Phenotypic plasticity: A missing element in the theory of vegetation pattern formation.

Regular spatial patterns of vegetation are a common sight in drylands. Their formation is a population-level response to water stress that increases water availability for the few via partial plant mortality. At the individual level, plants can also adapt to water stress by changing their phenotype.

Evidence for scale-dependent root-antation feedback and its role in halting the spread of a pantropical shrub into an endemic sedge.

Vegetation pattern formation is a widespread phenomenon in resource-limited environments, but the driving mechanisms are largely unconfirmed empirically. Combining results of field studies and mathematical modeling, empirical evidence for a generic pattern-formation mechanism is demonstrated with the clonal shrub L. (hereafter ) on the Brazilian island of Trindade.

Dryland mechanisms could widely control ecosystem functioning in a drier and warmer world.

Responses of terrestrial ecosystems to climate change have been explored in many regions worldwide. While continued drying and warming may alter process rates and deteriorate the state and performance of ecosystems, it could also lead to more fundamental changes in the mechanisms governing ecosystem functioning. Here we argue that climate change will induce unprecedented shifts in these mechanisms in historically wetter climatic zones, towards mechanisms currently prevalent in dry regions, which we refer to as 'dryland mechanisms'.

High-integrity human intervention in ecosystems: Tracking self-organization modes.

Humans play major roles in shaping and transforming the ecology of Earth. Unlike natural drivers of ecosystem change, which are erratic and unpredictable, human intervention in ecosystems generally involves planning and management, but often results in detrimental outcomes. Using model studies and aerial-image analysis, we argue that the design of a successful human intervention form calls for the identification of the self-organization modes that drive ecosystem change, and for studying their dynamics.

Linking spatial self-organization to community assembly and biodiversity.

Temporal shifts to drier climates impose environmental stresses on plant communities that may result in community reassembly and threatened ecosystem services, but also may trigger self-organization in spatial patterns of biota and resources, which act to relax these stresses. The complex relationships between these counteracting processes - community reassembly and spatial self-organization - have hardly been studied. Using a spatio-temporal model of dryland plant communities and a trait-based approach, we study the response of such communities to increasing water-deficit stress.

Belowground feedbacks as drivers of spatial self-organization and community assembly.

Vegetation patterning in water-limited and other resource-limited ecosystems highlights spatial self-organization processes as potentially key drivers of community assembly. These processes provide insight into predictable landscape-level relationships between organisms and their abiotic environment in the form of regular and irregular patterns of biota and resources. However, two aspects have largely been overlooked; the roles played by plant - soil-biota feedbacks and allelopathy in spatial self-organization, and their potential contribution, along with plant-resource feedbacks, to community assembly through spatial self-organization.

The role of spatial self-organization in the design of agroforestry systems.

The development of sustainable agricultural systems in drylands is currently a crucial issue in the context of mitigating the outcomes of population growth under the conditions of climatic changes. The need to meet the growing demand for food, fodder, and fuel, together with the hazards due to climate change, requires cross-disciplinary studies of ways to increase livelihood while minimizing the impact on the environment. Practices of agroforestry systems, in which herbaceous species are intercropped between rows of woody species plantations, have been shown to mitigate several of the predicaments of climatic changes.

Organizing principles for vegetation dynamics.

Plants and vegetation play a critical-but largely unpredictable-role in global environmental changes due to the multitude of contributing processes at widely different spatial and temporal scales. In this Perspective, we explore approaches to master this complexity and improve our ability to predict vegetation dynamics by explicitly taking account of principles that constrain plant and ecosystem behaviour: natural selection, self-organization and entropy maximization. These ideas are increasingly being used in vegetation models, but we argue that their full potential has yet to be realized.

A model study of terraced riverbeds as novel ecosystems.

Riverbed terracing has been introduced in ancient times to retain water and soil, to reduce hydrological connectivity and erosion and to increase primary and secondary productivity of agro-ecological systems. These presently abandoned human-made landscapes have become novel ecosystems and a potential source of ecosystem services to humans in drylands. We use a mathematical-modeling approach to study factors that regulate terraced riverbeds and affect community and ecosystem attributes such as productivity, functional diversity and resilience to droughts.

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

Frequency locking in forced oscillatory systems typically organizes 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.

Front Instabilities Can Reverse Desertification.

Degradation processes in living systems often take place gradually by front propagation. An important context of such processes is loss of biological productivity in drylands or desertification. Using a dryland-vegetation model, we analyze the stability and dynamics of desertification fronts, identify linear and nonlinear front instabilities, and highlight the significance of these instabilities in inducing self-recovery.

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 versus parametric forcing components; the stronger the parametric forcing, the more asymmetric is the response profile and the sharper is the traveling-wave front.

Implications of tristability in pattern-forming ecosystems.

Many ecosystems show both self-organized spatial patterns and multistability of possible states. The combination of these two phenomena in different forms has a significant impact on the behavior of ecosystems in changing environments. One notable case is connected to tristability of two distinct uniform states together with patterned states, which has recently been found in model studies of dryland ecosystems.

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.

Discovery of fairy circles in Australia supports self-organization theory.

Vegetation gap patterns in arid grasslands, such as the "fairy circles" of Namibia, are one of nature's greatest mysteries and subject to a lively debate on their origin. They are characterized by small-scale hexagonal ordering of circular bare-soil gaps that persists uniformly in the landscape scale to form a homogeneous distribution. Pattern-formation theory predicts that such highly ordered gap patterns should be found also in other water-limited systems across the globe, even if the mechanisms of their formation are different.

Pattern formation--A missing link in the study of ecosystem response to environmental changes.

Environmental changes can affect the functioning of an ecosystem directly, through the response of individual life forms, or indirectly, through interspecific interactions and community dynamics. The feasibility of a community-level response has motivated numerous studies aimed at understanding the mutual relationships between three elements of ecosystem dynamics: the abiotic environment, biodiversity and ecosystem function. Since ecosystems are inherently nonlinear and spatially extended, environmental changes can also induce pattern-forming instabilities that result in spatial self-organization of life forms and resources.

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.

Non-monotonic resonance in a spatially forced Lengyel-Epstein model.

We study resonant spatially periodic solutions of the Lengyel-Epstein model modified to describe the chlorine dioxide-iodine-malonic acid reaction under spatially periodic illumination. Using multiple-scale analysis and numerical simulations, we obtain the stability ranges of 2:1 resonant solutions, i.e.

Reversing desertification as a spatial resonance problem.

An important environmental application of pattern control by periodic spatial forcing is the restoration of vegetation patterns in water-limited ecosystems that went through desertification. Vegetation restoration is often based on periodic landscape modulations that intercept overland water flow and form favorable conditions for vegetation growth. Viewing this method as a spatial resonance problem, we show that plain realizations of this method, assuming a complete vegetation response to the imposed modulation pattern, suffer from poor resilience to rainfall variability.

Fronts and patterns in a spatially forced CDIMA reaction.

We use the CDIMA chemical reaction and the Lengyel-Epstein model of this reaction to study resonant responses of a pattern-forming system to time-independent spatial periodic forcing. We focus on the 2 : 1 resonance, where the wavenumber of a one-dimensional periodic forcing is about twice the wavenumber of the natural stripe pattern that the unforced system tends to form. Within this resonance, we study transverse fronts that shift the phase of resonant stripe patterns by π.

Spatial forcing of pattern-forming systems that lack inversion symmetry.

The entrainment of periodic patterns to spatially periodic parametric forcing is studied. Using a weak nonlinear analysis of a simple pattern formation model we study the resonant responses of one-dimensional systems that lack inversion symmetry. Focusing on the first three n:1 resonances, in which the system adjusts its wavenumber to one nth of the forcing wavenumber, we delineate commonalities and differences among the resonances.