From Hutchinsonian ratios to spatial scaling theory: the interplay among limiting similarity, body size and landscape structure

Abstract

Spatial scaling theory (SST) relates the physical structure of the environment to species coexistence and community assembly. Although SST is a recognized theory in ecology, few studies have evaluated its predictions, producing contradictory results and frequently failing to meet its assumptions. In addition, the ‘risk predictions’ of SST regarding an increase in species similarity with body size and the dependence of this pattern on the landscape and food fractal dimensions have not been evaluated. This study attempted to account for previous limitations, analyzing these predictions in coleopteran guilds that inhabit 18 temporary ponds. This metacommunity covers a large gradient of environmental variables, including food density, the landscape fractal dimension, the food fractal dimensions and other indicators of pond heterogeneity. Average similarity in carnivorous and herbivorous body sizes systematically increased with guild richness, fulfilling classical predictions of niche theory. Species similarity was associated with body size, but the association reverts from negative to positive as the landscape fractal dimension and heterogeneity increases, a pattern further supported by null model analyses. Several nonexclusive mechanisms may account for this pattern: 1) the body size‐dependent landscape perception, through which small animals detect more heterogeneity than larger animals; 2) the reaching of landscape limits by larger species, which prevents them from accessing novel largest clusters; 3) the large differences between the landscape and food fractal dimensions; and 4) the homogenization of the landscape when an integer fractal dimension is reached. These mechanisms may dictate that smaller organisms are more able to capitalize on heterogeneity or available resources than larger organisms, thus promoting increased similarity among smaller species. The presented results support the connection between landscape spatial structure and biodiversity and a mechanistic understanding of this connection from the SST.