Humans cooperate in large groups of unrelated individuals, and many authors have argued that such cooperation is sustained by contingent reward and punishment. However, such sanctioning systems can also stabilize a wide range of behaviours, including mutually deleterious behaviours. Moreover, it is very likely that large-scale cooperation is derived in the human lineage. Thus, understanding the evolution of mutually beneficial cooperative behaviour requires knowledge of when strategies that support such behaviour can increase when rare. Here, we derive a simple formula that gives the relatedness necessary for contingent cooperation in n-person iterated games to increase when rare. This rule applies to a wide range of pay-off functions and assumes that the strategies supporting cooperation are based on the presence of a threshold fraction of cooperators. This rule suggests that modest levels of relatedness are sufficient for invasion by strategies that make cooperation contingent on previous cooperation by a small fraction of group members. In contrast, only high levels of relatedness allow the invasion by strategies that require near universal cooperation. In order to derive this formula, we introduce a novel methodology for studying evolution in group structured populations including local and global group-size regulation and fluctuations in group size.

We examine bootstrap percolation on a regular (b+1)-ary tree with initial law given by Bernoulli(p). The sites are updated according to the usual rule: a vacant site becomes occupied if it has at least θ occupied neighbors, occupied sites remain occupied forever. It is known that, when b>θ≥2, the limiting density q=q(p) of occupied sites exhibits a jump at some p T=p T(b,θ)∈(0,1) from q T:=q(p T)<1 to q(p)=1 when p>p T. We investigate the metastable behavior associated with this transition. Explicitly, we pick p=p T+h with h>0 and show that, as h ↓0, the system lingers around the “critical” state for time order h −1/2 and then passes to fully occupied state in time O(1). The law of the entire configuration observed when the occupation density is q∈(q T,1) converges, as h ↓0, to a well-defined measure.

The ways in which natural selection can allow the proliferation of cooperative behavior have long been seen as a central problem in evolutionary biology. Most of the literature has focused on interactions between pairs of individuals and on linear public goods games. This emphasis has led to the conclusion that even modest levels of migration would pose a serious problem to the spread of altruism through population viscosity in group structured populations. Here we challenge this conclusion, by analyzing evolution in a framework which allows for complex group interactions and random migration among groups. We conclude that contingent forms of strong altruism that benefits equally all group members, regardless of kinship and without greenbeard effects, can spread when rare under realistic group sizes and levels of migration, due to the assortment of genes resulting only from population viscosity. Our analysis combines group-centric and gene-centric perspectives, allows for arbitrary strength of selection, and leads to extensions of Hamilton's rule for the spread of altruistic alleles, applicable under broad conditions.