The genetics, evolution, and maintenance of a biological rock-paper-scissors game

Side-blotched lizards (<i>Uta stansburiana</i>) play a biological rock-paper-scissors game in which three differently colored male morphs utilize alternative mating strategies. We identified the genetic basis of this polymorphism, which was previously posited to arise from three alleles at one locus. Orange usurper and blue mate-guarder morphs are associated with two divergent haplotypes in the regulatory region of the sepiapterin reductase gene, but yellow sneaker morphs appear to arise through phenotypic plasticity from the same genetic background as blue morphs. Our simulations show that rock-paper-scissors dynamics can better maintain a polymorphism with a genetic system of two alleles plus plasticity than with a three-allele system. This form of balancing selection that combines genetic determination with phenotypic plasticity expands the possibilities for how stable polymorphisms arise in nature.