Behavioural Ecology

Isabel Smallegange

Contest behaviour

Contest behaviour: competition

When many individuals exploit the same limited resources, they are
competitors.

exploitation competition
resource density decreases as
competitors exploit the resource
success rate decreases as a
result
no social interactions

interference
competition

individuals compete directly

outcome of social interactions
determines access to
resources
success rate decreases as a
result

Contest behaviour: game theory

When many individuals exploit the same limited resources, they are
competitors.
When such conflicts are resolved through a direct interaction, they are
contests
termed
Contests
represent the means by which individuals exclude others from
resources
Decisions made by one competitor may be influenced by what others
are doing
Contests are usually pairwise
Key elements in a contest are the resource, its value (V) and the costs
(C) associated with fighting over that resource
Contests are resolved when one of the opponents, the loser, is forced to
e theory:
How or
aggressive
should
an animal be when fighting over a disputed reso
withdraw
decides
to withdraw
Optimality models are inadequate to explain fighting decisions as the
actions of an individuals opponent will have a direct effect on the
utility of an agonistic behaviour that the individual might employ.

Game theory

Analyzes situations where one individuals

success
depends on the choices made by
others (Maynard
Smith 1974)
Characteristics:
A trait affects the survival of individuals
differently to that of whole populations
A trait affects the survival or reproduction
of the relatives of the individual
possessing it
This means that fitness of a genotype
depends upon
what other genotypes are
present in population and
in which
frequencies
Animal strategies are defined
mathematically and set
against one
another (often in simulations)
Differs from optimal foraging theory in
that currency
is usually fitness, not e.g.
energy intake
Doesnt usually explicitly consider

Game theory: evolutionarily stable strategy (ESS)

Strategy A is an ESS if (Maynard Smith & Price 1973):
When A is rare, it can invade a population in which
another
strategy is common
When A is common, it cannot be invaded by any other
strategy present in the game
Biological analogy: if A arises as a rare mutant, it will
increase in
frequency by selection. If A is the wild type in
the population, rare mutants will not increase in
frequency.
Mathematically: if EB(A) is the payoff to strategy A when
played against
strategy B, A is an ESS if:
For all B, EA(A) > EA(B)
If for any strategy B, EA(A) = EA(B), A is still an ESS if
EB(A)> EB(B)

Game theory: evolutionarily stable strategy (ESS)

In some games, no single, pure strategy is an ESS
Some games result in evolutionary equilibrium where
each strategy
maintained at some stable frequency
Example: t haplotype in mice predicted to stabilize at
70%
Example: negative frequency-dependent selection in
Mixture of pure strategies: any given individual plays only one
polymorphisms
strategy, percentages represent frequencies of those
individuals in the population
Biologically, represents highly canalized phenotypes
Mixed strategy: any given individual can play more than one
strategy
Biologically, represents plastic phenotypes
Percentages represent stable percentage of time an
individual should play each strategy
Mixed strategy can be an ESS even when no pure strategy is
When a mixed ESS exists, payoffs to the different strategies

Game theory: contest behaviour

Game theory applied to a wide range of decisions where there is
conflict of interest, but contests were the first example of game
theory applied to biology
Three main conceptual models for animal contests:
Hawk-Dove game (and bourgeois strategy and rock-scissor-paper
game)
War of Attrition model
game & War
of
Hawk-Dove
Sequential assessment
game
attrition
no information exchange
strategy depends on own characteristics or opponents characteristics

Sequential assessment game

opponents gain additional information about strength of the
opponent,
with each consecutive exchange of actions

Contest behaviour: hawk-dove game

Table showing the outcomes of conflicts
between strategies
Strategies present in population
placed
along sides of matrix
Each box shows outcome of an
encounter between individuals playing
each strategy
Units usually fitness payoff to
each
individual
Sometimes, only the payoffs to the
row
player (here, Individual 1)
are shown

Contest behaviour: hawk-dove game

Maynard Smith & Price
(1973)
Parameters:
V = value of the resource
C = cost of fighting
Strategies
Hawk: Always fights,
attempts to
kill/injure
opponent
Dove: Displays and then
retreats
p = frequency of Hawks,
1-p = frequency of Doves

Strategy A = Hawk
Strategy B = Dove

Contest behaviour: hawk-dove game

Whats the ESS?
If V > C:
If opponent plays Hawk, (V-C)/2
> 0,
so best to play Hawk
If opponent plays Dove, V > V/2,
so best to play Hawk
Hawk is the ESS
But if V < C:
If opponent plays Hawk, (V-C)/2
< 0,
so best to play Dove
If opponent plays Dove, V > V/2,
so best to play Hawk
Rare Hawks can invade common
Doves,
and vice versa

Contest behaviour: hawk-dove game

Is there an evolutionary equilibrium (a.k.a. a
mixed ESS)?
Find expressions for the fitnesses of Hawks (WH)
and Doves (WD), set them equal, and solve for
p:
WH = p[(V-C)/2] + (1p)V
WD = p(0) + (1p)(V/2)
Solve for p, get p = V/C
Equilibrium frequency of Hawks = V/C,
equilibrium frequency of Doves = 1 (V/C)

Contest behaviour: hawk-dove game

What does this mean
biologically?

In order for lethal fighting (extremely high value of C)

to be stable,
the value of the resource (V) must also
be very high
The greater the cost of fighting relative to the value
of the
resource, the more it pays a given
individual to display rather than
attempt to injure or
kill its opponent
A population with equilibrium frequencies of Hawks
and Doves can be stable
A population of individuals who play Hawk some
percentage of the time and Dove some percentage of
the time can also be stable

Contest behaviour: sex-ratio (example of mixed strategy)

In most species, one male can fertilize many females
eggs
Population growth rate would be higher if ratio of
males:females were lower
Why, then, do most species have sex ratios of
approximately 1:1?
Individual selection wins again!
In population with 2 females for each male, male has
twice the
expected reproductive success of a female
Mutant female producing malebiased sex ratio would
have more
offspring
But then population becomes male-biased...
And mutant female producing femalebiased sex ratio
is favoured

Contest behaviour: bourgeois strategy

Individuals dont behave the
same way in all circumstances
Animals sometimes seem to
exhibit
respect for
ownership
Without anthropomorphizing,
can we
determine if this is
evolutionarily
stable?
Bourgeois:
Play Hawk if territory owner,
play
Dove if intruder
assume that Bourgeois is a
territory
owner one half of
the time
Conditional strategy
Payoffs remain the same,
assume 50% probability of being
the territory owner

Contest behaviour: bourgeois strategy

When Hawks common:
EH(H) = (V-C)/2
EH(B) = (V-C)/4
For V < C, EH(B) > EH(H), so Bourgeois
invades
When Doves common:
ED(D) = V/2
ED(B) = (3V)/4
For V > 0, ED(B) > ED(D), so Bourgeois
invades
When Bourgeois common:
EB(B) = V/2
EB(H) = (3VC)/4
EB(D) = V/4
For V < C, EB(B) > EB(H), so Hawk cant

Contest behaviour: bourgeois strategy

Contest behaviour: bourgeois strategy

What could explain the sunspot butterfly observations?

Owners are physically stronger or otherwise of better
quality than individuals without territories
Weve been using only symmetrical games to model
interactions, but
this assumption is likely invalid
The value of the resource is higher for territory owners than
for intruders
Territory holder likely familiar with territory, whereas
intruder would
need to put in work to become familiar
Territory holder has already established territory
boundaries through
fights with neighbours, whereas
intruder would need to do so

Or bourgeois strategy wins if V < C

Contest behaviour: bourgeois strategy

What could explain the sunspot butterfly observations?

When two butterflies think they own a particular sunspot, they
fight to near death (high costs!), and in the meantime another
butterfly claims the open
sunspot.
Remember, if V < C, Bourgeois is the only ESS.
Although sunspots are a males only opportunity to mate, this
resource is not
fixed (i.e., the position of particular sunspots
change as the day progresses &
with cloud cover).
Therefore, although sunspots are an extremely valuable
resource, each
particular sunspot isn't all that valuable (there
will be other chances to own a sunspot if the butterflies wait)
relative to the cost of injury: V<C!

Contest behaviour: rock-scissor-paper game

Three strategies,
A,B,C.
The rules are: Rock
crushes scissors,
scissors cut paper, paper
covers rock

Nash Equilibria?
None

ESS?
None

Nash equilibrium (after John Nash) is a set of strategies, one for each
player, such that no player can obtain a higher payoff by unilaterally
changing her action. Players are in equilibrium if a change in strategies
by any one of them would lead to a lower payoff for that player than if
she remained with her current strategy.

Contest behaviour: rock-scissor-paper game

Lizard example (Sinervo and Lively 1996).

Contest behaviour: rock-scissor-paper game

Lizard example (Sinervo and Lively 1996).

Orange lizards are territory holders,

Yellow lizards are female mimics, and they can mate with the
females on the Orange-males territories.
Hence, Yellow can increase when rare if Orange is common.
Blue males hold territories, but, unlike the Orange males, Blue
males can recognize
Yellow males and kick them out.
So Blue can increase when rare if Yellow is common.
Finally, Orange can recognize Blue males, and since Orange is
more
aggressive, they can increase when rare when Blue is common.

Once Orange becomes common again, Yellow can increase.

The system is expect to cycle, and the data suggest that is does.

Contest behaviour: war of attrition

Different from previous games: no direct
interactions!

Yellow dung flies

Females lay eggs on fresh cow dung

Males wait near cowpats; first male to
encounter a
female mates with her
As cowpats dry out, they become less
favourable for eggs, so fewer females arrive
Males problem: How long to wait?
As time passes, number of competitors decreases, but so does
number of
females
Best strategy also depends on other males strategies
If most males wait a short time, a male who waits a little longer
will mate with
the later females
If most males wait to the bitter end, a male who moves to a
new cowpat a bit
earlier will mate with the early females there

Contest behaviour: war of attrition

What is the ESS?
V = value of the resource
m = cost of waiting (increases with waiting time)
xA = waiting time selected by individual A (i.e., strategy
played by individual A)
If xA > xB, individual A wins the resource
Payoff to A = V mA
Payoff to B = -mB
No pure strategy is an ESS, because any waiting time x can
be defeated by a slightly longer waiting time
ESS in this case is to play a random waiting time
May be a polymorphism, with different individuals
playing different
times drawn from a random distribution
Each individual could play a mixed strategy and wait a
random
amount of time for each female

Contest behaviour: war of attrition

Minutes after deposition

Reproductive success

No. of searching males

Do yellow dung flies play the war of attrition?

Male stay time

(min)

Males stay times followed negative exponential

distribution
Average reproductive success did not depend on
male stay time

Contest behaviour: other wars of attrition

Dung fly war of attrition is essentially a waiting game
Extensions of this basic model of a WOA exist:

Asymmetric WOA (A-WOA) (Hammerstein & Parker

1982)
- asymmetries in costs and resource value
- mutual assessment

WOA without assessment (WOA-WA) (MestertonGibbons et al. 1996))

- individuals have an energy-cost threshold
- the first individual that crosses this threshold loses
- no mutual assessment

energetic WOA (E-WOA) (Payne & Pagel 1996)

- energy expenditure may vary between individuals
so that rates of
costs accrual may differ between
opponents

Contest behaviour: sequential assessment game

Individuals benefit by avoiding fights they are
likely to lose
Essentially, assess whether V < C
Honest signals of size & strength
Parallel walk & roaring contests in stags
Headbutting in rams
Eyetoeye comparisons in flies
Wrestling matches and croak pitch in frogs &
toads
Pushup displays in lizards
Sequential assessment: Individuals start with the
least costly display and escalate until they have
enough information
about the opponent to decide

Contest behaviour: sequential assessment game

Probability of
winning??
correlated
with specific strategy or measurable individual character state;
resource holding power/potential (RHP)
experience
size
physiological state
animals should be able to measure RHP.

equal competitive ability = equal claw siz

Contest behaviour: sequential assessment game

Resource value
rich microsites

shore crab (Rovero et al. 2000)

access to females blue crab (Jackowski 1974)

velvet swimming crab (Smith et al. 1994)
shore crab (van der Meeren 1994)

food

blue crab (Jackowski 1974)

velvet swimming crab (Smith and Taylor 1993)
shore crab (Sneddon et al. 1997)

Contest behaviour: sequential assessment game

Predictions from sequential assessment game
fight duration increases, as contestants become more evenly matched
fight intensity increases, as contestants become more evenly matched
fight duration increases, as resource value increases
fight intensity increases, as resource value increases

fight

strike

ontest behaviour: testing the sequential assessment game

essed effects of variation in resource value and resource holding power

hting bout duration

ht intensity
hting rate

resource holding power

resource value
4
8
16

3 crab sizes

major claw length

(MCL)

Smallegange et al. 200

ontest behaviour: testing the sequential assessment game

d=56 cm

only males
one extra mussel added
total trial duration: till two mussels were
consumed by the crabs
no satiation
crabs were trained to break mussels

ontest behaviour: testing the sequential assessment game

Smallegange et al. 2007

fight duration was highest with equal

competitors
fight duration decreased with increasing
resource value
no effect of resource value on fighting rate

ontest behaviour: testing the sequential assessment game

Smallegange et al. 2007

fighting rate increased with increasing

competitor size
no effect on number of strikes!

ontest behaviour: testing the sequential assessment game

Summary:
effect of resource value on fight duration
effect of RHP on fight duration: size-matched crabs fight longer
no effect on fight intensity (number of strikes)

But...
SAM and WOAs and other games all predict:
contest duration

WOAs,
other games

Taylor and Elwood (2003)

SAM

absolute size

ontest behaviour: testing the sequential assessment game

contest duration

Smallegange et al. 2007

WOAs,
other games

SAM

absolute size

fight

ontest behaviour: testing the sequential assessment game

Summary:
effect of resource value on fight duration
effect of RHP on fight duration: size-matched crabs fight longer
no effect on fight intensity (number of strikes)
effect of absolute size on fight duration: none
push, hit, kick...
push, push, hit, hit, hit, kick...
push, push harder, push even harder...
continuous assessment game, not sequential

fight

Contest behaviour: one more model

Cumulative Assessment Model (Payne 1998)

Losers give up when accrued costs of fighting cross

threshold (cf. E-WOA)

Costs of fighting can accumulate in two ways:

- usual time and energy costs incurred by performing
agonistic
behaviours
- costs (injuries) can be inflicted directly by actions of
the opponent
- no mutual assessment

Difference with E-WOA is that in E-WOA an individuals

cost accrue from completely internal processes

Contest behaviour: how to distinguish between models?

Model:
Assumptions:
Decision rule
based on:
Type of
assessment
Function of
agonistic
behaviour
Predictions:
Makes
predictions
concerning:
Matching
between
opponents?
Contest
structure

Duration
correlates

Chapter 15, Westneat & Fox (201

Contest behaviour: not always straightforward

Larger blocks
equate to greater costs
Darker shading
equals a better
quality of information

Unlike current model predictions, the sequence of

behaviours used
during a contest both increased and
decreased in aggressiveness
These observations suggest that contests are a form of
unbalanced
partial assessment, corroborating other
recent empirical findings, and challenging theoreticians
to formalise such contest dynamics
Godsall & Smallegange, 2011

Contest behaviour: not always straightforward

Escalation within
a contest is always
initiated by
intruders
regardless of their
opponents size.
Residents are still
more successful in
contests.

It was found that contests dynamics in Uca

perplexa do not completely confirm with any of the
game theoretical models
Rebecca OBrien (2010) MSc thes

Contest behaviour: consequences for foraging behaviour

three competitor sizes:

35-45 mm CW 50-60 mm CW >65 mm CW

smaller

equal

larger

Smallegange & van der Meer 20

Contest behaviour: consequences for foraging behaviour

smaller

time to find
a mussel
equal

larger

Smallegange & van der Meer 20

References
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resident always
wins. Animal Behaviour 26: 138-147.

Godsall B, Smallegange IM. 2011. Assessment games in the mangrove tree-dwelling crab,
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Man, 1887). Crustaceana 84: 1697-1718
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of Theoretical Biology,
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Smallegange, I.M., Van der Meer, J., 2007. Interference from a game theoretical perspective:
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References
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