For Harn— iltonian loci, relatedness (r) to the offspring of potential helped individuals is calculated at the level of the entire genome. The inclusive fitness of. In contrast, at each greenbeard locus, there are four different possible greenbeard alleles (each producing a hypothetical unique phenotypic cue) that can. four genetically unlinked loci that are required for light -induced caroteno- genesis ; carAB, carC, carD and carR. Two classes of phenotype were identified: those.