Altruism presents a challenge to evolutionary theory because selection should favor selfish over caring strategies. Greenbeard altruism resolves this paradox by allowing cooperators to identify individuals carrying similar alleles producing a form of genic selection. In sideblotched lizards, genetically similar but unrelated blue male morphs settle on adjacent territories and cooperate. Here we show that payoffs of cooperation depend on asymmetric costs of orange neighbors. One blue male experiences low fitness and buffers his unrelated partner from aggressive orange males despite the potential benefits of defection. We show that recognition behavior is highly heritable in nature, and we map genetic factors underlying color and self-recognition behavior of genetic similarity in both sexes. Recognition and cooperation arise from genome-wide factors based on our mapping study of the location of genes responsible for self-recognition behavior, recognition of blue color, and the color locus. Our results provide an example of greenbeard interactions in a vertebrate that are typified by cycles of greenbeard mutualism interspersed with phases of transient true altruism. Such cycles provide a mechanism encouraging the origin and stability of true altruism.alternative strategies ͉ linkage map ͉ frequency-dependent selection ͉ evolutionarily stable strategy ͉ cooperation T he evolutionary stability of cooperative and altruistic behaviors requires that interindividual benefits be protected from competition, cheating, and defection (1-4). Without such safeguards, selfish strategies will eliminate altruistic strategies (5, 6). Hamilton (5) theorized that true altruism might evolve if a supergene simultaneously affected a signal and recognition of the signal and that signal recognition elicited social acts costly to donors but beneficial to recipients. Dawkins (6) coined Hamilton's social supergene a greenbeard in a hypothetical example of altruists that sported a green beard distinct in color from other beards sported by nonaltruists. Despite studies consistent with greenbeard altruism (7-12), few provide definitive evidence for greenbeard altruism.The annual side-blotched lizard, Uta stansburiana, exhibits six color genotypes (13, 14) (oo, bo, yo, bb, by, and yy), which serve as markers for three male strategies (15). Orange males (oo, bo, and yo) usurp territory. Blue males (bb) mate-guard. Yellow males (by and yy) are sneakers. Male competition drives rock-paper-scissors (RPS) cycles of three strategies: sneakers beat usurpers, mate guarders defeat sneakers, and usurpers prevail over mate guarders (13,(15)(16)(17)(18)(19). Previously, we showed that males with b alleles prefer to settle near non-kin but genetically similar bb males and cooperate in territory defense (15). Hereafter, we refer to bb males with genetically similar neighbors (based on allele sharing at nine microsatellite loci) as ''dyadic bb pairs'' (15). We contrast dyadic bb males with ''loner bb males'' that may have bb neighbors, but none are genetically...