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.     

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.     

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.     

What's going to be on the menu with global environmental changes?

Ongoing anthropogenic change is altering the planet at an unprecedented rate, threatening biodiversity, and ecosystem functioning. Species are responding to abiotic pressures at both individual and population levels, with changes affecting trophic interactions through consumptive pathways. Collectively, these impacts alter the goods and services that natural ecosystems will provide to society, as well as the persistence of all species. Here, we describe the physiological and behavioral responses of species to global changes on individual and population levels that result in detectable changes in diet across terrestrial and marine ecosystems. We illustrate shifts in the dynamics of food webs with implications for animal communities. Additionally, we highlight the myriad of tools available for researchers to investigate the dynamics of consumption patterns and trophic interactions, arguing that diet data are a crucial component of ecological studies on global change. We suggest that a holistic approach integrating the complexities of diet choice and trophic interactions with environmental drivers may be more robust at resolving trends in biodiversity, predicting food web responses, and potentially identifying early warning signs of diversity loss. Ultimately, despite the growing body of long-term ecological datasets, there remains a dearth of diet ecology studies across temporal scales, a shortcoming that must be resolved to elucidate vulnerabilities to changing biophysical conditions.     

Climate change, biotic interactions and ecosystem services

Climate change is real. The wrangling debates are over, and we now need to move onto a predictive ecology that will allow managers of landscapes and policy makers to adapt to the likely changes in biodiversity over the coming decades. There is ample evidence that ecological responses are already occurring at the individual species (population) level. The challenge is how to synthesize the growing list of such observations with a coherent body of theory that will enable us to predict where and when changes will occur, what the consequences might be for the conservation and sustainable use of biodiversity and what we might do practically in order to maintain those systems in as good condition as possible. It is thus necessary to investigate the effects of climate change at the ecosystem level and to consider novel emergent ecosystems composed of new species assemblages arising from differential rates of range shifts of species. Here, we present current knowledge on the effects of climate change on biotic interactions and ecosystem services supply, and summarize the papers included in this volume. We discuss how resilient ecosystems are in the face of the multiple components that characterize climate change, and suggest which current ecological theories may be used as a starting point to predict ecosystem-level effects of climate change.     

Impacts of recent climate change on terrestrial flora and fauna: Some emerging Australian examples

The effects of anthropogenic climate change on biodiversity are well known for some high‐profile Australian marine systems, including coral bleaching and kelp forest devastation. Less well‐published are the impacts of climate change being observed in terrestrial ecosystems, although ecological models have predicted substantial changes are likely. Detecting and attributing terrestrial changes to anthropogenic factors is difficult due to the ecological importance of extreme conditions, the noisy nature of short‐term data collected with limited resources, and complexities introduced by biotic interactions. Here, we provide a suite of case studies that have considered possible impacts of anthropogenic climate change on Australian terrestrial systems. Our intention is to provide a diverse collection of stories illustrating how Australian flora and fauna are likely responding to direct and indirect effects of anthropogenic climate change. We aim to raise awareness rather than be comprehensive. We include case studies covering canopy dieback in forests, compositional shifts in vegetation, positive feedbacks between climate, vegetation and disturbance regimes, local extinctions in plants, size changes in birds, phenological shifts in reproduction and shifting biotic interactions that threaten communities and endangered species. Some of these changes are direct and clear cut, others are indirect and less clearly connected to climate change; however, all are important in providing insights into the future state of terrestrial ecosystems. We also highlight some of the management issues relevant to conserving terrestrial communities and ecosystems in the face of anthropogenic climate change.     

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.     

Seagrass sediments reveal the long‐term deterioration of an estuarine ecosystem

The study of a Posidonia australis sediment archive has provided a record of ecosystem dynamics and processes over the last 600 years in Oyster Harbour (SW Australia). Ecosystem shifts are a widespread phenomenon in coastal areas, and this study identifies baseline conditions and the time‐course of ecological change (cycles, trends, resilience and thresholds of ecosystem change) under environmental stress in seagrass‐dominated ecosystem. The shifts in the concentrations of chemical elements, carbonates, sediments <0.125 mm and stable carbon isotope signatures (δ¹³C) of the organic matter were detected between 1850s and 1920s, whereas the shift detected in P concentration occurred several decades later (1960s). The first degradation phase (1850s–1950s) follows the onset of European settlement in Australia and was characterized by a strong increase in sediment accumulation rates and fine‐grained particles, driven primarily by enhanced run‐off due to land clearance and agriculture in the catchment. About 80% of total seagrass area at Oyster Harbour was lost during the second phase of environmental degradation (1960s until present). The sharp increase in P concentration and the increasing contribution of algae and terrestrial inputs into the sedimentary organic matter pool around 1960s provides compelling evidence of the documented eutrophication of the estuary and the subsequent loss of seagrass meadows. The results presented demonstrate the power of seagrass sedimentary archives to reconstruct the trajectories of anthropogenic pressures on estuarine ecosystem and the associated regime shifts, which can be used to improve the capacity of scientists and environmental managers to understand, predict and better manage ecological change in these ecosystems.     

Abrupt changes in Great Britain vegetation carbon projected under climate change

Past abrupt ‘regime shifts’ have been observed in a range of ecosystems due to various forcing factors. Large‐scale abrupt shifts are projected for some terrestrial ecosystems under climate change, particularly in tropical and high‐latitude regions. However, there is very little high‐resolution modelling of smaller‐scale future projected abrupt shifts in ecosystems, and relatively less focus on the potential for abrupt shifts in temperate terrestrial ecosystems. Here, we show that numerous climate‐driven abrupt shifts in vegetation carbon are projected in a high‐resolution model of Great Britain's land surface driven by two different climate change scenarios. In each scenario, the effects of climate and CO₂ combined are isolated from the effects of climate change alone. We use a new algorithm to detect and classify abrupt shifts in model time series, assessing the sign and strength of the non‐linear responses. The abrupt ecosystem changes projected are non‐linear responses to climate change, not simply driven by abrupt shifts in climate. Depending on the scenario, 374–1,144 grid cells of 1.5 km × 1.5 km each, comprising 0.5%–1.5% of Great Britain's land area show abrupt shifts in vegetation carbon. We find that abrupt ecosystem shifts associated with increases (rather than decreases) in vegetation carbon, show the greatest potential for early warning signals (rising autocorrelation and variance beforehand). In one scenario, 89% of abrupt increases in vegetation carbon show increasing autocorrelation and variance beforehand. Across the scenarios, 81% of abrupt increases in vegetation carbon have increasing autocorrelation and 74% increasing variance beforehand, whereas for decreases in vegetation carbon these figures are 56% and 47% respectively. Our results should not be taken as specific spatial or temporal predictions of abrupt ecosystem change. However, they serve to illustrate that numerous abrupt shifts in temperate terrestrial ecosystems could occur in a changing climate, with some early warning signals detectable beforehand.     

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.     

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).     

Spatio-temporal dynamics of multi-trophic communities reveal ecosystem-wide functional reorganization

Large-scale alterations in marine ecosystems as a response to environmental and anthropogenic pressures have been documented worldwide. Yet, these are primarily investigated by assessing abundance fluctuations of a few dominant species, which inadequately reflect ecosystem-wide changes. In addition, it is increasingly recognized that it is not species identity per se, but their traits that determine environmental responses, biological interactions and ecosystem functioning. In this study, we investigated long-term, spatio-temporal variability in trait composition across multiple organism groups to assess whether functional changes occur in a similar way across trophic levels and whether shifts in trait composition link to environmental change. We combined extensive trait datasets with long-term surveys (30–40 yr) of four organism groups (phytoplankton, zooplankton, benthic invertebrates and fish) in three environmentally distinct areas of a large marine ecosystem. We found similar temporal trajectories in the community weighted mean trait time-series of the different trophic groups, revealing ecosystem-wide functional changes. The traits involved and their dynamics differed between areas, concurrent with climate-driven changes in temperature and salinity, as well as more local dynamics in nutrients and oxygen. This finding highlights the importance of considering both global climate, as well as local external drivers when studying ecosystem changes. Using a multi-trophic trait-based approach, our study demonstrates the importance of integrating community functional dynamics across multiple trophic levels to capture ecosystem-wide responses which could, ultimately, help moving towards a holistic understanding, assessment and management of marine ecosystems.     

EFFECTS OF CLIMATE CHANGE ON GLOBAL SEAWEED COMMUNITIES

Seaweeds are ecologically important primary producers, competitors, and ecosystem engineers that play a central role in coastal habitats ranging from kelp forests to coral reefs. Although seaweeds are known to be vulnerable to physical and chemical changes in the marine environment, the impacts of ongoing and future anthropogenic climate change in seaweed‐dominated ecosystems remain poorly understood. In this review, we describe the ways in which changes in the environment directly affect seaweeds in terms of their physiology, growth, reproduction, and survival. We consider the extent to which seaweed species may be able to respond to these changes via adaptation or migration. We also examine the extensive reshuffling of communities that is occurring as the ecological balance between competing species changes, and as top‐down control by herbivores becomes stronger or weaker. Finally, we delve into some of the ecosystem‐level responses to these changes, including changes in primary productivity, diversity, and resilience. Although there are several key areas in which ecological insight is lacking, we suggest that reasonable climate‐related hypotheses can be developed and tested based on current information. By strategically prioritizing research in the areas of complex environmental variation, multiple stressor effects, evolutionary adaptation, and population, community, and ecosystem‐level responses, we can rapidly build upon our current understanding of seaweed biology and climate change ecology to more effectively conserve and manage coastal ecosystems.     

Synergistic Effects of Climate and Fishing in a Marine Ecosystem

Current climate change and overfishing are affecting the productivity and structure of marine ecosystems. This situation is unprecedented for the marine biosphere and it is essential to understand the mechanisms and pathways by which ecosystems respond. We report that climate change and overfishing are likely to be responsible for a rapid restructuring of a highly productive marine ecosystem with effects throughout the pelagos and the benthos. In the mid-1980s, climate change, consequent modifications in the North Sea plankton, and fishing, all reduced North Sea cod recruitment. In this region, production of many benthic species respond positively and immediately to temperature. Analysis of a long-term, spatially extensive biological (plankton and cod) and physical (sea surface temperature) dataset suggests that synchronous changes in cod numbers and sea temperature have established an extensive trophic cascade favoring lower trophic level groups over economic fisheries. A proliferation of jellyfish that we detect may signal the climax of these changes. This modified North Sea ecology may provide a clear indication of the synergistic consequences of coincident climate change and overfishing. The extent of the ecosystem restructuring that has occurred in the North Sea suggests we are unlikely to reverse current climate and human-induced effects through ecosystem resource management in the short term. Rather, we should understand and adapt to new ecological regimes. This implies that fisheries management policies will have to be fully integrated with the ecological consequences of climate change to prevent a similar collapse in an exploited marine ecosystem elsewhere. Author Contributions  RRK conceived the project and GB analysed the data. RRK, GB and JAL co-wrote the paper.     

Circumpolar dynamics of a marine top‐predator track ocean warming rates

Global warming is a nonlinear process, and temperature may increase in a stepwise manner. Periods of abrupt warming can trigger persistent changes in the state of ecosystems, also called regime shifts. The responses of organisms to abrupt warming and associated regime shifts can be unlike responses to periods of slow or moderate change. Understanding of nonlinearity in the biological responses to climate warming is needed to assess the consequences of ongoing climate change. Here, we demonstrate that the population dynamics of a long‐lived, wide‐ranging marine predator are associated with changes in the rate of ocean warming. Data from 556 colonies of black‐legged kittiwakes Rissa tridactyla distributed throughout its breeding range revealed that an abrupt warming of sea‐surface temperature in the 1990s coincided with steep kittiwake population decline. Periods of moderate warming in sea temperatures did not seem to affect kittiwake dynamics. The rapid warming observed in the 1990s may have driven large‐scale, circumpolar marine ecosystem shifts that strongly affected kittiwakes through bottom‐up effects. Our study sheds light on the nonlinear response of a circumpolar seabird to large‐scale changes in oceanographic conditions and indicates that marine top predators may be more sensitive to the rate of ocean warming rather than to warming itself.     

Robust ecological drought projections for drylands in the 21st century

Dryland ecosystems may be especially vulnerable to expected 21st century increases in temperature and aridity because they are tightly controlled by moisture availability. However, climate impact assessments in drylands are difficult because ecological dynamics are dictated by drought conditions that are difficult to define and complex to estimate from climate conditions alone. In addition, precipitation projections vary substantially among climate models, enhancing variation in overall trajectories for aridity. Here, we constrain this uncertainty by utilizing an ecosystem water balance model to quantify drought conditions with recognized ecological importance, and by identifying changes in ecological drought conditions that are robust among climate models, defined here as when >90% of models agree in the direction of change. Despite limited evidence for robust changes in precipitation, changes in ecological drought are robust over large portions of drylands in the United States and Canada. Our results suggest strong regional differences in long‐term drought trajectories, epitomized by chronic drought increases in southern areas, notably the Upper Gila Mountains and South‐Central Semi‐arid Prairies, and decreases in the north, particularly portions of the Temperate and West‐Central Semi‐arid Prairies. However, we also found that exposure to hot‐dry stress is increasing faster than mean annual temperature over most of these drylands, and those increases are greatest in northern areas. Robust shifts in seasonal drought are most apparent during the cool season; when soil water availability is projected to increase in northern regions and decrease in southern regions. The implications of these robust drought trajectories for ecosystems will vary geographically, and these results provide useful insights about the impact of climate change on these dryland ecosystems. More broadly, this approach of identifying robust changes in ecological drought may be useful for other assessments of climate impacts in drylands and provide a more rigorous foundation for making long‐term strategic resource management decisions.     

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).     

Antarctica: The final frontier for marine biological invasions

Antarctica is experiencing significant ecological and environmental change, which may facilitate the establishment of non‐native marine species. Non‐native marine species will interact with other anthropogenic stressors affecting Antarctic ecosystems, such as climate change (warming, ocean acidification) and pollution, with irreversible ramifications for biodiversity and ecosystem services. We review current knowledge of non‐native marine species in the Antarctic region, the physical and physiological factors that resist establishment of non‐native marine species, changes to resistance under climate change, the role of legislation in limiting marine introductions, and the effect of increasing human activity on vectors and pathways of introduction. Evidence of non‐native marine species is limited: just four marine non‐native and one cryptogenic species that were likely introduced anthropogenically have been reported freely living in Antarctic or sub‐Antarctic waters, but no established populations have been reported; an additional six species have been observed in pathways to Antarctica that are potentially at risk of becoming invasive. We present estimates of the intensity of ship activity across fishing, tourism and research sectors: there may be approximately 180 vessels and 500+ voyages in Antarctic waters annually. However, these estimates are necessarily speculative because relevant data are scarce. To facilitate well‐informed policy and management, we make recommendations for future research into the likelihood of marine biological invasions in the Antarctic region.     

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.