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Self-organized spatial patterns of vegetation are frequent in drylands and, because pattern shape correlates with water availability, they have been suggested as important indicators of ecosystem health. However, the mechanisms underlying pattern emergence remain unclear. Some theories hypothesize that patterns could result from a water-mediated scale-dependent feedback (SDF) whereby interactions favoring plant growth dominate at short distances and growth-inhibitory interactions dominate in the long range. However, we know little about how the presence of a focal plant affects the fitness of its neighbors as a function of the inter-individual distance, which is expected to be highly ecosystem-dependent. This lack of empirical knowledge and system dependency challenge the relevance of SDF as a unifying theory for vegetation pattern formation. Assuming that plant interactions are always inhibitory and only their intensity is scale-dependent, alternative theories also recover the typical vegetation patterns observed in nature. Importantly, although these alternative hypotheses lead to visually indistinguishable patterns, they predict contrasting desertification dynamics, which questions the potential use of vegetation patterns as ecosystem-state indicators. To help resolve this issue, we first review existing theories for vegetation self-organization and their conflicting predictions about desertification dynamics. Second, we discuss potential empirical tests via manipulative experiments to identify pattern-forming mechanisms and link them to specific desertification dynamics. A comprehensive view of models, the mechanisms they intend to capture, and experiments to test them in the field will help to better understand both how patterns emerge and improve predictions on the fate of the ecosystems where they form., Comment: 20 pages, 6 figures

A simplified model of clonal plant growth is formulated, motivated by observations of spatial structures in Posidonia oceanica meadows in the Mediterranean Sea. Two levels of approximation are considered for the scale-dependent feedback terms. Both take into account mortality and clonal, or vegetative, growth as well as competition and facilitation, but the first version is nonlocal in space while the second is local. Study of the two versions of the model in the one-dimensional case reveals that both cases exhibit qualitatively similar behavior (but quantitative differences) and describe the competition between three spatially extended states, the bare soil state, the populated state, and a pattern state, and the associated spatially localized structures. The latter are of two types, holes in the populated state and vegetation patches on bare ground, and are organized within distinct snaking bifurcation diagrams. Fronts between the three extended states are studied and a transition between pushed and pulled fronts identified. Numerical simulations in one spatial dimension are used to determine front speeds and confront the predictions from the marginal stability condition for pulled fronts., Comment: 14 pages, 18 figures. To appear in Physica D

Brownian particles interacting via repulsive soft-core potentials can spontaneously aggregate, despite repelling each other, and form periodic crystals of particle clusters. We study this phenomenon in low-dimensional situations (one and two dimensions) at two levels of description: performing numerical simulations of the discrete particle dynamics, and by linear and nonlinear analysis of the corresponding Dean-Kawasaki equation for the macroscopic particle density. Restricting to low dimensions and neglecting fluctuation effects we gain analytical insight into the mechanisms of the instability leading to clustering which turn out to be the interplay between diffusion, the intracluster forces and the forces between neighboring clusters. We show that the deterministic part of the Dean-Kawasaki equation provides a good description of the particle dynamics, including width and shape of the clusters, in a wide range of parameters, and analyze with weakly nonlinear techniques the nature of the pattern-forming bifurcation in one and two dimensions. Finally, we briefly discuss the case of attractive forces.

We study the spatial patterns formed by a system of interacting particles where the mobility of any individual is determined by the population crowding at two different spatial scales. In this way we model the behavior of some biological organisms (like mussels) that tend to cluster at short ranges as a defensive strategy, and strongly disperse if there is a high population pressure at large ranges for optimizing foraging. We perform stochastic simulations of a particle-level model of the system, and derive and analyze a continuous density description (a nonlinear diffusion equation). In both cases we show that this interplay of scale-dependent-behaviors gives rise to a rich formation of spatial patterns ranging from labyrinths to periodic cluster arrangements. In most cases these clusters have the very peculiar appearance of ring-like structures, i.e., organisms arranging in the perimeter of the clusters, that we discuss in detail., Comment: 13 pages, 9 figures

We study the conditions under which species interaction, as described by continuous versions of the competitive Lotka-Volterra model (namely the nonlocal Kolmogorov-Fisher model, and its differential approximation), can support the existence of localized states, i.e. patches of species with enhanced population surrounded in niche space by species at smaller densities. These states would arise from species interaction, and not by any preferred niche location or better fitness. In contrast to previous works we include only quadratic nonlinearities, so that the localized patches appear on a background of homogeneously distributed species coexistence, instead than on top of the no-species empty state. For the differential model we find and describe in detail the stable localized states. For the full nonlocal model, however competitive interactions alone do not allow the conditions for the observation of self-localized states, and we show how the inclusion of additional facilitative interactions lead to the appearance of them., Comment: 9 pages, 8 figures, to appear in Physical Review E

We review recent results obtained from simple individual-based models of biological competition in which birth and death rates of an organism depend on the presence of other competing organisms close to it. In addition the individuals perform random walks of different types (Gaussian diffusion and L\'{e}vy flights). We focus on how competition and random motions affect each other, from which spatial instabilities and extinctions arise. Under suitable conditions, competitive interactions lead to clustering of individuals and periodic pattern formation. Random motion has a homogenizing effect and then delays this clustering instability. When individuals from species differing in their random walk characteristics are allowed to compete together, the ones with a tendency to form narrower clusters get a competitive advantage over the others. Mean-field deterministic equations are analyzed and compared with the outcome of the individual-based simulations., Comment: 38 pages, including 6 figures

Regular vegetation patterns in semiarid ecosystems are believed to arise from the interplay between long-range competition and facilitation processes acting at smaller distances. We show that, under rather general conditions, long-range competition alone may be enough to shape these patterns. To this end we propose a simple, general model for the dynamics of vegetation, which includes only long-range competition between plants. Competition is introduced through a nonlocal term, where the kernel function quantifies the intensity of the interaction. We recover the full spectrum of spatial structures typical of vegetation models that also account for facilitation in addition to competition., Comment: 21 pages, 3 figures

We present properties of Lotka-Volterra equations describing ecological competition among a large number of competing species. First we extend to the case of a non-homogeneous niche space stability conditions for solutions representing species coexistence. Second, we discuss mechanisms leading to species clustering and obtain an analytical solution for a lumped state in a specific instance of the system. We also discuss how realistic ecological interactions may result in different types of competition coefficients., Comment: 9 pages, 4 figures. Replaced with published version. Freely available from the publisher site under the Creative Commons Attribution license

A central model in theoretical ecology considers the competition of a range of species for a broad spectrum of resources. Recent studies have shown that essentially two different outcomes are possible. Either the species surviving competition are more or less uniformly distributed over the resource spectrum, or their distribution is 'lumped' (or 'clumped'), consisting of clusters of species with similar resource use that are separated by gaps in resource space. Which of these outcomes will occur crucially depends on the competition kernel, which reflects the shape of the resource utilization pattern of the competing species. Most models considered in the literature assume a Gaussian competition kernel. This is unfortunate, since predictions based on such a Gaussian assumption are not robust. In fact, Gaussian kernels are a border case scenario, and slight deviations from this function can lead to either uniform or lumped species distributions. Here we illustrate the non-robustness of the Gaussian assumption by simulating different implementations of the standard competition model with constant carrying capacity. In this scenario, lumped species distributions can come about by secondary ecological or evolutionary mechanisms or by details of the numerical implementation of the model. We analyze the origin of this sensitivity and discuss it in the context of recent applications of the model., Comment: 11 pages, 3 figures, revised version

We study the properties of Lotka-Volterra competitive models in which the intensity of the interaction among species depends on their position along an abstract niche space through a competition kernel. We show analytically and numerically that the properties of these models change dramatically when the Fourier transform of this kernel is not positive definite, due to a pattern forming instability. We estimate properties of the species distributions, such as the steady number of species and their spacings, for different types of kernels., Comment: 4 pages, 3 figures

A model of interacting random walkers is presented and shown to give rise to patterns consisting in periodic arrangements of fluctuating particle clusters. The model represents biological individuals that die or reproduce at rates depending on the number of neighbors within a given distance. We evaluate the importance of the discrete and fluctuating character of this particle model on the pattern forming process. To this end, a deterministic mean-field description, including a linear stability and a weakly nonlinear analysis, is given and compared with the particle model. The deterministic approach is shown to reproduce some of the features of the discrete description, in particular, the existence of a finite-wavelength instability. Stochasticity in the particle dynamics, however, has strong effects in other important aspects such as the parameter values at which pattern formation occurs, or the nature of the homogeneous phase., Comment: 17 pages, 8 figures, elsart style; To appear in Physica D

We investigate the influence of walls and corners (with Dirichlet and Neumann boundary conditions) in the evolution of twodimensional autooscillating fields described by the complex Ginzburg-Landau equation. Analytical solutions are found, and arguments provided, to show that Dirichlet walls introduce strong selection mechanisms for the wave pattern. Corners between walls provide additional synchronization mechanisms and associated selection criteria. The numerical results fit well with the theoretical predictions in the parameter range studied., Comment: 10 pages, 9 figures; for related work visit http://www.nbi.dk/~martinez

We investigate the impact of the domain shape on wave propagation in excitable media. Channelled domains with sinusoidal boundaries are considered. Trains of fronts generated periodically at an extreme of the channel are found to adopt a quasiperiodic spatial configuration stroboscopically frozen in time. The phenomenon is studied in a model for the photo-sensitive Belousov-Zabotinsky reaction, but we give a theoretical derivation of the spatial return maps prescribing the height and position of the successive fronts that is valid for arbitrary excitable reaction-diffusion systems., Comment: 4 pages (figures included)

Coupled Complex Ginzburg-Landau equations describe generic features of the dynamics of coupled fields when they are close to a Hopf bifurcation leading to nonlinear oscillations. We study numerically this set of equations and find, within a particular range of parameters, the presence of uniformly propagating localized objects behaving as coherent structures. Some of these localized objects are interpreted in terms of exact analytical solutions., Comment: 7 pages, 3 postscript figures, uses the elsart style files. Related material availeble from http://www.imedea.uib.es/Nonlinear

Dynamical properties of topological defects in a twodimensional complex vector field are considered. These objects naturally arise in the study of polarized transverse light waves. Dynamics is modeled by a Vector Complex Ginzburg-Landau Equation with parameter values appropriate for linearly polarized laser emission. Creation and annihilation processes, and selforganization of defects in lattice structures, are described. We find "glassy" configurations dominated by vectorial defects and a melting process associated to topological-charge unbinding., Comment: 4 pages, 5 figures included in the text. To appear in Phys. Rev. Lett. (2000). Related material at http://www.imedea.uib.es/Nonlinear and http://www.imedea.uib.es/Photonics . In this new version, Fig. 3 has been replaced by a better one