Mechanisms of long-term decline in size of lesser sandeels in the North Sea explored using a growth and phenology model

The lesser sandeel Ammodytes marinus is a key species in the North Sea ecosystem, transferring energy from planktonic producers to top predators. Previous studies have shown a long-term decline in the size of 0-group sandeels in the western North Sea, but they were unable to pinpoint the mechanism (later hatching, slower growth or changes in size-dependent mortality) or cause. To investigate the first 2 possibilities we combined 2 independent time series of sandeel size, namely data from chick-feeding Atlantic puffins Fratercula arctica and from the Continuous Plankton Recorder (CPR), in a novel statistical model implemented using Markov Chain Monte Carlo (MCMC). The model estimated annual mean length on 1 July, as well as hatching date and growth rate for sandeels from 1973 to 2006. Mean length-at-date declined by 22% over this period, corresponding to a 60% decrease in energy content, with a sharper decline since 2002. Up to the mid-1990s, the decline was associated with a trend towards later hatching. Subsequently, hatching became earlier again, and the continued trend towards smaller size appears to have been driven by lower growth rates, particularly in the most recent years, although we could not rule out changes in size-dependent mortality. Our findings point to major changes in key aspects of sandeel life history, which we consider are most likely due to direct and indirect temperature-related changes over a range of biotic factors, including the seasonal distribution of copepods and intra- and inter-specific competition with planktivorous fish. The results have implications both for the many predators of sandeels and for age and size of maturation in this aggregation of North Sea sandeels.

Patterns of jellyfish abundance in the North Atlantic

A number of explanations have been advanced to account for the increased frequency and intensity at which jellyfish (pelagic cnidarians and ctenophores) blooms are being observed, most of which have been locally directed. Here, we investigate seasonal and inter-annual patterns in abundance and distribution of jellyfish in the North Atlantic Ocean to determine if there have been any system-wide changes over the period 1946–2005, by analysing records of the presence of coelenterates from the Continuous Plankton Recorder (CPR) survey. Peaks in jellyfish abundance are strongly seasonal in both oceanic and shelf areas: oceanic populations have a mid-year peak that is more closely related to peaks in phyto- and zooplankton, whilst the later peak of shelf populations mirrors changes in SST and reflects processes of advection and aggregation. There have been large amplitude cycles in the abundance of oceanic and shelf jellyfish (although not synchronous) over the last 60 years, with a pronounced synchronous increase in abundance in both areas over the last 10 years. Inter-annual variations in jellyfish abundance in oceanic areas are related to zooplankton abundance and temperature changes, but not to the North Atlantic Oscillation or to a chlorophyll index. The long-term inter-annual abundance of jellyfish on the shelf could not be explained by any environmental variables investigated. As multi-decadal cycles and more recent increase in jellyfish were obvious in both oceanic and shelf areas, we conclude that these are likely to reflect an underlying climatic signal (and bottom-up control) rather than any change in fishing pressure (top-down control). Our results also highlight the role of the CPR data in investigating long-term changes in jellyfish, and suggest that the cnidarians sampled by the CPR are more likely to be holoplanktic hydrozoans and not the much larger meroplanktic scyphozoans as has been suggested previously.

Synchronous response of marine plankton ecosystems to climate in the Northeast Atlantic and the North Sea

Over the last few decades, global warming has accelerated both the rate and magnitude of changes observed in many functional units of the Earth System. In this context, plankton are sentinel organisms because they are sensitive to subtle levels of changes in temperature and might help in identifying the current effects of climate change on pelagic ecosystems. In this paper, we performed a comparative approach in two regions of the North Atlantic (i.e. the Northeast Atlantic and the North Sea) to explore the relationships between changes in marine plankton, the regional physico-chemical environment and large-scale hydro-climatic forcing using four key indices: the North Atlantic Oscillation (NAO), the Atlantic Multidecadal Oscillation (AMO), the East Atlantic (EA) pattern and Northern Hemisphere Temperature (NHT) anomalies. Our analyses suggest that long-term changes in the states of the two ecosystems were synchronous and correlated to the same large-scale hydro-climatic variables: NHT anomalies, the AMO and to a lesser extent the EA pattern. No significant correlation was found between long-term ecosystem modifications and the state of the NAO. Our results suggest that the effect of climate on these ecosystems has mainly occurred in both regions through the modulation of the thermal regime.

Comparison of phosphate uptake rates by the smallest plastidic and aplastidic protists in the North Atlantic subtropical gyre

The smallest phototrophic protists (<3 μm) are important primary producers in oligotrophic subtropical gyres – the Earth's largest ecosystems. In order to elucidate how these protists meet their inorganic nutrient requirements, we compared the phosphate uptake rates of plastidic and aplastidic protists in the phosphate-depleted subtropical and tropical North Atlantic (4–29°N) using a combination of radiotracers and flow cytometric sorting on two Atlantic Meridional Transect cruises. Plastidic protists were divided into two groups according to their size (<2 and 2–3 μm). Both groups of plastidic protists showed higher phosphate uptake rates per cell than the aplastidic protists. Although the phosphate uptake rates of protist cells were on average seven times (P<0.001) higher than those of bacterioplankton, the biomass-specific phosphate uptake rates of protists were one fourth to one twentieth of an average bacterioplankton cell. The unsustainably low biomass-specific phosphate uptake by both plastidic and aplastidic protists suggests the existence of a common alternative means of phosphorus acquisition – predation on phosphorus-rich bacterioplankton cells.

Large bio-geographical shifts in the north-eastern Atlantic Ocean: From the subpolar gyre, via plankton, to blue whiting and pilot whales

Pronounced changes in fauna, extending from the English Channel in the south to the Barents Sea in the north-east and off Greenland in the north-west, have occurred in the late 1920s, the late 1960s and again in the late 1990s. We attribute these events to exchanges of subarctic and subtropical water masses in the north-eastern North Atlantic Ocean, associated with changes in the strength and extent of the subpolar gyre. These exchanges lead to variations in the influence exerted by the subarctic or Lusitanian biomes on the intermediate faunistic zone in the north-eastern Atlantic. This strong and persistent bottom-up bio-physical link is demonstrated using a numerical ocean general circulation model and data on four trophically connected levels in the food chain – phytoplankton, zooplankton, blue whiting, and pilot whales. The plankton data give a unique basin-scale depiction of these changes, and a long pilot whale record from the Faroe Islands offers an exceptional temporal perspective over three centuries. Recent advances in simulating the dynamics of the subpolar gyre suggests a potential for predicting the distribution of the main faunistic zones in the north-eastern Atlantic a few years into the future, which might facilitate a more rational management of the commercially important fisheries in this region.

The North Atlantic subpolar gyre regulates the spawning distribution of blue whiting (Micromesistius poutassou)

The spawning stock of blue whiting (Micromesistius poutassou), an economically important pelagic gadoid in the North Atlantic Ocean, increased threefold after 1995. The reproductive success of the stock is largely determined during the very early stages of life, but little is known about the spawning dynamics of this species. Here we show that the spawning distribution of blue whiting is variable, regulated by the hydrography west of the British Isles. When the North Atlantic subpolar gyre is strong and spreads its cold, fresh water masses east over Rockall Plateau, the spawning is constrained along the European continental slope and in a southerly position near Porcupine Bank. When the gyre is weak and conditions are relatively saline and warm, the spawning distribution moves northwards along the slope and especially westwards covering Rockall Plateau. The apparent link between the spawning distribution and the subpolar gyre is the first step towards understanding the reproduction variability, which currently is the main challenge for appropriate management of the blue whiting stock.

Macrophysiology of Calanus finmarchicus in the North Atlantic Ocean

Copepods represent the major part of the dry weight of the mesozooplankton in pelagic ecosystems and therefore have a central role in the secondary production of the North Atlantic Ocean. The calanoid copepod species Calanus finmarchicus is the main large copepod in subarctic waters of the North Atlantic, dominating the dry weight of the mesozooplankton in regions such as the northern North Sea and the Norwegian Sea. The objective of this work was to investigate the relationships between both the fundamental and realised niches of C. finmarchicus in order to better understand the future influence of global climate change on the abundance, the spatial distribution and the phenology of this key-structural species. Based on standardised Principal Component Analyses (PCAs), a macroecological approach was applied to determine factors affecting the spatial distribution of C. finmarchicus and to characterise its realised niche. Second, an ecophysiological model was used to calculate the Potential Egg Production Rate (PEPR) of C. finmarchicus and the centre of its fundamental niche. Relationships between the two niches were then investigated by correlation analysis. We found a close relationship between the fundamental and realised niches of C. finmarchicus at spatial, monthly and decadal scales. While the species is at the centre of its niche in the subarctic gyre, our joint macroecological and macrophysiological analyses show that it is at the edge of its niche in the North Sea, making the species in this region more vulnerable to temperature changes.

The meso-scale response of subarctic North Pacific seabird community structure to lower trophic level abundance and diversity

Understanding the mechanisms that structure communities and influence biodiversity are fundamental goals of ecology. To test the hypothesis that the abundance and diversity of upper-trophic level predators (seabirds) is related to the underlying abundance and diversity of their prey (zooplankton) and ecosystem-wide energy availability (primary production), we initiated a monitoring program in 2002 that jointly and repeatedly surveys seabird and zooplankton populations across a 7,500 km British Columbia-Bering Sea-Japan transect. Seabird distributions were recorded by a single observer (MH) using a strip-width technique, mesozooplankton samples were collected with a Continuous Plankton Recorder, and primary production levels were derived using the appropriate satellite parameters and the Vertically Generalized Production Model (Behrenfeld and Falkowski 1997). Each trophic level showed clear spatio-temporal patterns over the course of the study. The strongest relationship between seabird abundance and diversity and the lower trophic levels was observed in March/April ('spring') and significant relationships were also found through June/July ('summer'). No discernable relationships were observed during the September/October ('fall') months. Overall, mesozooplankton abundance and biomass explained the dominant portion of seabird abundance and diversity indices (richness, Simpson's Index, and evenness), while primary production was only related to seabird richness. These findings underscore the notion that perturbations of ocean productivity and lower trophic level ecosystem constituents influenced by climate change, such as shifts in timing (phenology) and synchronicity (match-mismatch), could impart far-reaching consequences throughout the marine food web.

Long-term changes in abundance and distribution of microzooplankton in the NE Atlantic and North Sea

Long-term changes in mesozooplankton and phytoplankton populations have been well documented in the North Atlantic region, whereas data for microzooplankton are scarce. This neglected component of the plankton is a vital link in marine food-webs, grazing on smaller flagellates and cyanobacteria and in turn providing food for the larger mesozooplankton. We use the latest tintinnid (Ciliophora, Protista) data from the Continuous Plankton Recorder (CPR) survey in the NE Atlantic and North Sea to examine the phenology, distribution and abundance of this important group of ciliates. Presence/absence data came from 167 122 CPR samples collected between 1960 and 2009 and abundance data from 49 662 samples collected between 1996 and 2009. In the North Atlantic the genus Dictyocysta spp. dominated and Parafavella gigantea showed an increase in abundance around Iceland and Greenland. In the North Sea higher densities of Tintinnopsis spp., Favella serrata and Ptychocylis spp. were found. The presence of tintinnids in CPR samples collected in the North Atlantic has increased over the last 50 years and the seasonal window of high abundance has lengthened. Conversely in the North Sea there has been an overall reduction in abundance. We discuss possible drivers for these long-term changes and point the way forward to more holistic studies that examine how ecosystems, rather than just selected taxa, are responding to climate change.

Temperature affects the timing of spawning and migration of North Sea mackerel

Climate change accentuates the need for knowing how temperature impacts the life history and productivity of economically and ecologically important species of fish. We examine the influence of temperature on the timing of the spawning and migrations of North Sea Mackerel using data from larvae CPR surveys, egg surveys and commercial landings from Danish coastal fisheries in the North Sea, Skagerrak, Kattegat and inner Danish waters. The three independent sources of data all show that there is a significant relationship between the timing of spawning and sea surface temperature. Large mackerel are shown to arrive at the feeding areas before and leave later than small mackerel and the sequential appearance of mackerel in each of the feeding areas studied supports the anecdotal evidence for an eastward post-spawning migration. Occasional commercial catches taken in winter in the Sound N, Kattegat and Skagerrak together with catches in the first quarter IBTS survey furthermore indicate some overwintering here. Significant relationships between temperature and North Sea mackerel spawning and migration have not been documented before. The results have implications for mackerel resource management and monitoring. An increase in temperature is likely to affect the timing and magnitude of the growth, recruitment and migration of North Sea mackerel with subsequent impacts on its sustainable exploitation.

Long-Term Retrospective Analysis of Mackerel Spawning in the North Sea: A New Time Series and Modeling Approach to CPR Data

We present a unique view of mackerel (Scomber scombrus) in the North Sea based on a new time series of larvae caught by the Continuous Plankton Recorder (CPR) survey from 1948-2005, covering the period both before and after the collapse of the North Sea stock. Hydrographic backtrack modelling suggested that the effect of advection is very limited between spawning and larvae capture in the CPR survey. Using a statistical technique not previously applied to CPR data, we then generated a larval index that accounts for both catchability as well as spatial and temporal autocorrelation. The resulting time series documents the significant decrease of spawning from before 1970 to recent depleted levels. Spatial distributions of the larvae, and thus the spawning area, showed a shift from early to recent decades, suggesting that the central North Sea is no longer as important as the areas further west and south. These results provide a consistent and unique perspective on the dynamics of mackerel in this region and can potentially resolve many of the unresolved questions about this stock

Update on the relationship between the North Atlantic Oscillation and Calanus finmarchicus

The relationship between climate, represented by the North Atlantic Oscillation (NAO), and the calanoid copepod Calanus finmarchicus has been extensively studied. The correlation between NAO and C. finmarchicus has broken down (post-1995). In the present study, we revisit the relationship between C. finmarchicus and the NAO. Our reanalysis shows that previous treatment of this data did not take into account 2 aspects of both the C. finmarchicus and NAO index time-series: (1) the presence of significant trends and (2) significant autocorrelation. Our analysis suggests that previously reported relationships between NAO and C. finmarchicus abundance can be explained largely by the trends in both data series. Removing the trend from both time-series resulted in a decrease in the amount of C. finmarchicus abundance variability explained by the NAO. Trend removal eliminated the autocorrelation from the NAO time-series, but not from the C. finmarchicus time-series. Partial autocorrelation analysis showed that the autocorrelation present in the C. finmarchicus time-series is only found at a lag of 1 yr, suggesting strong, year-to-year connectivity in this population. We included the lagged C. finmarchicus abundance into a regression with the NAO and found that C. finmarchicus variability is explained by the previous year’s abundance and, to a much smaller extent, by NAO variability. Limiting the time-series to the most recent 22 yr period (1981 to 2002) showed that the NAO is no longer correlated to C. finmarchicus abundance, and the autocorrelation in the C. finmarchicus abundance series also appears to be weakening.

Climate-induced effects on the meroplankton and the benthic-pelagic ecology of the North Sea

Analyses of long-term time series of North Sea plankton and sea surface temperature (SST) data reveal that the annual planktonic larval abundance of three benthic phyla, Echinodermata, Arthropoda, and Mollusca, responds positively and immediately to SST. Long-term outcomes for the planktonic abundance of these three phyla are different, however. The planktonic larvae of echinoderms and decapod crustaceans have increased in abundance from 1958 to 2005, and especially since the mid-1980s, as North Sea SST has increased. In contrast, the abundance of bivalve mollusc larvae has declined, despite the positive year-to-year relationship between temperature and bivalve larval abundance continuing to hold. We argue that the changes in meroplankton abundance, coincident with increased phytoplankton and declining holoplankton, reflect the synchronous effect of rising SST and related changes in the pelagic community on the reproduction and recruitment of many benthic marine invertebrates. Under this scenario, the long-term decline in bivalve mollusc larvae will reflect increased predation on the settled larvae and adults by benthic decapods. These alterations in the zooplankton may therefore describe an ecosystem-wide restructuring of North Sea trophic interactions.

Spatial Heterogeneity and Genetic Variation in the Copepod Neocalanus Cristatus Along Two Transects in the North Pacific Sampled By the Continuous Plankton Recorder

We present a macrogeographic study of spatial heterogeneity in an important subarctic Pacific copepod and describe the first genetic analysis of population structure using Continuous Plankton Recorder (CPR) samples. Samples of Neocalanus cristatus were collected at a constant depth of similar to 7 m from two CPR tow-routes, (i) an east-west similar to 6500-km transect from Vancouver Island, Canada to Hokkaido Island, Japan, and (ii) a north-south transect of similar to 2250 km from Anchorage, Alaska to Tacoma, Washington. Analysis of these samples revealed three features of the biology of N. cristatus. First, N. cristatus undergoes small-scale diel vertical migration that is larger among stages CV- adult (3-6 times more abundant at 7 m at night), than stages CI-CIV (only 2-4 times higher at night). Secondly, while there were no regions where N. cristatus did not appear, each transect sampled a few large-scale macrogeographic patches. Thirdly, an analysis of molecular variation, using a partial sequence of the N. cristatus cytochrome oxidase I gene, revealed that 7.3% (P < 0.0001) of the total genetic variation among N. cristatus sampled from macrogeographic patches by the CPR could be explained by spatial heterogeneity. We suggest that spatial heterogeneity at macrogeographic scales may be important in plankton evolution.

Decadal fluctuations in North Atlantic water inflow in the North Sea between 1958-2003: impacts on temperature and phytoplankton populations

The circulation of Atlantic water along the European continental slope, in particular the inflow into the North Sea, influences North Sea water characteristics with consequent changes in the environment affecting plankton community dynamics. The long-term effect of fluctuating oceanographic conditions oil the North Sea, pelagic ecosystem is assessed. It is shown that (i) there are similar regime shifts in the inflow through the northern North Sea and in Sea, Surface Temperature, (ii) long-term phytoplankton trends are influenced by the inflow only in some North Sea regions, and (iii) the spatial variability in chemicophysical and biological parameters highlight the influence of smaller scale processes.