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Ehud Meron

Publications and source records attributed to Ehud Meron.

At least 19 recordsLinked to original sources

Vegetation Patterning Can Both Impede and Trigger Critical Transitions from Savanna to Grassland

Tree-grass coexistence is a defining feature of savanna ecosystems, which play an important role in supporting biodiversity and human populations worldwide. While recent advances have clarified many of the underlying processes, how these mechanisms interact to shape ecosystem dynamics under environmental stress is not yet understood. Here, we present and analyze a minimalistic spatially extended model of tree-grass dynamics in dry savannas. We incorporate tree facilitation of grasses through shading and grass competing with trees for water, both varying with tree life stage. Our model shows that these mechanisms lead to grass-tree coexistence and bistability between savanna and grassland states. Moreover, the model predicts vegetation patterns consisting of trees and grasses, particularly under harsh environmental conditions, which can persist in situations where a non-spatial version of the model predicts ecosystem collapse from savanna to grassland instead (a phenomenon called ``Turing-evades-tipping''). Additionally, we identify a novel ``Turing-triggers-tipping'' mechanism, where unstable pattern formation drives tipping events that are overlooked when spatial dynamics are not included. These transient patterns act as early warning signals for ecosystem transitions, offering a critical window for intervention. Further theoretical and empirical research is needed to determine when spatial patterns prevent tipping or drive collapse.

q-bio.PE

Traveling vegetation-herbivore waves can sustain ecosystems threatened by droughts and population growth

Dryland vegetation can survive water stress by forming spatial patterns but is often subjected to herbivory as an additional stress that puts it at risk of desertification. Understanding the mutual relationships between vegetation patterning and herbivory is crucial for securing food production in drylands, which constitute the majority of rangelands worldwide. Here, we introduce a novel vegetation-herbivore model that captures pattern-forming feedbacks associated with water and herbivory stress and a behavioral aspect of herbivores representing an exploitation strategy.Applying numerical continuation methods, we analyze the bifurcation structure of uniform and patterned vegetation-herbivore solutions, and use direct numerical simulations to study various forms of collective herbivore dynamics. We find that herbivory stress can induce traveling vegetation-herbivore waves and uncover the ecological mechanism that drives their formation. In the traveling-wave state, the herbivore distribution is asymmetric with higher density on one side of each vegetation patch. At low precipitation values their distribution is localized, while at high precipitation the herbivores are spread over the entire landscape. Importantly, their asymmetric distribution results in uneven herbivory stress, strong on one side of each vegetation patch and weak on the opposing side - weaker than the stress exerted in spatially uniform herbivore distribution. Consequently, the formation of traveling waves results in increased sustainability to herbivory stress. We conclude that vegetation-herbivore traveling waves may play an essential role in sustaining herbivore populations under conditions of combined water and herbivory stress, thereby contributing to food security in endangered regions threatened by droughts and population growth.

q-bio.PE

The existence of localized vegetation patterns in a systematically reduced model for dryland vegetation

In this paper we consider the 2-component reaction-diffusion model that was recently obtained by a systematic reduction of the 3-component Gilad et al. model for dryland ecosystem dynamics. The nonlinear structure of this model is more involved than other more conceptual models, such as the extended Klausmeier model, and the analysis a priori is more complicated. However, the present model has a strong advantage over these more conceptual models in that it can be more directly linked to ecological mechanisms and observations. Moreover, we find that the model exhibits a richness of analytically tractable patterns that exceeds that of Klausmeier-type models. Our study focuses on the 4-dimensional dynamical system associated with the reaction-diffusion model by considering traveling waves in 1 spatial dimension. We use the methods of geometric singular perturbation theory to establish the existence of a multitude of heteroclinic/homoclinic/periodic orbits that `jump' between (normally hyperbolic) slow manifolds, representing various kinds of localized vegetation patterns. The basic 1-front invasion patterns and 2-front spot/gap patterns that form the starting point of our analysis have a direct ecological interpretation and appear naturally in simulations of the model. By exploiting the rich nonlinear structure of the model, we construct many multi-front patterns that are novel, both from the ecological and the mathematical point of view. In fact, we argue that these orbits/patterns are not specific for the model considered here, but will also occur in a much more general (singularly perturbed reaction-diffusion) setting. We conclude with a discussion of the ecological and mathematical implications of our findings.

math.DS

Spatial asymmetries of resonant oscillations in periodically forced heterogeneous media

Spatially localized oscillations in periodically forced systems are intriguing phenomena. They may occur in spatially homogeneous media (oscillons), but quite often emerge in heterogeneous media, such as the auditory system, where localized oscillations are believed to play an important role in frequency discrimination of incoming sound waves. In this paper, we use an amplitude-equation approach to study the spatial profile of the oscillations and the factors that affect it. More specifically, we use a variant of the forced complex Ginzburg-Landau (FCGL) equation to describes an oscillatory system below the Hopf bifurcation with space-dependent Hopf frequency, subject to both parametric and additive forcing. We show that spatial heterogeneity, combined with bistability of system states, results in spatial asymmetry of the localized oscillations. We further identify parameters that control that asymmetry, and characterize the spatial profile of the oscillations in terms of maximum amplitude, location, width and asymmetry. Our results bear qualitative similarities to empirical observation trends that have found in the auditory system.

nlin.PS

Spatial heterogeneity may form an inverse camel shape Arnol'd tongue in parametrically forced oscillations

Frequency locking in forced oscillatory systems typically occurs in 'V'-shaped domains in the plane spanned by the forcing frequency and amplitude, the so-called Arnol'd tongues. Here, we show that if the medium is spatially extended and monotonically heterogeneous, e.g., through spatially-dependent natural frequency, the resonance tongues can also display 'U' and 'W' shapes; to the latter, we refer as "inverse camel" shape. We study the generic forced complex Ginzburg-Landau equation for damped oscillations under parametric forcing and, using linear stability analysis and numerical simulations, uncover the mechanisms that lead to these distinct shapes. Additionally, we study the effects of discretization, by exploring frequency locking of oscillators chains. Since we study a normal-form equation, the results are model-independent near the onset of oscillations, and, therefore, applicable to inherently heterogeneous systems in general, such as the cochlea. The results are also applicable to controlling technological performances in various contexts, such as arrays of mechanical resonators, catalytic surface reactions, and nonlinear optics.

nlin.PS

Molding the asymmetry of localized frequency-locking waves by a generalized forcing and implications to the inner ear

Frequency locking to an external forcing frequency is a {well} known phenomenon. In the auditory system, it results in a localized traveling wave, the shape of which is essential for efficient discrimination between incoming frequencies. An amplitude equation approach is used to show that the shape of the localized traveling wave depends crucially on the relative strength of additive vs. parametric forcing components; the stronger the parametric forcing the more asymmetric the response profile and the sharper the traveling-wave front. The analysis captures the empirically observed regions of linear and nonlinear responses and highlights the significance of parametric forcing mechanisms in shaping the resonant response in the inner ear.

nlin.PS

Desertification by Front Propagation?

Understanding how desertification takes place in different ecosystems is an important step in attempting to forecast and prevent such transitions. Dryland ecosystems often exhibit patchy vegetation, which has been shown to be an important factor on the possible regime shifts that occur in arid regions in several model studies. In particular, both gradual shifts that occur by front propagation, and abrupt shifts where patches of vegetation vanish at once, are a possibility in dryland ecosystems due to their emergent spatial heterogeneity. However, recent theoretical work has suggested that the final step of desertification - the transition from spotted vegetation to bare soil - occurs only as an abrupt shift, but the generality of this result, and its underlying origin, remain unclear. We investigate two models that detail the dynamics of dryland vegetation using a markedly different functional structure, and find that in both models the final step of desertification can only be abrupt. Using a careful numerical analysis, we show that this behavior is associated with the disappearance of confined spot-pattern domains as stationary states, and identify the mathematical origin of this behavior. Our findings show that a gradual desertification to bare soil due to a front propagation process can not occur in these and similar models, and opens the question of whether these dynamics can take place in nature.

nlin.PS

Gradual Regime Shifts in Fairy Circles

Large responses of ecosystems to small changes in the conditions--regime shifts--are of great interest and importance. In spatially extended ecosystems, these shifts may be local or global. Using empirical data and mathematical modeling, we investigated the dynamics of the Namibian fairy circle ecosystem as a case study of regime shifts in a pattern-forming ecosystem. Our results provide new support, based on the dynamics of the ecosystem, for the view of fairy circles as a self-organization phenomenon driven by water-vegetation interactions. The study further suggests that fairy circle birth and death processes correspond to spatially confined transitions between alternative stable states. Cascades of such transitions, possible in various pattern-forming systems, result in gradual rather than abrupt regime shifts.

nlin.PS

Localized states qualitatively change the response of ecosystems to varying conditions and local disturbances

The response of dynamical systems to varying conditions and disturbances is a fundamental aspect of their analysis. In spatially extended systems, particularly in pattern-forming systems, there are many possible responses, including critical transitions, gradual transitions and locally confined responses. Here, we use the context of vegetation dynamics in drylands in order to study the response of pattern-forming ecosystems to oscillating precipitation and local disturbances. We focus on two precipitation ranges, a bistability range of bare soil with a patterned vegetation state, and a bistability range of uniform vegetation with a patterned vegetation state. In these ranges, there are many different stable states, which allow for both abrupt and gradual transitions between the system states to occur. We find that large amplitude oscillations of the precipitation rate can lead to a collapse of the vegetation in one range, while in the other range, they result in the convergence to a patterned state with a preferred wavelength. In addition, we show that a series of local disturbances results in the collapse of the vegetation in one range, while it drives the system toward fluctuations around a finite average biomass in the other range. Moreover, it is shown that under certain conditions, local disturbances can actually increase the overall vegetation density. These significant differences in the system response are attributed to the existence of localized states in one of the bistability ranges.

nlin.PS

Regime shifts in models of dryland vegetation

Drylands are pattern-forming systems showing self-organized vegetation patchiness, multiplicity of stable states and fronts separating domains of alternative stable states. Pattern dynamics, induced by droughts or disturbances, can result in desertification shifts from patterned vegetation to bare soil. Pattern-formation theory suggests various scenarios for such dynamics; an abrupt global shift involving a fast collapse to bare soil, a gradual global shift involving the expansion and coalescence of bare-soil domains, and an incipient shift to a hybrid state consisting of stationary bare-soil domains in an otherwise periodic pattern. Using models of dryland vegetation we address the question which of these scenarios can be realized. We found that the models can be split into two groups: models that exhibit multiplicity of periodic-pattern and bare-soil states, and models that exhibit, in addition, multiplicity of hybrid states. Furthermore, in all models we could not identify parameter regimes in which bare-soil domains expand into vegetated domains. The significance of these findings is that while models belonging to the first group can only exhibit abrupt shifts, model belonging to the second group can also exhibit gradual and incipient shifts. A discussion of open problems concludes the paper.

nlin.PS

A pattern-forming instability co-driven by distinct mechanisms increases pattern diversity

We use the context of dryland vegetation to study a general problem of complex pattern forming systems - multiple pattern-forming instabilities that are driven by distinct mechanisms but share the same spectral properties. We find that the co-occurrence of such instabilities results in the growth of a single mode rather than two interacting modes. The interplay between the two mechanisms, which promote or counteract each other, compensates for the simpler dynamics of a single mode by inducing higher pattern diversity. Possible implications to biodiversity of ecosystems are discussed.

nlin.PS

Biogenic crust dynamics on sand dunes

Sand dunes are often covered by vegetation and biogenic crusts. Despite their significant role in dune stabilization, biogenic crusts have rarely been considered in studies of dune dynamics. Using a simple model, we study the existence and stability ranges of different dune-cover states along gradients of rainfall and wind power. Two ranges of alternative stable states are identified: fixed crusted dunes and fixed vegetated dunes at low wind power, and fixed vegetated dunes and active dunes at high wind power. These results suggest a cross-over between two different forms of desertification.

physics.bio-ph

Resonance tongues and patterns in periodically forced reaction-diffusion systems

Various resonant and near-resonant patterns form in a light-sensitive Belousov-Zhabotinsky (BZ) reaction in response to a spatially-homogeneous time-periodic perturbation with light. The regions (tongues) in the forcing frequency and forcing amplitude parameter plane where resonant patterns form are identified through analysis of the temporal response of the patterns. Resonant and near-resonant responses are distinguished. The unforced BZ reaction shows both spatially-uniform oscillations and rotating spiral waves, while the forced system shows patterns such as standing-wave labyrinths and rotating spiral waves. The patterns depend on the amplitude and frequency of the perturbation, and also on whether the system responds to the forcing near the uniform oscillation frequency or the spiral wave frequency. Numerical simulations of a forced FitzHugh-Nagumo reaction-diffusion model show both resonant and near-resonant patterns similar to the BZ chemical system.

nlin.PS

Phase Dynamics of Nearly Stationary Patterns in Activator-Inhibitor Systems

The slow dynamics of nearly stationary patterns in a FitzHugh-Nagumo model are studied using a phase dynamics approach. A Cross-Newell phase equation describing slow and weak modulations of periodic stationary solutions is derived. The derivation applies to the bistable, excitable, and the Turing unstable regimes. In the bistable case stability thresholds are obtained for the Eckhaus and the zigzag instabilities and for the transition to traveling waves. Neutral stability curves demonstrate the destabilization of stationary planar patterns at low wavenumbers to zigzag and traveling modes. Numerical solutions of the model system support the theoretical findings.

nlin.PS

Stratified spatiotemporal chaos in anisotropic reaction-diffusion systems

Numerical simulations of two dimensional pattern formation in an anisotropic bistable reaction-diffusion medium reveal a new dynamical state, stratified spatiotemporal chaos, characterized by strong correlations along one of the principal axes. Equations that describe the dependence of front motion on the angle illustrate the mechanism leading to stratified chaos.

patt-sol

Multi-Phase Patterns in Periodically Forced Oscillatory Systems

Periodic forcing of an oscillatory system produces frequency locking bands within which the system frequency is rationally related to the forcing frequency. We study extended oscillatory systems that respond to uniform periodic forcing at one quarter of the forcing frequency (the 4:1 resonance). These systems possess four coexisting stable states, corresponding to uniform oscillations with successive phase shifts of $π/2$. Using an amplitude equation approach near a Hopf bifurcation to uniform oscillations, we study front solutions connecting different phase states. These solutions divide into two groups: $π$-fronts separating states with a phase shift of $π$ and $π/2$-fronts separating states with a phase shift of $π/2$. We find a new type of front instability where a stationary $π$-front ``decomposes'' into a pair of traveling $π/2$-fronts as the forcing strength is decreased. The instability is degenerate for an amplitude equation with cubic nonlinearities. At the instability point a continuous family of pair solutions exists, consisting of $π/2$-fronts separated by distances ranging from zero to infinity. Quintic nonlinearities lift the degeneracy at the instability point but do not change the basic nature of the instability. We conjecture the existence of similar instabilities in higher 2n:1 resonances (n=3,4,..) where stationary $π$-fronts decompose into n traveling $π/n$-fronts. The instabilities designate transitions from stationary two-phase patterns to traveling 2n-phase patterns. As an example, we demonstrate with a numerical solution the collapse of a four-phase spiral wave into a stationary two-phase pattern as the forcing strength within the 4:1 resonance is increased.

patt-sol

A Phase Front Instability in Periodically Forced Oscillatory Systems

Multiplicity of phase states within frequency locked bands in periodically forced oscillatory systems may give rise to front structures separating states with different phases. A new front instability is found within bands where $ω_{forcing}/ω_{system}=2n$ ($n>1$). Stationary fronts shifting the oscillation phase by $π$ lose stability below a critical forcing strength and decompose into $n$ traveling fronts each shifting the phase by $π/n$. The instability designates a transition from stationary two-phase patterns to traveling $n$-phase patterns.

patt-sol

Propagation Failure in Excitable Media

We study a mechanism of pulse propagation failure in excitable media where stable traveling pulse solutions appear via a subcritical pitchfork bifurcation. The bifurcation plays a key role in that mechanism. Small perturbations, externally applied or from internal instabilities, may cause pulse propagation failure (wave breakup) provided the system is close enough to the bifurcation point. We derive relations showing how the pitchfork bifurcation is unfolded by weak curvature or advective field perturbations and use them to demonstrate wave breakup. We suggest that the recent observations of wave breakup in the Belousov-Zhabotinsky reaction induced either by an electric field or a transverse instability are manifestations of this mechanism.

patt-sol