What determines the spatial extent of landscape effects on species?

Landscape ecologists are often interested in measuring the effects of an environmental variable on a biological response; however, the strength and direction of effect depend on the size of the area within which the environmental variable is measured. Thus a central objective is to identify the optimal spatial extent within which to measure the environmental variable, i.e. the "scale of effect". Our objectives are (1) to provide a comprehensive summary of the hypotheses concerning what determines the scale of effect, (2) to provide predictions that can be tested in empirical studies, and (3) to show, with a review of the literature, that most of these predictions have so far been inadequately tested. We propose 14 predictions derived from five hypotheses explaining what determines the scale of effect, and review the literature (if any) supporting each prediction. These predictions involve five types of factors: (A) species traits, (B) landscape variables, (C) biological responses (e.g. abundance vs. occurrence), (D) indirect influences, and (E) regional context of the study. We identify methodological issues that hinder estimation of the scale of effect. Of the 14 predictions, only nine have been tested empirically and only five have received some empirical support. Most support is from simulation studies. Empirical evidence usually does not support predictions. The study of the spatial scale at which landscape variables influence biological outcomes is in its infancy. We provide directions for future research by clarifying predictions concerning the determinants of the scale of effect.