Phase-separation physics underlies new theory for the resilience of patchy ecosystems.

Spatial self-organization of ecosystems into large-scale (from micron to meters) patterns is an important phenomenon in ecology, enabling organisms to cope with harsh environmental conditions and buffering ecosystem degradation. Scale-dependent feedbacks provide the predominant conceptual framework for self-organized spatial patterns, explaining regular patterns observed in, e.g., arid ecosystems or mussel beds. Here, we highlight an alternative mechanism for self-organized patterns, based on the aggregation of a biotic or abiotic species, such as herbivores, sediment, or nutrients. Using a generalized mathematical model, we demonstrate that ecosystems with aggregation-driven patterns have fundamentally different dynamics and resilience properties than ecosystems with patterns that formed through scale-dependent feedbacks. Building on the physics theory for phase-separation dynamics, we show that patchy ecosystems with aggregation patterns are more vulnerable than systems with patterns formed through scale-dependent feedbacks, especially at small spatial scales. This is because local disturbances can trigger large-scale redistribution of resources, amplifying local degradation. Finally, we show that insights from physics, by providing mechanistic understanding of the initiation of aggregation patterns and their tendency to coarsen, provide a new indicator framework to signal proximity to ecological tipping points and subsequent ecosystem degradation for this class of patchy ecosystems.

Evasion of tipping in complex systems through spatial pattern formation.

The concept of tipping points and critical transitions helps inform our understanding of the catastrophic effects that global change may have on ecosystems, Earth system components, and the whole Earth system. The search for early warning indicators is ongoing, and spatial self-organization has been interpreted as one such signal. Here, we review how spatial self-organization can aid complex systems to evade tipping points and can therefore be a signal of resilience instead. Evading tipping points through various pathways of spatial pattern formation may be relevant for many ecosystems and Earth system components that hitherto have been identified as tipping prone, including for the entire Earth system. We propose a systematic analysis that may reveal the broad range of conditions under which tipping is evaded and resilience emerges.

The effect of climate change on the resilience of ecosystems with adaptive spatial pattern formation.

In a rapidly changing world, quantifying ecosystem resilience is an important challenge. Historically, resilience has been defined via models that do not take spatial effects into account. These systems can only adapt via uniform adjustments. In reality, however, the response is not necessarily uniform, and can lead to the formation of (self-organised) spatial patterns - typically localised vegetation patches. Classical measures of resilience cannot capture the emerging dynamics in spatially self-organised systems, including transitions between patterned states that have limited impact on ecosystem structure and productivity. We present a framework of interlinked phase portraits that appropriately quantifies the resilience of patterned states, which depends on the number of patches, the distances between them and environmental conditions. We show how classical resilience concepts fail to distinguish between small and large pattern transitions, and find that the variance in interpatch distances provides a suitable indicator for the type of imminent transition. Subsequently, we describe the dependency of ecosystem degradation based on the rate of climatic change: slow change leads to sporadic, large transitions, whereas fast change causes a rapid sequence of smaller transitions. Finally, we discuss how pre-emptive removal of patches can minimise productivity losses during pattern transitions, constituting a viable conservation strategy.

Grazing Away the Resilience of Patterned Ecosystems.

Ecosystems' responses to changing environmental conditions can be modulated by spatial self-organization. A prominent example of this can be found in drylands, where formation of vegetation patterns attenuates the magnitude of degradation events in response to decreasing rainfall. In model studies, the pattern wavelength responds to changing conditions, which is reflected by a rather gradual decline in biomass in response to decreasing rainfall. Although these models are spatially explicit, they have adopted a mean-field approach to grazing. By taking into account spatial variability when modeling grazing, we find that (over)grazing can lead to a dramatic shift in biomass, so that degradation occurs at rainfall rates that would otherwise still maintain a relatively productive ecosystem. Moreover, grazing increases the resilience of degraded ecosystem states. Consequently, restoration of degraded ecosystems could benefit from the introduction of temporary small-scale exclosures to escape from the basin of attraction of degraded states.

Multistability of model and real dryland ecosystems through spatial self-organization.

Spatial self-organization of dryland vegetation constitutes one of the most promising indicators for an ecosystem's proximity to desertification. This insight is based on studies of reaction-diffusion models that reproduce visual characteristics of vegetation patterns observed on aerial photographs. However, until now, the development of reliable early warning systems has been hampered by the lack of more in-depth comparisons between model predictions and real ecosystem patterns. In this paper, we combined topographical data, (remotely sensed) optical data, and in situ biomass measurements from two sites in Somalia to generate a multilevel description of dryland vegetation patterns. We performed an in-depth comparison between these observed vegetation pattern characteristics and predictions made by the extended-Klausmeier model for dryland vegetation patterning. Consistent with model predictions, we found that for a given topography, there is multistability of ecosystem states with different pattern wavenumbers. Furthermore, observations corroborated model predictions regarding the relationships between pattern wavenumber, total biomass, and maximum biomass. In contrast, model predictions regarding the role of slope angles were not corroborated by the empirical data, suggesting that inclusion of small-scale topographical heterogeneity is a promising avenue for future model development. Our findings suggest that patterned dryland ecosystems may be more resilient to environmental change than previously anticipated, but this enhanced resilience crucially depends on the adaptive capacity of vegetation patterns.

Pulse dynamics in reaction-diffusion equations with strong spatially localized impurities.

In this article, a general geometric singular perturbation framework is developed to study the impact of strong, spatially localized, nonlinear impurities on the existence, stability and bifurcations of localized structures in systems of linear reaction-diffusion equations. By taking advantage of the multiple-scale nature of the problem, we derive algebraic conditions determining the existence and stability of pinned single- and multi-pulse solutions. Our methods enable us to explicitly control the spectrum associated with a (multi-)pulse solution. In the scalar case, we show how eigenvalues may move in and out of the essential spectrum and that Hopf bifurcations cannot occur. By contrast, even a pinned 1-pulse solution can undergo a Hopf bifurcation in a two-component system of linear reaction-diffusion equations with (only) one impurity.This article is part of the theme issue 'Stability of nonlinear waves and patterns and related topics'.

Phase separation explains a new class of self-organized spatial patterns in ecological systems.

The origin of regular spatial patterns in ecological systems has long fascinated researchers. Turing's activator-inhibitor principle is considered the central paradigm to explain such patterns. According to this principle, local activation combined with long-range inhibition of growth and survival is an essential prerequisite for pattern formation. Here, we show that the physical principle of phase separation, solely based on density-dependent movement by organisms, represents an alternative class of self-organized pattern formation in ecology. Using experiments with self-organizing mussel beds, we derive an empirical relation between the speed of animal movement and local animal density. By incorporating this relation in a partial differential equation, we demonstrate that this model corresponds mathematically to the well-known Cahn-Hilliard equation for phase separation in physics. Finally, we show that the predicted patterns match those found both in field observations and in our experiments. Our results reveal a principle for ecological self-organization, where phase separation rather than activation and inhibition processes drives spatial pattern formation.