1. Cooperation and Reciprocal Altruism
Definitions
Iterated prisoner’s dilemma
Examples
Food sharing
Alliance formation
Egg trading
Predator inspection
Social grooming

2. Evolution of cooperation
Mutualism
Kin selection
By-product mutualism
as a consequence of behaving selfishly, the donor inadvertantly benefits the recipient.
Example: cleaner wrasse selfishly consume ectoparasites of larger fish. The large fish cooperates by not eating the cleaner fish.
Cleaner wrasse have mimics that cheat!
Reciprocal altruism

3. Reciprocal Altruism
the trading of altruistic acts in which the benefit is larger than the cost so that over time participants enjoy a net gain.
Delay between donation cost and receipt of benefit separates mutualism from reciprocal altruism.
Delay allows for the possibility of cheating, thus cheaters must be detected and excluded
A sufficient number of interactions must occur to provide a net benefit to participants. Note that in many instances, the net benefit will increase with the number of exchanges. Thus, a large number of interactions favors reciprocity.

4. The prisoner’s dilemma
Two suspected criminals are jailed separately and encouraged to provide evidence that the other was involved in the crime
PD is defined by T > R > P > S and R > (T + S) / 2
ESS for single round of the game: always defect!

5. The Iterated Prisoner’s Dilemma
Iterating this game allows for cheating - the key distinction between mutualism and reciprocity
Iteration permits complicated strategies, e.g. one player can perform CDCDCCCD while another might do CCCCCCCC, etc.
TFT (cooperate on the first move and thereafter mimic your opponent) is the best strategy because
Outscored all other strategies in computer tournament (Axelrod)
Is an ESS if the probability of future encounter, w, meets these criteria:
w > (T - R)/(T - P) and w > (T - R)/(R - S)
Obtain these inequalities by applying 1, w, w2, w3,... to successive future payoffs and noting that w + w2 + w = 1/(1 - w)

6. Beyond tit-for-tat Once TFT evolves, can other strategies invade?
Subsequent work indicates that other trajectories may occur, e.g. TFT-> Generous TFT-> Pavlov-> cooperation (Nowak & Sigmund)
If mistakes are made, Generous-tit-for-tat does better than TFT (GTFT cooperates after opponent cooperates but also after opponent defects with some probability)
Pavlov - win-stay, lose-shift does better than TFT because it corrects occasional mistakes and exploits unconditional cooperators.

7. Vampire bat food sharing

8. Costs and benefits of food sharing

9. Survival gain of food sharing

10. Chimpanzee food sharing

11. Cotton-top tamarin food sharing

12. Alliance formation
Baboons
Vervet monkeys
Bottlenose dolphins

13. Egg-trading in polychaetes and bass

14. Predator inspection in fish

15. Predator inspection - mirror expt
But, same result is seen in
The absence of any predator!
Suggests that fish tend to school.
Fish with parallel mirror approached
closer than fish with oblique mirror

16. Social grooming in antelope
Females
Males

17. Implications for human behavior
Friendship formation
non-kin directed altruism
gift exchange ceremonies
Emotion evolution
Gratitude
guilt and reparative altruism
Justice
moralistic aggression
revenge
Reciprocal network size
cartel formation
dialects