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.     

Climate and cyclic hydrobiological changes of the Barents Sea from the twentieth to twenty-first centuries

The Barents Sea is a transition zone between North Atlantic and Arctic waters, so its marine ecosystem is highly sensitive to climate dynamics. Understanding of marine biota response to climate changes is necessary to assess the environmental stability and the state of marketable biological resources. These processes are analyzed using a database from the Murmansk Marine Biological Institute which holds oceanographic and hydrobiological data sets collected for more than 100?years along the meridional Kola Transect in the Barents Sea. The data demonstrate high variability in thermal state of the upper layer of the Barents Sea, which is regulated by varying the inflow of Atlantic water and by regional climate. At irregular intervals, cold periods with extended seasonal ice cover are followed by warm periods. The most recent warm period started in the late 1980s and reached its maximum from 2001 to 2006. These cyclic changes in hydrologic regime across the twentieth century and first decade of the twenty-first century are reflected (with a specific lag of 1–5?years) by changes in species composition, as well as abundance and distribution of boreal and arctic groups of macrozoobenthos and fish fauna. For instance, cod and cod fisheries in the Barents Sea are closely linked to the marine climate. Furthermore, Kamchatka crab stock recruitment benefited from the warm climate of 1989 and 1990. In general, studies in this region have shown that climatic dynamics may be assessed using biological indices of abundance, biomass, and migration of marine organisms, including commercial species.     

Climate and Demography Dictate the Strength of Predator-Prey Overlap in a Subarctic Marine Ecosystem

There is growing evidence that climate and anthropogenic influences on marine ecosystems are largely manifested by changes in species spatial dynamics. However, less is known about how shifts in species distributions might alter predator-prey overlap and the dynamics of prey populations. We developed a general approach to quantify species spatial overlap and identify the biotic and abiotic variables that dictate the strength of overlap. We used this method to test the hypothesis that population abundance and temperature have a synergistic effect on the spatial overlap of arrowtooth flounder (predator) and juvenile Alaska walleye pollock (prey, age-1) in the eastern Bering Sea. Our analyses indicate that (1) flounder abundance and temperature are key variables dictating the strength of flounder and pollock overlap, (2) changes in the magnitude of overlap may be largely driven by density-dependent habitat selection of flounder, and (3) species overlap is negatively correlated to juvenile pollock recruitment when flounder biomass is high. Overall, our findings suggest that continued increases in flounder abundance coupled with the predicted long-term warming of ocean temperatures could have important implications for the predator-prey dynamics of arrowtooth flounder and juvenile pollock. The approach used in this study is valuable for identifying potential consequences of climate variability and exploitation on species spatial dynamics and interactions in many marine ecosystems.     

Ecological Network Indicators of Ecosystem Status and Change in the Baltic Sea

Several marine ecosystems under anthropogenic pressure have experienced shifts from one ecological state to another. In the central Baltic Sea, the regime shift of the 1980s has been associated with food-web reorganization and redirection of energy flow pathways. These long-term dynamics from 1974 to 2006 have been simulated here using a food-web model forced by climate and fishing. Ecological network analysis was performed to calculate indices of ecosystem change. The model replicated the regime shift. The analyses of indicators suggested that the system’s resilience was higher prior to 1988 and lower thereafter. The ecosystem topology also changed from a web-like structure to a linearized food-web.     

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.     

Globalization,marine regime shifts and the Soviet Union

Regime shifts have been observed in marine ecosystems around the world, with climate and fishing suggested as major drivers of such shifts. The global and regional dynamics of the climate system have been studied in this context, and efforts to develop an analogous understanding of fishing activities are developing. Here, we investigate the timing of pelagic marine regime shifts in relation to the emergence of regional and global fishing activities of the Soviet Union. Our investigation of official catch statistics reflects that the Soviet Union was a major fishing actor in all large marine ecosystems where regime shifts have been documented, including in ecosystems where overfishing has been established as a key driver of these changes (in the Baltic and Black Seas and the Scotian Shelf). Globalization of Soviet Union fishing activities pushed exploitation to radically new levels and triggered regional and global governance responses for improved management. Since then, exploitation levels have remained and increased with new actors involved. Based on our exploratory work, we propose that a deeper understanding of the role of global fishing actors is central for improved management of marine ecosystems.     

Ecosystem regime shifts have not affected growth and survivorship of eastern Beaufort Sea belugas

Large-scale ocean-atmosphere physical dynamics can have profound impacts on the structure and organization of marine ecosystems. These changes have been termed “regime shifts”, and five different episodes have been detected in the North Pacific Ocean, with concurrent changes also occurring in the Bering and Beaufort Seas. Belugas from the Eastern Beaufort Sea (EBS) use the Bering Sea during winter and the Beaufort Sea during summer, yet the potential effects of regime shifts on belugas have not been assessed. We investigated whether body size and survivorship of EBS belugas harvested in the Mackenzie River delta region between 1993 and 2003 have been affected by previous purported regime shifts in the North Pacific. Residuals from the relationship between body length and age were calculated and compared among belugas born between 1932 and 1989. Residual body size was not significantly related to birth year for any regime, nor to the age group individuals belonged to during any regime. The percentage deviation in number of belugas born in any given year that survived to be included in the hunt (survivorship) did not show any significant trend within or between regimes. Accounting for lags of 1–5 years did not reveal any evidence of delayed effects. Furthermore, neither population index was significantly related to changes in major climatic variables that precede regime shifts. Our results suggest that EBS beluga body size and survivorship have not been affected by the major regime shifts of the North Pacific and the adjacent Bering and Beaufort Seas. EBS belugas may have been able to modify their diet without compromising their growth and survivorship. Diet and reproductive analyses over large and small time scales can help understand the mechanisms enabling belugas to avoid significant growth and reproductive effects of past regime shifts. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

North Sea ecosystem change from swimming crabs to seagulls

A recent increase in sea temperature has established a new ecosystem dynamic regime in the North Sea. Climate-induced changes in decapods have played an important role. Here, we reveal a coincident increase in the abundance of swimming crabs and lesser black-backed gull colonies in the North Sea, both in time and in space. Swimming crabs are an important food source for lesser black-backed gulls during the breeding season. Inhabiting the land, but feeding mainly at sea, lesser black-backed gulls provide a link between marine and terrestrial ecosystems, since the bottom-up influence of allochthonous nutrient input from seabirds to coastal soils can structure the terrestrial food web. We, therefore, suggest that climate-driven changes in trophic interactions in the marine food web may also have ensuing ramifications for the coastal ecology of the North Sea.     

The rise of a marine generalist predator and the fall of beta diversity

Determining the importance of physical and biological drivers in shaping biodiversity in diverse ecosystems remains a global challenge. Advancements have been made towards this end in large marine ecosystems with several studies suggesting environmental forcing as the primary driver. However, both empirical and theoretical studies point to additional drivers of changes in diversity involving trophic interactions and, in particular, predation. Moreover, a more integrated but less common approach to the assessment of biodiversity changes involves analyses of spatial β diversity, whereas most studies to date assess only changes in species richness (α diversity). Recent research has established that when cod, a dominant generalist predator, was overfished and collapsed in a northwest Atlantic food web, spatial β diversity increased; that is, the spatial structure of the fish assemblage became increasingly heterogeneous. If cod were to recover, would this situation be reversible, given the inherent complexity and non‐linear dynamics that typify such systems? A dramatic increase of cod in an ecologically similar large marine ecosystem may provide an answer. Here we show that spatial β diversity of fish assemblages in the Barents Sea decreased with increasing cod abundance, while decadal scale changes in temperature did not play a significant role. These findings indicate a reversibility of the fish assemblage structure in response to changing levels of an apex predator and highlight the frequently overlooked importance of trophic interactions in determining large‐scale biodiversity patterns. As increased cod abundance was largely driven by changes in fisheries management, our study also shows that management policies and practices, particularly those involving apex predators, can have a strong effect in shaping spatial diversity patterns, and one should not restrict the focus to effects of climate change alone.     

Future climate‐driven shifts in distribution of Calanus finmarchicus

Calanus finmarchicus is a key‐structural species of the North Atlantic polar biome. The species plays an important trophic role in subpolar and polar ecosystems as a grazer of phytoplankton and as a prey for higher trophic levels such as the larval stages of many fish species. Here, we used a recently developed ecological niche model to assess the ecological niche (sensu Hutchinson) of C. finmarchicus and characterize its spatial distribution. This model explained about 65% of the total variance of the observed spatial distribution inferred from an independent dataset (data of the continuous plankton recorder survey). Comparisons with other types of models (structured population and ecophysiological models) revealed a clear similarity between modeled spatial distributions at the scale of the North Atlantic. Contemporary models coupled with future projections indicated a progressive reduction of the spatial habitat of the species at the southern edge and a more pronounced one in the Georges Bank, the Scotian Shelf and the North Sea and a potential increase in abundance at the northern edge of its spatial distribution, especially in the Barents Sea. These major changes will probably lead to a major alteration of the trophodynamics of North Atlantic ecosystems affecting the trophodynamics and the biological carbon pump.     

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.     

土壤呼吸作用时空动态变化及其影响机制研究与展望

测定不同陆地生态系统土壤呼吸速率及其时空波动, 阐明其影响因子, 对于全球碳素平衡预算和全球变化潜在效应估计是最为基本的数据。然而, 有关土壤呼吸作用变异性及其影响因素的知识仍存在局限性, 一些关键的过程和机制还有待阐明。该文综述了近年来土壤呼吸作用时空动态规律、影响机制和模拟方面的研究进展, 指出环境因子和生物因子共同驱动着土壤呼吸作用的时间动态变化; 土壤呼吸作用在不同时间尺度上还具有明显的空间异质性, 这主要是植被覆盖、根系分布、主要的环境因素和土壤特性空间分布的异质性造成的。生物因子是影响土壤呼吸作用时空动态变化的主要因素之一。然而, 目前所使用的土壤呼吸作用经验模型通常利用土壤温度、土壤湿度或者两者的交互作用模拟土壤呼吸作用动态变化, 但没有考虑生物因子的影响, 这可能会导致明显的偏差和错误。因此, 为了精确估算土壤呼吸作用, 必须解决土壤呼吸作用小尺度上的空间变异性; 加强不同时间尺度上生物要素对土壤呼吸作用动态变化的影响研究; 除了气候因子外, 土壤呼吸作用经验模型应该纳入生物因子等其它影响因素作为变量, 用以提高模型模拟的正确性和准确性。     

Marine biological shifts and climate

Phenological, biogeographic and community shifts are among the reported responses of marine ecosystems and their species to climate change. However, despite both the profound consequences for ecosystem functioning and services, our understanding of the root causes underlying these biological changes remains rudimentary. Here, we show that a significant proportion of the responses of species and communities to climate change are deterministic at some emergent spatio-temporal scales, enabling testable predictions and more accurate projections of future changes. We propose a theory based on the concept of the ecological niche to connect phenological, biogeographic and long-term community shifts. The theory explains approximately 70% of the phenological and biogeographic shifts of a key zooplankton Calanus finmarchicus in the North Atlantic and approximately 56% of the long-term shifts in copepods observed in the North Sea during the period 1958–2009.     

New Directions for Understanding the Spatial Resilience of Social–Ecological Systems

The concept of spatial resilience has brought a new focus on the influence of multi-scale processes on the dynamics of ecosystems. Initial ideas about spatial resilience focused on coral reefs and emphasized escalating anthropogenic disturbances across the broader seascape. This perspective resonated with a new awareness of global drivers of change, such as growth in international trade and shifts in climate, and the need to respond by scaling up governance and management. We review recent trends and emerging ideas in spatial resilience, using coral reefs and dependent communities as exemplars of multi-scale social–ecological systems. Despite recent advances, management and governance of ecosystems remain spatially fragmented and constrained to small scales. Temporally, many interventions still miss or ignore warning signals and struggle to cope with history, politics, long-term cumulative pressures, feedbacks, and sudden surprises. Significant recent progress has been made in understanding the relevance of spatial and temporal scale, heterogeneity, networks, the importance of place, and multi-scale governance. Emerging themes include better integration of ecology and conservation with social and economic science, and incorporating temporal dynamics in spatial analyses. A better understanding of the multi-scale spatial and temporal processes that drive the resilience of linked social-ecosystems will help address the widespread mismatch between the scales of ongoing ecological change and effective long-term governance of land- and seascapes.     

Demersal fish assemblages and spatial diversity patterns in the Arctic-Atlantic transition zone in the Barents Sea

Direct and indirect effects of global warming are expected to be pronounced and fast in the Arctic, impacting terrestrial, freshwater and marine ecosystems. The Barents Sea is a high latitude shelf Sea and a boundary area between arctic and boreal faunas. These faunas are likely to respond differently to changes in climate. In addition, the Barents Sea is highly impacted by fisheries and other human activities. This strong human presence places great demands on scientific investigation and advisory capacity. In order to identify basic community structures against which future climate related or other human induced changes could be evaluated, we analyzed species composition and diversity of demersal fish in the Barents Sea. We found six main assemblages that were separated along depth and temperature gradients. There are indications that climate driven changes have already taken place, since boreal species were found in large parts of the Barents Sea shelf, including also the northern Arctic area. When modelling diversity as a function of depth and temperature, we found that two of the assemblages in the eastern Barents Sea showed lower diversity than expected from their depth and temperature. This is probably caused by low habitat complexity and the distance to the pool of boreal species in the western Barents Sea. In contrast coastal assemblages in south western Barents Sea and along Novaya Zemlya archipelago in the Eastern Barents Sea can be described as diversity "hotspots"; the South-western area had high density of species, abundance and biomass, and here some species have their northern distribution limit, whereas the Novaya Zemlya area has unique fauna of Arctic, coastal demersal fish. (see Information S1 for abstract in Russian).     

Quantifying heterogeneous responses of fish community size structure using novel combined statistical techniques

To understand changes in ecosystems, the appropriate scale at which to study them must be determined. Large marine ecosystems (LMEs) cover thousands of square kilometres and are a useful classification scheme for ecosystem monitoring and assessment. However, averaging across LMEs may obscure intricate dynamics within. The purpose of this study is to mathematically determine local and regional patterns of ecological change within an LME using empirical orthogonal functions (EOFs). After using EOFs to define regions with distinct patterns of change, a statistical model originating from control theory is applied (Nonlinear AutoRegressive Moving Average with eXogenous input – NARMAX) to assess potential drivers of change within these regions. We have selected spatial data sets (0.5° latitude × 1°longitude) of fish abundance from North Sea fisheries research surveys (spanning 1980–2008) as well as of temperature, oxygen, net primary production and a fishing pressure proxy, to which we apply the EOF and NARMAX methods. Two regions showed significant changes since 1980: the central North Sea displayed a decrease in community size structure which the NARMAX model suggested was linked to changes in fishing; and the Norwegian trench region displayed an increase in community size structure which, as indicated by NARMAX results, was primarily linked to changes in sea‐bottom temperature. These regions were compared to an area of no change along the eastern Scottish coast where the model determined the community size structure was most strongly associated to net primary production. This study highlights the multifaceted effects of environmental change and fishing pressures in different regions of the North Sea. Furthermore, by highlighting this spatial heterogeneity in community size structure change, important local spatial dynamics are often overlooked when the North Sea is considered as a broad‐scale, homogeneous ecosystem (as normally is the case within the political Marine Strategy Framework Directive).     

Reorganization of a large marine ecosystem due to atmospheric and anthropogenic pressure: a discontinuous regime shift in the Central Baltic Sea

Marine ecosystems such as the Baltic Sea are currently under strong atmospheric and anthropogenic pressure. Besides natural and human-induced changes in climate, major anthropogenic drivers such as overfishing and anthropogenic eutrophication are significantly affecting ecosystem structure and function. Recently, studies demonstrated the existence of alternative stable states in various terrestrial and aquatic ecosystems. These so-called ecosystem regime shifts have been explained mainly as a result of multiple causes, e.g. climatic regime shifts, overexploitation or a combination of both. The occurrence of ecosystem regime shifts has important management implications, as they can cause significant losses of ecological and economic resources. Because of hysteresis in ecosystem responses, restoring regimes considered as favourable may require drastic and expensive management actions. Also the Baltic Sea, the largest brackish water body in the world ocean, and its ecosystems are strongly affected by atmospheric and anthropogenic drivers. Here, we present results of an analysis of the state and development of the Central Baltic Sea ecosystem integrating hydroclimatic, nutrient, phyto- and zooplankton as well as fisheries data. Our analyses of 52 biotic and abiotic variables using multivariate statistics demonstrated a major reorganization of the ecosystem and identified two stable states between 1974 and 2005, separated by a transition period in 1988–1993. We show the change in Baltic ecosystem structure to have the characteristics of a discontinuous regime shift, initiated by climate-induced changes in the abiotic environment and stabilized by fisheries-induced feedback loops in the food web. Our results indicate the importance of maintaining the resilience of an ecosystem to atmospherically induced environmental change by reducing the anthropogenic impact.     

Fish and fisheries in hot water: What is happening and how do we adapt?

Rapid climate changes are currently driving substantial reorganizations of marine ecosystems around the world. A key question is how these changes will alter the provision of ecosystem services from the ocean, particularly from fisheries. To answer this question, we need to understand not only the ecological dynamics of marine systems, but also human adaptation and feedbacks between humans and the rest of the natural world. In this review, we outline what we have learned from research primarily in continental shelf ecosystems and fishing communities of North America. Key findings are that marine animals are highly sensitive to warming and are responding quickly to changes in water temperature, and that such changes are often happening faster than similar processes on land. Changes in species distributions and productivity are having substantial impacts on fisheries, including through changing catch compositions and longer distances traveled for fishing trips. Conflicts over access to fisheries have also emerged as species distributions are no longer aligned with regulations or catch allocations. These changes in the coupled natural-human system have reduced the value of ecosystem services from some fisheries and risk doing so even more in the future. Going forward, substantial opportunities for more effective fisheries management and operations, marine conservation, and marine spatial planning are likely possible through greater consideration of climate information over time-scales from years to decades.     

Footprints of climate change in the Arctic marine ecosystem

In this article, we review evidence of how climate change has already resulted in clearly discernable changes in marine Arctic ecosystems. After defining the term ‘footprint’ and evaluating the availability of reliable baseline information we review the published literature to synthesize the footprints of climate change impacts in marine Arctic ecosystems reported as of mid‐2009. We found a total of 51 reports of documented changes in Arctic marine biota in response to climate change. Among the responses evaluated were range shifts and changes in abundance, growth/condition, behaviour/phenology and community/regime shifts. Most reports concerned marine mammals, particularly polar bears, and fish. The number of well‐documented changes in planktonic and benthic systems was surprisingly low. Evident losses of endemic species in the Arctic Ocean, and in ice algae production and associated community remained difficult to evaluate due to the lack of quantitative reports of its abundance and distribution. Very few footprints of climate change were reported in the literature from regions such as the wide Siberian shelf and the central Arctic Ocean due to the limited research effort made in these ecosystems. Despite the alarming nature of warming and its strong potential effects in the Arctic Ocean the research effort evaluating the impacts of climate change in this region is rather limited.     

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.