Identity Signaling and Patterns of Cooperative Behavior

Recognition systems play a central role in mediating cooperative behavior among individuals in a population. Despite the importance of discriminating among potential recipients of cooperation, the evolutionary forces that maintain diversity in traits used for kin recognition are poorly understood. Greenbeard-based models of kin recognition in which alleles for cooperative behavior also control recognition of those alleles in potential cooperators suggest that discrimination based on a greenbeard locus leads to positive frequency dependence, eroding diversity at the very genes responsible for recognition. As a result, the phenotypic diversity used for kin recognition has been widely assumed to be cues rather than signals of genetic identity. Diversity in identity cues is maintained by selection on other traits for reasons unrelated to recognition. A major problem with greenbeard-based models is that greenbeard recognition systems are uncommon among animals, which tend to learn kin phenotypes. We develop a simple model showing that learning a kin recognition template is sufficient to increase and maintain diversity in genetic traits used for kin recognition. Thus, our results suggest that phenotypes used for recognition may be true signals of genetic identity. As such, phenotypes are expected to evolve to facilitate recognition. Increased diversity in genetically-based recognition signals is also predicted to initiate a positive feedback loop between recognition efficiency and levels of cooperation. Finally, we discuss how the genetic architecture of recognition traits may influence kin discrimination abilities.