the 2 choices

Gain from eating prey 2, once it is found is E2/h2

Gain from searching for and eating prey type 1:
E1/(S1 + h1)

•the denominator for the second equation

includes both search and handling time because
animal must find prey type 1 and then consume

•Both are rates of energy intake (calories/sec)

•One has to compare the decisions available to
the animal in the currency

•The forager should eat prey 2 when,

E2/h2 > E1/(S1 + h1)

How Long to Stay in a •To eat or not to eat is not the only
Patch problem facing foraging animals

•Food resources are patchy, so •If you picked berries, or when you have faced
foragers must decide how long to the same problem; should you search a bush so
stay in a patch with food resource, well so you get every last berry, or should you
leave and find a new patch move on to another bush?

Factors that affect the decision


•Richness of the patch – if •If berries are short supply, you might do better
resource is in short supply search to stay and search for every last one. If other
for every last bit. If other patches bushes have many berries, you will probably do
have more resource more to new better to move on.

•Distance between patches – near •If you can reach another bush by taking a few
patches as compared to far away steps, you will probably make a different
patches decision about leaving your patch than if you
have to walk half a mile.

Marginal Value Theorem

•The forager would maximise the

rate of energy intake – the currency

•The decision – stay or move to

new patch
•Constraint – searching for patch
and foraging within them, patches
are found one at a time
Central-Place Foraging • Animals that carry food back •Nesting birds and insects that live in hives
to a central location for storage are central-place foragers. The problem here
or for feeding to offspring. is not only when to leave the patch, but also
which and how much food to collect before
returning to home base.

• Time in the patch, load size and •As travel distance increases, pine squirrels
selectivity are predicted to increase gather cones with a larger number of seed
as travel distance increases. per cone (Elliot, 1988)

• Combination of central-place •Marine diving bird, the rhinoceros auklet

foraging and diet selection adult collected different prey items when
feeding young than when feeding
themselves. Most meals brought back to the
nest for chicks comprised one or two large
fish, which are more efficient to deliver,
whereas adults generally selected smaller
prey when foraging for themselves.
Risk Sensitive Foraging •The mean is not only characteristic • A small bird, a junco is given a choice
that may vary across patches between two trays of food. When it selects
one tray, the experimenter removes the
• If an animal can distinguish other. One tray always has four seeds in it.
between these two patches, we say The other tray varies; on half the trials, it has
they are risk sensitive. one seed, and the other half of the trials, it
has seven seeds. Thus, these two patches
have the same mean amount of seeds, but
the second patch has higher variance around
that mean. We say the second patch has
higher risk.
•One way to approach the Straight line utility function: for each seed
problem of why any animal ever that a bird finds, its fitness increases by the
be risk-sensitive is to consider same amount. This is unlikely to be
utility or the vale that resource common.
has to an animal
Concave down utility function: this might
occur if an animal can consume only a
certain number of seeds, and seeds found
after that are useless. This is the more
common function.

Concave up utility function: the resource

becomes more valuable to the animal as it
gains more of it. Here, a gamble may pay off
in a large win, or relatively small loss. This
animal should be risk-prone, and take the
Effects of Competitors on • The presence of competitors Competition by members of the same or
Foraging behaviour and predators may also be different species may force them to forage in
important in determining suboptimal habitat or to include food items they
foraging behaviour. would not otherwise consume.

Stickleback fish that had parasite infestations

feed on smaller-sized daphnia rather than
compete with infested stickleback for larger,
more profitable prey.

American crows change their behaviour in the

presence of conspecific competitors. Crows fly
up and drop walnuts onto the ground in order to
break them open. When the other crows are
present, they do not drop them as a great a
height, presumably so they have a better chance
of swooping down to pick up the nut before
competitors snatch it.
Effects on Predators on •Many foraging animals are
Foraging behaviour potential prey , and must balance
risk of predation with foraging
•Animal is less vigilant when
foraging and can blow cover
when attacking prey especially
those that rely on crypsis
(camouflage – blending with
background) – more vulnerable
to attack!
Effect of predator on diet
•When it is less dangerous to
capture more profitable prey
selectivity decreases
•When more dangerous to
capture less profitable prey
selectivity increases

•Risk of predation may also Desert baboons spent more times foraging in a
affect the choice of patch lower juvenile hoary mamots foraged close to
their burrows rather than risk predation by
eagles or coyotes in more distant but better-
quality patches.
State Sensitive Models of All members of a population may A very hungry animal may choose to go to a
Foraging not make the same foraging risky patch of food if the quantity of food is
decision. In fact the same higher, whereas a well-fed animal may give up
individual may make different food in favour of avoiding predators.
foraging decisions at different
times. Hungry scorpions are willing to forage under
bright moonlight when they are in danger from
predators, but well-fed scorpions hide when the
moon is bright


Modifying food supply Some animals modify their food Grazing animals in the grasslands stimulate the
supply so that is increases. growth of some species of grass and prevent
succession grassland communities to other
community types, such as forest.

The ultimate non-human Leaf sections are brought into the nest
manipulation of producers in an •Leaves are cut into smaller pieces with
ecosystem may be the “agriculture mandibles and becomes pulpy.
practiced by certain fungus- •Anal droplet is deposited on leaf before
growing ants. dropped into garden – these are processed leaves
•Then add fungal mycelium on the processed
leaves and fungus grows (ig 15.10)
•Ants feed on the tip of the fungus. This
arrangement is mutualistic association – fungus
breaks down cellulose to sugar while ants spread
and care for the fungus by weeding out alien
fungus, fertilising it, and producing antibiotics
that act against competing fungi and