28/1/2019

Behavioral Ecology 1
Evolutionarily Stable Strategies; Natural selection;
Optimality Theory in resource acquisition and defence

Animals demonstrate intriguing behaviour

Reproductive behaviour in lions
• Lion prides comprising related females and their cubs; females have long
reproductive years
• Males leave their natal pride at 3 years old, spend some time as nomads, then
attempt to take over a pride from old/weak males
• Males have short reproductive lives (2-3 years)
• Females in a pride all become oestrus at about the same time (pheromones)
youngs are born the same time – what are the advantages of this synchronized
reproduction?
• Females are receptive to mating (on heat) frequently but conception rate is low,
~1/3000 chance that copulation will produce an offspring reaching maturity
Why do females demonstrate such an inefficient reproductive behaviour?
good
• A new male taking over a pride will kill all cubs fathered by the previous male
𠠬

(Our Planet)

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This behaviour is adaptive

• physiological and functional
explanations for these features of
the lion’s reproductive behaviour
• Many of these features are
genetically heritable; the
acquisition of these features may
influence the reproductive success
and therefore fitness of the
individual
• At the species level, such features
are evolutionarily stable – the
“strategy” will remain adaptive
(and does not change) unless
parameters of the environment
change, e.g. food availability
(Krebs and Davis)

Evolutionarily stable strategies (ESSs)

• A strategy is an ESS if, when adopted by the majority of a population, it cannot be
invaded by the spread of any rare alternative strategy (Maynard Smith 1972)
• The reward (i.e. Darwinian fitness) for the strategy (i.e. a course of action chosen)
of an individual is dependent on the strategies taken by other individuals
• Competitive optima achieved through an ESS are different from simple optima (i.e.
the best strategy defined for a set of fixed environmental conditions, e.g. those
based on the “optimality” premise ) – fitness may not be the theoretical
maximum under an ESS
• An example of the application of the ESS concept is in analyzing animal contests,
e.g. the war of attrition model (whoever persists wins); the hawk-dove model (the
contestant who can injure his opponent wins)

Passengers “fight” over the armrest

on an airplane: which strategy is
optimal or stable?

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Behaviour is governed by genes

• Migratory birds exhibit restlessness when caged
• Activities (reflected by scratch marks) reflect distinct population-specific direction
of flight
• Austrian blackcaps SE to Israel and N Africa; German blackcaps SW to Spain
and N Africa; hybrids show intermediate flight directions
• Behaviour should be interpreted in ⼀一
the premise of natural selection

(RSPB)

Blackcap Sylvia atricapilla

Hypothesis testing in behavioural ecology

• Apply the scientific process in the study of behavioural ecology
• Observation: female lions mate with male lions in a pride frequently but rate of
successful offspring production is low (~1:3000)
• Hypothesis: females “devalue” each copulation with a male by becoming
receptive even when the chance of conception is low ( increases paternal
uncertainty), so as to reduce the chance that her cub may be killed by a male; less
male competition (and changeover of pride ownership)
• Alternative hypothesis: males have low fertility
• Prediction: females mate with multiple males in a pride; males will be less likely
to kill cubs; females and their cubs get better protection

Survivorship of litter significantly higher when reproduction was synchronized and

when no older cubs were present (Bertram 1975 J. Zool. 177:463-82)

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Optimality Theory (Robert H. MacArthur)

• All behaviour comes with costs and benefits, with proximal currencies such as
energy and time; with fitness (survivorship, reproductive success) as the ultimate
currency for the trade-off between cost and benefit
• Organisms are assumed to maximize the difference between cost and benefit so as
to achieve maximum fitness – the Optimality model
• This goal for optimization drives adaptive behaviour – organisms demonstrating
more optimal behaviour will be favoured by natural selection and the behaviour
persists in the population
• Examples: Blackheaded gull removing hatched shell to reduce risk of nest
predation (Tinbergen); crab choosing particular prey size while foraging
(Youtube) (Flickr)

Larus ridibundus removing reflective hatched Blue crab breaking ribbed mussel for food – what are
shell from nest to protect young from predators the costs and benefits of prey size selection?

“Economic” analysis of behaviour following the Optimality approach

• Usually more tangible currencies such as energy, time or nutrients are used in
analyses of behaviour, as fitness is more difficult to assess – Why? durationobservation
Long
• Usually only one currency is used to assess the costs and benefits of behaviour –
xexplain what are the potential drawbacks of this approach?fdynssesienergysoueanhaab
everything• Which other considerations animals must make before deciding on a certain hot a
behaviour? my be
maximizes
energy
Injury risk
Energy content
⻄西是timemine
Handling time

Interference competition

Predation risk Availability of

preferred prey size

Prey size selection by crab

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Examples of economic analysis of animal behaviour

Starling load carrying when foraging
• Starlings forage for leatherjacket (cranefly larvae) to feed their young
• Many trips (up to 400 per day) are necessary during peak breeding season
• Beak has finite/optimal holding capacity for larvae while continuing to probe
• There is a cost in making unnecessarily many trips (e.g. carrying only 1 larva each
time)
• How many larvae should a starling hold before flying back to nest in order to
maximize the food delivery rate to the nestlings?

Transport
load Probe, collect

(Sialis)

(Eylandt)

Optimal load for starling on foraging trips

• Decision rule: load is optimal when the rate of delivery of larvae is greatest, i.e.
(load/sum of travelling and searching times) is greatest
slope
• Travelling time is a constant for a particular foraging area (A)
• Searching time varies with load: short for initial catches, increases gradually
(curve) – actual relationship is not a smooth curve but stepped
• Optimal load value found by drawing straight lines relating A and search curve, e.g.
A,C for 8 larvae; A, D for 1 larva; and finding the load that gives the greatest slope
(max load per total time = line AB; i.e. 7 larvae per trip)
• Optimal load changes with the travelling time – smaller load for shorter travelling
time

l hadus 器
D

(Krebs and Davis)

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Time spent on copulation in dungfly males

• Male dungflies compete to fertilize females – males mating more recently fertilize
most of the eggs (80%) produced by a female (sperm competition)
• Males should hold on to a female after mating to ensure high fertilization rate
(benefit) but this risks reducing its own chance to fertilize additional females (cost)
• The gain in successful fertilization with increased copulation time diminishes
quickly ㄟ lose chaneooefevtiiiyotherfemaleeggs
• Q – how long should a male hold on to a female during copulation?
• “travelling time” (as in the starling example) in this case is the sum of searching
plus guarding time spent by the male (average = 156.5 minutes)
linewith
greatestslope
0.8

(Krebs and Davis)

Foraging decisions: optimal time in patch and optimal prey size

• Predators such as crabs foraging by searching for suitable prey and handling (e.g.
crushing) them for food
• Optimization here means maximizing the energy return per unit time foraging
(searching + handling)
• Continued foraging in a patch depletes prey of high value, resulting in a relationship
of diminishing marginal returns, similar to those of the starling and dungfly cases
• Optimal time spent on a patch (Topt) may be estimated in a similar way

Crab should leave the patch

at Topt

Time in patch
(Stiling)

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Marginal value theorem (Eric Charnov)

• Gain per unit time diminishes with time spent on patch (e.g. prey depletion, prey
evasion)
• Predator should leave patch when the marginal return drops to the average
energy return for the whole habitat
• The same decision rule (tangents parallel to that for the average patch) should be
applied to all patches to determine leaving time

• Slope of AB represents the

maximum rate of energy gain for

i
the average patch
• Same decision applies to abandon
more (i) and less (ii) productive
patches at different Topt

When should a predator switch prey?

Suppose a predator has two options of prey P1 and P2, with energy contents E1 and
E2; handling time h1 and h2; and searching time t1 and t2, respectively
When the predator is searching for P1, it encounters an individual of P2 – should the
predator switch to P2 or keep looking for P1?

The profitability of consuming the P2 individual is E2 / h2, while that for searching
and consuming another P1 is E1 / (h1 + t1), therefore

The predator should consume P2 if E2 / h2 > E1 / (h1 + t1), or

The predator should ignore P2 and keep looking for another P1 if

E2 / h2 < E1 / (h1 + t1)

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Prey size selection by a predator

• Predators invest energy in searching and handling prey
• Prey handling time (e.g. time taken by crab to crush a shell) often increases
exponentially with prey size
• Energy return (E) also changes with prey size but may not follow the same
relationship
• Profitability of a prey is the ratio of energy gained by eating the prey to energy spent
on finding, handling and digesting it
• Profitability often just simplified to handling time (Th), as energy spent, or searching
time (Ts) and digesting time are more difficult to measure
• Predators are expected to select prey with the highest profitability (E/T); T = Th+Ts
or just Th

(Stiling)

Selection of prey with different energy return and encounter rates

• A predator searches for a period of Ts and finds two prey types at rates of λ1 and λ2
(number per sec); energy contents E1 and E2 , and handling time and h2,
respectively
• Profitability of the two prey are therefore E1 / h1and E2 / h2, respectively
• If the predator eats both prey, in Ts seconds it will gain

E = Ts (λ1 E1 + λ2 E2)
Taking a total time of T = searching time + handling time = Ts+ Ts(λ1h1 + λ2h2)
The predator’s rate of energy intake is, cancelling out Ts ,
𝐸 λ1 E1 + λ2 E2
𝑇 1 λ1h1 + λ2h2

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ii
If prey 1 is more profitable than prey 2, predator should specialize on prey 1 if
λ1 E1 λ1 E1 + λ2 E2
1 λ1h1 1 λ1h1 + λ2h2
i.e. energy intake from specializing on prey 1 is greater than from consuming
both
Or, rearranging,
E1
(h – h )
λ1 E2 2 1

is the time taken to find an individual of prey 1.

λ1
(Fine Cooking)

Profitability
??

(tripadvisor)

Why simple optimality models are unrealistic

• Animals’ behaviour (e.g. foraging, mating) is not simply determined by
maximizing success (e.g. maximum energy yield per unit foraging time) but also
other selective pressures
• Time or risk minimization is often an important selective pressure influencing
behavioral choices (life-time reproductive success is the goal)
• Risks such as injury (e.g. tackling a large prey), predation (e.g. while mating,
handling prey) influence actual behaviour – trade-offs are common
• Juanes (1992) reported actual prey size range taken was significantly smaller
than the optimal size range predicted based on energy maximization - to avoid
injury

Predicted range

Actual range
taken
(shaded)

(Juanes 1992 Mar Ecol Prog Ser)

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Trade-offs: own predation risk may drive sub-optimal behaviour

Sticklebacks (Gasterosteus aculeatus) prefer
to attack high-density prey (water flea)
when hungry but low-density prey when
not: cost of picking out individual prey
higher in high-density prey patches

Using high-density prey patches also

reduce ability to detect its own predator –
higher predation risk only worth taking
when hungary

When a model kingfisher was flown over

the experimental tank of hungry
⼀一low-density
sticklebacks they switched to
prey patches

The degree of hunger of the predator is (Milinski and Heller 1978)

a determining factor in the optimality

model

What are optimality models good for?

• Provide testable, quantitative predictions for hypothesis testing about animal
behaviour, e.g. energy maximization in foraging, and the currency and constraints
on behaviour
• Allow explicit expressions of the assumptions underlying the currency and
constraints hypotheses, e.g. the starling can only encounter one foraging patch at a
time in the “load” model
• Emphasize the generality of simple decisions facing different animals in many
aspects of their daily lives

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Many experiments generating choice data are not context-sensitive

Laboratory data demonstrating optimal foraging in predators are often conducted in
unrealistic conditions, e.g.:
• Gregarious prey (e.g. mussels) were offered as isolated individuals – handling time
is greatly reduced
• Prey size may not be easily assessable by the predator in field situations
• Natural agents potentially interfering with the predator, e.g. competing
conspecifics, own predators, were ignored
• These and other constraints drive deviations from optimality models

(ThoughtCo)

What if animals do not “know-all”, after all?

• Optimality models assume the animal “knows” all about the energy content of its
prey, the quality of the patch (e.g. composition of the prey items), or the
searching time and handling time of different prey – unlikely true
• These quantities are often highly variable in nature, in stochastic (unpredictable)
ways
• The animal may be driven to deal with this unpredictability by (1) responding to
the risk of doing well or doing badly in a particular situation; and/or (2) deal with
the variability by sampling to acquire key information on patch quality
• The marginal value theorem predicts optimal outcomes when the animal already
has knowledge of patch quality, searching time, etc.

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Risk taking
• Decisions on the degree of risk associated with a particular situation changes
with the condition (e.g. physiological need, hunger level) of the animal
• Caraco (1981) provided choices of two feeding sites to the yellow-eyed junco: (1)
constant award; and (2) unpredictable award but with a similar mean award to (1)
• If juncos went for energy maximization, either options are fine no preference
• But: 1-hr starvation constant award site preferred
4-hr starvation unpredictable site preferred
• 1-hr starvation constant award site preferred: daily survival is guaranteed
risk aversion approach
• 4-hr starvation unpredictable site preferred: constant site is sure fail (not
enough for daily survival); unpredictable site offers a 50% chance of daily
survival risk prone approach

(E.J. Peiker) (The Deer Hunter)

Information
• A forager can obtain information about the quality of a patch by initial sampling
• A forager performing sampling should on average stay longer than period
predicted by the marginal value theorem because any extra time spent could
reveal that the patch is actually better than the current information suggests
• Worth sacrificing a little from the maximum energy intake rate predicted by the
deterministic optimal model to gain this extra beneficial information
• Patch sampling particularly useful when patch quality variance is high (but then
the foraging may be able to assess patch without sampling, e.g. olfactory signals)

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Territoriality
• Territorial behaviour is a result of intraspecific competition
• Individuals or groups of individuals occupy and defend exclusive habitats
(and their resources) through signals and /or aggression (interference
competition)
• Territoriality regulates population density
• There are costs associated with territorial behaviour – costs of defending
your turf

Competition for resource – the ideal free distribution

• Habitat use when there is no interference competition but only resource
exploitation will result in eventual equal rate of acquisition by all competitors
• Individuals will first occupy the rich habitat, no limit on how many can stay here
(no interference competition)
• Resource in rich habitat will be depleted with increasing occupants
• Late comers may achieve equal resource use in the poor habitat as benefit of the
rich habitat is reduced by increasing occupants
• Competitors then redistribute between the two habitats in relation to their quality
(the “ideal free distribution” – free movement to follow choice)
• Eventually profitability is the same for all competitors irrespective of habitat
Switching queues in supermarket?
Benefit of using rich habitat is the
same as switching to poor habitat
once no. of competitors reaches ‘a’

(The Mirror)

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Resource competition by defence – territoriality

• Late comers may be forced to occupy poor habitat when there is territorial
behaviour (interference competition) – resource defence by aggression to reduce
competitor’s ability /efficiency of searching / using resoruce
• Future comers may have no access to resources at all - floaters
• Vacant territories are quickly occupied
• Common behaviour among animals
• Often there is a mixture of both the ideal free distribution and resource defence

(Science Daily)
(Flickr)

Economic defendability of a territory

• Larger territories offer more resources but also incur higher costs of defence
• A territory is worth defending when the benefits (of having priority access to
resources) is greater than the cost of defending it
• Time spent on defending often used as a proxy for energy cost
• Optimal territory size = maximum net gain in energy
• Defence cost increases exponentially with territory size
• Benefit of territory is asymptotic – when resource level gets too high, animals
cannot fully utilize
The optimal territory sizes predicted
for the rich and poor habitats are X and
X’, respectively (= maximum difference
between benefit and cost)

Optimal size is larger for poorer

ǖi habitat – animals defend larger
territories in poor habitats

Any territory size between A and B is

economically defendable
(Krebs and Davis)

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