Non-equilibrium early-warning signals for critical transitions in ecological systems.

Complex systems can exhibit sudden transitions or regime shifts from one stable state to another, typically referred to as critical transitions. It becomes a great challenge to identify a robust warning sufficiently early that action can be taken to avert a regime shift. We employ landscape-flux theory from nonequilibrium statistical mechanics as a general framework to quantify the global stability of ecological systems and provide warning signals for critical transitions. We quantify the average flux as the nonequilibrium driving force and the dynamical origin of the nonequilibrium transition while the entropy production rate as the nonequilibrium thermodynamic cost and thermodynamic origin of the nonequilibrium transition. Average flux, entropy production, nonequilibrium free energy, and time irreversibility quantified by the difference in cross-correlation functions forward and backward in time can serve as early warning signals for critical transitions much earlier than other conventional predictors. We utilize a classical shallow lake model as an exemplar for our early warning prediction. Our proposed method is general and can be readily applied to assess the resilience of many other ecological systems. The early warning signals proposed here can potentially predict critical transitions earlier than established methods and perhaps even sufficiently early to avert catastrophic shifts.

Unifying deterministic and stochastic ecological dynamics via a landscape-flux approach.

The frequency distributions can characterize the population-potential landscape related to the stability of ecological states. We illustrate the practical utility of this approach by analyzing a forest-savanna model. Savanna and forest states coexist under certain conditions, consistent with past theoretical work and empirical observations. However, a grassland state, unseen in the corresponding deterministic model, emerges as an alternative quasi-stable state under fluctuations, providing a theoretical basis for the appearance of widespread grasslands in some empirical analyses. The ecological dynamics are determined by both the population-potential landscape gradient and the steady-state probability flux. The flux quantifies the net input/output to the ecological system and therefore the degree of nonequilibriumness. Landscape and flux together determine the transitions between stable states characterized by dominant paths and switching rates. The intrinsic potential landscape admits a Lyapunov function, which provides a quantitative measure of global stability. We find that the average flux, entropy production rate, and free energy have significant changes near bifurcations under both finite and zero fluctuation. These may provide both dynamical and thermodynamic origins of the bifurcations. We identified the variances in observed frequency time traces, fluctuations, and time irreversibility as kinematic measures for bifurcations. This framework opens the way to characterize ecological systems globally, to uncover how they change among states, and to quantify the emergence of quasi-stable states under stochastic fluctuations.

Tree clusters in savannas result from islands of soil moisture.

Tree clusters in savannas are commonly found in sizes that follow power laws with well-established exponents. We show that their size distributions could result from the space-time probabilistic structure of soil moisture, estimated over the range of rainfall observed in semiarid savannas; patterns of soil moisture display islands whose size, for moisture thresholds above the mean, follows power laws. These islands are the regions where trees are expected to exist and they have a fractal structure whose perimeter-area relationship is the same as observed in field data for the clustering of trees. When the impact of fire and herbivores is accounted for, as acting through the perimeter of the tree clusters, the power law of the soil moisture islands is transformed into a power law with the same exponents observed in the tree cluster data.

On the complex dynamics of savanna landscapes.

Simple mathematical models can exhibit rich and complex behaviors. Prototypical examples of these drawn from biology and other disciplines have provided insights that extend well beyond the situations that inspired them. Here, we explore a set of simple, yet realistic, models for savanna-forest vegetation dynamics based on minimal ecological assumptions. These models are aimed at understanding how vegetation interacts with both climate (a primary global determinant of vegetation structure) and feedbacks with chronic disturbances from fire. The model includes three plant functional types-grasses, savanna trees, and forest trees. Grass and (when they allow grass to persist in their subcanopy) savanna trees promote the spread of fires, which in turn, demographically limit trees. The model exhibits a spectacular range of behaviors. In addition to bistability, analysis reveals (i) that diverse cyclic behaviors (including limit and homo- and heteroclinic cycles) occur for broad ranges of parameter space, (ii) that large shifts in landscape structure can result from endogenous dynamics and not just from external drivers or from noise, and (iii) that introducing noise into this system induces resonant and inverse resonant phenomena, some of which have never been previously observed in ecological models. Ecologically, these results raise questions about how to evaluate complicated dynamics with data. Mathematically, they lead to classes of behaviors that are likely to occur in other models with similar structure.