Different Spatial Scales of Natural Selection and Gene Flow: The Evolution of Behavioral Geographic Variation and Phenotypic Plasticity

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The environmental variables hypothesized to cause behavioral adaptation are distributed across a wide array of spatial scales, from local variation in factors such as food sources and territorial encounters to regional or continental variation in factors such as seasonality and the presence of predators. Geographic variation in behavior, the topic of this book, is just one of the potential evolutionary responses to environmental variation. Because behavioral divergence among populations generated by disparate natural selection can be counterbalanced by the homogenizing influence of gene flow, adaptive geographic variation can evolve only if the spatial scale of variation in natural selection is greater than the scale of gene flow (Endler 1977, Slatkin 1978). If geographic variation does not evolve because the spatial scale of selection is smaller than the scale of gene flow, populations may instead evolve adaptive phenotypic plasticity (Bradshaw 1965); the expression, by a single genotype, of different fitness-enhancing phenotypes in different environments. Because the same evolutionary processes operating on different spatial scales can generate behavioral geographic variation, behavioral phenotypic plasticity, or geographic variation in phenotypic plasticity, I devote this chapter to development of a hierarchical perspective tor studying environmental variation and behavioral evolution. This perspective emphasizes the shared evolutionary processes and research methodologies common to different levels of spatial variation, such as the balance of gene flow, natural selection, and genetic drift, the relationship between environmental patch size and local adaptation, and the effects of historical contingencies and genetic constraints on behavioral adaptation and phenotypic plasticity. In what follows, I review behavioral research in two unrelated taxa to illustrate the range of possible evolutionary responses to different patterns of environmental variation. First, I discuss different spatial scales of adaptation in the climbing behavior of deer mice (Peromyscus maniculatus) and provide a hierarchical analysis of the effects of natural selection, genetic drift, and gene flow. Second, I discuss diet-induced phenotypic plasticity in the feeding behavior of acridid grasshoppers (Melanoplus femurrubrum and M. sanguinipes) and the evolution of behavioral norms of reaction in response to local spatial and temporal variation in plant environments.