“…This theory predicts that levels of energy limit richness: a minimum amount of energy must be available in the environment for rarer species to maintain viable populations (Hutchinson, 1959; Brown, 1981; Wright, 1983). Qualitative observations also suggest that habitat heterogeneity may be an important determinant of diversity (Simpson, 1964; Owen, 1990; Pagel, May, & Collie, 1991; Blackburn & Gaston, 1996). Relatively few studies (Bohning‐Gaese, 1997; Kerr & Packer, 1997), however, have attempted to quantify the large‐ scale influence of habitat variability, probably due in part to the difficulty in measuring macrospatial heterogeneity.…”