Spite and the Geometry of Negative Relatedness

AbstractSpite is the most surprising prediction of inclusive fitness theory because it suggests that a gene can be favored by natural selection despite causing harm to both the individuals that carry it and those around them. A gene for spite can only be favored because of negative relatedness, which means that the actor that carries the gene is less likely to share the gene for spite with the surrounding recipients than the random expectation. While positive relatedness can be simply reduced to the intuitive concept of kinship, negative relatedness is deeply counterintuitive. Here I clarify that negative relatedness is frequency dependent, and I identify a hidden assumption in its widely used formula. Accordingly, while the well-studied "lighter" side of inclusive fitness (with helping behaviors and positive relatedness) is dominated by traits that are favored under kin selection, I predict that the understudied "darker" side of inclusive fitness (with harming behaviors and negative relatedness) is dominated by traits that are favored under greenbeard/kind selection-and I discuss the existing evidence that tentatively supports this hypothesis.