The problem of scale has been a critical impediment to incorporating important fine-scale processes into global ecosystem models. Our knowledge of fine-scale physiological and ecological processes comes from a variety of measurements, ranging from forest plot inventories to remote sensing, made at spatial resolutions considerably smaller than the large scale at which global ecosystem models are defined. In this paper, we describe a new individual-based, terrestrial biosphere model, which we label the ecosystem demography model (ED). We then introduce a general method for scaling stochastic individual-based models of ecosystem dynamics (gap models) such as ED to large scales. The method accounts for the fine-scale spatial heterogeneity within an ecosystem caused by stochastic disturbance events, operating at scales down to individual canopy-tree-sized gaps. By conditioning appropriately on the occurrence of these events, we derive a size- and age-structured (SAS) approximation for the first moment of the stochastic ecosystem model. With this approximation, it is possible to make predictions about the large scales of interest from a description of the fine-scale physiological and population-dynamic processes without simulating the fate of every plant individually. We use the SAS approximation to implement our individual-based biosphere model over South America from 15° N to 15° S, showing that the SAS equations are accurate across a range of environmental conditions and resulting ecosystem types. We then compare the predictions of the biosphere model to regional data and to intensive data at specific sites. Analysis of the model at these sites illustrates the importance of fine-scale heterogeneity in governing large-scale ecosystem function, showing how population and community-level processes influence ecosystem composition and structure, patterns of aboveground carbon accumulation, and net ecosystem production.
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