Zero-sum games pit competitors against each other: your gain is my loss. In competition for a fixed resource—water, territory, mates—one organism's reproductive success comes at the expense of others. Natural selection predicts ruthlessness in zero-sum contexts.1
But many interactions are non-zero-sum: mutual cooperation produces more total benefit than either party could generate alone. Two hunters who coordinate can take down prey neither could kill solo. A mother and infant cooperate: the mother's investment in feeding the child is costly to her, but the benefit to the child (survival) exceeds the cost to the mother (she maximizes her genetic legacy through invested offspring). Coalition members cooperating against a rival can defeat a stronger individual who acts alone.2
Non-zero-sumness is the structural condition that makes cooperation evolutionarily possible. When A and B can both profit from cooperating (even if they profit unequally), cooperation becomes adaptive. When A profits more from exploiting B than from cooperating, exploitation is adaptive. Non-zero-sumness creates the motive; reciprocal altruism and kin selection provide the mechanisms.3
In non-zero-sum cooperation, there's a surplus—joint benefit minus individual gain if alone. Two hunters take down a 200-pound elk worth more than two solo hunters could separately obtain. The question then becomes: how is the surplus divided?
If both hunters split equally, each gets a better deal than hunting alone. But if one is much stronger and could take down larger prey alone, that hunter might demand more of the split, leaving the weaker hunter better off than solo (but worse off than the equal split). The division depends on bargaining power—alternatives available if the deal breaks down. If the strong hunter has many potential partners but the weak hunter has few, the strong hunter can demand a more favorable split.4
This economic principle appears everywhere in nature: a cleaner fish picks parasites off a larger fish, benefiting both (the large fish stays healthy, the cleaner gets food), but the large fish could simply kill the cleaner. The cleaner's bargaining power comes from being hard to find; the large fish's comes from being able to eat the cleaner. In equilibrium, they both benefit relative to their outside options.5