Despite their apparent simplicity, arid environments can be quite heterogeneous. From small-scale variation in substrate and slope to large-scale geographic variation in solar input and productivity, drylands and deserts provide organisms with a tremendous range of ecological challenges (Schmidt-Nielsen 1964, Huggett 1995). Any single species is unable to meet all of these challenges equally well. A species will do better in some environments than others because evolution in heterogeneous environments is constrained by fitness tradeoffs. Such tradeoffs prevent the evolution of a versatile species, competitively superior to all other species across the entire spectrum of heterogeneity (Rosenzweig 1987). Although fitness tradeoffs may hinder species’ evolution in heterogeneous environments, they are a blessing for biodiversity. The source of biodiversity that we address in this chapter is the interplay of heterogeneity, tradeoffs, and density dependence. While we focus on species interactions at the local scale, our presentation includes a model that predicts changes in local diversity as a function of climate. The model’s predictions are based on changes in the nature of competition wrought by changes in productivity levels and climatic regimes. Cast in terms of evolutionary stable strategies (ESSs), the predictions refer to evolutionary as well as ecological patterns. A mechanism of coexistence consists of an axis of environmental heterogeneity together with an axis that indicates a tradeoff in the abilities of species to exploit different parts of the axis. In the absence of some kind of heterogeneity, there is only one environmental type, and whatever species is best adapted to it will competitively exclude others. In the absence of a tradeoff, one species could evolve competitive superiority over the full range of heterogeneity, again resulting in a monomorphic community. Consider some examples of mechanisms of species’ coexistence from dryland communities (Kotler and Brown 1988, Brown et al. 1994). For many taxa, spatial heterogeneity in predation risk is a consequence of the pattern of bushy and open areas common in drylands. In certain rodent communities, some species are able to exploit the relatively riskier open microhabitats by virtue of antipredator morphologies (Kotler 1984).
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