2/23/15

Primate
Sociality
&
The Evolution
of
Cooperation

Primates are social

Todays Game Plan

Primate sociality and cooperative living
The evolution of cooperation
Kin Selection
Reciprocal Altruism

Humans are social animals. Why?

For this, it helps
to look at why
our primate
relatives are
social animals as
well.

Benefits of being social

More eyes to see predators detection

Benefits of being social

Less chance of being singled out by a predator
dilution

Gelada herds can number

over 800 individuals the largest
in the world.

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Benefits of being social

More to attack defense

Benefits of being social

Mates usually easy to find
More individuals to look for food
More individuals to defend food

Costs of being social

Social hierarchies produce subordinates

Stress hormone (cortisol)

Larger groups more visible to predators

Larger groups need more food, so more
competition for food.
Increased competition for mates

Costs of being social

Male dominance rank

Costs of being social

Increase in disease transmission

Primate sociality
When to be social?
When the benefits outweigh the costs.

Social animals are those that either

Rely on food that is hard to find but easy to share.
Rely on active (detection, defense) anti-predator
strategies.

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Cooperative Living

The evolution of cooperation

Social animals not only live in close proximity,

but live together cooperatively.
Examples:
Grooming
Alliance formation
Alarm calls and predator defense
Why are these two grooming?

How do alliances form?

These actions are costly to the participants, so why would they evolve?

Cooperative living

Altruism
Altruism: Taking a cost to benefit another
individual.
This is difficult to explain from an evolutionary
perspective.

+ Other

--

+ Self

--

Cooperative living

Cooperative living

+ Other

+ Other

++

-- +
--

+ Self
-- --

+ --

--

++
Mutualism

-- +
Altruistic
--

+ Self
-- -Spiteful

+ -Selfish
--

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Gene-level selection

Primate social groups

How do we explain altruism?

The genes are ultimately what is passed on to
offspring so it is the genes that must benefit.

Primate societies are kin-based

This is not the same at the level of

Kin selection and inclusive fitness theory

In principle, individuals could aggregate any way, including

randomlybut they dont.
We see vastly more cooperation/altruism/investment among
family members than among non-family members.

individual
population
species

Kin Selection
One of two explanations for apparent altruism
(the other is reciprocal altruism).
Kin selection is also referred to as Hamiltons
rule.

W. D. Hamilton
Observed that genes have 2 routes
to the next generation:
Direct: Producing more offspring by
meeting the challenges of the
environment.
Indirect: Aiming the benefits of
altruism at individuals that carry the
same genes you do.

Inclusive fitness: The combined

fitness benefits of direct and
indirect reproduction.

Hamiltons Rule

Coefficient of relatedness

A formal model of kin selection

Total advantage must take into account costs,
benefits, and the probability the same gene is in the
other individual.

rb > c
r: 0-1; coefficient of relatedness; the probability that two people
share a gene by virtue of their sharing a common ancestor.
b: benefit to other
c: cost to individual

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Coefficient of relatedness

Coefficient of relatedness
How related are Bart, Lisa and Maggie to each
other?
Same as asking how likely they are to share any
one particular allele.

50%

Coefficient of relatedness
Marge has Allele X. What is the probability it got
passed to both Lisa and Bart?
To Lisa

To Bart

Yes
No

Yes

No

YY

NY

YN

NN

Coefficient of relatedness
There is a 25% chance that an allele from Marge
makes it into both Lisa and Bart.
The same logic works for an allele from Homer.
Therefore, Lisa and Bart (and Maggie) share 50%
of their genes with each other.
25% + 25% = 50%

Coefficient of relatedness
Some common values for r:
.5 (50%): full siblings, parents
.25 (25%): half-siblings, g-parents, full aunt/uncle
.125 (12.5%): cousins, half aunt/uncle

Example of Hamiltons Rule

A and B are siblings
Cost to A: 3 fitness units
Benefit to B: 7 fitness units
rb > c

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Example of Hamiltons Rule

Example of Hamiltons Rule

A and B are siblings

Cost to A: 3 fitness units
Benefit to B: 7 fitness units

A and B are siblings

Cost to A: 3 fitness units
Benefit to B: 7 fitness units

rb > c
.5x7 ? 3

rb > c
3.5 > 3

Kin selection
Acts of support in wild baboons

Inclusive fitness
Inclusive fitness: It is the genes that benefit, not
necessarily the individual.

Beneficial for A to help.

Kin selection
Time spent
grooming in rhesus
monkeys

Parent-offspring conflict
Example: Weaning conflict
A and B are siblings, their mother
Cost to A: 3 fitness units
Benefit to B: 2 fitness units
Mother:
Cr < Br
3x.5 ? 2x.5
1.5 > 1

Adaptive, wants A to share, but A

does not want to share.

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Reciprocal Altruism
Robert Trivers
The other evolutionary
explanation for apparent
altruism.
You pay a cost now to
benefit me, I'll pay a cost
later to benefit you and we'll
both be better off in the long
run.

Reciprocal Altruism
Reciprocal altruism can only work if these
requirements are met:
Regular interaction between individuals
Recognition of individuals
Memory for others' past actions
Discrimination against non-reciprocators

Reciprocal Altruism
Bats must choose whether or not to help another
that did not eat.
Costs: Loose a portion of their nutrients.
Benefits: Other might help when they don't get to
eat.

Reciprocal Altruism

Me

Time 1

Time 2

Time 3

Time 4

You
-

+
+
-

Both parties have a net benefit from the interaction.

Reciprocal Altruism
Case Study: Vampire Bats
Basic ecology
Live in caves with hundreds of others
Hunt exclusively
If meal doesn't happen, severely impairs ability to get
meal next night
Die within three days of not eating
Can vomit portions of food at will

Reciprocal Altruism
A primate example
Vervet monkeys
repond more strongly
to a distress call if they
had previously
groomed the caller.

Observed behavior: Vampire bats give food to

those that have given food them in the past.

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Cooperation and Strategies

Cooperation can evolve under reciprocal altruism
only if 'cheaters' are not assisted.
Must discriminate against those that will not repay
the benefit.
How can you tell who this will be?

Cooperation and Strategies

Options are 'Cooperate' and 'Defect'
Tit-for-Tat Strategy
Always cooperate the first turn.
Do what the other player did in the previous turn.

Cooperation and Strategies

A rule of thumb is to withhold benefits from
those that have failed to provide previously and
continue the relationship with those that have
cooperated previously.

This is a strategy called Tit-for-Tat.

Cooperation and Strategies

Tit-for-Tat withholds cooperation from those
that do not provide benefits.
This protects from excessive losses.
Tit-for-Tat continues cooperation with those that
provide benefits.
Cooperating first allows Tit-for-Tat to play well
with itself.
Tit-for-Tat also 'forgives.

Cooperation and Strategies

Reciprocal Altruism

There are other strategies for how to play the

game.

Tit-for-Tat does better than Always Defect

Always Defect
Always Cooperate
Grim Trigger

Tit-for-Tat does better than Always Cooperate

Always Defect looses out on cooperative exchanges

when it becomes common
Always Cooperate plays well with Tit-for-Tat but is
vulnerable to Always Defect

Tit-for-Tat does better than Grim Trigger

Grim Trigger does not forgive so looses out on any
cooperation potential after first defection