Vulnerability of coastal ecosystems to changes in harmful algal bloom distribution in response to climate change: projections based on model analysis

Harmful algal blooms (HABs), those proliferations of algae that can cause fish kills, contaminate seafood with toxins, form unsightly scums, or detrimentally alter ecosystem function have been increasing in frequency, magnitude, and duration worldwide. Here, using a global modeling approach, we show, for three regions of the globe, the potential effects of nutrient loading and climate change for two HAB genera, pelagic Prorocentrum and Karenia, each with differing physiological characteristics for growth. The projections (end of century, 2090-2100) are based on climate change resulting from the A1B scenario of the Intergovernmental Panel on Climate Change Institut Pierre Simon Laplace Climate Model (IPCC, IPSL-CM4), applied in a coupled oceanographic-biogeochemical model, combined with a suite of assumed physiological 'rules' for genera-specific bloom development. Based on these models, an expansion in area and/or number of months annually conducive to development of these HABs along the NW European Shelf-Baltic Sea system and NE Asia was projected for both HAB genera, but no expansion (Prorocentrum spp.), or actual contraction in area and months conducive for blooms (Karenia spp.), was projected in the SE Asian domain. The implications of these projections, especially for Northern Europe, are shifts in vulnerability of coastal systems to HAB events, increased regional HAB impacts to aquaculture, increased risks to human health and ecosystems, and economic consequences of these events due to losses to fisheries and ecosystem services.

Projections of change in key ecosystem indicators for planning and management of marine protected areas: An example study for European seas

Marine Protected Areas (MPAs) are widely used as tools to maintain biodiversity, protect habitats and ensure that development is sustainable. If MPAs are to maintain their role into the future it is important for managers to understand how conditions at these sites may change as a result of climate change and other drivers, and this understanding needs to extend beyond temperature to a range of key ecosystem indicators. This case study demonstrates how spatially-aggregated model results for multiple variables can provide useful projections for MPA planners and managers. Conditions in European MPAs have been projected for the 2040s using unmitigated and globally managed scenarios of climate change and river management, and hence high and low emissions of greenhouse gases and riverborne nutrients. The results highlight the vulnerability of potential refuge sites in the north-west Mediterranean and the need for careful monitoring at MPAs to the north and west of the British Isles, which may be affected by changes in Atlantic circulation patterns. The projections also support the need for more MPAs in the eastern Mediterranean and Adriatic Sea, and can inform the selection of sites.

Projections of change in key ecosystem indicators for planning and management of marine protected areas: An example study for European seas

Marine Protected Areas (MPAs) are widely used as tools to maintain biodiversity, protect habitats and ensure that development is sustainable. If MPAs are to maintain their role into the future it is important for managers to understand how conditions at these sites may change as a result of climate change and other drivers, and this understanding needs to extend beyond temperature to a range of key ecosystem indicators. This case study demonstrates how spatially-aggregated model results for multiple variables can provide useful projections for MPA planners and managers. Conditions in European MPAs have been projected for the 2040s using unmitigated and globally managed scenarios of climate change and river management, and hence high and low emissions of greenhouse gases and riverborne nutrients. The results highlight the vulnerability of potential refuge sites in the north-west Mediterranean and the need for careful monitoring at MPAs to the north and west of the British Isles, which may be affected by changes in Atlantic circulation patterns. The projections also support the need for more MPAs in the eastern Mediterranean and Adriatic Sea, and can inform the selection of sites.

Macroecology of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas

Global climate change is expected to modify the spatial distribution of marine organisms. However, projections of future changes should be based on robust information on the ecological niche of species. This paper presents a macroecological study of the environmental tolerance and ecological niche (sensu Hutchinson 1957, i.e. the field of tolerance of a species to the principal factors of its environment) of Calanus finmarchicus and C. helgolandicus in the North Atlantic Ocean and adjacent seas. Biological data were collected by the Continuous Plankton Recorder (CPR) Survey, which samples plankton in the North Atlantic and adjacent seas at a standard depth of 7 m. Eleven parameters were chosen including bathymetry, temperature, salinity, nutrients, mixed-layer depth and an index of turbulence compiled from wind data and chlorophyll a concentrations (used herein as an index of available food). The environmental window and the optimum level were determined for both species and for each abiotic factor and chlorophyll concentration. The most important parameters that influenced abundance and spatial distribution were temperature and its correlates such as oxygen and nutrients. Bathymetry and other water-column-related parameters also played an important role. The ecological niche of C. finmarchicus was larger than that of C. helgolandicus and both niches were significantly separated. Our results have important implications in the context of global climate change. As temperature (and to some extent stratification) is predicted to continue to rise in the North Atlantic sector, changes in the spatial distribution of these 2 Calanus species can be expected. Application of this approach to the 1980s North Sea regime shift provides evidence that changes in sea temperature alone could have triggered the substantial and rapid changes identified in the dynamic regimes of these ecosystems. C. finmarchicus appears to be a good indicator of the Atlantic Polar Biome (mainly the Atlantic Subarctic and Arctic provinces) while C. helgolandicus is an indicator of more temperate waters (Atlantic Westerly Winds Biome) in regions characterised by more pronounced spatial changes in bathymetry.

Changes in pH at the exterior surface of plankton with ocean acidification

Anthropogenically released CO2 is dissolving in the ocean, causing a decrease in bulk-seawater pH (ocean acidification). Projections indicate that the pH will drop 0.3 units from its present value by 2100 (ref. 1). However, it is unclear how the growth of plankton is likely to respond. Using simulations we demonstrate how pH and carbonate chemistry at the exterior surface of marine organisms deviates increasingly from those of the bulk sea water as organism metabolic activity and size increases. These deviations will increase in the future as the buffering capacity of sea water decreases with decreased pH and as metabolic activity increases with raised seawater temperatures. We show that many marine plankton will experience pH conditions completely outside their recent historical range. However, ocean acidification is likely to have differing impacts on plankton physiology as taxon-specific differences in organism size, metabolic activity and growth rates during blooms result in very different microenvironments around the organism. This is an important consideration for future studies in ocean acidification as the carbonate chemistry experienced by most planktonic organisms will probably be considerably different from that measured in bulk-seawater samples. An understanding of these deviations will assist interpretation of the impacts of ocean acidification on plankton of different size and metabolic activity.

Potential Climate Change Effects on the Habitat of Antarctic Krill in the Weddell Quadrant of the Southern Ocean

Antarctic krill is a cold water species, an increasingly important fishery resource and a major prey item for many fish, birds and mammals in the Southern Ocean. The fishery and the summer foraging sites of many of these predators are concentrated between 0 degrees and 90 degrees W. Parts of this quadrant have experienced recent localised sea surface warming of up to 0.2 degrees C per decade, and projections suggest that further widespread warming of 0.27 degrees to 1.08 degrees C will occur by the late 21st century. We assessed the potential influence of this projected warming on Antarctic krill habitat with a statistical model that links growth to temperature and chlorophyll concentration. The results divide the quadrant into two zones: a band around the Antarctic Circumpolar Current in which habitat quality is particularly vulnerable to warming, and a southern area which is relatively insensitive. Our analysis suggests that the direct effects of warming could reduce the area of growth habitat by up to 20%. The reduction in growth habitat within the range of predators, such as Antarctic fur seals, that forage from breeding sites on South Georgia could be up to 55%, and the habitat's ability to support Antarctic krill biomass production within this range could be reduced by up to 68%. Sensitivity analysis suggests that the effects of a 50% change in summer chlorophyll concentration could be more significant than the direct effects of warming. A reduction in primary production could lead to further habitat degradation but, even if chlorophyll increased by 50%, projected warming would still cause some degradation of the habitat accessible to predators. While there is considerable uncertainty in these projections, they suggest that future climate change could have a significant negative effect on Antarctic krill growth habitat and, consequently, on Southern Ocean biodiversity and ecosystem services.

Projected impacts of climate change on marine fish and fisheries

This paper reviews current literature on the projected effects of climate change on marine fish and shellfish, their fisheries, and fishery-dependent communities throughout the northern hemisphere. The review addresses the following issues: (i) expected impacts on ecosystem productivity and habitat quantity and quality; (ii) impacts of changes in production and habitat on marine fish and shellfish species including effects on the community species composition, spatial distributions, interactions, and vital rates of fish and shellfish; (iii) impacts on fisheries and their associatedcommunities; (iv) implications for food security and associated changes; and (v) uncertainty andmodelling skill assessment. Climate change will impact fish and shellfish, their fisheries, and fishery-dependent communities through a complex suite of linked processes. Integrated interdisciplinary research teams are forming in many regions to project these complex responses. National and international marine research organizations serve a key role in the coordination and integration of research to accelerate the production of projections of the effects of climate change on marine ecosystems and to move towards a future where relative impacts by region could be compared on a hemispheric or global level. Eight research foci were identified that will improve the projections of climate impacts on fish, fisheries, and fishery-dependent communities.

Modeling marine ecosystem responses to global climate change: Where are we now and where should we be going?

Modeling of global climate change is moving from global circulation model (GCM)-type projections with coupled biogeochemical models to projections of ecological responses, including food web and upper trophic levels. Marine and coastal ecosystems are highly susceptible to the impacts of global climate change and also produce significant ecosystem services. The effects of global climate change on coastal and marine ecosystems involve a much wider array of effects than the usual temperature, sea level rise, and precipitation. This paper is an overview for a collection of 12 papers that examined various aspects of global climate change on marine ecosystems and comprise this special issue. We summarized the major features of the models and analyses in the papers to determine general patterns. A wide range of ecosystems were simulated using a diverse set of modeling approaches. Models were either 3-dimensional or used a few spatial boxes, and responses to global climate change were mostly expressed as changes from a baseline condition. Three issues were identified from the across-model comparison: (a) lack of standardization of climate change scenarios, (b) the prevalence of site-specific and even unique models for upper trophic levels, and (c) emphasis on hypothesis evaluation versus forecasting. We discuss why these issues are important as global climate change assessment continues to progress up the food chain, and, when possible, offer some initial steps for going forward.

Changes in significant and maximum wave heights in the Norwegian Sea

This paper analyses 10 years of in-situ measurements of significant wave height (Hs) and maximum wave height (Hmax) from the ocean weather ship Polarfront in the Norwegian Sea. The 30-minute Ship-Borne Wave Recorder measurements of Hmax and Hs are shown to be consistent with theoretical wave distributions. The linear regression between Hmax and Hs has a slope of 1.53. Neither Hs nor Hmax show a significant trend in the period 2000–2009. These data are combined with earlier observations. The long-term trend over the period 1980–2009 in annual Hs is 2.72 ± 0.88 cm/year. Mean Hs and Hmax are both correlated with the North Atlantic Oscillation (NAO) index during winter. The correlation with the NAO index is highest for the more frequently encountered (75th percentile) wave heights. The wave field variability associated with the NAO index is reconstructed using a 500-year NAO index record. Hs and H max are found to vary by up to 1.42 m and 3.10 m respectively over the 500-year period. Trends in all 30-year segments of the reconstructed wave field are lower than the trend in the observations during 1980–2009. The NAO index does not change significantly in 21st century projections from CMIP5 climate models under scenario RCP85, and thus no NAO-related changes are expected in the mean and extreme wave fields of the Norwegian Sea.

Physiology, ecological niches and species distribution

Although many studies have debated the theoretical links between physiology, ecological niches and species distribution, few studies have provided evidence for a tight empirical coupling between these concepts at a macroecological scale. We used an ecophysiological model to assess the fundamental niche of a key-structural marine species. We found a close relationship between its fundamental and realized niche. The relationship remains constant at both biogeographical and decadal scales, showing that changes in environmental forcing propagate from the physiological to the macroecological level. A substantial shift in the spatial distribution is detected in the North Atlantic and projections of range shift using IPCC scenarios suggest a poleward movement of the species of one degree of latitude per decade for the 21st century. The shift in the spatial distribution of this species reveals a pronounced alteration of polar pelagic ecosystems with likely implications for lower and upper trophic levels and some biogeochemical cycles.

Impacts of Climate Change on European Marine Ecosystems: Observations, Expectations and Indicators

The Northern Hemisphere has been warmer since 1980 than at any other time during the last 2000 years. The observed increase in temperature has been generally higher in northern than in southern European seas, and higher in enclosed than in open seas. Although European marine ecosystems are influenced by many other factors, such as nutrient enrichment and overfishing, every region has shown at least some changes that were most likely attributable to recent climate change. It is expected that within open systems there will generally be (further) northward movement of species, leading to a switch from polar to more temperate species in the northern seas such as the Arctic, Barents Sea and the Nordic Seas, and subtropical species moving northward to temperate regions such as the Iberian upwelling margin. For seas that are highly influenced by river runoff, such as the Baltic Sea, an increase in freshwater due to enhanced rainfall will lead to a shift from marine to more brackish and even freshwater species. If semi-enclosed systems such as the Mediterranean and the Black Sea lose their endemic species, the associated niches will probably be filled by species originating from adjacent waters and, possibly, with species transported from one region to another via ballast water and the Suez Canal. A better understanding of potential climate change impacts (scenarios) at both regional and local levels, the development of improved methods to quantify the uncertainty of climate change projections, the construction of usable climate change indicators, and an improvement of the interface between science and policy formulation in terms of risk assessment will be essential to formulate and inform better adaptive strategies to address the inevitable consequences of climate change.

Can we trust simple marine DMS parameterisations within complex climate models?

Dimethylsulphide (DMS) is a globally important aerosol precurser. In 1987 Charlson and others proposed that an increase in DMS production by certain phytoplankton species in response to a warming climate could stimulate increased aerosol formation, increasing the lower-atmosphere's albedo, and promoting cooling. Despite two decades of research, the global significance of this negative climate feedback remains contentious. It is therefore imperative that schemes are developed and tested, which allow for the realistic incorporation of phytoplankton DMS production into Earth System models. Using these models we can investigate the DMS-climate feedback and reduce uncertainty surrounding projections of future climate. Here we examine two empirical DMS parameterisations within the context of an Earth System model and find them to perform marginally better than the standard DMS climatology at predicting observations from an independent global dataset. We then question whether parameterisations based on our present understanding of DMS production by phytoplankton, and simple enough to incorporate into global climate models, can be shown to enhance the future predictive capacity of those models. This is an important question to ask now, as results from increasingly complex Earth System models lead us into the 5th assessment of climate science by the Intergovernmental Panel on Climate Change. Comparing observed and predicted inter-annual variability, we suggest that future climate projections may underestimate the magnitude of surface ocean DMS change. Unfortunately this conclusion relies on a relatively small dataset, in which observed inter-annual variability may be exaggerated by biases in sample collection. We therefore encourage the observational community to make repeat measurements of sea-surface DMS concentrations an important focus, and highlight areas of apparent high inter-annual variability where sampling might be carried out. Finally, we assess future projections from two similarly valid empirical DMS schemes, and demonstrate contrasting results. We therefore conclude that the use of empirical DMS parameterisations within simulations of future climate should be undertaken only with careful appreciation of the caveats discussed.

Biomass changes and trophic amplification of plankton in a warmer ocean

Ocean warming can modify the ecophysiology and distribution of marine organisms, and relationships between species, with nonlinear interactions between ecosystem components potentially resulting in trophic amplification. Trophic amplification (or attenuation) describe the propagation of a hydroclimatic signal up the food web, causing magnification (or depression) of biomass values along one or more trophic pathways. We have employed 3-D coupled physical-biogeochemical models to explore ecosystem responses to climate change with a focus on trophic amplification. The response of phytoplankton and zooplankton to global climate-change projections, carried out with the IPSL Earth System Model by the end of the century, is analysed at global and regional basis, including European seas (NE Atlantic, Barents Sea, Baltic Sea, Black Sea, Bay of Biscay, Adriatic Sea, Aegean Sea) and the Eastern Boundary Upwelling System (Benguela). Results indicate that globally and in Atlantic Margin and North Sea, increased ocean stratification causes primary production and zooplankton biomass to decrease in response to a warming climate, whilst in the Barents, Baltic and Black Seas, primary production and zooplankton biomass increase. Projected warming characterized by an increase in sea surface temperature of 2.29 ± 0.05 °C leads to a reduction in zooplankton and phytoplankton biomasses of 11% and 6%, respectively. This suggests negative amplification of climate driven modifications of trophic level biomass through bottom-up control, leading to a reduced capacity of oceans to regulate climate through the biological carbon pump. Simulations suggest negative amplification is the dominant response across 47% of the ocean surface and prevails in the tropical oceans; whilst positive trophic amplification prevails in the Arctic and Antarctic oceans. Trophic attenuation is projected in temperate seas. Uncertainties in ocean plankton projections, associated to the use of single global and regional models, imply the need for caution when extending these considerations into higher trophic levels.

Have we been underestimating the effects of ocean acidification in zooplankton?

Understanding how copepods may respond to ocean acidification (OA) is critical for risk assessments of ocean ecology and biogeochemistry. The perception that copepods are insensitive to OA is largely based on experiments with adult females. Their apparent resilience to increased carbon dioxide (pCO2 ) concentrations has supported the view that copepods are 'winners' under OA. Here, we show that this conclusion is not robust, that sensitivity across different life stages is significantly misrepresented by studies solely using adult females. Stage-specific responses to pCO2 (385-6000 μatm) were studied across different life stages of a calanoid copepod, monitoring for lethal and sublethal responses. Mortality rates varied significantly across the different life stages, with nauplii showing the highest lethal effects; nauplii mortality rates increased threefold when pCO2 concentrations reached 1000 μatm (year 2100 scenario) with LC50 at 1084 μatm pCO2 . In comparison, eggs, early copepodite stages, and adult males and females were not affected lethally until pCO2 concentrations ≥3000 μatm. Adverse effects on reproduction were found, with >35% decline in nauplii recruitment at 1000 μatm pCO2 . This suppression of reproductive scope, coupled with the decreased survival of early stage progeny at this pCO2 concentration, has clear potential to damage population growth dynamics in this species. The disparity in responses seen across the different developmental stages emphasizes the need for a holistic life-cycle approach to make species-level projections to climate change. Significant misrepresentation and error propagation can develop from studies which attempt to project outcomes to future OA conditions solely based on single life history stage exposures.

Response to Comments on "Impacts of Biodiversity Loss on Ocean Ecosystem Services"

We show that globally declining fisheries catch trends cannot be explained by random processes and are consistent with declining stock abundance trends. Future projections are inherently uncertain but may provide a benchmark against which to assess the effectiveness of conservation measures. Marine reserves and fisheries closures are among those measures and can be equally effective in tropical and temperate areas—but must be combined with catch-, effort-, and gear restrictions to meet global conservation objectives.