Robustness of early warning signals for catastrophic and non‐catastrophic transitions

Early warning signals (EWS) are statistical indicators that a rapid regime shift may be forthcoming. Their development has given ecologists hope of predicting rapid regime shifts before they occur. Accurate predictions, however, rely on the signals being appropriate to the system in question. Most of the EWS commonly applied in ecology have been studied in the context of one specific type of regime shift (the type brought on by a saddle‐node bifurcation, at which one stable equilibrium point collides with an unstable equilibrium and disappears) under one particular perturbation scheme (temporally uncorrelated noise that perturbs the net population growth rate in a density independent way). Whether and when these EWS can be applied to other ecological situations remains relatively unknown, and certainly underappreciated. We study a range of models with different types of dynamical transitions (including rapid regime shifts) and several perturbation schemes (density‐dependent uncorrelated or temporally‐correlated noise) and test the ability of EWS to warn of an approaching transition. We also test the sensitivity of our results to the amount of available pre‐transition data and various decisions that must be made in the analysis (i.e. the rolling window size and smoothing bandwidth used to compute the EWS). We find that EWS generally work well to signal an impending saddle‐node bifurcation, regardless of the autocorrelation or intensity of the noise. However, EWS do not reliably appear as expected for other types of transition. EWS were often very sensitive to the length of the pre‐transition time series analyzed, and usually less sensitive to other decisions. We conclude that the EWS perform well for saddle‐node bifurcation in a range of noise environments, but different methods should be used to predict other types of regime shifts. As a consequence, knowledge of the mechanism behind a possible regime shift is needed before EWS can be used to predict it.     

Severe drought drives novel community trajectories in desert stream pools

1. Ecological communities can be relatively stable for long periods of time, and then, often as a result of disturbance, transition rapidly to a novel state. When communities fail to recover to pre‐disturbance configurations, they are said to have experienced a regime shift or to be in an alternative stable state. 2. In this 8‐year study, we quantified the effects of complete water loss and subsequent altered disturbance regime on aquatic insect communities inhabiting a formerly perennial desert stream. We monitored two study pools seasonally for 4 years before and 4 years after the transition from perennial to intermittent flow to evaluate pre‐drying community dynamics and post‐drying recovery trajectories. 3. Mean species richness was not affected by the transition to intermittent flow, though seasonal patterns of richness did change. Sample densities were much higher in post‐drying samples. 4. The stream pool communities underwent a catastrophic regime shift after transition to intermittent flow, moving to an alternative stable state with novel seasonal trajectories, and did not recover to pre‐drying configurations after 4 years. Six invertebrate species were extirpated by the initial drying event, while other species were as much as 40 times more abundant in post‐drying samples. In general, large‐bodied top predators were extirpated from the system and replaced with high abundances of smaller‐bodied mesopredators. 5. Our results suggest that the loss of perennial flow caused by intensified droughts and water withdrawals could lead to significant changes in community structure and species composition at local and regional scales.     

Asymmetric responses of primary productivity to precipitation extremes: A synthesis of grassland precipitation manipulation experiments

Climatic changes are altering Earth's hydrological cycle, resulting in altered precipitation amounts, increased interannual variability of precipitation, and more frequent extreme precipitation events. These trends will likely continue into the future, having substantial impacts on net primary productivity (NPP) and associated ecosystem services such as food production and carbon sequestration. Frequently, experimental manipulations of precipitation have linked altered precipitation regimes to changes in NPP. Yet, findings have been diverse and substantial uncertainty still surrounds generalities describing patterns of ecosystem sensitivity to altered precipitation. Additionally, we do not know whether previously observed correlations between NPP and precipitation remain accurate when precipitation changes become extreme. We synthesized results from 83 case studies of experimental precipitation manipulations in grasslands worldwide. We used meta‐analytical techniques to search for generalities and asymmetries of aboveground NPP (ANPP) and belowground NPP (BNPP) responses to both the direction and magnitude of precipitation change. Sensitivity (i.e., productivity response standardized by the amount of precipitation change) of BNPP was similar under precipitation additions and reductions, but ANPP was more sensitive to precipitation additions than reductions; this was especially evident in drier ecosystems. Additionally, overall relationships between the magnitude of productivity responses and the magnitude of precipitation change were saturating in form. The saturating form of this relationship was likely driven by ANPP responses to very extreme precipitation increases, although there were limited studies imposing extreme precipitation change, and there was considerable variation among experiments. This highlights the importance of incorporating gradients of manipulations, ranging from extreme drought to extreme precipitation increases into future climate change experiments. Additionally, policy and land management decisions related to global change scenarios should consider how ANPP and BNPP responses may differ, and that ecosystem responses to extreme events might not be predicted from relationships found under moderate environmental changes.     

Tracking unstable states: ecosystem dynamics in a changing world

Ecological systems can show complex and sometimes abrupt responses to environmental change, with important implications for their resilience. Theories of alternate stable states have been used to predict regime shifts of ecosystems as equilibrium responses to sufficiently slow environmental change. The actual rate of environmental change is a key factor affecting the response, yet we are still lacking a non-equilibrium theory that explicitly considers the influence of this rate of environmental change. We present a metacommunity model of predator–prey interactions displaying multiple stable states, and we impose an explicit rate of environmental change in habitat quality (carrying capacity) and connectivity (dispersal rate). We study how regime shifts depend on the rate of environmental change and compare the outcome with a stability analysis in the corresponding constant environment. Our results reveal that in a changing environment, the community can track states that are unstable in the constant environment. This tracking can lead to regime shifts, including local extinctions, that are not predicted by alternative stable state theory. In our metacommunity, tracking unstable states also controls the maintenance of spatial heterogeneity and spatial synchrony. Tracking unstable states can also lead to regime shifts that may be reversible or irreversible. Our study extends current regime shift theories to integrate rate-dependent responses to environmental change. It reveals the key role of unstable states for predicting transient dynamics and long-term resilience of ecological systems to climate change.     

Net primary productivity and rain‐use efficiency as affected by warming,altered precipitation,and clipping in a mixed‐grass prairie

Grassland productivity in response to climate change and land use is a global concern. In order to explore the effects of climate change and land use on net primary productivity (NPP), NPP partitioning [f_BNPP, defined as the fraction of belowground NPP (BNPP) to NPP], and rain‐use efficiency (RUE) of NPP, we conducted a field experiment with warming (+3 °C), altered precipitation (double and half), and annual clipping in a mixed‐grass prairie in Oklahoma, USA since July, 2009. Across the years, warming significantly increased BNPP, f_BNPP, and RUE_BNPP by an average of 11.6%, 2.8%, and 6.6%, respectively. This indicates that BNPP was more sensitive to warming than aboveground NPP (ANPP) since warming did not change ANPP and RUE_ANPP much. Double precipitation stimulated ANPP, BNPP, and NPP but suppressed RUE_ANPP, RUE_BNPP, and RUE_NPP while half precipitation decreased ANPP, BNPP, and NPP but increased RUE_ANPP, RUE_BNPP, and RUE_NPP. Clipping interacted with altered precipitation in impacting RUE_ANPP, RUE_BNPP, and RUE_NPP, suggesting land use could confound the effects of precipitation changes on ecosystem processes. Soil moisture was found to be a main factor in regulating variation in ANPP, BNPP, and NPP while soil temperature was the dominant factor influencing f_BNPP. These findings suggest that BNPP is critical point to future research. Additionally, results from single‐factor manipulative experiments should be treated with caution due to the non‐additive interactive effects of warming with altered precipitation and land use (clipping).     

Recovery of understorey vegetation after release from a long history of sheep grazing in a herb‐rich woodland

The effects of stock grazing on native grassy ecosystems in temperate southern Australia are well documented. However, less is known about the potential of ecosystems to recover after a long history of stock grazing and, in particular, whether the removal of stock will have positive, negative or neutral impacts on biodiversity. We examined the response of understorey vegetation to the removal of sheep grazing in a herb‐rich Eucalyptus camaldulensis (red gum) woodland in western Victoria. Using a space‐for‐time chronosequence, woodlands were stratified into groups based on their time‐since‐grazing removal; these were long‐ungrazed (>20 years), intermediate‐time‐since‐grazing (9–14 years), recently ungrazed (5 years) and continuously grazed. We found significantly higher species density in long‐ungrazed sites relative to sites with a more recent grazing history. No differences were found in species density between continuously grazed sites and those ungrazed in the previous 14 years. Species composition differed with time‐since‐grazing removal and indicator species analysis detected several native species (including tall native geophytes and herbs) associated with long‐ungrazed sites that were absent or in low abundance in the more recently grazed sites. Seven of the eight species significantly associated with continuously grazed sites were exotic. Removal of sheep grazing in red gum woodlands can have positive benefits for understorey diversity but it is likely that recovery of key indicators such as native species will be slow.     

生态系统多稳态研究进展

在多稳态的生态系统中,外力可能导致生态系统状态突然之间发生不可逆转的转变,从而达到一个新的平衡状态。但目前对多稳态理论的系统研究很少,如何使用预警信号来预测生态系统的状态转变依旧是个难题。通过多稳态理论的梳理提出了一个更加综合的多稳态定义,并以放牧模型为例,系统总结了多稳态理论的相关概念,将多稳态理论应用在生态系统演替和扰沌理论的解释中;通过对生态系统稳态转换预警信号的原理、优缺点和应用条件的分析,对不同尺度下多稳态的研究方法进行了归纳;最后提出了目前多稳态领域的研究问题和未来的研究重点。结果表明:(1)将时间和空间预警信号结合在一起,并量化正确预警信号的概率,对错误预警信号的比例进行加权,可能会提供更准确的稳态转换的预报。(2)定量观测试验适用于小尺度的研究,而较大尺度的研究则采用简化的模型来模拟研究,选择正确的尺度极有可能改变预警信号的可靠性。(3)结合多稳态理论研究生态系统临界转换和反馈控制机制,并将基于性状的特征指标和进化动力学纳入其中,是生态系统修复实践的重要研究方向。(4)将多稳态相关理论和生态保护管理政策的实践相结合,是多稳态理论未来应用的前景。本研究为多稳态理论和实践的...     

Spatial early warning signals for impending regime shifts: A practical framework for application in real‐world landscapes

Prediction of ecosystem response to global environmental change is a pressing scientific challenge of major societal relevance. Many ecosystems display nonlinear responses to environmental change, and may even undergo practically irreversible ‘regime shifts’ that initiate ecosystem collapse. Recently, early warning signals based on spatiotemporal metrics have been proposed for the identification of impending regime shifts. The rapidly increasing availability of remotely sensed data provides excellent opportunities to apply such model‐based spatial early warning signals in the real world, to assess ecosystem resilience and identify impending regime shifts induced by global change. Such information would allow land‐managers and policy makers to interfere and avoid catastrophic shifts, but also to induce regime shifts that move ecosystems to a desired state. Here, we show that the application of spatial early warning signals in real‐world landscapes presents unique and unexpected challenges, and may result in misleading conclusions when employed without careful consideration of the spatial data and processes at hand. We identify key practical and theoretical issues and provide guidelines for applying spatial early warning signals in heterogeneous, real‐world landscapes based on literature review and examples from real‐world data. Major identified issues include (1) spatial heterogeneity in real‐world landscapes may enhance reversibility of regime shifts and boost landscape‐level resilience to environmental change (2) ecosystem states are often difficult to define, while these definitions have great impact on spatial early warning signals and (3) spatial environmental variability and socio‐economic factors may affect spatial patterns, spatial early warning signals and associated regime shift predictions. We propose a novel framework, shifting from an ecosystem perspective towards a landscape approach. The framework can be used to identify conditions under which resilience assessment with spatial remotely sensed data may be successful, to support well‐informed application of spatial early warning signals, and to improve predictions of ecosystem responses to global environmental change.     

青藏高原两种高寒草地植被变化及其水温驱动因素分析

高寒草甸和高寒草原作为青藏高原两种重要植被类型,研究其植被变化与气候变化相关性,有助于为青藏高原两种高寒草地生态系统应对全球气候变化管理提供参考。以位于同纬度的三江源高寒草甸和阿里高寒草原为研究对象,基于植被净初级生产力(Net Primary Productivity, NPP)变化表征植被变化,利用NPP数据和气象数据,分别分析两地2000—2017年植被NPP、降水和气温时空变化差异;利用Sen+Mann-Kendall趋势检验,研究两种高寒草地气候与植被净初级生产力变化趋势;以县域统计年鉴牛羊肉产量表征放牧强度,研究放牧活动对高寒草地植被变化的影响;通过Pearson相关和偏相关分析方法,分别研究降水和气温对两种高寒草地植被NPP变化影响差异。研究结果表明:(1)2000—2017年三江源高寒草甸和阿里高寒草原区年平均气温以0.085℃/a和0.084℃/a的趋势上升,降水以平均每年3.87 mm和2.23 mm的趋势增加,高寒草甸区变暖变湿速率较高寒草原区快。(2)三江源高寒草甸和阿里高寒草原植被NPP均呈现由东南向西北逐渐降低空间格局;2000—2017年高寒草甸区57.7...     

1982—2015年气候变化和人类活动对内蒙古草地净初级生产力的影响

研究1982—2015年气候变化和人类活动对内蒙古草地净初级生产力(NPP)的影响。结果表明: 1982—1998年和1999—2015年2个时期,内蒙古草地实际NPP(ANPP)增长速率分别为1.08和1.36 g C·m^-2·a^-1,草地以恢复为主,2个时期草地恢复面积分别占研究区总面积的81.6%和76.3%;草地退化面积有增加趋势,且气候变化和人类活动对不同类型草地的影响不同。2个时期气候变化对草地恢复贡献率分别为79.3%和94.1%,气候变化是草地恢复的主要因素,其中,ANPP与降水呈显著正相关,而与温度的相关性不显著,表明降水是影响草地恢复的主要气候因子。2个时期人类活动对草地退化的贡献率分别为83.3%和87.8%,说明人类活动是导致草地退化的主要原因。气候变化对内蒙古草地恢复起主导作用,而人类活动诸如放牧数量、耕地面积和造林面积的增加,加速了草地退化。     

Halting Regime Shifts in Floristically Intact Tropical Forests Deprived of Their Frugivores

Ecological restoration typically focuses on promoting vegetation recovery in degraded habitat or reintroducing endangered animals to enhance their regional or global persistence. Here, we argue that attention should also be devoted to vertebrate reintroductions in overhunted but floristically intact tropical forests in order to prevent insidious regime shifts in these systems. Growing evidence suggests that tropical forests deprived of seed‐dispersing animals exhibit replacement of fleshy fruiting trees by species with abiotic seed dispersal. Left unchecked, this process could eventually render the forest uninhabitable by frugivores through reduced density and diversity of their food plants. In tropical areas where hunting can be controlled, we contend that frugivore reintroduction, regulation of wild fruit harvest by humans, and outplanting of native fruiting trees should be deployed as management tools long before the systems are in need of traditional habitat restoration.     

2009-2011年我国西南地区旱灾程度及其对植被净初级生产力的影响

2009—2011年,我国西南地区遭受了极端干旱气候影响。利用1980—2011年气象站点观测数据和基于光能利用率的植被净初级生产力估算模型Glo PEM,研究了2009—2011年西南地区干旱灾害过程和程度及其对植被净初级生产力的影响,结果显示:2009—2011年西南地区年均降水量和湿润指数明显低于1980—2008年均值。受干旱气候影响,研究区植被净初级生产力比2001—2011年均值低12.55 g C m-2a-1,总计低0.017 Pg C/a,造成的碳损失约占我国总碳汇的7.91%。2001—2011年西南地区植被净初级生产力与蒸散量变化显著相关(R2=0.44,P0.05),而降水量和湿润指数变化过程与植被净初级生产力和蒸散量不同步,可能是由于该地区森林覆盖率较高,具有较强的涵养水源功能,导致土壤湿度变化滞后于降水量和湿润指数变化,从而使降水量变化过程与植被净初级生产力变化不同步。     

The effect of climate change on the resilience of ecosystems with adaptive spatial pattern formation

In a rapidly changing world, quantifying ecosystem resilience is an important challenge. Historically, resilience has been defined via models that do not take spatial effects into account. These systems can only adapt via uniform adjustments. In reality, however, the response is not necessarily uniform, and can lead to the formation of (self‐organised) spatial patterns – typically localised vegetation patches. Classical measures of resilience cannot capture the emerging dynamics in spatially self‐organised systems, including transitions between patterned states that have limited impact on ecosystem structure and productivity. We present a framework of interlinked phase portraits that appropriately quantifies the resilience of patterned states, which depends on the number of patches, the distances between them and environmental conditions. We show how classical resilience concepts fail to distinguish between small and large pattern transitions, and find that the variance in interpatch distances provides a suitable indicator for the type of imminent transition. Subsequently, we describe the dependency of ecosystem degradation based on the rate of climatic change: slow change leads to sporadic, large transitions, whereas fast change causes a rapid sequence of smaller transitions. Finally, we discuss how pre‐emptive removal of patches can minimise productivity losses during pattern transitions, constituting a viable conservation strategy.     

Drought effects on litterfall, wood production and belowground carbon cycling in an Amazon forest: results of a throughfall reduction experiment

The Amazon Basin experiences severe droughts that may become more common in the future. Little is known of the effects of such droughts on Amazon forest productivity and carbon allocation. We tested the prediction that severe drought decreases litterfall and wood production but potentially has multiple cancelling effects on belowground production within a 7-year partial throughfall exclusion experiment. We simulated an approximately 35-41% reduction in effective rainfall from 2000 through 2004 in a 1ha plot and compared forest response with a similar control plot. Wood production was the most sensitive component of above-ground net primary productivity (ANPP) to drought, declining by 13% the first year and up to 62% thereafter. Litterfall declined only in the third year of drought, with a maximum difference of 23% below the control plot. Soil CO2 efflux and its 14C signature showed no significant treatment response, suggesting similar amounts and sources of belowground production. ANPP was similar between plots in 2000 and declined to a low of 41% below the control plot during the subsequent treatment years, rebounding to only a 10% difference during the first post-treatment year. Live aboveground carbon declined by 32.5Mgha-1 through the effects of drought on ANPP and tree mortality. Results of this unreplicated, long-term, large-scale ecosystem manipulation experiment demonstrate that multi-year severe drought can substantially reduce Amazon forest carbon stocks.     

Sensitivity of plant species to warming and altered precipitation dominates the community productivity in a semiarid grassland on the Loess Plateau

Global warming and changes in precipitation patterns can critically influence the structure and productivity of terrestrial ecosystems. However, the underlying mechanisms are not fully understood. We conducted two independent but complementary experiments (one with warming and precipitation manipulation (+ or – 30%) and another with selective plant removal) in a semiarid grassland on the Loess Plateau, northwestern China, to assess how warming and altered precipitation affect plant community. Our results showed that warming and altered precipitation affected community aboveground net primary productivity (ANPP) through impacting soil moisture. Results of the removal experiment showed competitive relationships among dominant grasses, the dominant subshrub and nondominant species, which played a more important role than soil moisture in the response of plant community to warming and altered precipitation. Precipitation addition intensified the competition but primarily benefited the dominant subshrub. Warming and precipitation reduction enhanced water stresses but increased ANPP of the dominant subshrub and grasses, indicating that plant tolerance to drought critically meditated the community responses. These findings suggest that specie competitivity for water resources as well as tolerance to environmental stresses may dominate the responses of plant communities on the Loess Plateaus to future climate change factors.     

模拟夏季干旱对东灵山森林植被动态的影响

应用LPJ-GUESS植被动态模型, 在耦合不同物种的干旱响应策略的基础上, 研究了夏季干旱化对东灵山地区森林植被的物种组成及其功能的影响。结果表明, 在气候变暖、降水减少、CO₂浓度升高的情况下, 无论树种采取何种策略, 东灵山暖温带森林的总净初级生产力和生物量都有增加的趋势, 降水在未来近一个世纪内尚未成为本地区植被生长的限制因子。但森林植被的树种组成与树种的干旱响应策略密切相关, 不耐旱的物种核桃楸(Juglans mandshurica)的生物量水平在长期干旱条件下并没有降低, 而耐旱的物种辽东栎(Quercus liaotungensis)在受到干旱化长期影响时, 其生物量有下降的趋势。这种响应策略也会导致植被蒸散等生态系统水分循环过程的差异。因此, 降水变化对森林生态系统影响的长期模拟研究应该考虑物种对干旱的不同响应策略。