The Problem of Ecological Scaling in Spatially Complex, Nonequilibrium Ecological Systems

In the previous chapter we reviewed the challenges posed by spatial complexity and temporal disequilibrium to efforts to understand and predict the structure and dynamics of ecological systems. The central theme was that spatial variability in the environment and population processes fundamentally alters the interactions between species and their environments, largely invalidating the predictions of ideal models of community structure and population processes. In addition, we argued that temporal variability enormously amplifies the challenge of prediction, by altering and reversing species–species and species–environment relationships over time. Typically these fluctuations do not occur globally across space in synchrony; rather change in time is spatially dependent on location in the environment, and thus interacts in highly complex and nonlinear ways with spatial heterogeneity in influencing ecological processes. Given these challenges, we proposed focusing on the interactions between species and their immediate environments in the context of current and past conditions. However, given critical sensitivity of ecological processes to spatial and temporal factors, it is also necessary to consider their action within the context of a broader landscape of conditions, constraints and drivers. This therefore seems a catch-22, with fine-scale understanding of process required at the scale where ecological entities (e.g. organisms) directly interact with each other and their environments, and also integration of these fine-scale processes across complex and temporally varying broad-scale environments. This challenge fundamentally relates to scale and scaling ecological processes. One of the challenges that oft plagues efforts to discuss and integrate concepts related to multiple-scale analysis is inconsistency and contradiction of terminology and the resulting confusion of meanings (Allen and Hoekstra 1992; Schneider 1994; Peterson and Parker 1998). In this chapter we discuss concepts of scale and