Scale as a lurking factor: incorporating scale-dependence in experimental ecology

Ecologists have recognized for decades the importance of spatial scale in ecological processes and patterns, as well as the complications scale poses for understanding ecological mechanisms. Here we highlight the opportunity attention to scale offers experimental ecology. Despite many advantages to considering scale, a review of the literature indicates that multiscale experimental studies are rare. Although much work has focused on scale as a primary factor (e.g. island size), we draw attention to scale as a ‘lurking’ variable: one which influences the relationship between two or more variables that are not usually understood to be scale-dependent. We highlight three basic observations from which scale-dependence arises: abundance increases with area, environmental conditions vary across space, and the effect of an organism on its environment is spatially limited. From these arise first-order scale-dependence, which relates an ecological variable of interest to a measure of scale. Combining first-order relationships together, we can produce second-order scale-dependencies, which occur when the relationship between two or more variables is mediated by scale. It is these relationships that are of particular interest, as they have the potential to confound experimental results. Most ecological experiments have incorporated scale either implicitly or not at all. We suggest that an explicit consideration of scale could help resolve some long-standing debates when scale is turned from a lurking variable into a working variable. Finally, we review and evaluate four different experimental sampling designs and corresponding statistical analyses that can be used to address the effects of scale in ecological experiments.