Maximum entropy and the state-variable approach to macroecology.

The biodiversity scaling metrics widely studied in macroecology include the species-area relationship (SAR), the scale-dependent species-abundance distribution (SAD), the distribution of masses or metabolic energies of individuals within and across species, the abundance-energy or abundance-mass relationship across species, and the species-level occupancy distributions across space. We propose a theoretical framework for predicting the scaling forms of these and other metrics based on the state-variable concept and an analytical method derived from information theory. In statistical physics, a method of inference based on information entropy results in a complete macro-scale description of classical thermodynamic systems in terms of the state variables volume, temperature, and number of molecules. In analogy, we take the state variables of an ecosystem to be its total area, the total number of species within any specified taxonomic group in that area, the total number of individuals across those species, and the summed metabolic energy rate for all those individuals. In terms solely of ratios of those state variables, and without invoking any specific ecological mechanisms, we show that realistic functional forms for the macroecological metrics listed above are inferred based on information entropy. The Fisher log series SAD emerges naturally from the theory. The SAR is predicted to have negative curvature on a log-log plot, but as the ratio of the number of species to the number of individuals decreases, the SAR becomes better and better approximated by a power law, with the predicted slope z in the range of 0.14-0.20. Using the 3/4 power mass-metabolism scaling relation to relate energy requirements and measured body sizes, the Damuth scaling rule relating mass and abundance is also predicted by the theory. We argue that the predicted forms of the macroecological metrics are in reasonable agreement with the patterns observed from plant census data across habitats and spatial scales. While this is encouraging, given the absence of adjustable fitting parameters in the theory, we further argue that even small discrepancies between data and predictions can help identify ecological mechanisms that influence macroecological patterns.