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The response of species to perturbations strongly depends on spatial aspects in populations connected by dispersal. Asynchronous fluctuations in biomass among populations lower the risk of simultaneous local extinctions and thus reduce the regional extinction risk. However, dispersal is often seen as passive diffusion that balances species abundance between distant patches, whereas ecological constraints, such as predator avoidance or foraging for food, trigger the movement of individuals. Here, we propose a model in which dispersal rates depend on the abundance of the species interacting with the dispersing species (e.g., prey or predators) to determine how density-dependent dispersal shapes spatial synchrony in trophic metacommunities in response to stochastic perturbations. Thus, unlike those with passive dispersal, this model with density-dependent dispersal bypasses the classic vertical transmission of perturbations due to trophic interactions and deeply alters synchrony patterns. We show that the species with the highest coefficient of variation of biomass governs the dispersal rate of the dispersing species and determines the synchrony of its populations. In addition, we show that this mechanism can be modulated by the relative impact of each species on the growth rate of the dispersing species. Species affected by several constraints disperse to mitigate the strongest constraints (e.g., predation), which does not necessarily experience the highest variations due to perturbations. Our approach can disentangle the joint effects of several factors implied in dispersal and provides a more accurate description of dispersal and its consequences on metacommunity dynamics., Competing Interests: Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2021 Elsevier Ltd. All rights reserved.)

Within a global society there exist various land use patterns, inequality, and the movement of people and goods. The various practices and behaviours associated with our current society raise questions about the future sustainability of the human population and the natural environment. We derive a simplified model of the global socio-ecological system in an effort to explore the connections between human well-being and land resources, specifically looking at resource accessibility, conservation initiatives and human migration between two economically diverse regions. We find that the spatial aspect of a global system with two distinct regions allows for faster development of technology, higher peaks in population size, greater natural land degradation, and generally speaking lower population well-being in the long-term. The unequal access to resources and differences in technological progress, alter the outcome of land management (i.e., conservation) and social behaviours (i.e., migration). We conclude that any socio-ecological management practices should be conscientious of the diversity in land access, population size, population well-being and development within the global society, as the potential for unintended consequences is high., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

High species diversity is generally thought to be a requirement for sustaining forest multifunctionality. However, the degree to which the relationship between species-, structural-, and trait-diversity of forests and multifunctionality depend on the context (such as stand age or abiotic conditions) is not well studied. Here, we hypothesized that context-dependency of tree species diversity, functional trait composition and stand structural attributes promote temperate forest multifunctionality including above- and below-ground multiple and single functions. To do so, we used repeated forest inventory data, from temperate mixed forests of northeast China, to quantify two above-ground (i.e. coarse woody productivity and wild edible plant biomass), five below-ground (i.e. soil organic carbon, total soil nitrogen, potassium, phosphorus and sulfur) functions, tree species diversity, individual tree size variation (CV DBH ) and functional trait composition of specific leaf area (CWM SLA ) as well as stand age and abiotic conditions. We found that tree species diversity increased forest multifunctionality and most of the single functions. Below-ground single and multifunctionality were better explained by tree species diversity. In contrast, above-ground single and multifunctionality were better explained by CV DBH . However, CWM SLA was also an additional important driver for maintaining above- and below-ground forest multifunctionality through opposing plant functional strategies. Stand age markedly reduced forest multifunctionality, tree species diversity and CWM SLA but substantially increased CV DBH . Below-ground forest multifunctionality and tree species diversity decreased while above-ground forest multifunctionality increased on steep slopes. These results highlight that context-dependency of forest diversity attributes might regulate forest multifunctionality but may not have a consistent effect on above-ground and below-ground forest multifunctionality due to the fact that those functions were driven by varied functional strategies of different plant species. We argue that maximizing forest complexity could act as a viable strategy to maximizing forest multifunctionality, while also promoting biodiversity conservation to mitigate climate change effects., Competing Interests: Declaration of competing interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper., (Copyright © 2020 Elsevier B.V. All rights reserved.)

The mechanisms underlying nutrient-induced diversity-stability relationships have been examined extensively. However, the effects of nutrient-induced shifts of dominant species on ecosystem stability have rarely been evaluated. We compiled a dataset from a long-term nitrogen (N) and phosphorus (P) enrichment experiment conducted in an alpine grassland on the Tibetan Plateau to test the effects of nutrient-induced shifts of dominant species on stability. Our results show that N enrichment increased synchrony among the dominant species, which contributed to a significant increase in synchrony of the whole community. Meanwhile, N-induced shifts in dominant species composition significantly increased population variability. Increases in species synchrony and population variability resulted in a decline in ecosystem stability. Our study has important implications for progress in understanding the role of plant functional compensation in the stability of ecosystem functions, which is critical for better understanding the mechanisms driving both community assembly and ecosystem functions., (Copyright © 2019 Elsevier B.V. All rights reserved.)

The destruction of natural habitats for agricultural production results in local biodiversity loss. Biodiversity loss in turn affects agricultural production indirectly through a range of biodiversity-dependent ecosystem services. Land conversion thus results in a negative externality, mediated by changes in biodiversity. When the consequences of this externality are delayed in time, lack of internalization results in overshoot-and-collapse dynamics, which are undesirable from a sustainability perspective. Here, we emphasize the importance of forward-looking policies for the long-term sustainability of human-nature interactions. We show that the internalization of this externality through a land tax can result in several win-win effects in the long run. First, more biodiversity is preserved at equilibrium, which increases the carrying capacity and total well-being of the human population. Second, a taxation path that maximizes the discounted sum of human utilities prevents or greatly alleviates overshoot-and-collapse crises, thus increasing the sustainability of the system. In particular, this result holds in the case of imperfect information regarding the precise temporal dynamics of biodiversity loss, suggesting that the design of efficient land-use management policies is possible despite incomplete ecological data. This study highlights the need to internalize biodiversity-dependent externalities through economic incentives, especially under uncertainty regarding long-term ecological dynamics., Competing Interests: Competing Interests The authors have no competing interests.

The concept of the Anthropocene is based on the idea that human impacts are now the primary drivers of changes in the earth's systems, including ecological systems. In many cases, the behavior that causes ecosystem change is itself triggered by ecological factors. Yet most ecological models still treat human impacts as given, and frequently as constant. This undermines our ability to understand the feedbacks between human behavior and ecosystem change. Focusing on the problem of species dispersal, we evaluate the effect of dispersal on biodiversity in a system subject to predation by humans. People are assumed to obtain benefits from (a) the direct consumption of species (provisioning services), (b) the non-consumptive use of species (cultural services), and (c) the buffering effects of the mix of species (regulating services). We find that the effects of dispersal on biodiversity depend jointly on the competitive interactions among species, and on human preferences over species and the services they provide. We find that while biodiversity may be greatest at intermediate levels of dispersal, this depends on structure of preferences across the metacommunity., (Copyright © 2018 Elsevier Ltd. All rights reserved.)

The ability of the human population to continue growing depends strongly on the ecosystem services provided by nature. Nature, however, is becoming more and more degraded as the number of individuals increases, which could potentially threaten the future well-being of the human population. We use a dynamic model to conceptualise links between the global proportion of natural habitats and human demography, through four categories of ecosystem services (provisioning, regulating, cultural recreational and informational) to investigate the common future of nature and humanity in terms of size and well-being. Our model shows that there is generally a trade-off between the quality of life and human population size and identifies four short-term scenarios, corresponding to three long-term steady states of the model. First, human population could experience declines if nature becomes too degraded and regulating services diminish; second the majority of the population could be in a famine state, where the population continues to grow with minimal food provision. Between these scenarios, a desirable future scenario emerges from the model. It occurs if humans convert enough land to feed all the population, while maintaining biodiversity and ecosystem services. Finally, we find a fourth scenario, which combines famine and a decline in the population because of an overexploitation of land leading to a decrease in food production. Human demography is embedded in natural dynamics; the two factors should be considered together if we are to identify a desirable future for both nature and humans., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Forests play an important role in regulating the global carbon cycle. Yet, how abiotic (i.e. soil nutrients) and biotic (i.e. tree diversity, stand structure and initial biomass) factors simultaneously contribute to aboveground biomass (coarse woody) productivity, and how the relative importance of these factors changes over succession remain poorly studied. Coarse woody productivity (CWP) was estimated as the annual aboveground biomass gain of stems using 10-year census data in old growth and secondary forests (25-ha and 4.8-ha, respectively) in northeast China. Boosted regression tree (BRT) model was used to evaluate the relative contribution of multiple metrics of tree diversity (taxonomic, functional and phylogenetic diversity and trait composition as well as stand structure attributes), stand initial biomass and soil nutrients on productivity in the studied forests. Our results showed that community-weighted mean of leaf phosphorus content, initial stand biomass and soil nutrients were the three most important individual predictors for CWP in secondary forest. Instead, initial stand biomass, rather than diversity and functional trait composition (vegetation quality) was the most parsimonious predictor of CWP in old growth forest. By comparing the results from secondary and old growth forest, the summed relative contribution of trait composition and soil nutrients on productivity decreased as those of diversity indices and initial biomass increased, suggesting the stronger effect of diversity and vegetation quantity over time. Vegetation quantity, rather than diversity and soil nutrients, is the main driver of forest productivity in temperate mixed forest. Our results imply that diversity effect for productivity in natural forests may not be so important as often suggested, at least not during the later stage of forest succession. This finding suggests that as a change of the importance of different divers of productivity, the environmentally driven filtering decreases and competitively driven niche differentiation increases with forest succession., (Copyright © 2018 Elsevier B.V. All rights reserved.)

Quantifying stability properties of ecosystems is an important problem in ecology. A common approach is based on the recovery from pulse perturbations, and posits that the faster an ecosystem return to its pre-perturbation state, the more stable it is. Theoretical studies often collapse the recovery dynamics into a single quantity: the long-term rate of return, called asymptotic resilience. However, empirical studies typically measure the recovery dynamics at much shorter time scales. In this paper we explain why asymptotic resilience is rarely representative of the short-term recovery. First, we show that, in contrast to asymptotic resilience, short-term return rates depend on features of the perturbation, in particular on the way its intensity is distributed over species. We argue that empirically relevant predictions can be obtained by considering the median response over a set of perturbations, for which we provide explicit formulas. Next, we show that the recovery dynamics are controlled through time by different species: abundant species tend to govern the short-term recovery, while rare species often dominate the long-term recovery. This shift from abundant to rare species typically causes short-term return rates to be unrelated to asymptotic resilience. We illustrate that asymptotic resilience can be determined by rare species that have almost no effect on the observable part of the recovery dynamics. Finally, we discuss how these findings can help to better connect empirical observations and theoretical predictions., (Copyright © 2017 Elsevier Ltd. All rights reserved.)

In theoretical studies, the most commonly used measure of ecological stability is resilience: ecosystems asymptotic rate of return to equilibrium after a pulse-perturbation -or shock. A complementary notion of growing popularity is reactivity: the strongest initial response to shocks. On the other hand, empirical stability is often quantified as the inverse of temporal variability, directly estimated on data, and reflecting ecosystems response to persistent and erratic environmental disturbances. It is unclear whether and how this empirical measure is related to resilience and reactivity. Here, we establish a connection by introducing two variability-based stability measures belonging to the theoretical realm of resilience and reactivity. We call them intrinsic, stochastic and deterministic invariability; respectively defined as the inverse of the strongest stationary response to white-noise and to single-frequency perturbations. We prove that they predict ecosystems worst response to broad classes of disturbances, including realistic models of environmental fluctuations. We show that they are intermediate measures between resilience and reactivity and that, although defined with respect to persistent perturbations, they can be related to the whole transient regime following a shock, making them more integrative notions than reactivity and resilience. We argue that invariability measures constitute a stepping stone, and discuss the challenges ahead to further unify theoretical and empirical approaches to stability., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Accelerating rates of biodiversity loss have led ecologists to explore the effects of species richness on ecosystem functioning and the flow of ecosystem services. One explanation of the relationship between biodiversity and ecosystem functioning lies in the spatial insurance hypothesis, which centers on the idea that productivity and stability increase with biodiversity in a temporally varying, spatially heterogeneous environment. However, there has been little work on the impact of dispersal where environmental risks are more or less spatially correlated, or where dispersal rates are variable. In this paper, we extend the original Loreau model to consider stochastic temporal variation in resource availability, which we refer to as "environmental risk", and heterogeneity in species dispersal rates. We find that asynchronies across communities and species provide community-level stabilizing effects on productivity, despite varying levels of species richness. Although intermediate dispersal rates play a role in mitigating risk, they are less effective in insuring productivity against global (metacommunity-level) than local (individual community-level) risks. These results are particularly interesting given the emergence of global sources of risk such as climate change or the closer integration of world markets. Our results offer deeper insights into the Loreau model and new perspectives on the effectiveness of spatial insurance in the face of environmental risks., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

Cooperation plays a crucial role in many aspects of biology. We use the spatial ecological metrics of local densities to measure and model cooperative interactions. While local densities can be found as technical details in current theories, we aim to establish them as central to an approach that describes spatial effects in the evolution of cooperation. A resulting local interaction model neatly partitions various spatial and non-spatial selection mechanisms. Furthermore, local densities are shown to be fundamental for important metrics of game theory, multilevel selection theory and inclusive fitness theory. The corresponding metrics include structure coefficients, spatial variance, contextual covariance, relatedness, and inbreeding coefficient or F-statistics. Local densities serve as the basis of an emergent spatial theory that draws from and brings unity to multiple theories of cooperation., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

The debate between niche-based and neutral community theories centers around the question of which forces shape predominantly ecological communities. Niche theory attributes a central role to niche differences between species, which generate a difference between the strength of intra- and interspecific interactions. Neutral theory attributes a central role to migration processes and demographic stochasticity. One possibility to bridge these two theories is to combine them in a common mathematical framework. Here we propose a mathematical model that integrates the two perspectives. From a niche-based perspective, our model can be interpreted as a Lotka-Volterra model with symmetric interactions in which we introduce immigration and demographic stochasticity. From a neutral perspective, it can be interpreted as Hubbell's local community model in which we introduce a difference between intra- and interspecific interactions. We investigate the stationary species abundance distribution and other community properties as functions of the interaction coefficient, the immigration rate and the strength of demographic stochasticity., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

The influence of diversity on ecosystem functioning and ecosystem services is now well established. Yet predictive mechanistic models that link species traits and community-level processes remain scarce, particularly for multitrophic systems. Here we revisit MacArthur's classical consumer resource model and develop a trait-based approach to predict the effects of consumer diversity on cascading extinctions and aggregated ecosystem processes in a two-trophic-level system. We show that functionally redundant efficient consumers generate top-down cascading extinctions. This counterintuitive result reveals the limits of the functional redundancy concept to predict the consequences of species deletion. Our model also predicts that the biodiversity-ecosystem functioning relationship is different for different ecosystem processes and depends on the range of variation of consumer traits in the regional species pool, which determines the sign of selection effects. Lastly, competition among resources and consumer generalism both weaken complementarity effects, which suggests that selection effects may prevail at higher trophic levels. Our work emphasizes the potential of trait-based approaches for transforming biodiversity and ecosystem functioning research into a more predictive science., (Copyright © 2010 Elsevier Ltd. All rights reserved.)

Although density-dependent dispersal and relative dispersal (the difference in dispersal rates between species) have been documented in natural systems, their effects on the stability of metacommunities are poorly understood. Here we investigate the effects of intra- and interspecific density-dependent dispersal on the regional stability in a predator-prey metacommunity model. We show that, when the dynamics of the populations reach equilibrium, the stability of the metacommunity is not affected by density-dependent dispersal. However, the regional stability, measured as the regional variability or the persistence, can be modified by density-dependent dispersal when local populations fluctuate over time. Moreover these effects depend on the relative dispersal of the predator and the prey. Regional stability is modified through changes in spatial synchrony. Interspecific density-dependent dispersal always desynchronizses local dynamics, whereas intraspecific density-dependent dispersal may either synchronize or desynchronize it depending on dispersal rates. Moreover, intra- and interspecific density-dependent dispersal strengthen the top-down control of the prey by the predator at intermediate dispersal rates. As a consequence the regional stability of the metacommunity is increased at intermediate dispersal rates. Our results show that density-dependent dispersal and relative dispersal of species are keys to understanding the response of ecosystems to fragmentation., (2010 Elsevier Ltd. All rights reserved.)

Dispersal of organisms has large effects on the dynamics and stability of populations and communities. However, current metacommunity theory largely ignores how the flows of limiting nutrients across ecosystems can influence communities. We studied a meta-ecosystem model where two autotroph-consumer communities are spatially coupled through the diffusion of the limiting nutrient. We analyzed regional and local stability, as well as spatial and temporal synchrony to elucidate the impacts of nutrient recycling and diffusion on trophic dynamics. We show that nutrient diffusion is capable of inducing asynchronous local destabilization of biotic compartments through a diffusion-induced spatiotemporal bifurcation. Nutrient recycling interacts with nutrient diffusion and influences the susceptibility of the meta-ecosystem to diffusion-induced instabilities. This interaction between nutrient recycling and transport is further shown to depend on ecosystem enrichment. It more generally emphasizes the importance of meta-ecosystem theory for predicting species persistence and distribution in managed ecosystems., (Copyright 2010 Elsevier Ltd. All rights reserved.)

The potential consequences of plant-herbivore coevolution for ecosystem functioning are investigated using a simple nutrient-limited ecosystem model in which plant and herbivore traits are subject to adaptive dynamics. Although the ecological model is very simple and always reaches a stable equilibrium in the absence of evolution, coevolution can generate a great diversity of dynamical behaviors. The evolutionary dynamics can lead to a stable equilibrium. If the evolution of plants is fast enough, certain values of the trade-off parameters lead to complex evolutionary cycles bounded by physiological constraints. The dynamical behavior of the model is very different when the dynamics of inorganic nutrient is ignored and plant competition is modeled by a logistic growth function. This emphasizes the importance of including explicit nutrient dynamics in studies of plant-herbivore coevolution.