Abstract: Explaining altruistic cooperation is one of the greatest
challenges faced by sociologists, economists, and evolutionary biologists.
The problem is determining why an individual would carry
out a costly behavior that benefits another. Possible solutions to this
problem include kinship, repeated interactions, and policing. Another
solution that has recently received much attention is the threat
of punishment. However, punishing behavior is often costly for the
punisher, and so it is not immediately clear how costly punishment
could evolve. We use a direct (neighbor-modulated) fitness approach
to analyze when punishment is favored. This methodology reveals
that, contrary to previous suggestions, relatedness between interacting
individuals is not crucial to explaining cooperation through punishment.
In fact, increasing relatedness directly disfavors punishing
behavior. Instead, the crucial factor is a positive correlation between
the punishment strategy of an individual and the cooperation it
receives. This could arise in several ways, such as when facultative
adjustment of behavior leads individuals to cooperate more when
interacting with individuals who are more likely to punish. More
generally, our results provide a clear example of how the fundamental
factor driving the evolution of social traits is a correlation between
social partners and how this can arise for reasons other than genealogical
kinship.