The presence of heritable variation is a prerequisite for evolution, but natural selection typically reduces genetic variation. Variation can be maintained in traits under selection through spatial or temporal variation in Þtness surfaces, frequency-dependent selection, or disruptive selection. We evaluated the maintenance of variation in the enantiomeric blend of pheromones employed by the bark beetle Ips pini (Say). In natural populations, we quantiÞed Þtness surfaces for mating success and

more »

... ogeny production. We investigated the effects of paternal pheromone blend on offspring survival by comparing the spatial scales at which pheromone blends and larval mortality agents vary. Males with extreme pheromone blends obtained up to 1.8 times as many mates who each laid equivalent numbers of eggs, producing strong disruptive selection on male pheromone blend. In combination with imperfect assortative mating that continually produces intermediate genotypes, this Þtness surface is sufÞcient to maintain variation in a heritable trait that is strongly linked to Þtness. The ultimate explanation for female preference is unknown but could be because of selection for reduced mortality from specialist predators that prefer common prey pheromone blends. Selection is most likely occurring at the scale of small resource patches within pine stands. Selection at coarser scales (pine stands) is unlikely because pheromone blends did not vary among pine stands. Selection at Þner scales (within logs) is unlikely because males of similar enantiomeric blends were not aggregated on logs, and male pheromone blend did not affect the spacing to neighboring galleries. This study documents a rare case of diversifying selection in natural populations.