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The area-heterogeneity tradeoff (AHTO) implies that increasing environmental heterogeneity reduces the amount of effective area available for individual species (ref. 1 and Fig. 1). Theoretical models show that this tradeoff leads to a positive effect of heterogeneity on extinction rates and a unimodal effect on species richness (1–3). Our analysis of bird distribution in Catalonia using altitude range as a measure of heterogeneity (3) was consistent with both predictions. Main mechanisms of the AHTO. The population size of a species and its extinction risk are affected by the amount of suitable habitat available in the study area (the effective area). As heterogeneity increases, the proportion of some habitats might increase, but the average area of habitats decreases, leading, on average, …

Reply to Carnicer et al.: Environmental heterogeneity reduces breeding bird richness in Catalonia by increasing extinction rates