Game Theory

Game Theory is “a branch of

mathematics devoted to the study
of strategy in which players seek to
maximize their individual returns”
(Molles and Cahill).

Forms the basis of many economic

theories, war strategy, sociological
theories (and games)

John Maynard Smith emphasized

the potential for its use in
behavioral ecology
 Game Theory
What behavioral strategy should individuals adopt to
maximize their individual fitness?

Does the “best” strategy depend on the strategy of

other individuals?

Should individuals behave aggressively and fight for

resources or behave non-aggressively?
under what conditions should individuals
engage aggressively in conflict for limited
resources?

Can aggressive and non-aggressive

strategies coexist?
 Hawk vs. Dove Game

2 behavioral approaches for

interactions over a contested resource:

Hawks Doves
 The Rules: Hawks
Always attack other individuals, taking
the resource (R) if they win.
If a hawk encounters another hawk, it
will win only half the time.
When it loses, it will suffer an injury cost
(C).

AGRESSIVE
 The Rules: Doves
When a dove encounters an opponent, it may
put on an aggressive display, but it does NOT
fight.
When a dove encounters a dove, it will win
the resource one half the time.
When a dove encounters a hawk, the hawk
will always win the resource.
But since they don’t fight, they don’t incur any
injury costs (C).

NON-AGGRESSIVE
 Game The y
Hawk: player will continue to escalate until it
either is injured or the opponent gives up.

Like poker, the person who will always raise the

bet

Dove: player will attempt to escalate but will

retreat and give up if opponent escalates

Poker: the person who is “bluf ng”!

or
fi
 Conditions
V = value of the contested resource
(the “pot”)
C = cost of fighting (what you have to
lose)

4 possible combinations
Hawk-hawk
Hawk-dove
Dove-hawk
Dove-dove
attack other individuals, taking the
resource (R).
win only half the time (vs. hawk).
injury cost (C) when you lose.

AGRESSIVE

aggressive display, but does NOT fight.

dove vs dove, win half the time.
dove vs hawk, you lose.
No fight = no injury costs (C).

NON-AGGRESSIVE
Combinations and
outcomes Hawk Dove
hawk-dove
V

Hawk
0

Dove
V
Combinations and
outcomes Hawk Dove
dove-dove

Hawk

each “wins” half

the time V/2

Dove
No cost V/2
because other
leaves
Combinations and
outcomes Hawk Dove
hawk-hawk
(V – C)/2

only loser pays Hawk

the cost (V – C)/2

50% chance of
winning

50% chance of Dove

paying cost
(injury)
evolutionary stable strategy
What is evolutionary stable strategy?

Depends on relative value of V and C

If V > C, then being a hawk is the ESS

Hawk meets dove, hawk wins

Hawk meets hawk, win ½ time
dove wins anything only when it meets another dove
Reason: Price paid (C) is low compared to V and
you only pay it if you loose
 evolutionary stable strategy?
IF V < C, becomes more complex

Now not necessarily beneficial to be a hawk in a hawk

world. Price of fighting not worth the cost, which you
will pay ½ the time

Now some combination of hawks and doves

becomes the ESS.
 Real animals?
If value of resource is high, e.g. mating opportunities,
we can see how aggression would be selected for.
Fighting over the resource is the only way to win

If resource value is low, not worth fighting for, better

to be a dove
What behavioral strategy should individuals adopt to
maximize their individual fitness?
Depending on V and C

Does the “best” strategy depend on the strategy of

other individuals?
Apparently YES

Should individuals behave aggressively and fight for

resources or behave non-aggressively?
Depending on V and C
 under what conditions should individuals
engage aggressively in conflict for limited
resources?
If V > C

Can aggressive and non-aggressive strategies

coexist?
YES
the idea conveyed by the notion of ESS is that
sometimes what is better for an individual
depends on what the other individuals of the
group do.

An ESS is not stable because it is the best

possible strategy but simply because, once it
has been adopted by the majority of the
individuals in the population no other
alternative strategies can develop
successfully
 Ghirlanda and Vallortigara (2004) examined whether behavioral
lateralization can arise as an ESS in the context of prey-predator
interactions

They considered predators and group-living prey meeting in contests

in which the prey have two lateralization strategies available, “left” and
“right”

First prey lateralized in the same direction have a greater chance of

keeping together as a group. Second, predators are better at
capturing the prey type that they meet more often
 For instance, predators may learn to anticipate prey escape
movements, or to approach prey from a given direction. If p(x) is
the survival probability of a prey when a proportion x of its
groupmates employ the same lateralization strategy, a general
way of writing p(x) is:

p(x)= p0 + g(x) - l(x)

where p0 is a baseline escape probability, g(x) represents the

bene t gained, under attack, by keeping together with a
proportion x of fellow prey, and l(x) represents the cost of having
the same directional bias as a proportion x of other
fi
 When g becomes larger, stable populations consist of
left- and right- type prey in unequal numbers

The intuitive content of such a situation is that the majority

of prey gain protection by keeping together but pay a
cost because predators are better at handling them

A minority of prey manages to enjoy the same probability

of escaping by trading off protection from the group
with an advantage in the face of predators because….
 “frequency dependent selection”

This refers to an advantage of the minority group

which depends on the frequency of these individuals
(i.e., an advantage that disappears when these
individuals increase in number)
 Fish Behavior
• Shoals and Other Aggregations
• Forms of fish grouping
– Solitary
– Shoal
– School
– Pod
• Reasons for grouping
– Traveling
– Feeding
– Dealing with predators
– Reproduction
Schooling
not all fish aggregations are schools

aggregation: group created by the attractiveness of

some feature of the environment.
shoal: group of fish remaining together for social reasons
school: polarized, synchronized shoal of fish that swim
together in the same direction, at a similar speed, with a
similar distance between one another

school shoal
“dilution” effect “confusion” effect
 9.1 n Living in groups x 407

Table 9.1 Some suggested costs and benefits of social aggregation. See text.

Benefits Costs

Aggregative response to prey density Greater interference from competitors

means that groups indicate rich sources (Forrester 1991)
of food (Krebs 1974)
Individual risk of predation diluted by Greater risk of contracting disease
joining a group (Treherne & Foster 1981) (Hoogland 1979)
Groups can tackle larger prey than Greater chance of being cuckolded, or
single individuals (Kruuk 1972) mistakenly feeding someone else’s
offspring (Bray et al. 1975; McCracken
Grouping confuses predators, making
1984)
it harder for them to target prey
(Milinski 1977) Investment in foraging, courtship, or other
activities exploited by other group
Huddling in groups helps
members (Barnard & Sibly 1981)
thermoregulation (Trune &
Slobodchikoff 1976) Young may be cannibalised by neighbours
(Sherman 1981)
Energetic advantages to swimming or
flying in a group through ‘slipstreaming’ Greater risk of inbreeding
(Weihs 1973, but see Partridge & (Faulkes & Bennett 2001)
Pitcher 1979)

microorganisms can show a degree of sociality (Oleskin 1994; Crespi 2001). The costs and
benefits of aggregating are thus likely to be diverse (Krause & Ruxton 2002). Table 9.1
 Types of social acti
Recipient’s fitness
increases decreases
Actor’s fitness

increases Cooperation Selfishness

decreases Altruism Spite

o
 Altruism
Natural selection favors behavior that maximizes an
individual’s survival and reproduction

These behaviors are often selfish

some animals behave in ways that reduce their individual

fitness but increase the fitness of others

This kind of behavior is called altruism, or selflessness

Altruism
33% of young bats fail to feed each night
Only 7% of adults fail to feed.
Chronic threat of starvation among vampire bats
since they can only survive 3 days without a meal
Successful bats will regurgitate part of their meal
for group members that were not successful – but
not randomly
Bats only give to those from whom they have
received blood in the past

Wilkinson, G.W. 1984. Reciprocal food sharing in the vampire bat. Nature. 308:181-184)
 Al uism

If co-operation and altruism have evolved, then they must have

some adaptive benefits, researchers have analysed the conditions
under which adaptations for engaging in such behaviour can be
expected to evolve.
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 Al uism

Ground squirrels will warn others of the presence of a

predator, even though making such a call may draw the
attention of the predator to itself (Sherman, 1977).

EUSOCIALITY: In many species of social insects, workers

avoid reproduction entirely (they are sterile) in order to help
raise their sisters (Wilson, 1971).
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in naked mole rat populations, nonreproductive
individuals may sacrifice their lives protecting
their reproductive queen and kings from
predators
the prisoner’s dilemma
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