Phenotypic plasticity and the ecotypic differentiation of aggressive behavior in threespine stickleback

The threespine stickleback fish, Gasterosteus aculeatus, has undergone a remarkable postglacial adaptive radiation in which an ancient oceanic ancestor has given rise to uncountable freshwater populations. The radiation is characterized by repeated, independent evolution of similar derived phenotypes under similar environmental conditions. A common pattern of divergence is caused by differences in habitat that favor morphological and behavioral features that enhance efficiency of feeding on plankton (limnetic ecotypes) vs. those that enhance efficiency of feeding on benthic invertebrates (benthic ecotypes). These two ecotypes exhibit consistently different patterns of courtship and of foraging and cannibalistic behavior (divergent behavioral syndromes). Here, we demonstrate that there also exist differences in aggression toward conspecifics that are likely to be characteristic of the ecotypes. We report differences in patterns of aggression toward rivals between the ecotypes and offer evidence of differences in the patterns of phenotypic plasticity (norms of reaction) for these traits across population types, and of differences in the incorporation of aggressive elements of behavior in courtship. These data support an earlier suggestion that differences in aggressive tendencies could have facilitated assortative mating between the four benthic-limnetic species pairs found in British Columbia lakes, and they demonstrate the need to evaluate divergent behavioral phenotypes in this radiation as phenotypic norms of reaction rather than as fixed traits.