Spatial and temporal scaling in benthic ecology

Using modern computer based search engines we were able to assemble, review, and document the rise of the concept of scale in benthic ecology. In the early literature the term appears to have been most commonly used to verbally qualify a result as being found at a particular spatial scale. However we document a shift from verbal expression to quantitative methods that explicitly incorporate scale into the design of the study. Examples include the development of nested multi-scale studies and experiments conducted over large spatio-temporal scales at the level of estuary or region. We also review novel methods for assessing scale effects in benthic ecosystems. First, gradient or correlative study designs have been developed in order to assess spatial and temporal heterogeneity in benthic ecosystems. These designs involve the careful selection of study site locations along gradients, or recommend nesting small-scale experiments within a broad-scale correlative framework. Secondly, we briefly assess the development of statistical analytical techniques to analyze spatial data and formal mathematical techniques for scaling functional relationships. We believe that the design and analytical advances identified in this review provide a framework for ecologists to address pressing large-scale questions.