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The Maximum Entropy Theory of Ecology (METE) predicts the shapes of macroecological metrics in relatively static ecosystems, across spatial scales, taxonomic categories and habitats, using constraints imposed by static state variables. In disturbed ecosystems, however, with time‐varying state variables, its predictions often fail. We extend macroecological theory from static to dynamic by combining the MaxEnt inference procedure with explicit mechanisms governing disturbance. In the static limit, the resulting theory, DynaMETE, reduces to METE but also predicts a new scaling relationship among static state variables. Under disturbances, expressed as shifts in demographic, ontogenic growth or migration rates, DynaMETE predicts the time trajectories of the state variables as well as the time‐varying shapes of macroecological metrics such as the species abundance distribution and the distribution of metabolic rates over individuals. An iterative procedure for solving the dynamic theory is presented. Characteristic signatures of the deviation from static predictions of macroecological patterns are shown to result from different kinds of disturbance. By combining MaxEnt inference with explicit dynamical mechanisms of disturbance, DynaMETE is a candidate theory of macroecology for ecosystems responding to anthropogenic or natural disturbances.

 

Accumulating evidence suggests that ecological communities undergoing change in response to either anthropogenic or natural disturbances exhibit macroecological patterns that differ from those observed in similar types of communities in relatively undisturbed sites. In contrast to such cross‐site comparisons, however, there are few empirical studies of shifts over time in the shapes of macroecological patterns. Here, we provide a dramatic example of a plant community in which the species–area relationship and the species‐abundance distribution change markedly over a period of six years. These patterns increasingly deviate from the predictions of the maximum entropy theory of ecology (METE), which successfully predicts macroecological patterns in relatively static systems. The error in the species–area relationship prediction additionally correlates over time with increased stress measured as mortality minus recruitment, providing a link between demography and the failure of macroecological theory. Information on the dynamic state of an ecosystem inferred from snapshot measurements of macroecological community structure can potentially assist in identifying causes and consequences of disturbance and extending the domain of current theories and models to disturbed ecosystems.

 

Spatial patterns in ecology contain useful information about underlying mechanisms and processes. Although there are many summary statistics used to quantify these spatial patterns, there are far fewer models that directly link explicit ecological mechanisms to observed patterns easily derived from available data. We present a model of intraspecific spatial aggregation that quantitatively relates static spatial patterning to negative density dependence. Individuals are placed according to the colonization rule consistent with the Maximum Entropy Theory of Ecology (METE), and die with probability proportional to their abundance raised to a power α, a parameter indicating the degree of density dependence. This model can therefore be interpreted as a hybridization of MaxEnt and mechanism. Our model shows quantitatively and generally that increasing density dependence randomizes spatial patterning. α = 1 recovers the strongly aggregated METE distribution that is consistent with many ecosystems empirically, and as α → 2 our prediction approaches the binomial distribution consistent with random placement. For 1 < α < 2, our model predicts more aggregation than random placement but less than METE. We additionally relate our mechanistic parameter α to the statistical aggregation parameter k in the negative binomial distribution, giving it an ecological interpretation in the context of density dependence. We use our model to analyze two contrasting datasets, a 50 ha tropical forest and a 64 m 2 serpentine grassland plot. For each dataset, we infer α for individual species as well as a community α parameter. We find that α is generally larger in the tightly packed forest than the sparse grassland, and the degree of density dependence increases at smaller scales. These results are consistent with current understanding in both ecosystems, and we infer this underlying density dependence using only empirical spatial patterns. Our model can easily be applied to other datasets where spatially explicit data are available.