Climate change and marine plankton

Understanding how climate change will affect the planet is a key issue worldwide. Questions concerning the pace and impacts of climate change are thus central to many ecological and biogeochemical studies, and addressing the consequences of climate change is now high on the list of priorities for funding agencies. Here, we review the interactions between climate change and plankton communities, focusing on systematic changes in plankton community structure, abundance, distribution and phenology over recent decades. We examine the potential socioeconomic impacts of these plankton changes, such as the effects of bottom-up forcing on commercially exploited fish stocks (i.e. plankton as food for fish). We also consider the crucial roles that plankton might have in dictating the future pace of climate change via feedback mechanisms responding to elevated atmospheric CO sub(2) levels. An important message emerges from this review: ongoing plankton monitoring programmes worldwide will act as sentinels to identify future changes in marine ecosystems.

Impacts of fisheries on plankton community structure

There has been much debate on the extent to which resource availability (bottom-up) versus predation pressure from fish (top-down) modulates the dynamics of plankton in marine systems. Physico/chemical bottom-up forcing has been considered to be the main mechanism structuring marine ecosystems, although some field observations and empirical correlations support top-down modulation. Models have indicated possible feedback loops to the plankton and other studies have interpreted a grazing impact from long-term changes in fish stocks. In freshwater systems, evidence for top-down forcing by fish and trophic cascading is well documented. First, evidence for equivalent top-down effects in the marine environment is presented, with an overview of relevant publications. In the second part, time series, averaged for the North Sea (when possible from 1948 to 1997), of fish catch, recruitment, and spawning stock biomass are related to the abundance of species or larger groupings of zooplankton and phytoplankton from the Continuous Plankton Recorder survey and selected environmental parameters. Preliminary analysis suggests that there is strong interaction between different fish species and the plankton and that the fishery, through top-down control, may at times be an important contributor to changes in the North Sea ecosystem.

Long-term changes in the pelagos, benthos and fisheries of the North Sea

Pronounced changes have occurred in the fisheries, plankton and benthos of the North Sea over the last five decades. Attribution of the relative contribution of anthropogenic versus natural hydrometeorological modulation to these changes is still unclear. As a background a summary history of our understanding of the state of health of the North Sea is outlined. We then focus on two contrasting periods in the North Sea, one between 1978-82 (cold) and the other post 1987 (warm) when pronounced alterations in many ecosystem characteristics occurred. The scale of the changes in the second of these periods is sufficiently large and wide ranging for it to have been termed a regime shift. A combination of local, regional and far field hydrometeorological forcing, and in particular variability in oceanic inflow, is believed to be responsible for the observed changes. Finally attention is drawn to the poor status of North Sea fish stocks where 7 stocks are documented as being fished outside safe biological limits. This situation is primarily believed to be a consequence of overfishing, but may have been exacerbated by environmental change.

From plankton to top predators: bottom-up control of a marine food web across four trophic levels

1. Abundant mid-trophic pelagic fish often play a central role in marine ecosystems, both as links between zooplankton and top predators and as important fishery targets. In the North Sea, the lesser sandeel occupies this position, being the main prey of many bird, mammal and fish predators and the target of a major industrial fishery. However, since 2003, sandeel landings have decreased by > 50%, and many sandeel-dependent seabirds experienced breeding failures in 2004. 2. Despite the major economic implications, current understanding of the regulation of key constituents of this ecosystem is poor. Sandeel abundance may be regulated 'bottom-up' by food abundance, often thought to be under climatic control, or 'top-down' by natural or fishery predation. We tested predictions from these two hypotheses by combining unique long-term data sets (1973–2003) on seabird breeding productivity from the Isle of May, SE Scotland, and plankton and fish larvae from the Continuous Plankton Recorder survey. We also tested whether seabird breeding productivity was more tightly linked to sandeel biomass or quality (size) of individual fish. 3. The biomass of larval sandeels increased two- to threefold over the study period and was positively associated with proxies of the abundance of their plankton prey. Breeding productivity of four seabirds bringing multiple prey items to their offspring was positively related to sandeel larval biomass with a 1-year lag, indicating dependence on 1-year-old fish, but in one species bringing individual fish it was strongly associated with the size of adult sandeels. 4. These links are consistent with bottom-up ecosystem regulation and, with evidence from previous studies, indicate how climate-driven changes in plankton communities can affect top predators and potentially human fisheries through the dynamics of key mid-trophic fish. However, the failing recruitment to adult sandeel stocks and the exceptionally low seabird breeding productivity in 2004 were not associated with low sandeel larval biomass in 2003, so other mechanisms (e.g. predation, lack of suitable food after metamorphosis) must have been important in this case. Understanding ecosystem regulation is extremely important for predicting the fate of keystone species, such as sandeels, and their predators.

Continuous Plankton Records - Seasonal-Variations In The Distribution And Abundance Of Plankton In The North-Atlantic Ocean And The North-Sea

Charts are presented of the seasonal variations in the distribution of four phytoplankton and five zooplankton taxa in the North Atlantic and the North Sea. The main factors determining the seasonal variations appear to be the distribution of the main overwintering stocks, the current system and, in some instances, temperature control of the rate of population increase. Information is presented about the variation with latitude (over the range from 34° N to 65 ° N) of the seasonal regime of the plankton. On the assumption that there is a relationship between nutrient supply and vertical temperature stratification the main features of this variability can be interpreted. In the south (to about 43° N) nutrient limitation plus grazing appear to be dominant, resulting in a bimodal seasonal cycle of phytoplankton. North of about 60° N the system appears to be limited by the size of the phytoplankton stocks being grazed primarily by Calanus Finmarchicus and Euphausiacea. In an extensive zone, from about 44° N to 60° N, it would appear that the spring bloom of phytoplankton is under-exploited by grazing while in summer the zooplankton graze the daily production of the phytoplankton, the stocks of which are probably maintained by in situ nutrient regeneration. The implications, for at least this mid-latitude zone, that rates and fluxes of processes, as opposed to density dependent interactions between stocks, play a major role in the dynamics of the seasonal cycle is consistent with previously reported observations suggesting that physical environmental factors play a major role in determining year-to-year fluctuations in the abundance of the plankton.

Continuous Plankton Records - Persistence In Time-Series And The Population-Dynamics Of Pseudocalanus-Elongatus And Acartia-Clausi

Rates of population increase in early spring and the sizes of overwintering stocks were calculated for the planktonic copepods Pseudocalanus elongatus and Acartia clausi for a set of areas covering the open waters of the north-east Atlantic Ocean and the North Sea for the period 1948 to 1979. For both species, the rates of population increase were higher in the open ocean than in the North Sea and appear to be related to temperature. The overwintering stocks in the North Sea were larger than those in the open ocean and are probably related to phytoplanton concentration. P. elongatus shows higher overwintering stocks and lower rates of population increase than A. clausi, resulting in different levels of persistence in the stocks of the two species. It is suggested that this difference in persistence is responsible for differences between the two species with respect to geographical distribution in summer and different patterns of year-to-year fluctuations in abundance.

Continuous Plankton Records - Phytoplankton Zooplankton And Environment Northeast Atlantic And North-Sea 1958-1980

Year-to-year fluctuations in the abundance of phytoplankton in the North-East Atlantic and the North Sea for the period 1958 to 1980 are described. Based on similarities between their annual fluctuations in abundance, the taxa may be divided into two groups, one of 12 species of diatoms and 1 species of Ceratium, the other of 5 species of Ceratium. The annual fluctuations in abundance of the Ceratium group is negatively correlated with a component of sea surface temperature (representing changes in the open ocean) and with the frequency of cyclonic weather over the United Kingdom. The Diatom group shows very similar annual fluctuations to those of most of the zooplankton species. Both groups show a high ·proportion of long wavelength variability in the form of a more less linear downward trend in abundance over the whole period. There is evidence to suggest that the high proportion of long wavelength variability shown by the zooplankton is influenced by inherent persistence in stocks from year-to year. The phytoplankton show little or no persistence. The close relationship between zooplankton and phytoplankton may, therefore, involve feed-back through nutrient recycling so influencing the annual levels of abundance of phytoplankton.

Continuous Plankton Records - Persistence In Time-Series Of Annual Means Of Abundance Of Zooplankton

Time-series of annual means of abundance of zooplankton of the north-east Atlantic Ocean and the North Sea, for the period 1948 to 1977, show considerable associations between successive years. The seasonal dynamics of the stocks appear to be consistent with at least a proportion of this being due to inherent persistence from year-to-year. Experiments with a simple model suggest that the observed properties of the time-series cannot be reproduced as a response to simple random forcing. The extent of trends and long wavelength variations can be simulated by introducing fairly extensive persistence into the perturbations, but this underestimates the extent of shorter wavelength variability in the observed time-series. The effect of persistence is to increase the proportion of trend and long wavelength variability in time-series of annual means, but stocks can respond to short wavelength perturbations provided these have a clearly defined frequency.

Towards an integrated forecasting system for pelagic fisheries

First results of a coupled modeling and forecasting system for the pelagic fisheries are being presented. The system consists currently of three mathematically fundamentally different model subsystems: POLCOMS-ERSEM providing the physical-biogeochemical environment implemented in the domain of the North-West European shelf and the SPAM model which describes sandeel stocks in the North Sea. The third component, the SLAM model, connects POLCOMS-ERSEM and SPAM by computing the physical-biological interaction. Our major experience by the coupling model subsystems is that well-defined and generic model interfaces are very important for a successful and extendable coupled model framework. The integrated approach, simulating ecosystem dynamics from physics to fish, allows for analysis of the pathways in the ecosystem to investigate the propagation of changes in the ocean climate and lower trophic levels to quantify the impacts on the higher trophic level, in this case the sandeel population, demonstrated here on the base of hindcast data. The coupled forecasting system is tested for some typical scientific questions appearing in spatial fish stock management and marine spatial planning, including determination of local and basin scale maximum sustainable yield, stock connectivity and source/sink structure. Our presented simulations indicate that sandeels stocks are currently exploited close to the maximum sustainable yield, but large uncertainty is associated with determining stock maximum sustainable yield due to stock eigen dynamics and climatic variability. Our statistical ensemble simulations indicates that the predictive horizon set by climate interannual variability is 2–6 yr, after which only an asymptotic probability distribution of stock properties, like biomass, are predictable.

Review of climate change impacts on marine fisheries in the UK and Ireland

1.Commercial fishing is an important socio-economic activity in coastal regions of the UK and Ireland. Ocean–atmospheric changes caused by greenhouse gas emissions are likely to affect future fish and shellfish production, and lead to increasing challenges in ensuring long-term sustainable fisheries management. 2.The paper reviews existing knowledge and understanding of the exposure of marine ecosystems to ocean-atmospheric changes, the consequences of these changes for marine fisheries in the UK and Ireland, and the adaptability of the UK and Irish fisheries sector. 3.Ocean warming is resulting in shifts in the distribution of exploited species and is affecting the productivity of fish stocks and underlying marine ecosystems. In addition, some studies suggest that ocean acidification may have large potential impacts on fisheries resources, in particular shell-forming invertebrates. 4.These changes may lead to loss of productivity, but also the opening of new fishing opportunities, depending on the interactions between climate impacts, fishing grounds and fleet types. They will also affect fishing regulations, the price of fish products and operating costs, which in turn will affect the economic performance of the UK and Irish fleets. 5.Key knowledge gaps exist in our understanding of the implications of climate and ocean chemistry changes for marine fisheries in the UK and Ireland, particularly on the social and economic responses of the fishing sectors to climate change. However, these gaps should not delay climate change mitigation and adaptation policy actions, particularly those measures that clearly have other ‘co-benefits’.

Severe tissue damage in Atlantic cod larvae under increasing ocean acidification

Ocean acidification, caused by increasing atmospheric concentrations of CO2 (refs 1-3), is one of the most critical anthropogenic threats to marine life. Changes in seawater carbonate chemistry have the potential to disturb calcification, acid-base regulation, blood circulation and respiration, as well as the nervous system of marine organisms, leading to long-term effects such as reduced growth rates and reproduction(4,5). In teleost fishes, early life-history stages are particularly vulnerable as they lack specialized internal pH regulatory mechanisms(6,7). So far, impacts of relevant CO2 concentrations on larval fish have been found in behaviour(8,9) and otolith size(10,11), mainly in tropical, non-commercial species. Here we show detrimental effects of ocean acidification on the development of a mass-spawning fish species of high. commercial importance. We reared Atlantic cod larvae at three levels of CO2, (1) present day, (2) end of next century and (3) an extreme, coastal upwelling scenario, in a long-term (2; months) mesocosm experiment. Exposure to CO2 resulted in severe to lethal tissue damage in many internal organs, with the degree of damage increasing with CO2 concentration. As larval survival is the bottleneck to recruitment, ocean acidification has the potential to act as an additional source of natural mortality, affecting populations of already exploited fish stocks.

Causes and prediction 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.

Spatial changes in the sensitivity of Atlantic cod to climate-driven effects in the plankton

Recent strategies to sustain fish stocks have suggested a move towards an ecosystem based fisheries management (EBFM) approach. While EBFM considers the effect of fishing at the ecosystem level, it generally struggles with climate-driven environmental variability. In this study we show that the position of a fish stock within its distributional range or thermal niche (we use Icelandic and North Sea cod as examples of stocks at the centre and edge of their niche, respectively) will influence the relative importance of fishing and climate on abundance. At the warmer edge of the thermal niche of cod in the North Sea, we show a prominent influence of climate on the cod stock that is mediated through temperature effects on the plankton. In contrast, the influence of climate through its effects on plankton appears much less important at the present centre of the niche around Iceland. Recognising the potentially strong effect of climate on fish stocks, at a time of rapid global climate change, is probably an important prerequisite towards the synthesis of a cod management strategy.

Climate, plankton and cod

While a few North Atlantic cod stocks are stable, none have increased and many have declined in recent years. Although overfishing is the main cause of most observed declines, this study shows that in some regions, climate by its influence on plankton may exert a strong control on cod stocks, complicating the management of this species that often assumes a constant carrying capacity. First, we investigate the likely drivers of changes in the cod stock in the North Sea by evaluating the potential relationships between climate, plankton and cod. We do this by deriving a Plankton Index that reflects the quality and quantity of plankton food available for larval cod. We show that this Plankton Index explains 46.24% of the total variance in cod recruitment and 68.89% of the variance in total cod biomass. Because the effects of climate act predominantly through plankton during the larval stage of cod development, our results indicate a pronounced sensitivity of cod stocks to climate at the warmer, southern edge of their distribution, for example in the North Sea. Our analyses also reveal for the first time, that at a large basin scale, the abundance of Calanus finmarchicus is associated with a high probability of cod occurrence, whereas the genus Pseudocalanus appears less important. Ecosystem-based fisheries management (EBFM) generally considers the effect of fishing on the ecosystem and not the effect of climate-induced changes in the ecosystem state for the living resources. These results suggest that EBFM must consider the position of a stock within its ecological niche, the direct effects of climate and the influence of climate on the trophodynamics of the ecosystem.

A synthesis of large-scale patterns in the planktonic prey of larval and juvenile cod (Gadus morhua)

Data from 40 published studies of the diet composition of larval and juvenile cod (Gadus morhua) from around the northern North Atlantic were summarized to assess generic patterns in ontogenetic and regional variability in the key prey. The results showed that larvae at the northern edge of the latitudinal range of cod depend primarily on development stages of the copepod Calanus finmarchicus, whilst those at the southern edge depend on Para- and Pseudocalanus species. Juvenile cod preyed on a wider range of taxa than larvae, but euphausiids were the main target prey. Analysis of regional variations in the relative abundances of C. finmarchicus and Para/Pseudocalanus spp. in the plankton, as estimated by the continuous plankton recorder (CPR) surveys, showed a similar geographical pattern to the larval cod stomach contents. Comparison of CPR data from the 1960s and 70s with data from the 1990s showed that the boundary between C. finmarchicus and Para/Pseudocalanus spp. dominance has shifted northwards on both sides of the Atlantic, whilst the abundance of euphausiids in the southern cod stock regions has declined. The results are discussed in relation to regional differences in the response of cod stocks to climate variability.