Regime Shifts in the Anthropocene: Drivers,Risks, and Resilience

Many ecosystems can experience regime shifts: surprising, large and persistent changes in the function and structure of ecosystems. Assessing whether continued global change will lead to further regime shifts, or has the potential to trigger cascading regime shifts has been a central question in global change policy. Addressing this issue has, however, been hampered by the focus of regime shift research on specific cases and types of regime shifts. To systematically assess the global risk of regime shifts we conducted a comparative analysis of 25 generic types of regime shifts across marine, terrestrial and polar systems; identifying their drivers, and impacts on ecosystem services. Our results show that the drivers of regime shifts are diverse and co-occur strongly, which suggests that continued global change can be expected to synchronously increase the risk of multiple regime shifts. Furthermore, many regime shift drivers are related to climate change and food production, whose links to the continued expansion of human activities makes them difficult to limit. Because many regime shifts can amplify the drivers of other regime shifts, continued global change can also be expected to increase the risk of cascading regime shifts. Nevertheless, the variety of scales at which regime shift drivers operate provides opportunities for reducing the risk of many types of regime shifts by addressing local or regional drivers, even in the absence of rapid reduction of global drivers.     

Comparative analysis of European wide marine ecosystem shifts: a large-scale approach for developing the basis for ecosystem-based management

Abrupt and rapid ecosystem shifts (where major reorganizations of food-web and community structures occur), commonly termed regime shifts, are changes between contrasting and persisting states of ecosystem structure and function. These shifts have been increasingly reported for exploited marine ecosystems around the world from the North Pacific to the North Atlantic. Understanding the drivers and mechanisms leading to marine ecosystem shifts is crucial in developing adaptive management strategies to achieve sustainable exploitation of marine ecosystems. An international workshop on a comparative approach to analysing these marine ecosystem shifts was held at Hamburg University, Institute for Hydrobiology and Fisheries Science, Germany on 1-3 November 2010. Twenty-seven scientists from 14 countries attended the meeting, representing specialists from seven marine regions, including the Baltic Sea, the North Sea, the Barents Sea, the Black Sea, the Mediterranean Sea, the Bay of Biscay and the Scotian Shelf off the Canadian East coast. The goal of the workshop was to conduct the first large-scale comparison of marine ecosystem regime shifts across multiple regional areas, in order to support the development of ecosystem-based management strategies.     

Trophic amplification of climate warming

Ecosystems can alternate suddenly between contrasting persistent states due to internal processes or external drivers. It is important to understand the mechanisms by which these shifts occur, especially in exploited ecosystems. There have been several abrupt marine ecosystem shifts attributed either to fishing, recent climate change or a combination of these two drivers. We show that temperature has been an important driver of the trophodynamics of the North Sea, a heavily fished marine ecosystem, for nearly 50 years and that a recent pronounced change in temperature established a new ecosystem dynamic regime through a series of internal mechanisms. Using an end-to-end ecosystem approach that included primary producers, primary, secondary and tertiary consumers, and detritivores, we found that temperature modified the relationships among species through nonlinearities in the ecosystem involving ecological thresholds and trophic amplifications. Trophic amplification provides an alternative mechanism to positive feedback to drive an ecosystem towards a new dynamic regime, which in this case favours jellyfish in the plankton and decapods and detritivores in the benthos. Although overfishing is often held responsible for marine ecosystem degeneration, temperature can clearly bring about similar effects. Our results are relevant to ecosystem-based fisheries management (EBFM), seen as the way forward to manage exploited marine ecosystems.     

Marine and freshwater regime changes impact a community of migratory Pacific salmonids in decline

Marine and freshwater ecosystems are increasingly at risk of large and cascading changes from multiple human activities (termed “regime shifts”), which can impact population productivity, resilience, and ecosystem structure. Pacific salmon exhibit persistent and large fluctuations in their population dynamics driven by combinations of intrinsic (e.g., density dependence) and extrinsic factors (e.g., ecosystem changes, species interactions). In recent years, many Pacific salmon have declined due to regime shifts but clear understanding of the processes driving these changes remains elusive. Here, we unpacked the role of density dependence, ecosystem trends, and stochasticity on productivity regimes for a community of five anadromous Pacific salmonids (Steelhead, Coho Salmon, Pink Salmon, Dolly Varden, and Coastal Cutthroat Trout) across a rich 40-year time-series. We used a Bayesian multivariate state-space model to examine whether productivity shifts had similarly occurred across the community and explored marine or freshwater changes associated with those shifts. Overall, we identified three productivity regimes: an early regime (1976–1990), a compensatory regime (1991–2009), and a declining regime (since 2010) where large declines were observed for Steelhead, Dolly Varden, and Cutthroat Trout, intermediate declines in Coho and no change in Pink Salmon. These regime changes were associated with multiple cumulative effects across the salmon life cycle. For example, increased seal densities and ocean competition were associated with lower adult marine survival in Steelhead. Watershed logging also intensified over the past 40 years and was associated with (all else equal) ≥97% declines in freshwater productivity for Steelhead, Cutthroat, and Coho. For Steelhead, marine and freshwater dynamics played approximately equal roles in explaining trends in total productivity. Collectively, these changing environments limited juvenile production and lowered future adult returns. These results reveal how changes in freshwater and marine environments can jointly shape population dynamics among ecological communities, like Pacific salmon, with cascading consequences to their resilience.     

Causes and projections of abrupt climate-driven ecosystem shifts in the North Atlantic

Warming of the global climate is now unequivocal and its impact on Earth' functional units has become more apparent. Here, we show that marine ecosystems are not equally sensitive to climate change and reveal a critical thermal boundary where a small increase in temperature triggers abrupt ecosystem shifts seen across multiple trophic levels. This large-scale boundary is located in regions where abrupt ecosystem shifts have been reported in the North Atlantic sector and thereby allows us to link these shifts by a global common phenomenon. We show that these changes alter the biodiversity and carrying capacity of ecosystems and may, combined with fishing, precipitate the reduction of some stocks of Atlantic cod already severely impacted by exploitation. These findings offer a way to anticipate major ecosystem changes and to propose adaptive strategies for marine exploited resources such as cod in order to minimize social and economic consequences.     

Interdisciplinary knowledge exchange across scales in a globally changing marine environment

The effects of anthropogenic global environmental change on biotic and abiotic processes have been reported in aquatic systems across the world. Complex synergies between concurrent environmental stressors and the resilience of the system to regime shifts, which vary in space and time, determine the capacity for marine systems to maintain structure and function with global environmental change. Consequently, an interdisciplinary approach that facilitates the development of new methods for the exchange of knowledge between scientists across multiple scales is required to effectively understand, quantify and predict climate impacts on marine ecosystem services. We use a literature review to assess the limitations and assumptions of current pathways to exchange interdisciplinary knowledge and the transferability of research findings across spatial and temporal scales and levels of biological organization to advance scientific understanding of global environmental change in marine systems. We found that species‐specific regional scale climate change research is most commonly published, and “supporting” is the ecosystem service most commonly referred to in publications. In addition, our paper outlines a trajectory for the future development of integrated climate change science for sustaining marine ecosystem services such as investment in interdisciplinary education and connectivity between disciplines.     

Modelling marine community responses to climate‐driven species redistribution to guide monitoring and adaptive ecosystem‐based management

As a consequence of global climate‐driven changes, marine ecosystems are experiencing polewards redistributions of species – or range shifts – across taxa and throughout latitudes worldwide. Research on these range shifts largely focuses on understanding and predicting changes in the distribution of individual species. The ecological effects of marine range shifts on ecosystem structure and functioning, as well as human coastal communities, can be large, yet remain difficult to anticipate and manage. Here, we use qualitative modelling of system feedback to understand the cumulative impacts of multiple species shifts in south‐eastern Australia, a global hotspot for ocean warming. We identify range‐shifting species that can induce trophic cascades and affect ecosystem dynamics and productivity, and evaluate the potential effectiveness of alternative management interventions to mitigate these impacts. Our results suggest that the negative ecological impacts of multiple simultaneous range shifts generally add up. Thus, implementing whole‐of‐ecosystem management strategies and regular monitoring of range‐shifting species of ecological concern are necessary to effectively intervene against undesirable consequences of marine range shifts at the regional scale. Our study illustrates how modelling system feedback with only limited qualitative information about ecosystem structure and range‐shifting species can predict ecological consequences of multiple co‐occurring range shifts, guide ecosystem‐based adaptation to climate change and help prioritise future research and monitoring.     

Nutrient reduction and climate change cause a potential shift from pelagic to benthic pathways in a eutrophic marine ecosystem

The degree to which marine ecosystems may support the pelagic or benthic food chain has been shown to vary across natural and anthropogenic gradients for e.g., in temperature and nutrient availability. Moreover, such external forcing may not only affect the flux of organic matter but could trigger large and abrupt changes, i.e., trophic cascades and ecological regime shifts, which once having occurred may prove potentially irreversible. In this study, we investigate the state and regulatory pathways of the Kattegat; a eutrophied and heavily exploited marine ecosystem, specifically testing for the occurrence of regime shifts and the relative importance of multiple drivers, e.g., climate change, eutrophication and commercial fishing on ecosystem dynamics and trophic pathways. Using multivariate statistics and nonlinear regression on a comprehensive data set, covering abiotic factors and biotic variables across all trophic levels, we here propose a potential regime shift from pelagic to benthic regulatory pathways; a possible first sign of recovery from eutrophication likely triggered by drastic nutrient reductions (involving both nitrogen and phosphorus), in combination with climate‐driven changes in local environmental conditions (e.g., temperature and oxygen concentrations).     

Region‐wide changes in marine ecosystem dynamics: state‐space models to distinguish trends from step changes

Regime shifts are sudden changes in ecosystem structure that can be detected across several ecosystem components. The concept that regime shifts are common in marine ecosystems has gained popularity in recent years. Many studies have searched for the step‐like changes in ecosystem state expected under a simple interpretation of this idea. However, other kinds of change, such as pervasive trends, have often been ignored. We assembled over 300 ecological time series from seven UK marine regions, covering two to three decades. We developed state‐space models for the first principal component of the time series in each region, a common measure of ecosystem state. Our models allowed both trends and step changes, possibly in combination. We found trends in three of seven regions and step changes in two of seven regions. Gradual and sudden changes are therefore important trajectories to consider in marine ecosystems.     

草地和荒漠生态系统服务功能的形成与调控机制

生态系统服务功能的形成和维持机制是近年来生态学研究的热点领域。该文从生态系统服务功能的概念与内涵入手, 分析了生态系统服务功能研究的科学前沿和热点领域, 重点介绍了草地和荒漠生态系统服务功能形成与调控机制联网研究的思路、关键科学问题、研究目标、实验设计、观测指标体系, 以及联网研究取得的重要进展。在此基础上, 对未来草地和荒漠生态系统服务功能研究的重点进行了展望。     

A niche for ecosystem multifunctionality in global change research

Concern about human modification of Earth's ecosystems has recently motivated ecologists to address how global change drivers will impact the simultaneous provisioning of multiple functions, termed ecosystem multifunctionality (EMF). However, metrics of EMF have often been applied in global change studies with little consideration of the information they provide beyond single functions, or how and why EMF may respond to global change drivers. Here, we critically review the current state of this rapidly expanding field and provide a conceptual framework to guide the effective incorporation of EMF in global change research. In particular, we emphasize the need for a priori identification and explicit testing of the biotic and abiotic mechanisms through which global change drivers impact EMF, as well as assessing correlations among multiple single functions because these patterns underlie shifts in EMF. While the role of biodiversity in mediating global change effects on EMF has justifiably received much attention, empirical support for effects via other biotic and physicochemical mechanisms are also needed. Studies also frequently stated the importance of measuring EMF responses to global change drivers to understand the potential consequences for multiple ecosystem services, but explicit links between measured functions and ecosystem services were missing from many such studies. While there is clear potential for EMF to provide novel insights to global change research, predictive understanding will be greatly improved by insuring future research is strongly hypothesis‐driven, is designed to explicitly test multiple abiotic and biotic mechanisms, and assesses how single functions and their covariation drive emergent EMF responses to global change drivers.     

Indicators of biodiversity and ecosystem services: a synthesis across ecosystems and spatial scales

According to the Millennium Ecosystem Assessment, common indicators are needed to monitor the loss of biodiversity and the implications for the sustainable provision of ecosystem services. However, a variety of indicators are already being used resulting in many, mostly incompatible, monitoring systems. In order to synthesise the different indicator approaches and to detect gaps in the development of common indicator systems, we examined 531 indicators that have been reported in 617 peer‐reviewed journal articles between 1997 and 2007. Special emphasis was placed on comparing indicators of biodiversity and ecosystem services across ecosystems (forests, grass‐ and shrublands, wetlands, rivers, lakes, soils and agro‐ecosystems) and spatial scales (from patch to global scale). The application of biological indicators was found most often focused on regional and finer spatial scales with few indicators applied across ecosystem types. Abiotic indicators, such as physico‐chemical parameters and measures of area and fragmentation, are most frequently used at broader (regional to continental) scales. Despite its multiple dimensions, biodiversity is usually equated with species richness only. The functional, structural and genetic components of biodiversity are poorly addressed despite their potential value across habitats and scales. Ecosystem service indicators are mostly used to estimate regulating and supporting services but generally differ between ecosystem types as they reflect ecosystem‐specific services. Despite great effort to develop indicator systems over the past decade, there is still a considerable gap in the widespread use of indicators for many of the multiple components of biodiversity and ecosystem services, and a need to develop common monitoring schemes within and across habitats. Filling these gaps is a prerequisite for linking biodiversity dynamics with ecosystem service delivery and to achieving the goals of global and sub‐global initiatives to halt the loss of biodiversity.     

Marine ecosystem regime shifts: challenges and opportunities for ecosystem-based management

Regime shifts have been observed in marine ecosystems around the globe. These phenomena can result in dramatic changes in the provision of ecosystem services to coastal communities. Accounting for regime shifts in management clearly requires integrative, ecosystem-based management (EBM) approaches. EBM has emerged as an accepted paradigm for ocean management worldwide, yet, despite the rapid and intense development of EBM theory, implementation has languished, and many implemented or proposed EBM schemes largely ignore the special characteristics of regime shifts. Here, we first explore key aspects of regime shifts that are of critical importance to EBM, and then suggest how regime shifts can be better incorporated into EBM using the concept of integrated ecosystem assessment (IEA). An IEA uses approaches that determine the likelihood that ecological or socio-economic properties of systems will move beyond or return to acceptable bounds as defined by resource managers and policy makers. We suggest an approach for implementing IEAs for cases of regime shifts where the objectives are either avoiding an undesired state or returning to a desired condition. We discuss the suitability and short-comings of methods summarizing the status of ecosystem components, screening and prioritizing potential risks, and evaluating alternative management strategies. IEAs are evolving as an EBM approach that can address regime shifts; however, advances in statistical, analytical and simulation modelling are needed before IEAs can robustly inform tactical management in systems characterized by regime shifts.     

Evaluating Ecosystem Services Provided by Non-Native Species: An Experimental Test in California Grasslands

The concept of ecosystem services – the benefits that nature provides to human''s society – has gained increasing attention over the past decade. Increasing global abiotic and biotic change, including species invasions, is threatening the secure delivery of these ecosystem services. Efficient evaluation methods of ecosystem services are urgently needed to improve our ability to determine management strategies and restoration goals in face of these new emerging ecosystems. Considering a range of multiple ecosystem functions may be a useful way to determine such strategies. We tested this framework experimentally in California grasslands, where large shifts in species composition have occurred since the late 1700''s. We compared a suite of ecosystem functions within one historic native and two non-native species assemblages under different grazing intensities to address how different species assemblages vary in provisioning, regulatory and supporting ecosystem services. Forage production was reduced in one non-native assemblage (medusahead). Cultural ecosystem services, such as native species diversity, were inherently lower in both non-native assemblages, whereas most other services were maintained across grazing intensities. All systems provided similar ecosystem services under the highest grazing intensity treatment, which simulated unsustainable grazing intensity. We suggest that applying a more comprehensive ecosystem framework that considers multiple ecosystem services to evaluate new emerging ecosystems is a valuable tool to determine management goals and how to intervene in a changing ecosystem.     

Marine biodiversity and ecosystem functioning: what's known and what's next?

Marine ecosystems are experiencing rapid and pervasive changes in biodiversity and species composition. Understanding the ecosystem consequences of these changes is critical to effectively managing these systems. Over the last several years, numerous experimental manipulations of species richness have been performed, yet existing quantitative syntheses have focused on a just a subset of processes measured in experiments and, as such, have not summarized the full data available from marine systems. Here, we present the results of a meta‐analysis of 110 marine experiments from 42 studies that manipulated the species richness of organisms across a range of taxa and trophic levels and analysed the consequences for various ecosystem processes (categorised as production, consumption or biogeochemical fluxes). Our results show that, generally, mixtures of species tend to enhance levels of ecosystem function relative to the average component species in monoculture, but have no effect or a negative effect on functioning relative to the ‘highest‐ performing’ species. These results are largely consistent with those from other syntheses, and extend conclusions to ecological functions that are commonly measured in the marine realm (e.g. nutrient release from sediment bioturbation). For experiments that manipulated three or more levels of richness, we attempted to discern the functional form of the biodiversity–ecosystem functioning relationship. We found that, for response variables related to consumption, a power‐function best described the relationship, which is also consistent with previous findings. However, we identified a linear relationship between richness and production. Combined, our results suggest that changes in the number of species will, on average, tend to alter the functioning of marine ecosystems. We outline several research frontiers that will allow us to more fully understand how, why, and when diversity may drive the functioning of marine ecosystems. Synthesis The oceans host an incredible number and variety of species. However, human activities are driving rapid changes in the marine environment. It is imperative we understand ecosystem consequences of any associated loss of species. We summarized data from 110 experiments that manipulated species diversity and evaluated resulting changes to a range of ecosystem responses. We show that losing species, on average, decreases productivity, growth, and a myriad of other processes related to how marine organisms capture and utilize resources. Finally, we suggest that the loss of species may have stronger consequences for some processes than others.     

Latitudinal patterns of forest ecosystem stability across spatial scales as affected by biodiversity and environmental heterogeneity

Our planet is facing a variety of serious threats from climate change that are unfolding unevenly across the globe. Uncovering the spatial patterns of ecosystem stability is important for predicting the responses of ecological processes and biodiversity patterns to climate change. However, the understanding of the latitudinal pattern of ecosystem stability across scales and of the underlying ecological drivers is still very limited. Accordingly, this study examines the latitudinal patterns of ecosystem stability at the local and regional spatial scale using a natural assembly of forest metacommunities that are distributed over a large temperate forest region, considering a range of potential environmental drivers. We found that the stability of regional communities (regional stability) and asynchronous dynamics among local communities (spatial asynchrony) both decreased with increasing latitude, whereas the stability of local communities (local stability) did not. We tested a series of hypotheses that potentially drive the spatial patterns of ecosystem stability, and found that although the ecological drivers of biodiversity, climatic history, resource conditions, climatic stability, and environmental heterogeneity varied with latitude, latitudinal patterns of ecosystem stability at multiple scales were affected by biodiversity and environmental heterogeneity. In particular, α diversity is positively associated with local stability, while β diversity is positively associated with spatial asynchrony, although both relationships are weak. Our study provides the first evidence that latitudinal patterns of the temporal stability of naturally assembled forest metacommunities across scales are driven by biodiversity and environmental heterogeneity. Our findings suggest that the preservation of plant biodiversity within and between forest communities and the maintenance of heterogeneous landscapes can be crucial to buffer forest ecosystems at higher latitudes from the faster and more intense negative impacts of climate change in the future.     

Aquatic biodiversity in forests: a weak link in ecosystem services resilience

The diversity of aquatic ecosystems is being quickly reduced on many continents, warranting a closer examination of the consequences for ecological integrity and ecosystem services. Here we describe intermediate and final ecosystem services derived from aquatic biodiversity in forests. We include a summary of the factors framing the assembly of aquatic biodiversity in forests in natural systems and how they change with a variety of natural disturbances and human-derived stressors. We consider forested aquatic ecosystems as a multi-state portfolio, with diverse assemblages and life-history strategies occurring at local scales as a consequence of a mosaic of habitat conditions and past disturbances and stressors. Maintaining this multi-state portfolio of assemblages requires a broad perspective of ecosystem structure, various functions, services, and management implications relative to contemporary stressors. Because aquatic biodiversity provides multiple ecosystem services to forests, activities that compromise aquatic ecosystems and biodiversity could be an issue for maintaining forest ecosystem integrity. We illustrate these concepts with examples of aquatic biodiversity and ecosystem services in forests of northwestern North America, also known as Northeast Pacific Rim. Encouraging management planning at broad as well as local spatial scales to recognize multi-state ecosystem management goals has promise for maintaining valuable ecosystem services. Ultimately, integration of information from socio-ecological ecosystems will be needed to maintain ecosystem services derived directly and indirectly from forest aquatic biota.     

Agricultural modifications of hydrological flows create ecological surprises

Agricultural expansion and intensification have altered the quantity and quality of global water flows. Research suggests that these changes have increased the risk of catastrophic ecosystem regime shifts. We identify and review evidence for agriculture-related regime shifts in three parts of the hydrological cycle: interactions between agriculture and aquatic systems, agriculture and soil, and agriculture and the atmosphere. We describe the processes that shape these regime shifts and the scales at which they operate. As global demands for agriculture and water continue to grow, it is increasingly urgent for ecologists to develop new ways of anticipating, analyzing and managing nonlinear changes across scales in human-dominated landscapes.     

干旱区生态系统稳态转换及其预警信号——基于景观格局特征的识别方法

在日益加剧的气候变化和土地开垦、放牧等人类活动干扰下,具有多稳态特征的干旱区生态系统可能会经历从相对健康状态到退化状态的稳态转换,导致生态系统的功能下降。早期预警信号的识别是生态系统稳态转换研究的热点,也是管理实践中防止生态系统退化的关键环节。以往预警信号研究聚焦于通用信号如自相关性、方差等统计学指标,然而这些指标对于具有特定机制的干旱区生态系统可能并不适用。基于干旱区景观格局特征所发展起来的空间指标为生态系统稳态转换提供了独特的空间视角,对于理解干旱区生态系统退化过程和机理具有科学意义和实践价值。介绍了干旱区生态系统稳态转换现象及其转换机制;聚焦景观生态学的指标和方法,从空间视角总结基于干旱区景观格局特征的关键预警指标(植被覆盖度、植被斑块形态、植被斑块大小频率分布和水文连通性等),重点剖析这些关键指标的概念、量化方法、识别特征及其实践应用;最后针对指标的优势和局限性对未来的研究方向进行展望,包括发掘潜在景观指标,加强干旱区生态系统变化的多种驱动要素的相互作用机制研究,开展多时空尺度的实证研究,构建生态系统稳态转换预警信号的整体分析框架,以及加强指标阈值的量化研究等方面。