Jvancleve Evol2017

Stags, hawks, and doves:
Individual variation in helping in social evolution theory
Jeremy Van Cleve

University of Kentucky
June 24th, 2017
Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Variation and the four games of cooperation

Consider a social interaction with

two individuals where each can
“cooperate” (C) or “not cooperate / defect” (D)

Consider a social interaction with

two individuals where each can
“cooperate” (C) or “not cooperate / defect” (D)
Game Equilibrium Type of variation
Prisoner’s dilemma No cooperation No variation
Mutualism game Full cooperation No variation
Stag-hunt game Full or no cooperation Coarse-scale variation
Hawk-dove game Mixed cooperation Fine-scale variation

The four basic games of cooperation
Cooperation produces additive Benefit (B)

to partner at Cost (C) to self. ( B > C )
Cooperate (C) Defect (D)

Payoff to
B–C+D –C
Cooperate (C)
Payoff to
B 0
Defect (D)


D = extra benefit of two cooperative acts

D = “synergy”
Payoff to
B–C+D –C
Cooperate (C)
Payoff to
B 0
Defect (D)



D = “synergy”
If C > 0 : (cooperation individually costly)
Payoff to
D < C : Prisoner’s dilemma B–C+D –C
Cooperate (C)
D > C : Stag-hunt game
Payoff to
B 0
Defect (D)



D = “synergy”
If C > 0 : (cooperation individually costly)
Payoff to
D < C : Prisoner’s dilemma B–C+D –C
Cooperate (C)
D > C : Stag-hunt game
Payoff to
B 0
If C < 0 : (cooperation individually helpful) Defect (D)
D < C : Hawk dove game
D > C : Mutualism game

Moving between the four games
Moving between games can occur by altering the fitness costs and
benefits of the behavior.

I. Demographic factors that affect scaled relatedness

I. Demographic factors that affect scaled relatedness
II. Behavioral factors that affect responsiveness (reciprocity)

Shifting games with relatedness
Hamilton’s rule: –C+Bκ>0
payoffs scaled relatedness
dispersal
N N
N N


If Bκ>C full cooperation
dispersal
(mutualism game)
N N
N N


dispersal
(mutualism game)
N N
If Bκ<C no cooperation
(prisoner’s dilemma)
N N


dispersal
(mutualism game)
N N
N N
Neither outcome selects for variation.


dispersal
(mutualism game)
N N
N N
Neither outcome selects for variation.
Need synergy ( D ≠ 0 ) for either

stag-hunt or hawk dove games

Start with the payoff matrix with synergy

Payoff
(1+ κ)(B – C + D) – C + κ B
to C
Payoff
B–κC 0
to D

Add relatedness to the payoff matrix

Payoff
(1+ κ)(B – C + D) – C + κ B
to C
Payoff
B–κC 0
to D

Which of the four games the payoff

matrix represents depends on Cooperate (C) Defect (D)
both D and κ. Payoff
(1+ κ)(B – C + D) – C + κ B
to C
Payoff
B–κC 0
to D

Which of the four games the payoff

matrix represents depends on Cooperate (C) Defect (D)
both D and κ. Payoff
(1+ κ)(B – C + D) – C + κ B
to C
Assume costly cooperation ( – C < 0 ) Payoff
B–κC 0
to D

Stag Hunt Mutualism

C
Synergy, D
0 Payoff
(1+ κ)(B – C + D) – C + κ B
to C
Payoff
B–κC 0
Prisoner's to D
Dilemma Hawk Dove
0 C/B 1
Relatedness, κ

Shifting games with responsiveness
Interactions can be persistent (i.e., repeated)

Animals may respond to one another as they choose

how much to cooperate


“Reciprocity” or “reciprocal altruism”


Can allow cooperation even in prisoner’s dilemma

ρ

Can allow cooperation even in prisoner’s dilemma
Measure with responsiveness ( ρ ), which is how

much one responds to the past action of its partner.

ρ
Start with the payoff matrix

๣ ߼ #߼ ๣ $
#๣$ ๣ ߼ #߼ӝ ๣ $
Payoff
#๣$ ๣߼
๣ ߼ # ๣ $߼ ๣ ߼ӝ
to C
๣߼ #๣ $߼
Payoff ๣ ߼ӝ
๣ ߼ӝ
to D ๣߼

ρ
Add average payoffs due to level

of responsiveness ( ρ ).

๣ ߼ #߼ ๣ $
#๣$ ๣ ߼ #߼ӝ ๣ $
Payoff
#๣$ ๣߼
๣ ߼ # ๣ $߼ ๣ ߼ӝ
to C
๣߼ #๣ $߼
Payoff ๣ ߼ӝ
๣ ߼ӝ
to D ๣߼

ρ

If Bρ>C full cooperation Cooperate (C) Defect (D)

๣ ߼ #߼ ๣ $
#๣$
(mutualism game)
๣ ߼ #߼ӝ ๣ $
Payoff
#๣$ ๣߼
๣ ߼ # ๣ $߼ ๣ ߼ӝ
to C
๣߼ #๣ $߼
Payoff ๣ ߼ӝ
๣ ߼ӝ
to D ๣߼

ρ


๣ ߼ #߼ ๣ $
#๣$
(mutualism game)
๣ ߼ #߼ӝ ๣ $
Payoff
#๣$ ๣߼
๣ ߼ # ๣ $߼ ๣ ߼ӝ
to C
๣߼ #๣ $߼
If Bρ<C no cooperation
Payoff ๣ ߼ӝ
๣ ߼ӝ
(prisoner’s dilemma) to D ๣߼

ρ


๣ ߼ #߼ ๣ $
#๣$
(mutualism game)
๣ ߼ #߼ӝ ๣ $
Payoff
#๣$ ๣߼
๣ ߼ # ๣ $߼ ๣ ߼ӝ
to C
๣߼ #๣ $߼
If Bρ<C no cooperation
Payoff ๣ ߼ӝ
๣ ߼ӝ
(prisoner’s dilemma) to D ๣߼
Again, need synergy ( D ≠ 0 ) for either

stag-hunt or hawk dove games

ρ
Adding synergy to the payoff matrix

๣ ߼ #߼ ๣ $
#๣$ ๣ ߼ #߼ӝ ๣ $
Payoff
#๣$ ๣߼
๣ ߼ # ๣ $߼ ๣ ߼ӝ
to C
๣߼ #๣ $߼
Payoff ๣ ߼ӝ
๣ ߼ӝ
to D ๣߼

ρ
Cooperate
Cooperate
(C) (C) Defect
Defect
(D)(D)
๣ ߼ ߼ ๣
๣ ߼ %#%߼ ๣ ߼๣
#๣ #๣$ $ % %# ๣ $ ๣ ߼ ߼๣
๣ ߼% ߼ӝ ๣
߼๣
Payoff Payoff ## $$
#๣#߼๣ ߼
๣๣ $ $ %%# ๣ $ ๣߼ ӝ#%ӝ $$
๣๣߼ ߼# # ๣$ $
๣ ๣%߼ %߼#߼๣ $߼ ๣๣߼๣ӝ߼߼
to C to C
ӝӝ
##
Payoff ๣
๣߼Payoff
߼##๣ ߼๣ӝ$ӝ$
๣ ๣%߼ %߼#߼๣ $߼
๣߼ ๣߼ ӝ

to D to ๣
๣
D ߼ ߼ӝ ๣ ߼
ӝ ӝ

ρ
Which of the four games the payoff matrix

represents depends on both D and ρ.
Cooperate
Cooperate
(C) (C) Defect
Defect
(D)(D)
๣ ߼ ߼ ๣
๣ ߼ %#%߼ ๣ ߼๣
#๣ #๣$ $ % %# ๣ $ ๣ ߼ ߼๣
๣ ߼% ߼ӝ ๣
߼๣
Payoff Payoff ## $$
#๣#߼๣ ߼
๣๣ $ $ %%# ๣ $ ๣߼ ӝ#%ӝ $$
๣๣߼ ߼# # ๣$ $
๣ ๣%߼ %߼#߼๣ $߼ ๣๣߼๣ӝ߼߼
to C to C
ӝӝ
##
Payoff ๣
๣߼Payoff
߼##๣ ߼๣ӝ$ӝ$
๣ ๣%߼ %߼#߼๣ $߼
๣߼ ๣߼ ӝ

to D to ๣
๣
D ߼ ߼ӝ ๣ ߼
ӝ ӝ

ρ
Stag
Hunt Mutualism
C
Synergy, D

๣
๣߼ ߼ # # % %߼ ๣߼๣
#๣ #๣ ๣
๣߼ ߼ #๣#߼% %߼ ๣
߼๣
$$
๣ӝ߼ӝ
0
Hawk Dove ๣ ߼ ###๣ ๣$ $๣$% %% ߼
$ $ %% $$
๣߼ # ๣$ % ߼ ๣ ๣߼ӝ߼ӝ
๣ ๣߼ ߼##๣ ߼๣ӝ$๣ӝ$ % %߼ ߼
๣߼
Prisoner's ๣
๣߼ӝ߼ӝ
Dilemma
0 C/B 1
Responsiveness, ρ

Shifting games with relatedness & responsiveness
Combine scaled relatedness & responsiveness into one payoff matrix

ρ=0
Synergy, D
Stag Hunt Mutualism

C
Prisoner's
Dilemma Hawk Dove
0 C/B 1
Relatedness, κ

ρ=0 ρ = 0.1 ρ = 0.3 ρ = 0.5
Stag Stag
Synergy, D
Stag Hunt Mutualism Mutualism Mutualism Mutualism

Hunt Hunt
C
Prisoner's Prisoner's Prisoner's

Dilemma Hawk Dove Dilemma Hawk Dove Dilemma Hawk Dove PD Hawk Dove
0 C/B 10 C/B 1 0 C/B 1 0 C/B 1
Relatedness, κ

κ=0
Stag
Synergy, D
Hunt Mutualism
C
0
Hawk Dove
Prisoner's
Dilemma
0 C/B 1
Responsiveness, ρ

κ=0 κ = 0.1 κ = 0.3 κ = 0.5
Stag Stag Stag

Synergy, D
Hunt Mutualism Hunt Mutualism Hunt Mutualism Mutualism

C
0
Hawk Dove Hawk Dove Hawk Dove
Prisoner's Prisoner's Prisoner's
Dilemma Dilemma Dilemma Hawk Dove
0 C/B 10 C/B 1 0 C/B 1 0 C/B 1
Responsiveness, ρ

General patterns for variation in social behavior
Squinting a little…
Variation most likely when…

Fine-scale variation (polymorphism / division of labor):

~ negative synergy, and some relatedness ( κ ) or moderate
responsiveness ( ρ )


Coarse-scale variation (variation among groups / across time):

~ strong positive synergy, and zero to moderate relatedness ( κ ) or


Coarse-scale variation (variation among groups / across time):

~ strong positive synergy, and zero to moderate relatedness ( κ ) or
Thus, comparative patterns should depend on synergy, relatedness,

and responsiveness (and other factors such as groups size)
Questions?
Van Cleve, J. Integr. Comp. Biol. 2017.

Peña, J., Nöldeke, G., & Lehmann, L. 2015. J Theor Biol 382:122--136. doi:10.1016/j.jtbi.2015.06.039
Van Cleve, J. 2015. Theor Popul Biol 103:2--26. doi:10.1016/j.tpb.2015.05.002
Van Cleve, J. & Akçay, E. 2014. Evolution 68:2245--2258. doi:10.1111/evo.12438
Acknowledgements
SICB
NSF
National Academies Keck Futures Initiative

Shifting games and group size
Social interactions among >2 individuals can be highly complex

How do the benefits of cooperation vary from 1 to n cooperators???

Try to match pairwise interaction as closely and “simply” as possible

Try to match pairwise interaction as closely and “simply” as possible
n=2 n=3 n=5 n = 10
Stag Stag Stag

Synergy, D
Stag Hunt Mutualism Mutualism Mutualism Mutualism

Hunt Hunt Hunt
C
Prisoner's Prisoner's Prisoner's Prisoner's

Dilemma Hawk Dove Dilemma Hawk Dove Dilemma Hawk Dove Dilemma Hawk Dove
0 C/B 10 C/B 10 C/B 10 C/B 1
Relatedness, κ

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Stags, hawks, and doves:

Individual variation in helping in social evolution theory

Jeremy Van Cleve

June 24th, 2017

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Consider a social interaction with

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Consider a social interaction with

Game Equilibrium Type of variation

Prisoner’s dilemma No cooperation No variation

Mutualism game Full cooperation No variation

Stag-hunt game Full or no cooperation Coarse-scale variation

Hawk-dove game Mixed cooperation Fine-scale variation

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Cooperation produces additive Benefit (B)

Cooperate (C) Defect (D)

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Cooperation produces additive Benefit (B)

D = extra benefit of two cooperative acts

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Cooperation produces additive Benefit (B)

D = extra benefit of two cooperative acts

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Cooperation produces additive Benefit (B)

D = extra benefit of two cooperative acts

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Game Equilibrium Type of variation

Prisoner’s dilemma No cooperation No variation

Mutualism game Full cooperation No variation

Stag-hunt game Full or no cooperation Coarse-scale variation

Hawk-dove game Mixed cooperation Fine-scale variation

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Game Equilibrium Type of variation

Prisoner’s dilemma No cooperation No variation

Mutualism game Full cooperation No variation

Stag-hunt game Full or no cooperation Coarse-scale variation

Hawk-dove game Mixed cooperation Fine-scale variation

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Game Equilibrium Type of variation

Prisoner’s dilemma No cooperation No variation

Mutualism game Full cooperation No variation

Stag-hunt game Full or no cooperation Coarse-scale variation

Hawk-dove game Mixed cooperation Fine-scale variation

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Hamilton’s rule: –C+Bκ>0

payoffs scaled relatedness

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Hamilton’s rule: –C+Bκ>0

payoffs scaled relatedness

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Hamilton’s rule: –C+Bκ>0

payoffs scaled relatedness

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Hamilton’s rule: –C+Bκ>0

payoffs scaled relatedness

Neither outcome selects for variation.

Individual variation in helping in social evolution theory UNIVERSITY OF KENTUCKY

Hamilton’s rule: –C+Bκ>0

payoffs scaled relatedness

Neither outcome selects for variation.

Need synergy ( D ≠ 0 ) for either