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The hypothesis that living systems can benefit from operating at the vicinity of critical points has gained momentum in recent years. Criticality may confer an optimal balance between exceedingly ordered and too noisy states. We here present a model, based on information theory and statistical mechanics, illustrating how and why a community of agents aimed at understanding and communicating with each other converges to a globally coherent state in which all individuals are close to an internal critical state, i.e. at the borderline between order and disorder. We study --both analytically and computationally-- the circumstances under which criticality is the best possible outcome of the dynamical process, confirming the convergence to critical points under very generic conditions. Finally, we analyze the effect of cooperation (agents try to enhance not only their fitness, but also that of other individuals) and competition (agents try to improve their own fitness and to diminish those of competitors) within our setting. The conclusion is that, while competition fosters criticality, cooperation hinders it and can lead to more ordered or more disordered consensual solutions., Comment: 20 pages, 5 figures. Supplementary Material: 8 pages

The role of species interactions in controlling the interplay between the stability of an ecosystem and its biodiversity is still not well understood. The ability of ecological communities to recover after a small perturbation of the species abundances (local asymptotic stability) has been well studied, whereas the likelihood of a community to persist when the interactions are altered (structural stability) has received much less attention. Our goal is to understand the effects of diversity, interaction strenghts and ecological network structure on the volume of parameter space leading to feasible equilibria, i.e., ones in which all populations have positive abundances. We develop a geometrical framework to study the range of conditions necessary for feasible coexistence in both mutualistic and consumer-resource systems. Using analytical and numerical methods, we show that feasibility is determined by just a handful of quantities describing the interactions, yielding a nontrivial complexity-feasibility relationship. Analyzing more than 100 empirical networks, we show that the range of coexistence conditions in mutualistic systems can be analytically predicted by means of a null model of random interactions, whereas food webs are characterized by smaller coexistence domains than those expected by chance. Finally, we characterize the geometric shape of the feasibility domain, thereby identifying the direction of perturbations that are more likely to cause extinctions. Interestingly, the structure of mutualistic interactions leads to very heterogeneous responses to perturbations, making those systems more fragile than expected by chance., Comment: 61 pages, 17 figures

The simplest theories often have much merit and many limitations, and in this vein, the value of Neutral Theory (NT) has been the subject of much debate over the past 15 years. NT was proposed at the turn of the century by Stephen Hubbell to explain pervasive patterns observed in the organization of ecosystems. Its originally tepid reception among ecologists contrasted starkly with the excitement it caused among physicists and mathematicians. Indeed, NT spawned several theoretical studies that attempted to explain empirical data and predicted trends of quantities that had not yet been studied. While there are a few reviews of NT oriented towards ecologists, our goal here is to review the quantitative results of NT and its extensions for physicists who are interested in learning what NT is, what its successes are and what important problems remain unresolved. Furthermore, we hope that this review could also be of interest to theoretical ecologists because many potentially interesting results are buried in the vast NT literature. We propose to make these more accessible by extracting them and presenting them in a logical fashion. We conclude the review by discussing how one might introduce realistic non-neutral elements into the current models., Comment: 55 pages, 8 figures

Neutral dynamics, where taxa are assumed to be demographically equivalent and their abundance is governed solely by the stochasticity of the underlying birth-death process, has proved itself as an important minimal model that accounts for many empirical datasets in genetics and ecology. However, the restriction of the model to demographic [${\cal{O}} ({\sqrt N})$] noise yields relatively slow dynamics that appears to be in conflict with both short-term and long-term characteristics of the observed systems. Here we analyze two of these problems - age size relationships and species extinction time - in the framework of a neutral theory with both demographic and environmental stochasticity. It turns out that environmentally induced variations of the demographic rates control the long-term dynamics and modify dramatically the predictions of the neutral theory with demographic noise only, yielding much better agreement with empirical data. We consider two prototypes of "zero mean" environmental noise, one which is balanced with regard to the arithmetic abundance, another balanced in the logarithmic (fitness) space, study their species lifetime statistics and discuss their relevance to realistic models of community dynamics.

Understanding the assembly of ecosystems to estimate the number of species at different spatial scales is a challenging problem. Until now, maximum entropy approaches have lacked the important feature of considering space in an explicit manner. We propose a spatially explicit maximum entropy model suitable to describe spatial patterns such as the species area relationship and the endemic area relationship. Starting from the minimal information extracted from presence/absence data, we compare the behavior of two models considering the occurrence or lack thereof of each species and information on spatial correlations. Our approach uses the information at shorter spatial scales to infer the spatial organization at larger ones. We also hypothesize a possible ecological interpretation of the effective interaction we use to characterize spatial clustering.

Social ecological systems are often difficult to investigate and manage because of their inherent complexity1. Small variations in external drivers can lead to abrupt changes associated with instabilities and bifurcations in the underlying dynamics2-4. Anticipating critical transitions and divergence from the present state of the system is particularly crucial to the prevention or mitigation of the effects of unwanted and irreversible changes5-10. Recent research in ecology has focused on leading indicators of regime shift in ecosystems characterized by one state variable5,7,11,12. The case of systems with several mutually interacting components, however, has remained poorly investigated13, while the connection between network stability and research on indicators for loss of resilience has been elusive14. Here we develop a theoretical framework to analyze early warning signs of instability and regime shift in social ecological networks. We provide analytical expressions for a set of precursors of instability in social ecological systems with additive noise for a variety of network structures. In particular, we show that the covariance matrix of the dynamics can effectively anticipate the emergence of instability. We also compare signals of early warning based on the dynamics of suitably selected nodes, to indicators based on the integrated behavior of the whole network. We find that the performances of these indicators are affected by the network structure and the type of interaction among nodes. These results provide new advances in multidimensional early warning analysis and offer a framework to evaluate the resilience of social ecological networks., Comment: 14 pages, 4 figures. Supplementary Information available upon request

Mutualistic networks are formed when the interactions between two classes of species are mutually beneficial. They are important examples of cooperation shaped by evolution. Mutualism between animals and plants plays a key role in the organization of ecological communities. Such networks in ecology have generically evolved a nested architecture independent of species composition and latitude - specialists interact with proper subsets of the nodes with whom generalists interact. Despite sustained efforts to explain observed network structure on the basis of community-level stability or persistence, such correlative studies have reached minimal consensus. Here we demonstrate that nested interaction networks could emerge as a consequence of an optimization principle aimed at maximizing the species abundance in mutualistic communities. Using analytical and numerical approaches, we show that because of the mutualistic interactions, an increase in abundance of a given species results in a corresponding increase in the total number of individuals in the community, as also the nestedness of the interaction matrix. Indeed, the species abundances and the nestedness of the interaction matrix are correlated by an amount that depends on the strength of the mutualistic interactions. Nestedness and the observed spontaneous emergence of generalist and specialist species occur for several dynamical implementations of the variational principle under stationary conditions. Optimized networks, while remaining stable, tend to be less resilient than their counterparts with randomly assigned interactions. In particular, we analytically show that the abundance of the rarest species is directly linked to the resilience of the community. Our work provides a unifying framework for studying the emergent structural and dynamical properties of ecological mutualistic networks., Comment: 10 pages, 4 figures

Empirical evidence suggesting that living systems might operate in the vicinity of critical points, at the borderline between order and disorder, has proliferated in recent years, with examples ranging from spontaneous brain activity to flock dynamics. However, a well-founded theory for understanding how and why interacting living systems could dynamically tune themselves to be poised in the vicinity of a critical point is lacking. Here we employ tools from statistical mechanics and information theory to show that complex adaptive or evolutionary systems can be much more efficient in coping with diverse heterogeneous environmental conditions when operating at criticality. Analytical as well as computational evolutionary and adaptive models vividly illustrate that a community of such systems dynamically self-tunes close to a critical state as the complexity of the environment increases while they remain non-critical for simple and predictable environments. A more robust convergence to criticality emerges in co-evolutionary and co-adaptive set-ups in which individuals aim to represent other agents in the community with fidelity, thereby creating a collective critical ensemble and providing the best possible trade-off between accuracy and flexibility. Our approach provides a parsimonious and general mechanism for the emergence of critical-like behavior in living systems needing to cope with complex environments or trying to efficiently coordinate themselves as an ensemble., Comment: Main text: 9 pages, 3 figures. Supplementary Information: 22 pages, 14 figures, 3 videos

Modern ecology has re-emphasized the need for a quantitative understanding of the original 'survival of the fittest theme' based on analyzis of the intricate trade-offs between competing evolutionary strategies that characterize the evolution of life. This is key to the understanding of species coexistence and ecosystem diversity under the omnipresent constraint of limited resources. In this work we propose an agent based model replicating a community of interacting individuals, e.g. plants in a forest, where all are competing for the same finite amount of resources and each competitor is characterized by a specific growth-reproduction strategy. We show that such an evolution dynamics drives the system towards a stationary state characterized by an emergent optimal strategy, which in turn depends on the amount of available resources the ecosystem can rely on. We find that the share of resources used by individuals is power-law distributed with an exponent directly related to the optimal strategy. The model can be further generalized to devise optimal strategies in social and economical interacting systems dynamics., Comment: 10 pages, 5 figures

Population growth is in general constrained by food production, which in turn depends on the access to water resources. At a country level, some populations use more water than they control because of their ability to import food and the virtual water required for its production. Here, we investigate the dependence of demographic growth on available water resources for exporting and importing nations. By quantifying the carrying capacity of nations based on calculations of the virtual water available through the food trade network, we point to the existence of a global water unbalance. We suggest that current export rates will not be maintained and consequently we question the long-run sustainability of the food trade system as a whole. Water rich regions are likely to soon reduce the amount of virtual water they export, thus leaving import-dependent regions without enough water to sustain their populations. We also investigate the potential impact of possible scenarios that might mitigate these effects through (1) cooperative interactions among nations whereby water rich countries maintain a tiny fraction of their food production available for export; (2) changes in consumption patterns; and (3) a positive feedback between demographic growth and technological innovations. We find that these strategies may indeed reduce the vulnerability of water-controlled societies., Comment: 11 pages, 3 figures

In the last years, a remarkable theoretical effort has been made in order to understand stability and complexity in ecological communities. The non-random structures of real ecological interaction networks has been recognized as one key ingredient contributing to the coexistence between high complexity and stability in real ecosystems. However most of the theoretical studies have considered communities with only one interaction type (either antagonistic, competitive, or mutualistic). Recently it has been proposed a theoretical analysis on multiple interaction types in ecological systems, concluding that: a) Mixture of antagonistic and mutualistic interactions stabilize the system with respect to the less realistic case of only one interaction type; b) Complexity, measured in terms of the number of interactions and the number of species, increases stability in systems with different types of interactions. By introducing new theoretical investigations and analyzing 21 empirical data sets representing mutualistic plant-pollinator networks, we show that that conclusions are incorrect. We will prove that the positive complexity-stability effect observed in systems with different kind of interactions is a mere consequence of a rescaling of the interaction strengths, and thus unrelated to the mixing of interaction types., Comment: 12 pages, 5 figures

We present novel analytical results about ecosystem species diversity that stem from a proposed coarse grained neutral model based on birth-death processes. The relevance of the problem lies in the urgency for understanding and synthesizing both theoretical results of ecological neutral theory and empirical evidence on species diversity preservation. Neutral model of biodiversity deals with ecosystems in the same trophic level where per-capita vital rates are assumed to be species-independent. Close-form analytical solutions for neutral theory are obtained within a coarse-grained model, where the only input is the species persistence time distribution. Our results pertain: the probability distribution function of the number of species in the ecosystem both in transient and stationary states; the n-points connected time correlation function; and the survival probability, definned as the distribution of time-spans to local extinction for a species randomly sampled from the community. Analytical predictions are also tested on empirical data from a estuarine fish ecosystem. We find that emerging properties of the ecosystem are very robust and do not depend on specific details of the model, with implications on biodiversity and conservation biology., Comment: 20 pages, 4 figures. To appear in Journal of Statistichal Mechanics

Natural ecosystems are characterized by striking diversity of form and functions and yet exhibit deep symmetries emerging across scales of space, time and organizational complexity. Species-area relationships and species-abundance distributions are examples of emerging patterns irrespective of the details of the underlying ecosystem functions. Here we present empirical and theoretical evidence for a new macroecological pattern related to the distributions of local species persistence times, defined as the timespans between local colonizations and extinctions in a given geographic region. Empirical distributions pertaining to two different taxa, breeding birds and herbaceous plants, analyzed in a new framework that accounts for the finiteness of the observational period, exhibit power-law scaling limited by a cut-off determined by the rate of emergence of new species. In spite of the differences between taxa and spatial scales of analysis, the scaling exponents are statistically indistinguishable from each other and significantly different from those predicted by existing models. We theoretically investigate how the scaling features depend on the structure of the spatial interaction network and show that the empirical scaling exponents are reproduced once a two-dimensional isotropic texture is used, regardless of the details of the ecological interactions. The framework developed here also allows to link the cut-off timescale with the spatial scale of analysis, and the persistence-time distribution to the species-area relationship. We conclude that the inherent coherence obtained between spatial and temporal macroecological patterns points at a seemingly general feature of the dynamical evolution of ecosystems., Comment: 5 pages, 5 figures. Supplementary materials avaliable on http://www.pnas.org/content/108/11/4346

Generalized Langevin equations (GLE) with multiplicative white Poisson noise pose the usual prescription dilemma leading to different evolution equations (master equations) for the probability distribution. Contrary to the case of multiplicative gaussian white noise, the Stratonovich prescription does not correspond to the well known mid-point (or any other intermediate) prescription. By introducing an inertial term in the GLE we show that the Ito and Stratonovich prescriptions naturally arise depending on two time scales, the one induced by the inertial term and the other determined by the jump event. We also show that when the multiplicative noise is linear in the random variable one prescription can be made equivalent to the other by a suitable transformation in the jump probability distribution. We apply these results to a recently proposed stochastic model describing the dynamics of primary soil salinization, in which the salt mass balance within the soil root zone requires the analysis of different prescriptions arising from the resulting stochastic differential equation forced by multiplicative white Poisson noise whose features are tailored to the characters of the daily precipitation. A method is finally suggested to infer the most appropriate prescription from the data.

Higher temperatures expected by midcentury increase the risk of shocks to crop production, while the interconnected nature of the current global food system functions to spread the impact of localized production shocks throughout the world. In this study, we analyze the global potential impact of a present-day event of equivalent magnitude to the US Dust Bowl, modeling the ways in which a sudden decline in US wheat production could cascade through the global network of agricultural trade. We use observations of country-level production, reserves, and trade data in a Food Shock Cascade model to explore trade adjustments and country-level inventory changes in response to a major, multiyear production decline. We find that a 4-year decline in wheat production of the same proportional magnitude as occurred during the Dust Bowl greatly reduces both wheat supply and reserves in the United States and propagates through the global trade network. By year 4 of the event, US wheat exports fall from 90.5 trillion kcal before the drought to 48 trillion to 52 trillion kcal, and the United States exhausts 94% of its reserves. As a result of reduced US exports, other countries meet their needs by leveraging their own reserves, leading to a 31% decline in wheat reserves globally. These findings demonstrate that an extreme production decline would lead to substantial supply shortfalls in both the United States and in other countries, where impacts outside the United States strongly depend on a country's reserves and on its relative position in the global trade network.