First derivations of the functional response were mechanistic, but subsequent uses of these funct... more First derivations of the functional response were mechanistic, but subsequent uses of these functions tended to be phenomenological. Further understanding of the mechanisms underpinning predator-prey relationships might lead to novel insights into functional response in natural systems. Because recent consideration of the physical properties of the environment has improved our understanding of predator-prey interactions, we advocate the use of physics-based approaches for the derivation of the functional response from first principles. These physical factors affect the functional response by constraining the ability of both predators and prey to move according to their size. A physics-based derivation of the functional response should thus consider the movement of organisms in relation to their physical environment. One recent article presents a model along these criteria. As an initial validation of our claim, we use a slightly modified version of this model to derive the classical parameters of the functional response (i.e., attack rate and handling time) of aquatic organisms, as affected by body size, buoyancy, water density and viscosity. We compared the predictions to relevant data. Our model provided good fit for most parameters, but failed to predict handling time. Remarkably, this is the only parameter whose derivation did not rely on physical principles. Parameters in the model were not estimated from observational data. Hence, systematic discrepancies between predictions and real data point immediately to errors in the model. An added benefit to functional response derivation from physical principles is thus to provide easy ways to validate or falsify hypotheses about predator-prey relationships.
and stable crop production depends on biodiversity and the spatial fragmentation of agricultural ... more and stable crop production depends on biodiversity and the spatial fragmentation of agricultural landscapes, and by revealing the ecological mechanisms of food security in crop pollination systems. Management for food security should consider factors such as pollinators' spillover, the amount and spatial aggregation of seminatural habitat and the animal pollination-dependence of crops. This information would be useful to design agricultural landscapes for high Landscape Pollination Potential. These results are highly relevant in the global change context, and given the worldwide trends in agriculture, which shifts towards more animal-dependent crop production.
Changes in land use generate trade-offs in the delivery of ecosystem services in agricultural lan... more Changes in land use generate trade-offs in the delivery of ecosystem services in agricultural landscapes. However, we know little about how the stability of ecosystem services responds to landscape composition, and what ecological mechanisms underlie these trade-offs. Here, we develop a model to investigate the dynamics of three ecosystem services in intensively managed agroecosystems, i.e., pollination-independent crop yield, crop pollination, and biodiversity. Our model reveals trade-offs and synergies imposed by landscape composition that affect not only the magnitude but also the stability of ecosystem service delivery. Trade-offs involving crop pollination are strongly affected by the degree to which crops depend on pollination and by their relative requirement for pollinator densities. We show conditions for crop production to increase with biodiversity and decreasing crop area, reconciling farmers' profitability and biodiversity conservation. Our results further suggest that, for pollination-dependent crops, management strategies that focus on maximizing yield will often overlook its stability. Given that agriculture has become more pollination-dependent over time, it is essential to understand the mechanisms driving these trade-offs to ensure food security.
Agricultural management should consider multiple services and stakeholders. Yet, it remains uncle... more Agricultural management should consider multiple services and stakeholders. Yet, it remains unclear how to guarantee the provision of ecosystem services that reaches stakeholders' demands, especially considering the observed biodiversity decline and the current global change predictions that may affect food security. Here, we use a model to examine how landscape compositionfraction of seminatural habitat (SNH)affects biodiversity and crop production services in intensively-managed agricultural systems. We analyse three groups of stakeholders assumed to value different ecosystem services mostindividual farmers (crop yield per area), agricultural unions (landscape production) and conservationists (biodiversity). We find that trade-offs among stakeholders' demands strongly depend on the degree of pollination dependence of crops, the strength of environmental and demographic stochasticity, and the relative amount of an ecosystem service demanded by each stakeholder, i.e. function thresholds. Intermediate amounts of SNH can allow for the delivery of relatively high levels of the three ecosystem services. Our analysis further suggests that the current levels of SNH protection lie below these intermediate amounts of SNH in intensively-managed agricultural landscapes. Given the worldwide trends in agriculture and global change, these results suggest ways of managing landscapes to reconcile the demands of several actors and ensure for biodiversity conservation and food production.
Robert MacArthur's niche theory makes explicit predictions on how community function should chang... more Robert MacArthur's niche theory makes explicit predictions on how community function should change over time in a competitive community. A key prediction is that succession progressively minimizes the energy wasted by a community, but this minimization is a trade-off between energy losses from unutilised resources and costs of maintenance. By predicting how competition determines community efficiency over time MacArthur's theory may inform on the impacts of disturbance on community function and invasion risk. We provide a rare test of this theory using phytoplankton communities, and find that older communities wasted less energy than younger ones but that the reduction in energy wastage was not monotonic over time. While community structure followed consistent and clear trajectories, community function was more idiosyncratic among adjoining successional stages and driven by total community biomass rather than species composition. Our results suggest that subtle shifts in successional sequence can alter community efficiency and these effects determine community function independently of individual species membership. We conclude that, at least in phytoplankton communities, general trends in community function are predictable over time accordingly to MacArthur's theory. Tests of MacArthur's minimization principle across very different systems should be a priority given the potential of this theory to inform on the functional properties of communities.
, et al.. How to estimate complementarity and selection effects from an incomplete sample of spec... more , et al.. How to estimate complementarity and selection effects from an incomplete sample of species.
Understanding the effects of plant species diversity and trait composition on aboveground biomass... more Understanding the effects of plant species diversity and trait composition on aboveground biomass is a central focus of ecology and has important implications for biodiversity conservation. However, the simultaneous direct and indirect effects of soil nutrients, species asynchrony, functional trait diversity, and trait composition for explaining the community temporal stability of aboveground biomass remain underrepresented in natural forests. Here, we hypothesized that species asynchrony relative to soil nutrients, functional trait diversity, and trait composition plays a central role in stabilizing the community temporal stability of natural forests. We tested this hypothesis using a structural equation model based on 10-year continuous monitoring data (i.e., three-time repeated forest inventories) in both second-growth and old-growth temperate forests in northeast China. Our results showed that the community temporal stability of aboveground biomass was driven by a strong direct positive effect of species asynchrony in both second-growth and old-growth temperate forests, whereas functional trait diversity and composition (i.e. community-weighted mean of leaf nitrogen content
While previous studies have evaluated the change in stability for the addition or removal of indi... more While previous studies have evaluated the change in stability for the addition or removal of individual species from trophic food chains and food webs, we know of no study that presents a general theory for how stability changes with the addition or removal of trophic levels. In this study, we present a simple model of a linear food chain and systematically evaluate how stability-measured as invariability-changes with the addition or removal of trophic levels. We identify the presence of trophic cascades in the stability of species. Owing to top-down control by predation and bottom-up regulation by prey, we find that stability of a species is highest when it is at the top of the food chain and lowest when it is just under the top of the food chain. Thus, stability shows patterns identical to those of mean biomass with the addition or removal of trophic levels in food chains. Our results provide a baseline towards a general theory of the effect of adding or removing trophic levels on stability, which can be used to inform empirical studies.
Aim-Ecosystem stability and its link with biodiversity have mainly been studied at the local scal... more Aim-Ecosystem stability and its link with biodiversity have mainly been studied at the local scale. Here we present a simple theoretical model to address the joint dependence of diversity and stability on spatial scale, from local to continental. Methods-The notion of stability we use is based on the temporal variability of an ecosystemlevel property, such as primary productivity. In this way, our model integrates the well-known species-area relationship (SAR) with a recent proposal to quantify the spatial scaling of stability, called the invariability-area relationship (IAR). Results-We show that the link between the two relationships strongly depends on whether the temporal fluctuations of the ecosystem property of interest are more correlated within than between species. If fluctuations are correlated within species but not between them, then the IAR is strongly constrained by the SAR. If instead individual fluctuations are only correlated by spatial proximity, then the IAR is unrelated to the SAR. We apply these two correlation assumptions to explore the effects of species loss and habitat destruction on stability, and find a rich variety of multi-scale spatial dependencies, with marked differences between the two assumptions. Main conclusions-The dependence of ecosystem stability on biodiversity across spatial scales is governed by the spatial decay of correlations within and between species. Our work provides a point of reference for mechanistic models and data analyses. More generally, it illustrates the relevance of macroecology for ecosystem functioning and stability.
The question whether communities should be viewed as superorganisms or loose collections of indiv... more The question whether communities should be viewed as superorganisms or loose collections of individual species has been the subject of a long-standing debate in ecology. Each view implies different spatiotemporal community patterns. Along spatial environmental gradients, the organismic view predicts that species turnover is discontinuous, with sharp boundaries between communities, while the individualistic view predicts gradual changes in species composition. Using a spatially explicit multispecies competition model, we show that organismic and individualistic forms of community organisation are two limiting cases along a continuum of outcomes. A high variance of competition strength leads to the emergence of organism-like communities due to the presence of alternative stable states, while weak and uniform interactions induce gradual changes in species composition. Dispersal can play a confounding role in these patterns. Our work highlights the critical importance of considering species interactions to understand and predict the responses of species and communities to environmental changes.
Climate change involves modifications in both the mean and the variability of temperature and pre... more Climate change involves modifications in both the mean and the variability of temperature and precipitation. According to global warming projections, both the magnitude and the frequency of extreme weather events are increasing, thereby increasing climate variability. The previous studies have reported that climate warming tends to decrease biodiversity and the temporal stability of community primary productivity (i.e., community stability), but the effects of the variability of temperature and precipitation on biodiversity, community stability, and their relationship have not been clearly explored. We used a long-term (from 1982 to 2014) field data set from a temperate grassland in northern China to explore the effects of the variability of mean temperature and total precipitation on species richness, community stability, and their relationship. Results showed that species richness promoted community stability through increases in asynchronous dynamics across species (i.e., species asynchrony). Both species richness and species asynchrony were positively associated with the residuals of community stability after controlling for its dependence on the variability of mean temperature and total precipitation. Furthermore, the variability of mean temperature reduced species richness, while the variability of total precipitation decreased species asynchrony and community stability. Overall, the present study revealed that species richness and species asynchrony promoted community stability, but increased climate variability may erode these positive effects and thereby threaten community stability.
Anthropogenic activities heavily affect biogeochemical cycles at global scales; thus it is critic... more Anthropogenic activities heavily affect biogeochemical cycles at global scales; thus it is critical to understand the degree to which these cycles can be regulated by organisms. Autotrophs can regulate nutrient abundance through resource consumption, but their growth should not be affected by changes in the supply of non-limiting nutrients. Here we present a model where autotrophs consume two nutrients-one limiting and one non-limiting nutrient-and access only part of the nutrients available in the environment. We apply our model to the oceanic cycles of iron and phosphorus to examine whether phytoplankton can regulate the concentrations of these key nutrients and how interactions between the two cycles affect their regulation efficiency. Our model predicts that autotrophs cannot efficiently regulate concentrations of the non-limiting nutrient. We show that changes in the supply of the limiting nutrient affect the concentrations of the non-limiting nutrient, and that the two nutrients vary in opposite directions. Our results suggest that interactions between biogeochemical cycles can result either in an increase or in a decrease in the regulation efficiency of nutrient concentrations, depending on whether the supplies of the limiting and non-limiting nutrients vary in the same or opposite directions due to anthropogenic activities.
bioRxiv (Cold Spring Harbor Laboratory), Oct 12, 2021
Ecology is a science of scale, which guides our description of both ecological processes and patt... more Ecology is a science of scale, which guides our description of both ecological processes and patterns, but we lack a systematic understanding of how process scale and pattern scale are connected. Recent calls for a synthesis between population ecology, community ecology, and ecosystem ecology motivate the integration of phenomena at multiple levels of organization. Furthermore, many studies leave out the scaling of a critical process: species interactions, which may be non-local through mobility or vectors (resources or species) and must be distinguished from dispersal scales. Here, we use simulations to explore the consequences of different process scales (i.e. species interactions, dispersal, and the environment) on emergent patterns of biodiversity, ecosystem functioning, and their relationship, in a spatially-explicit landscape. A major result of our study is that the spatial scales of dispersal and species interactions have opposite effects: a larger dispersal scale homogenizes spatial biomass patterns, while a larger interaction scale amplifies their heterogeneity. We find that an interesting interplay between process scales occurs when the spatial distribution of species is heterogeneous at large scales, i.e., when the environment is not too uniform and dispersal not very strong. Interestingly, the specific scale at which scales of dispersal and interactions begin to influence landscape patterns depends on the environmental heterogeneity of the landscape-in other words, the scale of one process allows important scales to emerge in other processes. Finally, contrary to our expectations, we observe that the spatial scale of ecological processes is more clearly reflected in landscape patterns (i.e. distribution of local outcomes) than in global patterns such as Species-Area Relationships or large-scale biodiversity-functioning relationships.
Rapid changes in species composition, also known as ecotones, can result from various causes incl... more Rapid changes in species composition, also known as ecotones, can result from various causes including rapid changes in environmental conditions, or physiological thresholds. The possibility that ecotones arise from ecological niche construction by ecosystem engineers has received little attention. In this study, we investigate how the diversity of ecosystem engineers, and their interactions, can give rise to ecotones. We build a spatially explicit dynamical model that couples a multispecies community and its abiotic environment. We use numerical simulations and analytical techniques to determine the biotic and abiotic conditions under which ecotone emergence is expected to occur, and the role of biodiversity therein. We show that the diversity of ecosystem engineers can lead to indirect interactions through the modification of their shared environment. These interactions, which can be either competitive or mutualistic, can lead to the emergence of discrete communities in space, separated by sharp ecotones where a high species turnover is observed. Considering biodiversity is thus critical when studying the influence of species-environment interactions on the emergence of ecotones. This is especially true for the wide range of species that have small to moderate effects on their environment. Our work highlights new mechanisms by which biodiversity loss could cause significant changes in spatial community patterns in changing environments.
bioRxiv (Cold Spring Harbor Laboratory), Nov 11, 2016
Ecological stability is a bewildering broad concept. The most common stability measures are asymp... more Ecological stability is a bewildering broad concept. The most common stability measures are asymptotic resilience, widely used in theoretical studies, and measures based on temporal variability, commonly used in empirical studies. We construct measures of invariability, defined as the inverse of variability, that can be directly compared with asymptotic resilience. We show that asymptotic resilience behaves like the invariability of the most variable species, which is often a rare species close to its extinction boundary. Therefore, asymptotic resilience displays complete loss of stability with changes in community composition. In contrast, mean population invariability and ecosystem invariability are insensitive to rare species and quantify stability consistently whether details of species composition are considered or not. Invariability provides a consistent framework to predict diversitystability relationships that agree with empirical data at population and ecosystem levels. Our findings can enhance the dialogue between theoretical and empirical stability studies. 1
Food chain theory is one of the cornerstones of ecology, providing many of its basic predictions,... more Food chain theory is one of the cornerstones of ecology, providing many of its basic predictions, such as biomass pyramids, trophic cascades and predator-prey oscillations. Yet, ninety years into this theory, the conditions under which these patterns may occur and persist in nature remain subject to debate. Rather than address each pattern in isolation, we propose that they must be understood together, calling for synthesis in a fragmented landscape of theoretical and empirical results. As a first step, we propose a minimal theory that combines the long-standing energetic and dynamical approaches of food chains. We chart theoretical predictions on a concise map, where two main regimes emerge: across various functioning and stability metrics, one regime is characterized by pyramidal patterns, the other by cascade patterns. The axes of this map combine key physiological and ecological variables, such as metabolic rates and self-regulation. A quantitative comparison with data sheds light on conflicting theoretical predictions and empirical puzzles, from size spectra to causes of trophic cascade strength. We conclude that drawing systematic connections between various existing approaches to food chains, and between their predictions on functioning and stability, is a crucial step in confronting this theory to real ecosystems.
bioRxiv (Cold Spring Harbor Laboratory), Sep 30, 2018
Empirical knowledge of ecosystem stability and diversity-stability relationships is mostly based ... more Empirical knowledge of ecosystem stability and diversity-stability relationships is mostly based on the analysis of temporal variability of population and ecosystem properties. Variability, however, often depends on external factors that act as disturbances, making it difficult to compare its value across systems and relate it to other stability concepts. Here we show how variability, when viewed as a response to stochastic perturbations, can reveal inherent stability properties of ecological communities, with clear connections with other stability notions. This requires abandoning one-dimensional representations, in which a single variability measurement is taken as a proxy for how stable a system is, and instead consider the whole set of variability values associated to a given community, reflecting the whole set of perturbations that can generate variability. Against the vertiginous dimensionality of the perturbation set, we show that a generic variability-abundance pattern emerges from community assembly, which relates variability to the abundance of perturbed species. As a consequence, the response to stochastic immigration is governed by rare species while common species drive the response to environmental perturbations. In particular, the contrasting contributions of different species abundance classes can lead to opposite diversity-stability patterns, which can be understood from basic statistics of the abundance distribution. Our work shows that a multidimensional perspective on variability allows one to better appreciate the dynamical richness of ecological systems and the underlying meaning of their stability patterns.
Predation often deviates from the law of mass action: many micro-and meso-scale experiments have ... more Predation often deviates from the law of mass action: many micro-and meso-scale experiments have shown that consumption saturates with resource abundance, and decreases due to interference between consumers. But does this observation hold at macro-ecological scales, spanning many species and orders of magnitude in biomass? If so, what are its consequences for large-scale ecological patterns and dynamics? We perform a meta-analysis of predator-prey pairs of mammals, birds and reptiles, and show that predation losses appear to increase, not as the product of predator and prey densities following the Lotka-Volterra (mass action) model, but rather as the square root of that product. This suggests a phenomenological power-law expression of the effective cross-ecosystem functional response. We discuss whether the same power-law may hold dynamically within an ecosystem, and assuming that it does, we explore its consequences in a simple food chain model. The empirical exponents fall close to the boundary between regimes of donor and consumer limitation. Exponents on this boundary are singular in multiple ways. First, they maximize predator abundance and some stability metrics. Second, they create proportionality relations between biomass and productivity, both within and between trophic levels. These intuitive relations do not hold in general in mass action models, yet they are widely observed empirically. These results provide evidence of mechanisms limiting predation across multiple ecological scales. Some of this evidence was previously associated with donor control, but we show that it supports a wider range of possibilities, including forms of consumer control. As limiting consumption counter-intuitively allows larger populations, it is worthwhile to reconsider whether the observed functional response arises from microscopic mechanisms, or could hint at selective pressure at the population level.
First derivations of the functional response were mechanistic, but subsequent uses of these funct... more First derivations of the functional response were mechanistic, but subsequent uses of these functions tended to be phenomenological. Further understanding of the mechanisms underpinning predator-prey relationships might lead to novel insights into functional response in natural systems. Because recent consideration of the physical properties of the environment has improved our understanding of predator-prey interactions, we advocate the use of physics-based approaches for the derivation of the functional response from first principles. These physical factors affect the functional response by constraining the ability of both predators and prey to move according to their size. A physics-based derivation of the functional response should thus consider the movement of organisms in relation to their physical environment. One recent article presents a model along these criteria. As an initial validation of our claim, we use a slightly modified version of this model to derive the classical parameters of the functional response (i.e., attack rate and handling time) of aquatic organisms, as affected by body size, buoyancy, water density and viscosity. We compared the predictions to relevant data. Our model provided good fit for most parameters, but failed to predict handling time. Remarkably, this is the only parameter whose derivation did not rely on physical principles. Parameters in the model were not estimated from observational data. Hence, systematic discrepancies between predictions and real data point immediately to errors in the model. An added benefit to functional response derivation from physical principles is thus to provide easy ways to validate or falsify hypotheses about predator-prey relationships.
and stable crop production depends on biodiversity and the spatial fragmentation of agricultural ... more and stable crop production depends on biodiversity and the spatial fragmentation of agricultural landscapes, and by revealing the ecological mechanisms of food security in crop pollination systems. Management for food security should consider factors such as pollinators' spillover, the amount and spatial aggregation of seminatural habitat and the animal pollination-dependence of crops. This information would be useful to design agricultural landscapes for high Landscape Pollination Potential. These results are highly relevant in the global change context, and given the worldwide trends in agriculture, which shifts towards more animal-dependent crop production.
Changes in land use generate trade-offs in the delivery of ecosystem services in agricultural lan... more Changes in land use generate trade-offs in the delivery of ecosystem services in agricultural landscapes. However, we know little about how the stability of ecosystem services responds to landscape composition, and what ecological mechanisms underlie these trade-offs. Here, we develop a model to investigate the dynamics of three ecosystem services in intensively managed agroecosystems, i.e., pollination-independent crop yield, crop pollination, and biodiversity. Our model reveals trade-offs and synergies imposed by landscape composition that affect not only the magnitude but also the stability of ecosystem service delivery. Trade-offs involving crop pollination are strongly affected by the degree to which crops depend on pollination and by their relative requirement for pollinator densities. We show conditions for crop production to increase with biodiversity and decreasing crop area, reconciling farmers' profitability and biodiversity conservation. Our results further suggest that, for pollination-dependent crops, management strategies that focus on maximizing yield will often overlook its stability. Given that agriculture has become more pollination-dependent over time, it is essential to understand the mechanisms driving these trade-offs to ensure food security.
Agricultural management should consider multiple services and stakeholders. Yet, it remains uncle... more Agricultural management should consider multiple services and stakeholders. Yet, it remains unclear how to guarantee the provision of ecosystem services that reaches stakeholders' demands, especially considering the observed biodiversity decline and the current global change predictions that may affect food security. Here, we use a model to examine how landscape compositionfraction of seminatural habitat (SNH)affects biodiversity and crop production services in intensively-managed agricultural systems. We analyse three groups of stakeholders assumed to value different ecosystem services mostindividual farmers (crop yield per area), agricultural unions (landscape production) and conservationists (biodiversity). We find that trade-offs among stakeholders' demands strongly depend on the degree of pollination dependence of crops, the strength of environmental and demographic stochasticity, and the relative amount of an ecosystem service demanded by each stakeholder, i.e. function thresholds. Intermediate amounts of SNH can allow for the delivery of relatively high levels of the three ecosystem services. Our analysis further suggests that the current levels of SNH protection lie below these intermediate amounts of SNH in intensively-managed agricultural landscapes. Given the worldwide trends in agriculture and global change, these results suggest ways of managing landscapes to reconcile the demands of several actors and ensure for biodiversity conservation and food production.
Robert MacArthur's niche theory makes explicit predictions on how community function should chang... more Robert MacArthur's niche theory makes explicit predictions on how community function should change over time in a competitive community. A key prediction is that succession progressively minimizes the energy wasted by a community, but this minimization is a trade-off between energy losses from unutilised resources and costs of maintenance. By predicting how competition determines community efficiency over time MacArthur's theory may inform on the impacts of disturbance on community function and invasion risk. We provide a rare test of this theory using phytoplankton communities, and find that older communities wasted less energy than younger ones but that the reduction in energy wastage was not monotonic over time. While community structure followed consistent and clear trajectories, community function was more idiosyncratic among adjoining successional stages and driven by total community biomass rather than species composition. Our results suggest that subtle shifts in successional sequence can alter community efficiency and these effects determine community function independently of individual species membership. We conclude that, at least in phytoplankton communities, general trends in community function are predictable over time accordingly to MacArthur's theory. Tests of MacArthur's minimization principle across very different systems should be a priority given the potential of this theory to inform on the functional properties of communities.
, et al.. How to estimate complementarity and selection effects from an incomplete sample of spec... more , et al.. How to estimate complementarity and selection effects from an incomplete sample of species.
Understanding the effects of plant species diversity and trait composition on aboveground biomass... more Understanding the effects of plant species diversity and trait composition on aboveground biomass is a central focus of ecology and has important implications for biodiversity conservation. However, the simultaneous direct and indirect effects of soil nutrients, species asynchrony, functional trait diversity, and trait composition for explaining the community temporal stability of aboveground biomass remain underrepresented in natural forests. Here, we hypothesized that species asynchrony relative to soil nutrients, functional trait diversity, and trait composition plays a central role in stabilizing the community temporal stability of natural forests. We tested this hypothesis using a structural equation model based on 10-year continuous monitoring data (i.e., three-time repeated forest inventories) in both second-growth and old-growth temperate forests in northeast China. Our results showed that the community temporal stability of aboveground biomass was driven by a strong direct positive effect of species asynchrony in both second-growth and old-growth temperate forests, whereas functional trait diversity and composition (i.e. community-weighted mean of leaf nitrogen content
While previous studies have evaluated the change in stability for the addition or removal of indi... more While previous studies have evaluated the change in stability for the addition or removal of individual species from trophic food chains and food webs, we know of no study that presents a general theory for how stability changes with the addition or removal of trophic levels. In this study, we present a simple model of a linear food chain and systematically evaluate how stability-measured as invariability-changes with the addition or removal of trophic levels. We identify the presence of trophic cascades in the stability of species. Owing to top-down control by predation and bottom-up regulation by prey, we find that stability of a species is highest when it is at the top of the food chain and lowest when it is just under the top of the food chain. Thus, stability shows patterns identical to those of mean biomass with the addition or removal of trophic levels in food chains. Our results provide a baseline towards a general theory of the effect of adding or removing trophic levels on stability, which can be used to inform empirical studies.
Aim-Ecosystem stability and its link with biodiversity have mainly been studied at the local scal... more Aim-Ecosystem stability and its link with biodiversity have mainly been studied at the local scale. Here we present a simple theoretical model to address the joint dependence of diversity and stability on spatial scale, from local to continental. Methods-The notion of stability we use is based on the temporal variability of an ecosystemlevel property, such as primary productivity. In this way, our model integrates the well-known species-area relationship (SAR) with a recent proposal to quantify the spatial scaling of stability, called the invariability-area relationship (IAR). Results-We show that the link between the two relationships strongly depends on whether the temporal fluctuations of the ecosystem property of interest are more correlated within than between species. If fluctuations are correlated within species but not between them, then the IAR is strongly constrained by the SAR. If instead individual fluctuations are only correlated by spatial proximity, then the IAR is unrelated to the SAR. We apply these two correlation assumptions to explore the effects of species loss and habitat destruction on stability, and find a rich variety of multi-scale spatial dependencies, with marked differences between the two assumptions. Main conclusions-The dependence of ecosystem stability on biodiversity across spatial scales is governed by the spatial decay of correlations within and between species. Our work provides a point of reference for mechanistic models and data analyses. More generally, it illustrates the relevance of macroecology for ecosystem functioning and stability.
The question whether communities should be viewed as superorganisms or loose collections of indiv... more The question whether communities should be viewed as superorganisms or loose collections of individual species has been the subject of a long-standing debate in ecology. Each view implies different spatiotemporal community patterns. Along spatial environmental gradients, the organismic view predicts that species turnover is discontinuous, with sharp boundaries between communities, while the individualistic view predicts gradual changes in species composition. Using a spatially explicit multispecies competition model, we show that organismic and individualistic forms of community organisation are two limiting cases along a continuum of outcomes. A high variance of competition strength leads to the emergence of organism-like communities due to the presence of alternative stable states, while weak and uniform interactions induce gradual changes in species composition. Dispersal can play a confounding role in these patterns. Our work highlights the critical importance of considering species interactions to understand and predict the responses of species and communities to environmental changes.
Climate change involves modifications in both the mean and the variability of temperature and pre... more Climate change involves modifications in both the mean and the variability of temperature and precipitation. According to global warming projections, both the magnitude and the frequency of extreme weather events are increasing, thereby increasing climate variability. The previous studies have reported that climate warming tends to decrease biodiversity and the temporal stability of community primary productivity (i.e., community stability), but the effects of the variability of temperature and precipitation on biodiversity, community stability, and their relationship have not been clearly explored. We used a long-term (from 1982 to 2014) field data set from a temperate grassland in northern China to explore the effects of the variability of mean temperature and total precipitation on species richness, community stability, and their relationship. Results showed that species richness promoted community stability through increases in asynchronous dynamics across species (i.e., species asynchrony). Both species richness and species asynchrony were positively associated with the residuals of community stability after controlling for its dependence on the variability of mean temperature and total precipitation. Furthermore, the variability of mean temperature reduced species richness, while the variability of total precipitation decreased species asynchrony and community stability. Overall, the present study revealed that species richness and species asynchrony promoted community stability, but increased climate variability may erode these positive effects and thereby threaten community stability.
Anthropogenic activities heavily affect biogeochemical cycles at global scales; thus it is critic... more Anthropogenic activities heavily affect biogeochemical cycles at global scales; thus it is critical to understand the degree to which these cycles can be regulated by organisms. Autotrophs can regulate nutrient abundance through resource consumption, but their growth should not be affected by changes in the supply of non-limiting nutrients. Here we present a model where autotrophs consume two nutrients-one limiting and one non-limiting nutrient-and access only part of the nutrients available in the environment. We apply our model to the oceanic cycles of iron and phosphorus to examine whether phytoplankton can regulate the concentrations of these key nutrients and how interactions between the two cycles affect their regulation efficiency. Our model predicts that autotrophs cannot efficiently regulate concentrations of the non-limiting nutrient. We show that changes in the supply of the limiting nutrient affect the concentrations of the non-limiting nutrient, and that the two nutrients vary in opposite directions. Our results suggest that interactions between biogeochemical cycles can result either in an increase or in a decrease in the regulation efficiency of nutrient concentrations, depending on whether the supplies of the limiting and non-limiting nutrients vary in the same or opposite directions due to anthropogenic activities.
bioRxiv (Cold Spring Harbor Laboratory), Oct 12, 2021
Ecology is a science of scale, which guides our description of both ecological processes and patt... more Ecology is a science of scale, which guides our description of both ecological processes and patterns, but we lack a systematic understanding of how process scale and pattern scale are connected. Recent calls for a synthesis between population ecology, community ecology, and ecosystem ecology motivate the integration of phenomena at multiple levels of organization. Furthermore, many studies leave out the scaling of a critical process: species interactions, which may be non-local through mobility or vectors (resources or species) and must be distinguished from dispersal scales. Here, we use simulations to explore the consequences of different process scales (i.e. species interactions, dispersal, and the environment) on emergent patterns of biodiversity, ecosystem functioning, and their relationship, in a spatially-explicit landscape. A major result of our study is that the spatial scales of dispersal and species interactions have opposite effects: a larger dispersal scale homogenizes spatial biomass patterns, while a larger interaction scale amplifies their heterogeneity. We find that an interesting interplay between process scales occurs when the spatial distribution of species is heterogeneous at large scales, i.e., when the environment is not too uniform and dispersal not very strong. Interestingly, the specific scale at which scales of dispersal and interactions begin to influence landscape patterns depends on the environmental heterogeneity of the landscape-in other words, the scale of one process allows important scales to emerge in other processes. Finally, contrary to our expectations, we observe that the spatial scale of ecological processes is more clearly reflected in landscape patterns (i.e. distribution of local outcomes) than in global patterns such as Species-Area Relationships or large-scale biodiversity-functioning relationships.
Rapid changes in species composition, also known as ecotones, can result from various causes incl... more Rapid changes in species composition, also known as ecotones, can result from various causes including rapid changes in environmental conditions, or physiological thresholds. The possibility that ecotones arise from ecological niche construction by ecosystem engineers has received little attention. In this study, we investigate how the diversity of ecosystem engineers, and their interactions, can give rise to ecotones. We build a spatially explicit dynamical model that couples a multispecies community and its abiotic environment. We use numerical simulations and analytical techniques to determine the biotic and abiotic conditions under which ecotone emergence is expected to occur, and the role of biodiversity therein. We show that the diversity of ecosystem engineers can lead to indirect interactions through the modification of their shared environment. These interactions, which can be either competitive or mutualistic, can lead to the emergence of discrete communities in space, separated by sharp ecotones where a high species turnover is observed. Considering biodiversity is thus critical when studying the influence of species-environment interactions on the emergence of ecotones. This is especially true for the wide range of species that have small to moderate effects on their environment. Our work highlights new mechanisms by which biodiversity loss could cause significant changes in spatial community patterns in changing environments.
bioRxiv (Cold Spring Harbor Laboratory), Nov 11, 2016
Ecological stability is a bewildering broad concept. The most common stability measures are asymp... more Ecological stability is a bewildering broad concept. The most common stability measures are asymptotic resilience, widely used in theoretical studies, and measures based on temporal variability, commonly used in empirical studies. We construct measures of invariability, defined as the inverse of variability, that can be directly compared with asymptotic resilience. We show that asymptotic resilience behaves like the invariability of the most variable species, which is often a rare species close to its extinction boundary. Therefore, asymptotic resilience displays complete loss of stability with changes in community composition. In contrast, mean population invariability and ecosystem invariability are insensitive to rare species and quantify stability consistently whether details of species composition are considered or not. Invariability provides a consistent framework to predict diversitystability relationships that agree with empirical data at population and ecosystem levels. Our findings can enhance the dialogue between theoretical and empirical stability studies. 1
Food chain theory is one of the cornerstones of ecology, providing many of its basic predictions,... more Food chain theory is one of the cornerstones of ecology, providing many of its basic predictions, such as biomass pyramids, trophic cascades and predator-prey oscillations. Yet, ninety years into this theory, the conditions under which these patterns may occur and persist in nature remain subject to debate. Rather than address each pattern in isolation, we propose that they must be understood together, calling for synthesis in a fragmented landscape of theoretical and empirical results. As a first step, we propose a minimal theory that combines the long-standing energetic and dynamical approaches of food chains. We chart theoretical predictions on a concise map, where two main regimes emerge: across various functioning and stability metrics, one regime is characterized by pyramidal patterns, the other by cascade patterns. The axes of this map combine key physiological and ecological variables, such as metabolic rates and self-regulation. A quantitative comparison with data sheds light on conflicting theoretical predictions and empirical puzzles, from size spectra to causes of trophic cascade strength. We conclude that drawing systematic connections between various existing approaches to food chains, and between their predictions on functioning and stability, is a crucial step in confronting this theory to real ecosystems.
bioRxiv (Cold Spring Harbor Laboratory), Sep 30, 2018
Empirical knowledge of ecosystem stability and diversity-stability relationships is mostly based ... more Empirical knowledge of ecosystem stability and diversity-stability relationships is mostly based on the analysis of temporal variability of population and ecosystem properties. Variability, however, often depends on external factors that act as disturbances, making it difficult to compare its value across systems and relate it to other stability concepts. Here we show how variability, when viewed as a response to stochastic perturbations, can reveal inherent stability properties of ecological communities, with clear connections with other stability notions. This requires abandoning one-dimensional representations, in which a single variability measurement is taken as a proxy for how stable a system is, and instead consider the whole set of variability values associated to a given community, reflecting the whole set of perturbations that can generate variability. Against the vertiginous dimensionality of the perturbation set, we show that a generic variability-abundance pattern emerges from community assembly, which relates variability to the abundance of perturbed species. As a consequence, the response to stochastic immigration is governed by rare species while common species drive the response to environmental perturbations. In particular, the contrasting contributions of different species abundance classes can lead to opposite diversity-stability patterns, which can be understood from basic statistics of the abundance distribution. Our work shows that a multidimensional perspective on variability allows one to better appreciate the dynamical richness of ecological systems and the underlying meaning of their stability patterns.
Predation often deviates from the law of mass action: many micro-and meso-scale experiments have ... more Predation often deviates from the law of mass action: many micro-and meso-scale experiments have shown that consumption saturates with resource abundance, and decreases due to interference between consumers. But does this observation hold at macro-ecological scales, spanning many species and orders of magnitude in biomass? If so, what are its consequences for large-scale ecological patterns and dynamics? We perform a meta-analysis of predator-prey pairs of mammals, birds and reptiles, and show that predation losses appear to increase, not as the product of predator and prey densities following the Lotka-Volterra (mass action) model, but rather as the square root of that product. This suggests a phenomenological power-law expression of the effective cross-ecosystem functional response. We discuss whether the same power-law may hold dynamically within an ecosystem, and assuming that it does, we explore its consequences in a simple food chain model. The empirical exponents fall close to the boundary between regimes of donor and consumer limitation. Exponents on this boundary are singular in multiple ways. First, they maximize predator abundance and some stability metrics. Second, they create proportionality relations between biomass and productivity, both within and between trophic levels. These intuitive relations do not hold in general in mass action models, yet they are widely observed empirically. These results provide evidence of mechanisms limiting predation across multiple ecological scales. Some of this evidence was previously associated with donor control, but we show that it supports a wider range of possibilities, including forms of consumer control. As limiting consumption counter-intuitively allows larger populations, it is worthwhile to reconsider whether the observed functional response arises from microscopic mechanisms, or could hint at selective pressure at the population level.
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Papers by Michel Loreau