Local ecosystem feedbacks and critical transitions in the climate

Global and regional climate models, such as those used in IPCC assessments, are the best tools available for climate predictions. Such models typically account for large-scale land-atmosphere feedbacks. However, these models omit local vegetation-environment feedbacks that may be crucial for critical transitions in ecosystems at larger scales. In this viewpoint paper, we propose the hypothesis that, if the balance of feedbacks is positive at all scales, local vegetation-environment feedbacks may trigger a cascade of amplifying effects, propagating from local to large scale, possibly leading to critical transitions in the large-scale climate. We call for linking local ecosystem feedbacks with large-scale land-atmosphere feedbacks in global and regional climate models in order to improve climate predictions.     

Foreseeing the future of mutualistic communities beyond collapse

Changing conditions may lead to sudden shifts in the state of ecosystems when critical thresholds are passed. Some well‐studied drivers of such transitions lead to predictable outcomes such as a turbid lake or a degraded landscape. Many ecosystems are, however, complex systems of many interacting species. While detecting upcoming transitions in such systems is challenging, predicting what comes after a critical transition is terra incognita altogether. The problem is that complex ecosystems may shift to many different, alternative states. Whether an impending transition has minor, positive or catastrophic effects is thus unclear. Some systems may, however, behave more predictably than others. The dynamics of mutualistic communities can be expected to be relatively simple, because delayed negative feedbacks leading to oscillatory or other complex dynamics are weak. Here, we address the question of whether this relative simplicity allows us to foresee a community's future state. As a case study, we use a model of a bipartite mutualistic network and show that a network's post‐transition state is indicated by the way in which a system recovers from minor disturbances. Similar results obtained with a unipartite model of facilitation suggest that our results are of relevance to a wide range of mutualistic systems.     

Global negative vegetation feedback to climate warming responses of leaf litter decomposition rates in cold biomes

Whether climate change will turn cold biomes from large long-term carbon sinks into sources is hotly debated because of the great potential for ecosystem-mediated feedbacks to global climate. Critical are the direction, magnitude and generality of climate responses of plant litter decomposition. Here, we present the first quantitative analysis of the major climate-change-related drivers of litter decomposition rates in cold northern biomes worldwide. Leaf litters collected from the predominant species in 33 global change manipulation experiments in circum-arctic-alpine ecosystems were incubated simultaneously in two contrasting arctic life zones. We demonstrate that longer-term, large-scale changes to leaf litter decomposition will be driven primarily by both direct warming effects and concomitant shifts in plant growth form composition, with a much smaller role for changes in litter quality within species. Specifically, the ongoing warming-induced expansion of shrubs with recalcitrant leaf litter across cold biomes would constitute a negative feedback to global warming. Depending on the strength of other (previously reported) positive feedbacks of shrub expansion on soil carbon turnover, this may partly counteract direct warming enhancement of litter decomposition.     

Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change

Global environmental change, related to climate change and the deposition of airborne N‐containing contaminants, has already resulted in shifts in plant community composition among plant functional types in Arctic and temperate alpine regions. In this paper, we review how key ecosystem processes will be altered by these transformations, the complex biological cascades and feedbacks that might result, and some of the potential broader consequences for the earth system. Firstly, we consider how patterns of growth and allocation, and nutrient uptake, will be altered by the shifts in plant dominance. The ways in which these changes may disproportionately affect the consumer communities, and rates of decomposition, are then discussed. We show that the occurrence of a broad spectrum of plant growth forms in these regions (from cryptogams to deciduous and evergreen dwarf shrubs, graminoids and forbs), together with hypothesized low functional redundancy, will mean that shifts in plant dominance result in a complex series of biotic cascades, couplings and feedbacks which are supplemental to the direct responses of ecosystem components to the primary global change drivers. The nature of these complex interactions is highlighted using the example of the climate‐driven increase in shrub cover in low‐Arctic tundra, and the contrasting transformations in plant functional composition in mid‐latitude alpine systems. Finally, the potential effects of the transformations on ecosystem properties and processes that link with the earth system are reviewed. We conclude that the effects of global change on these ecosystems, and potential climate‐change feedbacks, cannot be predicted from simple empirical relationships between processes and driving variables. Rather, the effects of changes in species distributions and dominances on key ecosystem processes and properties must also be considered, based upon best estimates of the trajectories of key transformations, their magnitude and rates of change.     

Abrupt shifts in African savanna tree cover along a climatic gradient

Aim To describe patterns of tree cover in savannas over a climatic gradient and a range of spatial scales and test if there are identifiable climate‐related mean structures, if tree cover always increases with water availability and if there is a continuous trend or a stepwise trend in tree cover. Location Central Tropical Africa. Methods We compared a new analysis of satellite tree cover data with botanical, phytogeographical and environmental data. Results Along the climatic transect, six vegetation structures were distinguished according to their average tree cover, which can co‐occur as mosaics. The resulting abrupt shifts in tree cover were not correlated to any shifts in either environmental variables or in tree species distributions. Main conclusions A strong contrast appears between fine‐scale variability in tree cover and coarse‐scale structural states that are stable over several degrees of latitude. While climate parameters and species pools display a continuous evolution along the climatic gradient, these stable structural states have discontinuous transitions, resulting in regions containing mosaics of alternative stable states. Soils appear to have little effect inside the climatic stable state domains but a strong action on the location of the transitions. This indicates that savannas are patch dynamics systems, prone to feedbacks stabilizing their coarse‐scale structure over wide ranges of environmental conditions.     

Evolutionary regime shifts in simulated ecosystems

Catastrophic regime shifts in ecosystems occur when the system is tipped into a new attractor state under some external forcing. Here we consider whether evolutionary adaptations within ecosystems can trigger similar transitions. We use an individual‐based, evolutionary model of interconnected ecosystems to analyze nonlinear changes in global state resulting from local adaptations. Transitions between periods of stability occur when new traits arise that allow exploitation of under‐utilized resources. Subsequent rapid growth of the population carrying the new trait causes abrupt environmental change that drives incumbent species extinct. We call these transitions ‘evolutionary regime shifts’. These internally generated perturbations can result in ecosystem collapse, followed by recovery to an alternate stable state, or occasionally system‐wide extinction. While these disruptions may have a negative impact on ecosystem productivity in individual simulation runs, mean results over many simulations show a trend for increasing ecosystem productivity and stability over time. Feedback between life and the abiotic environment in the model creates a ‘long‐tailed’ distribution of extinction sizes without any external trigger for large extinction events.     

Ecosystems off track: rate‐induced critical transitions in ecological models

Theory suggests that gradual environmental change may erode the resilience of ecosystems and increase their susceptibility to critical transitions. This notion has received a lot of attention in ecology in recent decades. An important question receiving far less attention is whether ecosystems can cope with the rapid environmental changes currently imposed. The importance of this question was recently highlighted by model studies showing that elevated rates of change may trigger critical transitions, whereas slow environmental change would not. This paper aims to provide a mechanistic understanding of these rate‐induced critical transitions to facilitate identification of rate sensitive ecosystems. Analysis of rate sensitive ecological models is challenging, but we demonstrate how rate‐induced transitions in an elementary model can still be understood. Our analyses reveal that rate‐induced transitions 1) occur if the rate of environmental change is high compared to the response rate of ecosystems, 2) are driven by rates, rather than magnitudes, of change and 3) occur once a critical rate of change is exceeded. Disentangling rate‐induced transitions from classical transitions in observations would be challenging. However, common features of rate‐sensitive models suggest that ecosystems with coupled fast–slow dynamics, exhibiting repetitive catastrophic shifts or displaying periodic spatial patterns are more likely to be rate sensitive. Our findings are supported by experimental studies showing rate‐dependent outcomes. Rate sensitivity of models suggests that the common definition of ecological resilience is not suitable for a subset of real ecosystems and that formulating limits to magnitudes of change may not always safeguard against ecosystem degradation. Synthesis Understanding and predicting ecosystem response to environmental change is one of the key challenges in ecology. Model studies have suggested that slow, gradual environmental change beyond some critical threshold can trigger so‐called critical transitions and abrupt ecosystem degradation. An important question remains however whether ecosystems can cope with the ongoing rapid anthropogenic environmental changes to which they are currently imposed. In this study we demonstrate that in some ecological models elevated rates of change can trigger critical transitions even if slow environmental change of the same magnitude would not. Such rateinduced critical transitions in models suggest that concepts like resilience and planetary boundaries may not always be sufficient to explain and prevent ecosystem degradation.     

Negative feedback processes following drainage slow down permafrost degradation

The sustainability of the vast Arctic permafrost carbon pool under climate change is of paramount importance for global climate trajectories. Accurate climate change forecasts, therefore, depend on a reliable representation of mechanisms governing Arctic carbon cycle processes, but this task is complicated by the complex interaction of multiple controls on Arctic ecosystem changes, linked through both positive and negative feedbacks. As a primary example, predicted Arctic warming can be substantially influenced by shifts in hydrologic regimes, linked to, for example, altered precipitation patterns or changes in topography following permafrost degradation. This study presents observational evidence how severe drainage, a scenario that may affect large Arctic areas with ice‐rich permafrost soils under future climate change, affects biogeochemical and biogeophysical processes within an Arctic floodplain. Our in situ data demonstrate reduced carbon losses and transfer of sensible heat to the atmosphere, and effects linked to drainage‐induced long‐term shifts in vegetation communities and soil thermal regimes largely counterbalanced the immediate drainage impact. Moreover, higher surface albedo in combination with low thermal conductivity cooled the permafrost soils. Accordingly, long‐term drainage effects linked to warming‐induced permafrost degradation hold the potential to alleviate positive feedbacks between permafrost carbon and Arctic warming, and to slow down permafrost degradation. Self‐stabilizing effects associated with ecosystem disturbance such as these drainage impacts are a key factor for predicting future feedbacks between Arctic permafrost and climate change, and, thus, neglect of these mechanisms will exaggerate the impacts of Arctic change on future global climate projections.     

Nutrient enrichment weakens the stabilizing effect of species richness

With global freshwater biodiversity declining at an even faster rate than in the most disturbed terrestrial ecosystems, understanding the effects of changing environmental conditions on relationships between biodiversity and the variability of community and population processes in aquatic ecosystems is of significant interest. Evidence is accumulating that biodiversity loss results in more variable communities; however, the mechanisms underlying this effect have been the subject of considerable debate. We manipulated species richness and nutrients in outdoor aquatic microcosms composed of naturally occurring assemblages of zooplankton and benthic invertebrates to determine how the relationship between species richness and variability might change under different nutrient conditions. Temporal variability of populations and communities decreased with increasing species richness in low nutrient microcosms. In contrast, we found no relationship between species richness and either population or community variability in nutrient enriched microcosms. Of the different mechanisms we investigated (e.g. overyielding, statistical averaging, insurance effects, and the stabilizing effect of species richness on populations) the only one that was consistent with our results was that increases in species richness led to more stable community abundances through the stabilizing effect of species richness on the component populations. While we cannot conclusively determine the mechanism(s) by which species richness stabilized populations, our results suggest that more complete resource-use in the more species-rich low nutrient communities may have dampened population fluctuations.     

Allometric scaling enhances stability in complex food webs

Classic local stability theory predicts that complex ecological networks are unstable and are unlikely to persist despite empiricists' abundant documentation of such complexity in nature. This contradiction has puzzled biologists for decades. While some have explored how stability may be achieved in small modules of a few interacting species, rigorous demonstrations of how large complex and ecologically realistic networks dynamically persist remain scarce and inadequately understood. Here, we help fill this void by combining structural models of complex food webs with nonlinear bioenergetic models of population dynamics parameterized by biological rates that are allometrically scaled to populations' average body masses. Increasing predator–prey body mass ratios increase population persistence up to a saturation level that is reached by invertebrate and ectotherm vertebrate predators when being 10 or 100 times larger than their prey respectively. These values are corroborated by empirical predator–prey body mass ratios from a global data base. Moreover, negative effects of diversity (i.e. species richness) on stability (i.e. population persistence) become neutral or positive relationships at these empirical ratios. These results demonstrate that the predator–prey body mass ratios found in nature may be key to enabling persistence of populations in complex food webs and stabilizing the diversity of natural ecosystems.     

植物——土壤反馈研究进展

该文综述了植物—土壤反馈研究的定义、途经、方法和国内外的研究现状以及存在的问题。植物—土壤反馈是指植物改变根际土壤的生物和非生物特征,同时被改变的也能提高或降低该植物的生长,形成正的或负的反馈,从而影响植物群落组成及植物间相互作用。植物—土壤反馈研究对于理解植物群落演替、生态系统多样性与生产力形成与维持机制,认识生态系统对气候变化和生物入侵等全球生态事件的响应具有重要的理论意义。外来物种快速生长和繁殖及其可能的负反馈可能会导致本地种被竞争排除,未来气候变化可能导致物种组成发生变化及生物多样性丢失,但资源互补和植物—土壤反馈效应则可能使植物群落具有较高的生产力和多样性。因此,未来植物—土壤反馈关系应该加强以下几方面研究:(1)开展不同生态系统植物—土壤反馈关系的比较研究;(2)植物—土壤及土壤—植物等群落水平的反馈研究;(3)特别是要加强分子和基因工具在植物土壤—反馈关系中的应用,揭示植物—土壤反馈关系的分子机理。     

Competing associations in bacterial warfare with two toxins

Simple combinations of common competitive mechanisms can easily result in cyclic competitive dominance relationships between species. The topological features of such competitive networks allow for complex spatial coexistence patterns. We investigate self-organization and coexistence in a lattice model, describing the spatial population dynamics of competing bacterial strains. With increasing diffusion rate the community of the nine possible toxicity/resistance types undergoes two phase transitions. Below a critical level of diffusion, the system exhibits expanding domains of three different defensive alliances, each consisting of three cyclically dominant species. Due to the neutral relationship between these alliances and the finite system size effect, ultimately only one of them remains. At large diffusion rates the system admits three coexisting domains, each containing mutually neutral species. Because of the cyclical dominance between these domains, a long term stable coexistence of all species is ensured. In the third phase at intermediate diffusion the spatial structure becomes even more complicated with domains of mutually neutral species persisting along the borders of defensive alliances. The study reveals that cyclic competitive relationships may produce a large variety of complex coexistence patterns, exhibiting common features of natural ecosystems, like hierarchical organization, phase transitions and sudden, large-scale fluctuations.     

From Transient to Steady-state Response of Ecosystems to Atmospheric CO2-enrichment and Global Climate Change: Conceptual Challenges and Need for an Integrated Approach

Evidence continues to accumulate that humans are significantly increasing atmospheric CO₂ concentrations, resulting in unprecedented changes in the global climate system. Experimental manipulations of terrestrial ecosystems and their components have greatly increased our understanding of short-term responses to these global perturbations and have provided valuable input to ecosystem, dynamic vegetation, and global scale models. However, concerns exist that these initial experimental responses may be transitory, thereby limiting our ability to extrapolate short-term experimental responses to infer longer-term effects. To do these extrapolations, it will be necessary to understand changes in response patterns over time, including alterations in the magnitude, direction, and rate of change of the responses. These issues represent one of our largest challenges in accurately predicting longer-term changes in ecosystems and associated feedbacks to the climate system. Key issues that need to be considered when designing future experiments or refining models include: linear vs. non-linear responses, direct vs. indirect effects, lags in response, acclimation, resource limitation, homeostasis, buffers, thresholds, ecosystem stoichiometry, turnover rates and times, and alterations in species composition. Although experimental and landscape evidence for these response patterns exist, extrapolating longer-term response patterns from short-term experiments will ultimately require a unified multidisciplinary approach, including better communication and collaboration between theoreticists, experimentalists and modelers.     

Coastal eutrophication research: a new awareness

An analysis of the contents and conclusions of the papers contained in this issue (Hydrobiologia Volume xxx) suggests that a new vision is taking shape that may correspond to an emerging new paradigm in the way we understand and manage coastal eutrophication. This new paradigm emphasizes its global dimension and the connections with other global environmental pressures, and re-evaluates the targets of remedial actions and policies. Eutrophication research must evolve toward a more integrative, ecosystem perspective which requires that it be extended to include impacts beyond primary producers and to examine possible cascading effects and feedbacks involving other components of the ecosystem. A quantitative framework that incorporates the interacting top-down and bottom-up effects in eutrophication models must be urgently developed to guide diagnostics and establish targets to mitigate coastal eutrophication. The required macroscopic view must also be extended to the managerial and policy frameworks addressing eutrophication, through the development of policies that examine activities in the environment in an integrative, rather than sectorial, manner. Recent evidence of complex responses of coastal ecosystems to nutrient reduction requires that management targets, and the policies that support them, be reconsidered to recognize the complexities of the responses of coastal ecosystems to reduced nutrient inputs, including non-linear responses and associated thresholds. While a predictive framework for the complex trajectories of coastal ecosystems subject to changes in nutrient inputs is being developed, the assessment of managerial actions should be reconsidered to focus on the consideration of the status achieved as the outcome of nutrient reduction plans against that possibly derived from a ‘do nothing’ scenario. A proper assessment of eutrophication and the efforts to mitigate it also requires that eutrophication be considered as a component of global change, in addressing both its causes and its consequences, and that the feedbacks between other components of global change (e.g., climate change, overfishing, altered biogeochemical cycles, etc.) be explicitly considered in designing eutrophication research and in managing the problem. Guest editors: J. H. Andersen & D. J. Conley Eutrophication in Coastal Ecosystems: Selected papers from the Second International Symposium on Research and Management of Eutrophication in Coastal Ecosystems, 20–23 June 2006, Nyborg, Denmark     

Effects of plant species on nutrient cycling

Plant species create positive feedbacks to patterns of nutrient cycling in natural ecosystems. For example, in nutrient-poor ecosystems, plants grow slowly, use nutrients efficiently and produce poor-quality litter that decomposes slowly and deters herbivores. /n contrast, plant species from nutrient-rich ecosystems grow rapidly, produce readily degradable litter and sustain high rates of herbivory, further enhancing rates of nutrient cycling. Plants may also create positive feedbacks to nutrient cycling because of species' differences in carbon deposition and competition with microbes for nutrients in the rhizosphere. New research is showing that species' effects can be as or more important than abiotic factors, such as climate, in controlling ecosystem fertility.     

Climate change feedbacks to microbial decomposition in boreal soils

Boreal ecosystems store 10–20 % of global soil carbon and may warm by 4–7 °C over the next century. Higher temperatures could increase the activity of boreal decomposers and indirectly affect decomposition through other ecosystem feedbacks. For example, permafrost melting will likely alleviate constraints on microbial decomposition and lead to greater soil CO₂ emissions. However, wet boreal ecosystems underlain by permafrost are often CH₄ sources, and permafrost thaw could ultimately result in drier soils that consume CH₄, thereby offsetting some of the greenhouse warming potential of soil CO₂ emissions. Climate change is also likely to increase winter precipitation and snow depth in boreal regions, which may stimulate decomposition by moderating soil temperatures under the snowpack. As temperatures and evapotranspiration increase in the boreal zone, fires may become more frequent, leading to additional permafrost loss from burned ecosystems. Although post-fire decomposition could also increase due to higher soil temperatures, reductions in microbial biomass and activity may attenuate this response. Other feedbacks such as soil drying, increased nutrient mineralization, and plant species shifts are either weak or uncertain. We conclude that strong positive feedbacks to decomposition will likely depend on permafrost thaw, and that climate feedbacks will probably be weak or negative in boreal ecosystems without permafrost. However, warming manipulations should be conducted in a broader range of boreal systems to validate these predictions.     

Multiple feedbacks and the prevalence of alternate stable states on coral reefs

The prevalence of alternate stable states on coral reefs has been disputed, although there is universal agreement that many reefs have experienced substantial losses of coral cover. Alternate stable states require a strong positive feedback that causes self-reinforcing runaway change when a threshold is passed. Here we use a simple model of the dynamics of corals, macroalgae and herbivores to illustrate that even weak positive feedbacks that individually cannot lead to alternate stable states can nonetheless do so if they act in concert and reinforce each other. Since the strength of feedbacks varies over time and space, our results imply that we should not reject or accept the general hypothesis that alternate stable states occur in coral reefs. Instead, it is plausible that shifts between alternate stable states can occur sporadically, or on some reefs but not others depending on local conditions. Therefore, we should aim at a better mechanistic understanding of when and why alternate stable states may occur. Our modelling results point to an urgent need to recognize, quantify, and understand feedbacks, and to reorient management interventions to focus more on the mechanisms that cause abrupt transitions between alternate states.     

Gradual regime shifts in spatially extended ecosystems

Ecosystem regime shifts are regarded as abrupt global transitions from one stable state to an alternative stable state, induced by slow environmental changes or by global disturbances. Spatially extended ecosystems, however, can also respond to local disturbances by the formation of small domains of the alternative state. Such a response can lead to gradual regime shifts involving front propagation and the coalescence of alternative-state domains. When one of the states is spatially patterned, a multitude of intermediate stable states appears, giving rise to step-like gradual shifts with extended pauses at these states. Using a minimal model, we study gradual state transitions and show that they precede abrupt transitions. We propose indicators to probe gradual regime shifts, and suggest that a combination of abrupt-shift indicators and gradual-shift indicators might be needed to unambiguously identify regime shifts. Our results are particularly relevant to desertification in drylands where transitions to bare soil take place from spotted vegetation, and the degradation process appears to involve step-like events of local vegetation mortality caused by repeated droughts.     

Resource subsidies between stream and terrestrial ecosystems under global change

Streams and adjacent terrestrial ecosystems are characterized by permeable boundaries that are crossed by resource subsidies. Although the importance of these subsidies for riverine ecosystems is increasingly recognized, little is known about how they may be influenced by global environmental change. Drawing from available evidence, in this review we propose a conceptual framework to evaluate the effects of global change on the quality and spatiotemporal dynamics of stream–terrestrial subsidies. We illustrate how changes to hydrological and temperature regimes, atmospheric CO₂ concentration, land use and the distribution of nonindigenous species can influence subsidy fluxes by affecting the biology and ecology of donor and recipient systems and the physical characteristics of stream–riparian boundaries. Climate‐driven changes in the physiology and phenology of organisms with complex life cycles will influence their development time, body size and emergence patterns, with consequences for adjacent terrestrial consumers. Also, novel species interactions can modify subsidy dynamics via complex bottom‐up and top‐down effects. Given the seasonality and pulsed nature of subsidies, alterations of the temporal and spatial synchrony of resource availability to consumers across ecosystems are likely to result in ecological mismatches that can scale up from individual responses, to communities, to ecosystems. Similarly, altered hydrology, temperature, CO₂ concentration and land use will modify the recruitment and quality of riparian vegetation, the timing of leaf abscission and the establishment of invasive riparian species. Along with morphological changes to stream–terrestrial boundaries, these will alter the use and fluxes of allochthonous subsidies associated with stream ecosystems. Future research should aim to understand how subsidy dynamics will be affected by key drivers of global change, including agricultural intensification, increasing water use and biotic homogenization. Our conceptual framework based on the match–mismatch between donor and recipient organisms may facilitate understanding of the multiple effects of global change and aid in the development of future research questions.