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Unified spatial scaling of species and their trophic interactions


Two largely independent bodies of scaling theory address the quantitative relationships between habitat area, species diversity and trophic interactions. Spatial theory within macroecology addresses how species richness scales with area in landscapes, while typically ignoring interspecific interactions1,2,3,4,5,6. Complexity theory within community ecology addresses how trophic links scale with species richness in food webs, while typically ignoring spatial considerations7,8,9,10,11,12. Recent studies suggest unifying these theories by demonstrating how spatial patterns influence food-web structure13,14,15,16 and vice versa17. Here, we follow this suggestion by developing and empirically testing a more unified scaling theory. On the basis of power-law species–area relationships, we develop link–area and non-power-law link–species models that accurately predict how trophic links scale with area and species richness of microcosms, lakes and streams from community to metacommunity levels. In contrast to previous models that assume that species richness alone determines the number of trophic links7,8, these models include the species' spatial distribution, and hence extend the domain of complexity theory to metacommunity scales. This generality and predictive success shows how complexity theory and spatial theory can be unified into a much more general theory addressing new domains of ecology.

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Figure 1: Scaling of links (filled circles) and species (open circles) in aquatic microcosms, lakes and streams with fitted regressions (solid lines), the unified scaling model (equation (5); dashed lines) and prior link–species models (dotted lines).


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We thank J. Harte and J. Green for comments on the manuscript. U.B. is supported by the German Academy of Naturalists Leopoldina and by funds from the German Federal Ministry of Education and Science. A.O. is supported by the National Center for Environmental Research (NCER) STAR Program, EPA. N.D.M is supported by the Cornell Telluride Association, Cornell Center for Applied Mathematics, and the US National Science Foundation Biocomplexity, Biological Databases and Informatics, Information Technology Research, and Interdisciplinary Graduate Education and Research Training programs.

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Correspondence to Ulrich Brose.

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Describes details of the unified spatial scaling model. (DOC 89 kb)

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Brose, U., Ostling, A., Harrison, K. et al. Unified spatial scaling of species and their trophic interactions. Nature 428, 167–171 (2004).

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