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.     

River ecosystem resilience to extreme flood events

Floods have a major influence in structuring river ecosystems. Considering projected increases in high‐magnitude rainfall events with climate change, major flooding events are expected to increase in many regions of the world. However, there is uncertainty about the effect of different flooding regimes and the importance of flood timing in structuring riverine habitats and their associated biotic communities. In addition, our understanding of community response is hindered by a lack of long‐term datasets to evaluate river ecosystem resilience to flooding. Here we show that in a river ecosystem studied for 30 years, a major winter flood reset the invertebrate community to a community similar to one that existed 15 years earlier. The community had not recovered to the preflood state when recurrent summer flooding 9 years later reset the ecosystem back to an even earlier community. Total macroinvertebrate density was reduced in the winter flood by an order of magnitude more than the summer flood. Meiofaunal invertebrates were more resilient to the flooding than macroinvertebrates, possibly due to their smaller body size facilitating greater access to in‐stream refugia. Pacific pink salmon escapement was markedly affected by the winter flood when eggs were developing in redds, compared to summer flooding, which occurred before the majority of eggs were laid. Our findings inform a proposed conceptual model of three possible responses to flooding by the invertebrate community in terms of switching to different states and effects on resilience to future flooding events. In a changing climate, understanding these responses is important for river managers to mitigate the biological impacts of extreme flooding effects.     

Quantifying resilience of multiple ecosystem services and biodiversity in a temperate forest landscape

Resilience is increasingly being considered as a new paradigm of forest management among scientists, practitioners, and policymakers. However, metrics of resilience to environmental change are lacking. Faced with novel disturbances, forests may be able to sustain existing ecosystem services and biodiversity by exhibiting resilience, or alternatively these attributes may undergo either a linear or nonlinear decline. Here we provide a novel quantitative approach for assessing forest resilience that focuses on three components of resilience, namely resistance, recovery, and net change, using a spatially explicit model of forest dynamics. Under the pulse set scenarios, we explored the resilience of nine ecosystem services and four biodiversity measures following a one‐off disturbance applied to an increasing percentage of forest area. Under the pulse + press set scenarios, the six disturbance intensities explored during the pulse set were followed by a continuous disturbance. We detected thresholds in net change under pulse + press scenarios for the majority of the ecosystem services and biodiversity measures, which started to decline sharply when disturbance affected >40% of the landscape. Thresholds in net change were not observed under the pulse scenarios, with the exception of timber volume and ground flora species richness. Thresholds were most pronounced for aboveground biomass, timber volume with respect to the ecosystem services, and ectomycorrhizal fungi and ground flora species richness with respect to the biodiversity measures. Synthesis and applications. The approach presented here illustrates how the multidimensionality of stability research in ecology can be addressed and how forest resilience can be estimated in practice. Managers should adopt specific management actions to support each of the three components of resilience separately, as these may respond differently to disturbance. In addition, management interventions aiming to deliver resilience should incorporate an assessment of both pulse and press disturbances to ensure detection of threshold responses to disturbance, so that appropriate management interventions can be identified.     

A new approach for rapid detection of nearby thresholds in ecosystem time series

Massive changes to ecosystems sometimes cross thresholds from which recovery can be difficult, expensive and slow. These thresholds are usually discovered in post hoc analyses long after the event occurred. Anticipating these changes prior to their occurrence could give managers a chance to intervene. Here we present a novel approach for anticipating ecosystem thresholds that combines resilience indicators with Quickest detection of change points. Unlike existing methods, the Quickest detection method is updated every time a data point arrives, and minimizes the time to detect an approaching threshold given the users’ tolerance for false alarms. The procedure accurately detected an impending regime shift in an experimentally manipulated ecosystem. An ecosystem model was used to determine if the method can detect an approaching threshold soon enough to prevent a regime shift. When the monitored variable was directly involved in the interaction that caused the regime shift, detection was quick enough to avert collapse. When the monitored variable was only indirectly linked to the critical transition, detection came too late. The procedure is useful for assessing changes in resilience as ecosystems approach thresholds. However some thresholds cannot be detected in time to prevent regime shifts, and surprises will be inevitable in ecosystem management.     

区县尺度下的河流生态系统健康评价——以北京房山区为例

伴随郊区城市化的快速发展,高强度人工管控对河流生态系统产生持续胁迫作用,河流生态系统健康受到严重威胁甚至损害。以北京房山区河流水系为例,于2016—2017年在区域内的31处样点进行野外调查。选取河流的水环境功能、防洪效益功能、生态效益功能和支持利用功能4方面16个具体评价指标,并通过层次分析法计算权重,构建区县尺度下的河流生态系统健康评价指标体系,并进行健康状况评价。结果表明:房山区河流的防洪效益功能与生态效益功能中"健康与亚健康"状态比例分别为58.0%与41.9%,而水环境功能与和支持利用功能中"较差与差"状态比例为71.0%与41.9%;河流生态系统健康整体水平较差,勉强满足河流生态系统一般需求。同时,房山区境内的拒马河与小清河的河流生态系统健康状态远优于大石河与永定河。河流生态基流是影响河流生态系统健康的关键因素,适当释放库区截留水量与调配区外清洁水体,已成为恢复房山区河流生态系统健康的前提与基础。     

火干扰与生态系统的碳循环

火干扰是陆地生态系统碳循环的重要影响因子。它改变着整个系统的碳循环过程与碳分布格局。正确评估火干扰在碳循环过程中的作用,对推进全球碳循环研究有着重要的意义。从4个方面系统的回顾了火干扰对碳循环的影响过程及其研究方法:(1)火烧过程中含碳痕量气体排放的估算;(2)火烧迹地恢复过程中净第一性生产力(NPP)与土壤呼吸的变化;(3)火干扰对生态系统碳源/汇的影响;(4)模型方法在火干扰与生态系统碳循环研究中的应用。目前火灾碳排量的估算方法业已成熟,但进行更精确的估算必须基于对受干扰生态系统的性质以及火势的时空变异性质的准确理解;相比之下,对于间接的、更为重要的影响,即对火烧迹地恢复过程中碳循环变化的研究则显不足。由于数据缺乏,现有研究大多限于对碳循环某一方面的观测与定量描述,缺乏全面的机理性分析。对此,实地观测、模型模拟与遥感观测的跨尺度集成将成为未来火干扰研究的一个主要方向。     

Biodiversity and ecosystem productivity: implications for carbon storage

Recent experiments have found that Net Primary Productivity (NPP) can often be a positive saturating function of plant species and functional diversity. These findings raised the possibility that more diverse ecosystems might store more carbon as a result of increased photosynthetic inputs. However, carbon inputs will not only remain in plant biomass, but will be translocated to the soil via root exudation, fine root turnover, and litter fall. Thus, we must consider not just plant productivity (NPP), but also net productivity of the whole ecosystem (NEP), which itself measures net carbon storage. We currently know little about how plant diversity could influence soil processes that return carbon back to the atmosphere, such as heterotrophic respiration and decomposition of organic matter. Nevertheless, it is clear that any effects on such processes could make NPP a poor predictor of whole-ecosystem productivity, and potentially the ability of the ecosystem to store carbon. We examine the range of mechanisms by which plant diversity could influence net ecosystem productivity, incorporating processes involved with carbon uptake (productivity), loss (autotrophic and heterotrophic respiration), and residence time within the system (decomposition rate). Understanding the relationship between plant diversity and ecosystem carbon dynamics must be made a research priority if we wish to provide information relevant to global carbon policy decisions. This goal is entirely feasible if we utilize some basic methods for measuring the major fluxes of carbon into and out of the ecosystem.     

Disturbance frequency and functional identity mediate ecosystem processes in prairie streams

A major consequence of climate change will be the alteration of precipitation patterns and concomitant changes in the flood frequencies in streams. Species losses or introductions will accompany these changes, which necessitates understanding the interactions between altered disturbance regimes and consumer functional identity to predict dynamics of streams. We used experimental mesocosms and field enclosures to test the interactive effects of flood frequency and two fishes from distinct consumer groups (benthic grazers and water-column minnows) on recovery of stream ecosystem properties (algal form and biomass, invertebrate densities, metabolism and nutrient uptake rates). Our results generally suggest that periphyton communities under nutrient limitation are likely to recover more quickly when grazing and water-column minnows are present and these effects can diminish or reverse with time since the disturbance. We hypothesized that increased periphyton production and biomass was the result of increased nutrient turnover, but decreased light limitation and indirect effects on other trophic levels are alternative explanations. Recovery of stream ecosystem properties after a natural flood differed from mesocosms (e.g. lower algal biomass and no long algal filaments present) and species manipulations did not explain recovery of ecosystem properties; rather, ecosystem processes varied along a downstream gradient of increasing temperature and nutrient concentrations. Different results between field enclosures and experimental mesocosms are attributable to a number of factors including differences in algal and invertebrate communities in the natural stream and relatively short enclosure lengths (mean area=35.8 m²) compared with recirculating water in the experimental mesocosms. These differences may provide insight into conditions necessary to elicit a strong interaction between consumers and ecosystem properties.     

Mechanistic insights into the role of large carnivores for ecosystem structure and functioning

Large carnivores can exert top–down effects in ecosystems, but the size of these effects are largely unknown. Empirical investigation on the importance of large carnivores for ecosystem structure and functioning presents a number of challenges due to the large spatio-temporal scale and the complexity of such dynamics. Here, we applied a mechanistic global ecosystem model to investigate the influence of large-carnivore removal from undisturbed ecosystems. First, we simulated large-carnivore removal on the global scale to inspect the geographic pattern of top–down control and to disentangle the functional role of large carnivores in top–down control in different environmental contexts. Second, we conducted four small-scale ecosystem simulation experiments to understand direct and indirect changes in food-web structure under different environmental conditions. We found that the removal of top–down control exerted by large carnivores (> 21 kg) can trigger large trophic cascades, leading to an overall decrease in autotroph biomass globally. Furthermore, the loss of large carnivores resulted in an increase of mesopredators. The magnitude of these changes was positively related to primary productivity (NPP), in line with the ‘exploitation ecosystem hypothesis’. In addition, we found that seasonality in NPP dampened the magnitude of change following the removal of large carnivores. Our results reinforce the idea that large carnivores play a fundamental role in shaping ecosystems, and further declines and extinctions can trigger substantial ecosystem responses. Our findings also support previous studies suggesting that natural ecosystem dynamics have been severely modified and are still changing as a result of the widespread decline and extinction of large carnivores.     

典型森林和草地生态系统呼吸各组分间的相互关系

生态系统呼吸是陆地生态系统碳收支的重要组成部分,分析其组分间的相互关系对理解生态系统呼吸过程和精确评价生态系统碳收支具有重要意义,也是当前碳循环研究工作的一大难点。本研究利用ChinaFLUX的长白山温带针阔混交林（CBS）,鼎湖山亚热带常绿阔叶林（DHS）和海北灌丛草甸（HBGC）三个典型生态系统的通量观测数据,采用经验统计方法,分析了其在中国典型生态系统中的适用性及敏感性,揭示了生态系统呼吸各组分的动态变化特征及相互关系。结果表明：采用本研究中的呼吸组分拆分方法所获结果与理论推测及实测数据大致相同,拆分结果对净初级生产力与总初级生产力的比值（NPP/GPP）较为敏感,NPP/GPP变化0.1时,自养呼吸在生态系统呼吸中的比例（Ra/RE）改变0.05。各生态系统中,生态系统呼吸及其组分在年内均表现出明显的单峰型变化特征,在夏季生长旺盛的时节达到最大值。异养呼吸与生态系统呼吸的比值（Rh/RE）也具有明显的季节变化,但在生态系统间表现出明显差异,CBS和HBGC分别表现出先增大后减小和先减小后增大的变化趋势,DHS则相对稳定,在0.5附近波动, Ra/RE的季节动态与Rh/RE相反。在年总量上,HBGC主要通过异养呼吸向大气排放CO2,异养呼吸占生态系统呼吸的60%,而CBS和DHS的自养呼吸和异养呼吸所占比重大致相似,异养呼吸占生态系统呼吸的49%。这说明,该统计学模型可以用来进行生态系统呼吸组分的拆分,进而可以为生态系统碳循环过程的精细研究提供参考数据,但今后应加强NPP/GPP的测定,以提高生态系统呼吸拆分的精度。     

River beads as a conceptual framework for building carbon storage and resilience to extreme climate events into river management

River beads refer to retention zones within a river network that typically occur within wider, lower gradient segments of the river valley. In lowland, floodplain rivers that have been channelized and leveed, beads can also be segments of the river in which engineering has not reduced lateral channel mobility and channel-floodplain connectivity. Decades of channel engineering and flow regulation have reduced the spatial heterogeneity and associated ecosystem functions of beads occurring throughout river networks from headwaters to large, lowland rivers. We discuss the processes that create and maintain spatial heterogeneity within river beads, including examples of beads along mountain streams of the Southern Rockies in which large wood and beaver dams are primary drivers of heterogeneity. We illustrate how spatial heterogeneity of channels and floodplains within beads facilitates storage of organic carbon; retention of water, solutes, sediment, and particulate organic matter; nutrient uptake; biomass and biodiversity; and resilience to disturbance. We conclude by discussing the implications of river beads for understanding solute and particulate organic matter dynamics within river networks and the implications for river management. We also highlight gaps in current understanding of river form and function related to river beads. River beads provide an example of how geomorphic understanding of river corridor form and process can be used to restore retention and resilience within human-altered river networks.     

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.     

Ecological effects of river ice break-up: a review and perspective

1. Abiotic disturbances strongly modify spatial and temporal patterns of lotic ecosystem community structure and function. Such effects are produced because disturbances alter organic matter, nutrient and contaminant dynamics and the distribution and abundance of bacterial, algal, macroinvertebrate and fish communities. 2. River ice break-up is a seasonal disturbance in rivers at high altitudes and latitudes world-wide and is characterized, in part, by large increases in current velocity, stage, water temperature, concentrations of suspended materials and substrate scouring. 3. These abiotic factors are likely to have important effects on primary producers, consumers, and food-web dynamics of river biota. Despite the potential importance of river ice break-up on community structure and function, detailed information describing the magnitude of their effects and underlying causal mechanisms is scarce. 4. The objective of this paper is to provide a hydrological and ecological review and perspective on the potential effects of ice break-up on lotic ecosystems. Specifically, the potential importance of break-up on water temperature, river sediments and geomorphology, riverine energy sources, contaminants, and its effects on river biota and food-web dynamics are evaluated.     

Demographic noise and resilience in a semi-arid ecosystem model

The scarcity of water characterising drylands forces vegetation to adopt appropriate survival strategies. Some of these generate water–vegetation feedback mechanisms that can lead to spatial self-organisation of vegetation, as it has been shown with models representing plants by a density of biomass, varying continuously in time and space. However, although plants are usually quite plastic they also display discrete qualities and stochastic behaviour. These features may give rise to demographic noise, which in certain cases can influence the qualitative dynamics of ecosystem models. In the present work we explore the effects of demographic noise on the resilience of a model semi-arid ecosystem. We introduce a spatial stochastic eco-hydrological hybrid model in which plants are modelled as discrete entities subject to stochastic dynamical rules, while the dynamics of surface and soil water are described by continuous variables. The model has a deterministic approximation very similar to previous continuous models of arid and semi-arid ecosystems. By means of numerical simulations we show that demographic noise can have important effects on the extinction and recovery dynamics of the system. In particular we find that the stochastic model escapes extinction under a wide range of conditions for which the corresponding deterministic approximation predicts absorption into desert states.     

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.     

Productivity and Connectivity in Tropical Riverscapes of Northern Australia: Ecological Insights for Management

Flow regimes are fundamental to sustaining ecological characteristics of rivers worldwide, including their associated floodplains. Recent advances in understanding tropical river–floodplain ecosystems suggest that a small set of basic ecological concepts underpins their biophysical characteristics, especially the high levels of productivity, biodiversity and natural resilience. The concepts relate to (1) river-specific flow patterns, (2) processes ‘fuelled’ by a complex of locally generated carbon and nutrients seasonally mixed with carbon and nutrients from floodplains and catchments, (3) seasonal movements of biota facilitated by flood regimes, (4) food webs and overall productivity sustained by hydrological connectivity, (5) fires in the wet/dry tropical floodplains and riparian zones being major consumers of carbon and a key factor in the subsequent redistribution of nutrients, and (6) river–floodplains having inherent resilience to natural variability but only limited resilience to artificial modifications. Understanding these concepts is particularly timely in anticipating the effects of impending development that may affect tropical river–floodplains at the global scale. Australia, a region encompassing some of the last relatively undisturbed tropical riverine landscapes in the world, provides a valuable case study for understanding the productivity, diversity and resilience of tropical river–floodplain systems. However, significant knowledge gaps remain. Despite substantial recent advances in understanding, present knowledge of these highly complex tropical rivers is insufficient to predict many ecological responses to either human-generated or climate-related changes. The major research challenges identified herein (for example, those related to food web structure, nutrient transfers, productivity, connectivity and resilience), if accomplished in the next decade, will offer substantial insights toward assessing and managing ecological changes associated with human alterations to rivers and their catchments.     

The relationship between biodiversity and ecosystem functioning in food webs

Recent theoretical and experimental work provides clear evidence that biodiversity loss can have profound impacts on functioning of natural and managed ecosystems and the ability of ecosystems to deliver ecological services to human societies. Work on simplified ecosystems in which the diversity of a single trophic level is manipulated shows that diversity can enhance ecosystem processes such as primary productivity and nutrient retention. Theory also strongly suggests that biodiversity can act as biological insurance against potential disruptions caused by environmental changes. However, these studies generally concern a single trophic level, primary producers for the most part. Changes in biodiversity also affect ecosystem functioning through trophic interactions. Here we review, through the analysis of a simple ecosystem model, several key aspects inherent in multitrophic systems that may strongly affect the relationship between diversity and ecosystem processes. Our analysis shows that trophic interactions have a strong impact on the relationships between diversity and ecosystem functioning, whether the ecosystem property considered is total biomass or temporal variability of biomass at the various trophic levels. In both cases, food-web structure and trade-offs that affect interaction strength have major effects on these relationships. Multitrophic interactions are expected to make biodiversity–ecosystem functioning relationships more complex and non-linear, in contrast to the monotonic changes predicted for simplified systems with a single trophic level.     

Stability of ecosystem properties in response to above-ground functional group richness and composition

While there has been a rapidly increasing research effort focused on understanding whether and how composition and richness of species and functional groups may determine ecosystem properties, much remains unknown about how these community attributes affect the dynamic properties of ecosystems. We conducted an experiment in 540 mini‐ecosystems in glasshouse conditions, using an experimental design previously shown to be appropriate for testing for functional group richness and composition effects in ecosystems. Artificial communities representing 12 different above‐ground community structures were assembled. These included treatments consisting of monoculture and two‐ and four‐species mixtures from a pool of four plant species; each plant species represented a different functional group. Additional treatments included two herbivore species, either singly or in mixture, and with or without top predators. These experimental units were then either subjected to an experimentally imposed disturbance (drought) for 40 d or left undisturbed. Community composition and drought both had important effects on plant productivity and biomass, and on several below‐ground chemical and biological properties, including those linked to the functioning of the decomposer subsystem. Many of these compositional effects were due to effects both of plant and of herbivore species. Plant functional group richness also exerted positive effects on plant biomass and productivity, but not on any of the below‐ground properties. Above‐ground composition also had important effects on the response of below‐ground properties to drought and thus influenced ecosystem stability (resistance); effects of composition on drought resistance of above‐ground plant response variables and soil chemical properties were weaker and less consistent. Despite the positive effects of plant functional group richness on some ecosystem properties, there was no effect of richness on the resistance of any of the ecosystem properties we considered. Although herbivores had detectable effects on the resistance of some ecosystem properties, there were no effects of the mixed herbivore species treatment on resistance relative to the single species herbivore treatments. Increasing above‐ground food chain length from zero to three trophic levels did not have any consistent effect on the stability of ecosystem properties. There was no evidence of either above‐ground composition or functional group richness affecting the recovery rate of ecosystem properties from drought and hence ecosystem resilience. Our data collectively point to the role of composition (identity of functional group), but not functional group richness, in determining the stability (resistance to disturbance) of ecosystem properties, and indicates that the nature of the above‐ground community can be an important determinant of the consistency of delivery of ecosystem services.