Confronting Feedbacks of Degraded Marine Ecosystems

In many coastal areas, marine ecosystems have shifted into contrasting states having reduced ecosystem services (hereafter called degraded). Such degraded ecosystems may be slow to revert to their original state due to new ecological feedbacks that reinforce the degraded state. A better understanding of the way human actions influence the strength and direction of feedbacks, how different feedbacks could interact, and at what scales they operate, may be necessary in some cases for successful management of marine ecosystems. Here we synthesize interactions of critical feedbacks of the degraded states from six globally distinct biomes: coral reefs, kelp forests, seagrass beds, shallow soft sediments, oyster reefs, and coastal pelagic food webs. We explore to what extent current management captures these feedbacks and propose strategies for how and when (that is, windows of opportunity) to influence feedbacks in ways to break the resilience of the degraded ecosystem states. We conclude by proposing some challenges for future research that could improve our understanding of these issues and emphasize that management of degraded marine states will require a broad social–ecological approach to succeed.     

Contrasting mechanisms of resilience at mesic and semi-arid boundaries of fynbos,a mega-diverse heathland of South Africa

Biome boundaries are expected to be sensitive to changes in climate and disturbance, because it is here that ecological communities are at environmental, ecological or disturbance limits. Using palaeoecology to study ecosystem dynamics at biome boundaries provides opportunities for understanding ecosystem resilience or sensitivity at ecologically meaningful timescales, and under varying climatic and disturbance conditions.The fynbos biome is a megadiverse Mediterranean type shrubland, found only in South Africa, that is threatened by climate change, land-use change and invasion by alien species. We used palaeoecological records from the semi-arid and mesic boundaries of the fynbos biome to test hypotheses regarding ecosystem resilience over timescales of centuries to millennia. We hypothesised that fynbos would expand at its mesic boundary at the expense of afrotemperate forest under drier and / or more fire prone conditions. In contrast, we hypothesised that at the semi-arid boundary, fynbos would expand at the expense of succulent karoo under wetter and cooler and / or more fire-prone conditions. Contrary to our expectations, the fossil pollen record at both biome boundaries showed remarkable stability at centennial - millennial timescales. To explain our results, we generated new hypotheses exploring possible mechanisms that might confer resilience.At the mesic (temperate) boundary, we suggest that decreased seasonality of rainfall during drier phases favoured fire and fynbos persistence, while in wetter periods, increased seasonality of rainfall resulted in enhanced summer drought stress, inhibiting forest expansion. At this boundary, internal reorganisation from grassy to proteoid fynbos states conferred resilience through resistance. At the succulent karoo boundary, we suggest that increased aridity was offset by less seasonality of rainfall, which enhanced biomass and allowed fire to persist, favouring persistence of fynbos. At this boundary, fynbos sensu stricto retreated during arid phases but recovered during climate amelioration, consistent with resilience through recovery. In both cases, this mega-diverse, disturbance-adapted flora provided a range of traits that enabled fynbos to persist despite environmental perturbation. Our findings agree with general observations that for ecosystems in regions of ample resource availability (i.e. at the mesic boundary), biotic interactions and disturbance tend to become more important in ecosystem dynamics, whereas in regions of scarce resources (in this case water scarcity at the semi-arid boundary) abiotic stress is more important. Our findings contribute to debates over the mechanisms that confer resistance and resilience to environmental change. Understanding and conserving the processes and mechanisms underpinning its resilience will be critical to effective conservation planning.     

Scaling up the diversity–resilience relationship with trait databases and remote sensing data: the recovery of productivity after wildfire

Understanding the mechanisms underlying ecosystem resilience – why some systems have an irreversible response to disturbances while others recover – is critical for conserving biodiversity and ecosystem function in the face of global change. Despite the widespread acceptance of a positive relationship between biodiversity and resilience, empirical evidence for this relationship remains fairly limited in scope and localized in scale. Assessing resilience at the large landscape and regional scales most relevant to land management and conservation practices has been limited by the ability to measure both diversity and resilience over large spatial scales. Here, we combined tools used in large‐scale studies of biodiversity (remote sensing and trait databases) with theoretical advances developed from small‐scale experiments to ask whether the functional diversity within a range of woodland and forest ecosystems influences the recovery of productivity after wildfires across the four‐corner region of the United States. We additionally asked how environmental variation (topography, macroclimate) across this geographic region influences such resilience, either directly or indirectly via changes in functional diversity. Using path analysis, we found that functional diversity in regeneration traits (fire tolerance, fire resistance, resprout ability) was a stronger predictor of the recovery of productivity after wildfire than the functional diversity of seed mass or species richness. Moreover, slope, elevation, and aspect either directly or indirectly influenced the recovery of productivity, likely via their effect on microclimate, while macroclimate had no direct or indirect effects. Our study provides some of the first direct empirical evidence for functional diversity increasing resilience at large spatial scales. Our approach highlights the power of combining theory based on local‐scale studies with tools used in studies at large spatial scales and trait databases to understand pressing environmental issues.     

Trophic networks: How do theories link ecosystem structure and functioning to stability properties? A review

In the context of present global changes, interest in understanding how systems respond to anthropogenic environmental pressures and stress has increased. Indices that characterize ecosystem state are helpful tools for the interpretation of ecosystem responses. The central question is how to link these responses to ecosystem structure and functioning and to quantify ecosystem persistence, resistance or resilience. Quantification and characterization of trophic networks by ecological network analysis (ENA) indices is proceeding rapidly, especially in the field of coastal ecology. In this contribution, we review several theories that relate ecosystem structure and function to stability. The structure and functioning of ecosystems change during the maturation of ecosystems. In the first section, the maturation of ecosystems is described using thermodynamics. In the second and third parts of this paper, we define some concepts for analysing structure and functioning of food webs and discuss their relation to stability. In the last section, we describe three ENA indices and their link to stability. We demonstrate that ENA provides powerful tools for describing local stability, combining quantitative and qualitative concepts. However, it remains incomplete for describing real conservation cases that combine local and global stability.     

What is a healthy ecosystem?

Rapid deterioration of the world's major ecosystems has intensified the need for effective environmental monitoring and the development of operational indicators of ecosystem health. Ecosystem health represents a desired endpoint of environmental management, but it requires adaptive, ongoing definition and assessment. We propose that a healthy ecosystem is one that is sustainable – that is, it has the ability to maintain its structure (organization) and function (vigor) over time in the face of external stress (resilience). Various methods to quantify these three ecosystem attributes (vigor, organization, and resilience) are discussed. These attributes are then folded into a comprehensive assessment of ecosystem health. A network analysis based ecosystem health assessment is developed and tested using trophic exchange networks representing several different aquatic ecosystems. Results indicate the potential of such an ecosystem health assessment for evaluating the relative health of similar ecosystems, and quantifying the effects of natural or anthropogenic stress on the health of a particular ecosystem over time.     

土壤微生物群落抵抗力和恢复力研究进展

人类活动的不断加剧使得土壤生态系统承受着环境干扰压力。土壤微生物受到环境干扰的响应程度（抵抗力）及恢复至原来状态的能力（恢复力）决定着土壤生态系统的可持续性。梳理和总结了土壤微生物群落对环境干扰的抵抗力和恢复力方面的研究进展。首先,在介绍土壤微生物群落抵抗力和恢复力概念的基础上,阐述了通过评估微生物群落的结构和功能的变化来系统表征抵抗力和恢复力;随后,分析了最近十年（2012-2021年）有关文献,发现土壤微生物群落的结构和（或）功能在环境干扰后的恢复力总体较弱,但耕作、有机物料添加和轮作等农田管理措施下的响应趋势表现出一定的规律性;继而,从个体水平的休眠和胁迫忍耐、种群水平的生存策略、群落水平的多样性和相互作用以及生态系统水平的历史遗留效应等方面分析了土壤微生物群落抵抗力和恢复力的维持机制;最后,从功能性状、多功能性和植物-土壤微生物整体性对未来研究做出了展望,以期为构建土壤健康评价体系及预测环境干扰对土壤功能的影响提供科学依据。     

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.     

Perturbation and abrupt shift in trophic control of biodiversity and productivity

Ecology is founded on the view that ecosystem properties like biodiversity and productivity change smoothly with changing environmental conditions. However, emerging theory predicts that environmental change may cause abrupt shifts to alternate states. In many ecosystems, top predators play a pivotal role in controlling plant productivity and diversity. Yet it remains uncertain if altering this control shifts systems to alternate states. I report on a test of the hypothesis that loss of predator control of ecosystem function causes abrupt state changes in diversity and productivity. In this meadow ecosystem, predators enhance plant diversity by causing a highly productive, competitively dominant plant species to be suppressed by herbivores. Experimental predator removal caused rapid proliferation of the competitively dominant plant. Moreover, temporally staggered predator reintroductions failed to restore the ecosystem. This loss of resilience confirmed that the ecosystem crossed a critical threshold and entrained into an alternate state.     

Macrofaunal Functional Diversity Provides Resilience to Nutrient Enrichment in Coastal Sediments

The degradation of ecosystems is often associated with losses of large organisms and the concomitant losses of the ecological functions they mediate. Conversely, the resilience of ecosystems to stress is strongly influenced by faunal communities and their impacts on processes. Denitrification in coastal sediments is a process that may provide ecosystem resilience to eutrophication by removing excess bioavailable nitrogen. Here, we conducted a large-scale field experiment to test the effect of macrofaunal community composition on denitrification in response to two levels of nutrient enrichment at 28 sites across a biologically heterogeneous sandflat. After 7 weeks of enrichment, we measured denitrification enzyme activity (DEA) along with benthic macrofaunal community composition and environmental variables. We normalised treatment site specific DEA values by those in ambient sediments (DEA_CN) to reveal the underlying response across the heterogeneous landscape. Nutrient enrichment caused reductions in DEA_CN as well as functional changes in the community; these were both more pronounced under the highest level of nutrient loading (on average DEA_CN was reduced by 34%). The degree of suppression of DEA_CN following moderate nitrogen loading was mitigated by a key bioturbating species, but following high nitrogen loading (which reduced the key species density) the abundance and diversity of other nutrient processing species were the most important factors alleviating negative effects. This study provides a prime example of the context-dependent role of biodiversity in maintaining ecosystem functioning, underlining that different elements of biodiversity can become important as stress levels increase. Our results emphasise that management and conservation strategies require a real-world understanding of the community attributes that facilitate nutrient processing and maintain resilience in coastal ecosystems.     

Ecosystemic structural–functional approach of the state and transition model

Objective: To contribute to the integration of key ecological concepts such as dynamic equilibrium, critical threshold, resistance and resilience to the ‘State and Transition Model’ (STM), in order to apply them in a more feasible way for rangeland management. Methods: Review and discussion of conceptual models and applied literature, including examples of rangeland dynamics. Results and Conclusions: We propose to enhance the STM considering two principal axes: (a) the x axis determined by structural ecosystem changes (vegetation and soil) and (b) the y axis determined by ecosystem functions and/or processes (recruitment, rain use efficiency). These axes define what we will call Structural–Functional State and Transition Model (SFSTM). Both axes of SFSTM make it possible to determine and quantify states and transitions, critical thresholds and to evaluate the resistance and resilience of an ecosystem to a given disturbance. The critical threshold is identified by structural and functional thresholds (x and y axes), thus defining the point where the ecosystem loses its resilience. Furthermore, in the supplementary file we provide examples with field data from Patagonia to illustrate the SFSTM. The proposed SFSTM has large implications for rangeland research and management, facilitating the understanding and integration of key concepts to enhance the STM. The identification of variables to assess structure and processes makes the model more useful.     

Vulnerability of the global terrestrial ecosystems to climate change

Climate change has far‐reaching impacts on ecosystems. Recent attempts to quantify such impacts focus on measuring exposure to climate change but largely ignore ecosystem resistance and resilience, which may also affect the vulnerability outcomes. In this study, the relative vulnerability of global terrestrial ecosystems to short‐term climate variability was assessed by simultaneously integrating exposure, sensitivity, and resilience at a high spatial resolution (0.05°). The results show that vulnerable areas are currently distributed primarily in plains. Responses to climate change vary among ecosystems and deserts and xeric shrublands are the most vulnerable biomes. Global vulnerability patterns are determined largely by exposure, while ecosystem sensitivity and resilience may exacerbate or alleviate external climate pressures at local scales; there is a highly significant negative correlation between exposure and sensitivity. Globally, 61.31% of the terrestrial vegetated area is capable of mitigating climate change impacts and those areas are concentrated in polar regions, boreal forests, tropical rainforests, and intact forests. Under current sensitivity and resilience conditions, vulnerable areas are projected to develop in high Northern Hemisphere latitudes in the future. The results suggest that integrating all three aspects of vulnerability (exposure, sensitivity, and resilience) may offer more comprehensive and spatially explicit adaptation strategies to reduce the impacts of climate change on terrestrial ecosystems.     

Assessment of ecosystem resilience to hydroclimatic disturbances in India

Recent studies have shown an increasing trend in hydroclimatic disturbances like droughts, which are anticipated to become more frequent and intense under global warming and climate change. Droughts adversely affect the vegetation growth and crop yield, which enhances the risks to food security for a country like India with over 1.2 billion people to feed. Here, we compared the response of terrestrial net primary productivity (NPP) to hydroclimatic disturbances in India at different scales (i.e., at river basins, land covers, and climate types) to examine the ecosystems’ resilience to such adverse conditions. The ecosystem water use efficiency (WUE_e: NPP/Evapotranspiration) is an effective indicator of ecosystem productivity, linking carbon (C) and water cycles. We found a significant difference (p < .05) in WUE_e across India at different scales. The ecosystem resilience analysis indicated that most of the river basins were not resilient enough to hydroclimatic disturbances. Drastic reduction in WUE_e under dry conditions was observed for some basins, which highlighted the cross‐biome incapability to withstand such conditions. The ecosystem resilience at land cover and climate type scale did not completely relate to the basin‐scale ecosystem resilience, which indicated that ecosystem resilience at basin scale is controlled by some other ecohydrological processes. Our results facilitate the identification of the most sensitive regions in the country for ecosystem management and climate policy making, and highlight the need for taking sufficient adaptation measures to ensure sustainability of ecosystems.     

Anthropogenic modifications to fire regimes in the wider Serengeti‐Mara ecosystem

Fire is a key driver in savannah systems and widely used as a land management tool. Intensifying human land uses are leading to rapid changes in the fire regimes, with consequences for ecosystem functioning and composition. We undertake a novel analysis describing spatial patterns in the fire regime of the Serengeti‐Mara ecosystem, document multidecadal temporal changes and investigate the factors underlying these patterns. We used MODIS active fire and burned area products from 2001 to 2014 to identify individual fires; summarizing four characteristics for each detected fire: size, ignition date, time since last fire and radiative power. Using satellite imagery, we estimated the rate of change in the density of livestock bomas as a proxy for livestock density. We used these metrics to model drivers of variation in the four fire characteristics, as well as total number of fires and total area burned. Fires in the Serengeti‐Mara show high spatial variability—with number of fires and ignition date mirroring mean annual precipitation. The short‐term effect of rainfall decreases fire size and intensity but cumulative rainfall over several years leads to increased standing grass biomass and fuel loads, and, therefore, in larger and hotter fires. Our study reveals dramatic changes over time, with a reduction in total number of fires and total area burned, to the point where some areas now experience virtually no fire. We suggest that increasing livestock numbers are driving this decline, presumably by inhibiting fire spread. These temporal patterns are part of a global decline in total area burned, especially in savannahs, and we caution that ecosystem functioning may have been compromised. Land managers and policy formulators need to factor in rapid fire regime modifications to achieve management objectives and maintain the ecological function of savannah ecosystems.     

Perspectives for ecosystem management based on ecosystem resilience and ecological thresholds against multiple and stochastic disturbances

Ecosystem resilience is the inherent ability to absorb various disturbances and reorganize while undergoing state changes to maintain critical functions. When ecosystem resilience is sufficiently degraded by disturbances, ecosystem is exposed at high risk of shifting from a desirable state to an undesirable state. Ecological thresholds represent the points where even small changes in environmental conditions associated with disturbances lead to switch between ecosystem states. There is a growing body of empirical evidence for such state transitions caused by anthropogenic disturbances in a variety of ecosystems. However, fewer studies addressed the interaction of anthropogenic and natural disturbances that often force an ecosystem to cross a threshold which an anthropogenic disturbance or a natural disturbance alone would not have achieved. This fact highlights how little is known about ecosystem dynamics under uncertainties around multiple and stochastic disturbances. Here, we present two perspectives for providing a predictive scientific basis to the management and conservation of ecosystems against multiple and stochastic disturbances. The first is management of predictable anthropogenic disturbances to maintain a sufficient level of biodiversity for ensuring ecosystem resilience (i.e., resilience-based management). Several biological diversity elements appear to confer ecosystem resilience, such as functional redundancy, response diversity, a dominant species, a foundation species, or a keystone species. The greatest research challenge is to identify key elements of biodiversity conferring ecosystem resilience for each context and to examine how we can manage and conserve them. The second is the identification of ecological thresholds along existing or experimental disturbance gradients. This will facilitate the development of indicators of proximity to thresholds as well as the understanding of threshold mechanisms. The implementation of forewarning indicators will be critical particularly when resilience-based management fails. The ability to detect an ecological threshold along disturbance gradients should therefore be essential to establish a backstop for preventing the threshold from being crossed. These perspectives can take us beyond simply invoking the precautionary principle of conserving biodiversity to a predictive science that informs practical solutions to cope with uncertainties and ecological surprises in a changing world.     

火在生态系统中的作用

前言火对植被的影响是在近十多年来才被完全确认的。在二十世纪的大部分时间内,火常常被认为是破坏生态系统的非自然因子;然而,植被学家、人类学家、地理学家和其他许多学者都认为北美的土族印地安人和其他许多民族曾广泛地利用火作为土地管理的工具(Van-     

Recent exposure to environmental stochasticity does not determine the demographic resilience of natural populations

Escalating climatic and anthropogenic pressures expose ecosystems worldwide to increasingly stochastic environments. Yet, our ability to forecast the responses of natural populations to this increased environmental stochasticity is impeded by a limited understanding of how exposure to stochastic environments shapes demographic resilience. Here, we test the association between local environmental stochasticity and the resilience attributes (e.g. resistance, recovery) of 2242 natural populations across 369 animal and plant species. Contrary to the assumption that past exposure to frequent environmental shifts confers a greater ability to cope with current and future global change, we illustrate how recent environmental stochasticity regimes from the past 50 years do not predict the inherent resistance or recovery potential of natural populations. Instead, demographic resilience is strongly predicted by the phylogenetic relatedness among species, with survival and developmental investments shaping their responses to environmental stochasticity. Accordingly, our findings suggest that demographic resilience is a consequence of evolutionary processes and/or deep-time environmental regimes, rather than recent-past experiences.     

Species richness facilitates ecosystem resilience in aquatic food webs

1. Many studies indicate that biodiversity in ecosystems affects stability, either by promoting temporal stability of ecosystem attributes or by enhancing ecosystem resistance and resilience to perturbation. The effects on temporal stability are reasonably well understood and documented but effects on resistance and resilience are not. 2. Here, we report results from an aquatic mesocosm experiment in which we manipulated the species richness and composition of aquatic food webs (macrophytes, macro‐herbivores and invertebrate predators), imposed a pulse disturbance (acidification), and monitored the resistance (initial response) and resilience (recovery) of ecosystem productivity and respiration. 3. We found that species‐rich macroinvertebrate communities had higher resilience of whole‐ecosystem respiration, but were not more resistant to perturbations. We also found that resilience and resistance were unaffected by species composition, despite the strong role composition is known to play in determining mean levels of function in these communities. 4. Biodiversity’s effects on resilience were probably mediated through complex pathways affecting phytoplankton and microbial communities (e.g. via changes in nutrient regeneration, grazing or compositional changes) rather than through simpler effects (e.g. insurance effects, enhanced facilitation) although these simpler mechanisms probably played minor roles in enhancing respiration resilience. 5. Current mechanisms for understanding biodiversity’s effects on ecosystem stability have been developed primarily in the context of single‐trophic level communities. These mechanisms may be overly simplistic for understanding the consequences of species richness on ecosystem stability in complex, multi‐trophic food webs where additional factors such as indirect effects and highly variable life‐history traits of species may also be important.     

陆地生态系统临界转换理论及其生态学机制研究进展

随着气候变化和人类活动对陆地生态系统双重扰动的不断加剧,越来越多的研究已经意识到生态系统结构和功能会发生难以预知的突变,并且恢复起来需要很长时间.开发判别典型生态系统临界转换的早期预警模型及理解其生态学机制成为生态学研究的热点.目前,基于跨越多个时空尺度的理论和实验研究,提出了多种预警陆地生态系统临界转换的理论框架和指...