Emergent encoding of dispersal network topologies in spatial metapopulation models.

We address a generalization of the concept of metapopulation capacity for trees and networks acting as the template for ecological interactions. The original measure had been derived from an insightful phenomenological model and is based on the leading eigenvalue of a suitable landscape matrix. It yields a versatile predictor of metapopulation persistence through a threshold value of the eigenvalue determined by ecological features of the focal species. Here, we present an analytical solution to a fundamental microscopic model that incorporates key ingredients of metapopulation dynamics and explicitly distinguishes between individuals comprising the "settled population" and "explorers" seeking colonization. Our approach accounts for general network characteristics (in particular graph-driven directional dispersal which is known to significantly constrain many ecological estimates) and yields a generalized version of the original model, to which it reduces for particular cases. Through examples, including real landscapes used as the template, we compare the predictions from our approach with those of the standard model. Results suggest that in several cases of practical interest, differences are significant. We also examine, with both models, how changes in habitat fragmentation, including removal, addition, or alteration in size, affect metapopulation persistence. The current approach demonstrates a high level of flexibility, enabling the incorporation of diverse "microscopic" elements and their impact on the resulting biodiversity landscape pattern.

Delay effects on the stability of large ecosystems.

The common intuition among the ecologists of the midtwentieth century was that large ecosystems should be more stable than those with a smaller number of species. This view was challenged by Robert May, who found a stability bound for randomly assembled ecosystems; they become unstable for a sufficiently large number of species. In the present work, we show that May's bound greatly changes when the past population densities of a species affect its own current density. This is a common feature in real systems, where the effects of species' interactions may appear after a time lag rather than instantaneously. The local stability of these models with self-interaction is described by bounds, which we characterize in the parameter space. We find a critical delay curve that separates the region of stability from that of instability, and correspondingly, we identify a critical frequency curve that provides the characteristic frequencies of a system at the instability threshold. Finally, we calculate analytically the distributions of eigenvalues that generalize Wigner's as well as Girko's laws. Interestingly, we find that, for sufficiently large delays, the eigenvalues of a randomly coupled system are complex even when the interactions are symmetric.

Generalized Dynamical Mean Field Theory for Non-Gaussian Interactions.

We present a generalized dynamical mean field theory for studying the effects of non-Gaussian quenched noise in a general set of dynamical systems. We apply the framework to the generalized Lotka-Volterra equations, a central model in theoretical ecology, where species interactions are fixed over time and heterogeneous. Our results show that the new mean field equations have solutions that depend on all cumulants of the distribution of species interactions. We obtain an analytic solution when the interaction couplings are α-stable distributed and find a relationship between the abundance distribution of species and the statistics of microscopic interactions. In the case of sparse interactions, which we investigate analytically, we establish a simple relationship between the distribution of interactions and the one of population densities.

Effect of delay on the emergent stability patterns in generalized Lotka-Volterra ecological dynamics.

Understanding the conditions of feasibility and stability in ecological systems is a major challenge in theoretical ecology. The seminal work of May in 1972 and recent developments based on the theory of random matrices have shown the existence of emergent universal patterns of both stability and feasibility in ecological dynamics. However, only a few studies have investigated the role of delay coupled with population dynamics in the emergence of feasible and stable states. In this work, we study the effects of delay on generalized Loka-Volterra population dynamics of several interacting species in closed ecological environments. First, we investigate the relation between feasibility and stability of the modelled ecological community in the absence of delay and find a simple analytical relation when intra-species interactions are dominant. We then show how, by increasing the time delay, there is a transition in the stability phases of the population dynamics: from an equilibrium state to a stable non-point attractor phase. We calculate analytically the critical delay of that transition and show that it is in excellent agreement with numerical simulations. Finally, following a similar approach to characterizing stability in empirical studies, we investigate the coefficient of variation, which quantifies the magnitude of population fluctuations. We show that in the oscillatory regime induced by the delay, the variability at community level decreases for increasing diversity. This article is part of the theme issue 'Emergent phenomena in complex physical and socio-technical systems: from cells to societies'.

Effective Resource Competition Model for Species Coexistence.

Local coexistence of species in large ecosystems is traditionally explained within the broad framework of niche theory. However, its rationale hardly justifies rich biodiversity observed in nearly homogeneous environments. Here we consider a consumer-resource model in which a coarse-graining procedure accounts for a variety of ecological mechanisms and leads to effective spatial effects which favor species coexistence. Herein, we provide conditions for several species to live in an environment with very few resources. In fact, the model displays two different phases depending on whether the number of surviving species is larger or smaller than the number of resources. We obtain conditions whereby a species can successfully colonize a pool of coexisting species. Finally, we analytically compute the distribution of the population sizes of coexisting species. Numerical simulations as well as empirical distributions of population sizes support our analytical findings.

Redundancy-selection trade-off in phenotype-structured populations.

Realistic fitness landscapes generally display a redundancy-fitness trade-off: highly fit trait configurations are inevitably rare, while less fit trait configurations are expected to be more redundant. The resulting sub-optimal patterns in the fitness distribution are typically described by means of effective formulations, where redundancy provided by the presence of neutral contributions is modelled implicitly, e.g. with a bias of the mutation process. However, the extent to which effective formulations are compatible with explicitly redundant landscapes is yet to be understood, as well as the consequences of a potential miss-match. Here we investigate the effects of such trade-off on the evolution of phenotype-structured populations, characterised by continuous quantitative traits. We consider a typical replication-mutation dynamics, and we model redundancy by means of two dimensional landscapes displaying both selective and neutral traits. We show that asymmetries of the landscapes will generate neutral contributions to the marginalised fitness-level description, that cannot be described by effective formulations, nor disentangled by the full trait distribution. Rather, they appear as effective sources, whose magnitude depends on the geometry of the landscape. Our results highlight new important aspects on the nature of sub-optimality. We discuss practical implications for rapidly mutant populations such as pathogens and cancer cells, where the qualitative knowledge of their trait and fitness distributions can drive disease management and intervention policies.

Modelling determinants of extinction across two Mesozoic hyperthermal events.

The Late Triassic and Early Toarcian extinction events are both associated with greenhouse warming events triggered by massive volcanism. These Mesozoic hyperthermals were responsible for the mass extinction of marine organisms and resulted in significant ecological upheaval. It has, however, been suggested that these events merely involved intensification of background extinction rates rather than significant shifts in the macroevolutionary regime and extinction selectivity. Here, we apply a multivariate modelling approach to a vast global database of marine organisms to test whether extinction selectivity varied through the Late Triassic and Early Jurassic. We show that these hyperthermals do represent shifts in the macroevolutionary regime and record different extinction selectivity compared to background intervals of the Late Triassic and Early Jurassic. The Late Triassic mass extinction represents a more profound change in selectivity than the Early Toarcian extinction but both events show a common pattern of selecting against pelagic predators and benthic photosymbiotic and suspension-feeding organisms, suggesting that these groups of organisms may be particularly vulnerable during episodes of global warming. In particular, the Late Triassic extinction represents a macroevolutionary regime change that is characterized by (i) the change in extinction selectivity between Triassic background intervals and the extinction event itself; and (ii) the differences in extinction selectivity between the Late Triassic and Early Jurassic as a whole.

The effect of environmental stochasticity on species richness in neutral communities.

Environmental stochasticity is known to be a destabilizing factor, increasing abundance fluctuations and extinction rates of populations. However, the stability of a community may benefit from the differential response of species to environmental variations due to the storage effect. This paper provides a systematic and comprehensive discussion of these two contradicting tendencies, using the metacommunity version of the recently proposed time-average neutral model of biodiversity which incorporates environmental stochasticity and demographic noise and allows for extinction and speciation. We show that the incorporation of demographic noise into the model is essential to its applicability, yielding realistic behavior of the system when fitness variations are relatively weak. The dependence of species richness on the strength of environmental stochasticity changes sign when the correlation time of the environmental variations increases. This transition marks the point at which the storage effect no longer succeeds in stabilizing the community.

Quantifying the Resilience of Coal Energy Supply in China Toward Carbon Neutrality.

Facing the challenge of achieving the goal of carbon neutrality, China is decoupling the currently close dependence of its economy on coal use. The energy supply and demand decarbonization has substantial influence on the resilience of the coal supply. However, a general understanding of the precise impact of energy decarbonization on the resilience of the coal energy supply is still lacking. Here, from the perspective of network science, we propose a theoretical framework to explore the resilience of the coal market of China. We show that the processes of increasing the connectivity and the competition between the coal enterprises, which are widely believed to improve the resilience of the coal market, can undermine the sustainability of the coal supply. Moreover, our results reveal that the policy of closing small-sized coal mines may not only reduce the safety accidents in the coal production but also improve the resilience of the coal market network. Using our model, we also suggest a few practical policies for minimizing the systemic risk of the coal energy supply.

Effects of noise on the critical points of Turing instability in complex ecosystems.

Noise is ubiquitous in natural and artificial systems. In a noisy environment, the interactions among nodes may fluctuate randomly, leading to more complicated interactions. In this paper we focus on the effects of noise and network topology on the Turing pattern of ecological networks with activator-inhibitor structure, which may be interpreted as prey-predator interactions. Based on the stability theory of stochastic differential equations, a sufficient condition for the uniform state is derived. The analytical results indicate that noise is beneficial for the uniform state. When the ratio between the diffusion coefficients of the predator and prey increases, the ecosystems can exhibit a transition from a uniform stable state to a Turing pattern, while when the ratio decreases, the ecosystems transit from a Turing pattern to a uniform stable state. There are two crucial critical points in Turing patterns, forward and backward. We find that both forward and backward critical points increase as the noise intensity increases. This means that noise favors a stable homogeneous state compared to a state with a heterogeneous pattern, which is consistent with the analytical results. In addition, noise can weaken the hysteresis phenomenon and even eliminate it in some cases. Furthermore, we report that network topology plays an important role in modulating the uniform state of ecosystems, such as the size of prey-predator systems, the network connectivity, and the strength of interaction.

Spatial aggregation and the species-area relationship across scales.

There has been a considerable effort to understand and quantify the spatial distribution of species across different ecosystems. Relative species abundance (RSA), beta diversity and species-area relationship (SAR) are among the most used macroecological measures to characterize plants communities in forests. In this paper we introduce a simple phenomenological model based on Poisson cluster processes which allows us to exactly link RSA and beta diversity to SAR. The framework is spatially explicit and accounts for the spatial aggregation of conspecific individuals. Under the simplifying assumption of neutral theory, we derive an analytical expression for the SAR which reproduces tri-phasic behavior as sample area increases from local to continental scales, explaining how the tri-phasic behavior can be understood in terms of simple geometric arguments. We also find an expression for the endemic area relationship (EAR) and for the scaling of the RSA.

Downscaling species occupancy from coarse spatial scales.

The measurement and prediction of species' populations at different spatial scales is crucial to spatial ecology as well as conservation biology. An efficient yet challenging goal to achieve such population estimates consists of recording empirical species' presence and absence at a specific regional scale and then trying to predict occupancies at finer scales. So far the majority of the methods have been based on particular species' distributional features deemed to be crucial for downscaling occupancy. However, only a minority of them have dealt explicitly with specific spatial features. Here we employ a wide class of spatial point processes, the shot noise Cox processes (SNCP), to model species occupancies at different spatial scales and show that species' spatial aggregation is crucial for predicting population estimates at fine scales starting from coarser ones. These models are formulated in continuous space and locate points regardless of the arbitrary resolution that one employs to study the spatial pattern. We compare the performances of nine models, calibrated at regional scales and demonstrate that a very simple class of SNCP, the Thomas process, is able to outperform other published models in predicting occupancies down to areas four orders of magnitude smaller than the ones employed for the parameterization. We conclude by explaining the ability of the approach to infer spatially explicit information from spatially implicit measures, the potential of the framework to combine niche and spatial models, and the possibility of reversing the method to allow upscaling.

Dynamical evolution of ecosystems.

The assembly of an ecosystem such as a tropical forest depends crucially on the species interaction network, and the deduction of its rules is a formidably complex problem. In spite of this, many recent studies using Hubbell's neutral theory of biodiversity and biogeography have demonstrated that the resulting emergent macroscopic behaviour of the ecosystem at or near a stationary state shows a surprising simplicity reminiscent of many physical systems. Indeed the symmetry postulate, that the effective birth and death rates are species-independent within a single trophic level, allows one to make analytical predictions for various static distributions such as the relative species abundance, beta-diversity and the species-area relationship. In contrast, there have only been a few studies of the dynamics and stability of tropical rain forests. Here we consider the dynamical behaviour of a community, and benchmark it against the exact predictions of a neutral model near or at stationarity. In addition to providing a description of the relative species abundance, our analysis leads to a quantitative understanding of the species turnover distribution and extinction times, and a measure of the temporal scales of neutral evolution. Our model gives a very good description of the large quantity of data collected in Barro Colorado Island in Panama in the period 1990-2000 with just three ecologically relevant parameters and predicts the dynamics of extinction of the existing species.

Predicting spatial similarity of freshwater fish biodiversity.

A major issue in modern ecology is to understand how ecological complexity at broad scales is regulated by mechanisms operating at the organismic level. What specific underlying processes are essential for a macroecological pattern to emerge? Here, we analyze the analytical predictions of a general model suitable for describing the spatial biodiversity similarity in river ecosystems, and benchmark them against the empirical occurrence data of freshwater fish species collected in the Mississippi-Missouri river system. Encapsulating immigration, emigration, and stochastic noise, and without resorting to species abundance data, the model is able to reproduce the observed probability distribution of the Jaccard similarity index at any given distance. In addition to providing an excellent agreement with the empirical data, this approach accounts for heterogeneities of different subbasins, suggesting a strong dependence of biodiversity similarity on their respective climates. Strikingly, the model can also predict the actual probability distribution of the Jaccard similarity index for any distance when considering just a relatively small sample. The proposed framework supports the notion that simplified macroecological models are capable of predicting fundamental patterns-a theme at the heart of modern community ecology.

Evolution of dispersal in explicitly spatial metacommunities.

We apply an evolutionary game theoretic approach to the evolution of dispersal in explicitly spatial metacommunities, using a flexible parametric class of dispersal kernels, namely 2Dt kernels, and study the resulting evolutionary dynamics and outcomes. We observe strong selective pressure on mean dispersal distance (i.e., the first moment), and weaker, but significant, one on the shape of dispersal kernel (i.e., higher moments). We investigate the effects of landscape topology and spatial heterogeneity on the resulting 'optimal' dispersal kernels. The shape-importantly the tail structure-and stability of evolutionarily optimal dispersal strategies are strongly affected by landscape topology or connectivity. Specifically, the results suggest that the optimal dispersal kernels in the river network topology have heavier tails and are stable, while those in the direct topology, where organisms are allowed to travel directly from one location to another, have relatively thin tails and may be unstable. We also find that habitat spatial heterogeneity enables coexistence and controls spatial distribution of distinct groups of dispersal strategies and that alteration in topology alone may not be sufficient to change such coexistence. This work provides a tool to translate environmental changes such as global climate change and human intervention into changes in dispersal behavior, which in turn may lead to important alterations of biodiversity and biological invasion patterns.

Ginzburg-Landau amplitude equation for nonlinear nonlocal models.

Regular spatial structures emerge in a wide range of different dynamics characterized by local and/or nonlocal coupling terms. In several research fields this has spurred the study of many models, which can explain pattern formation. The modulations of patterns, occurring on long spatial and temporal scales, cannot be captured by linear approximation analysis. Here, we show that, starting from a general model with long range couplings displaying patterns, the spatiotemporal evolution of large-scale modulations at the onset of instability is ruled by the well-known Ginzburg-Landau equation, independently of the details of the dynamics. Hence, we demonstrate the validity of such equation in the description of the behavior of a wide class of systems. We introduce a mathematical framework that is also able to retrieve the analytical expressions of the coefficients appearing in the Ginzburg-Landau equation as functions of the model parameters. Such framework can include higher order nonlocal interactions and has much larger applicability than the model considered here, possibly including pattern formation in models with very different physical features.

Asymmetric stochastic resetting: Modeling catastrophic events.

In the classical stochastic resetting problem, a particle, moving according to some stochastic dynamics, undergoes random interruptions that bring it to a selected domain, and then the process recommences. Hitherto, the resetting mechanism has been introduced as a symmetric reset about the preferred location. However, in nature, there are several instances where a system can only reset from certain directions, e.g., catastrophic events. Motivated by this, we consider a continuous stochastic process on the positive real line. The process is interrupted at random times occurring at a constant rate, and then the former relocates to a value only if the current one exceeds a threshold; otherwise, it follows the trajectory defined by the underlying process without resetting. An approach to obtain the exact nonequilibrium steady state of such systems and the mean first passage time to reach the origin is presented. Furthermore, we obtain the explicit solutions for two different model systems. Some of the classical results found in symmetric resetting, such as the existence of an optimal resetting, are strongly modified. Finally, numerical simulations have been performed to verify the analytical findings, showing an excellent agreement.

Upscaling species richness and abundances in tropical forests.

The quantification of tropical tree biodiversity worldwide remains an open and challenging problem. More than two-fifths of the number of worldwide trees can be found either in tropical or in subtropical forests, but only ≈0.000067% of species identities are known. We introduce an analytical framework that provides robust and accurate estimates of species richness and abundances in biodiversity-rich ecosystems, as confirmed by tests performed on both in silico-generated and real forests. Our analysis shows that the approach outperforms other methods. In particular, we find that upscaling methods based on the log-series species distribution systematically overestimate the number of species and abundances of the rare species. We finally apply our new framework on 15 empirical tropical forest plots and quantify the minimum percentage cover that should be sampled to achieve a given average confidence interval in the upscaled estimate of biodiversity. Our theoretical framework confirms that the forests studied are comprised of a large number of rare or hyper-rare species. This is a signature of critical-like behavior of species-rich ecosystems and can provide a buffer against extinction.

System size expansion for systems with an absorbing state.

The well-known van Kampen system size expansion, while of rather general applicability, is shown to fail to reproduce some qualitative features of the time evolution for systems with an absorbing state, apart from a transient initial time interval. We generalize the van Kampen ansatz by introducing a new prescription leading to non-Gaussian fluctuations around the absorbing state. The two expansion predictions are explicitly compared for the infinite range voter model with speciation as a paradigmatic model with an absorbing state. The new expansion, both for a finite size system in the large time limit and at finite time in the large size limit, converges to the exact solution as obtained in a numerical implementation using the Gillespie algorithm. Furthermore, the predicted lifetime distribution is shown to have the correct asymptotic behavior.

Stochastic dynamics of cholera epidemics.

We describe the predictions of an analytically tractable stochastic model for cholera epidemics following a single initial outbreak. The exact model relies on a set of assumptions that may restrict the generality of the approach and yet provides a realm of powerful tools and results. Without resorting to the depletion of susceptible individuals, as usually assumed in deterministic susceptible-infected-recovered models, we show that a simple stochastic equation for the number of ill individuals provides a mechanism for the decay of the epidemics occurring on the typical time scale of seasonality. The model is shown to provide a reasonably accurate description of the empirical data of the 2000/2001 cholera epidemic which took place in the Kwa Zulu-Natal Province, South Africa, with possibly notable epidemiological implications.