Dynamics of air–sea CO2 fluxes in the northwestern European shelf based on voluntary observing ship and satellite observations

From January 2011 to December 2013, we constructed a comprehensive pCO2 data set based on voluntary observing ship (VOS) measurements in the western English Channel (WEC). We subsequently estimated surface pCO2 and air–sea CO2 fluxes in northwestern European continental shelf waters using multiple linear regressions (MLRs) from remotely sensed sea surface temperature (SST), chlorophyll a concentration (Chl a), wind speed (WND), photosynthetically active radiation (PAR) and modeled mixed layer depth (MLD). We developed specific MLRs for the seasonally stratified northern WEC (nWEC) and the permanently well-mixed southern WEC (sWEC) and calculated surface pCO2 with uncertainties of 17 and 16 μatm, respectively. We extrapolated the relationships obtained for the WEC based on the 2011–2013 data set (1) temporally over a decade and (2) spatially in the adjacent Celtic and Irish seas (CS and IS), two regions which exhibit hydrographical and biogeochemical characteristics similar to those of WEC waters. We validated these extrapolations with pCO2 data from the SOCAT and LDEO databases and obtained good agreement between modeled and observed data. On an annual scale, seasonally stratified systems acted as a sink of CO2 from the atmosphere of −0.6 ± 0.3, −0.9 ± 0.3 and −0.5 ± 0.3 mol C m−2 yr−1 in the northern Celtic Sea, southern Celtic sea and nWEC, respectively, whereas permanently well-mixed systems acted as source of CO2 to the atmosphere of 0.2 ± 0.2 and 0.3 ± 0.2 mol C m−2 yr−1 in the sWEC and IS, respectively. Air–sea CO2 fluxes showed important inter-annual variability resulting in significant differences in the intensity and/or direction of annual fluxes. We scaled the mean annual fluxes over these provinces for the last decade and obtained the first annual average uptake of −1.11 ± 0.32 Tg C yr−1 for this part of the northwestern European continental shelf. Our study showed that combining VOS data with satellite observations can be a powerful tool to estimate and extrapolate air–sea CO2 fluxes in sparsely sampled area.

How does Calanus helgolandicus maintain its population in a variable environment? Analysis of a 25-year time series from the English Channel

Calanus helgolandicus is a key copepod of the NE Atlantic and fringing shelves, with a distribution that is expanding northwards with oceanic warming. The Plymouth L4 site has warmed over the past 25-years, and experiences large variations in the timing and availability of food for C. helgolandicus. Here we examine the degree to which these changes translate into variation in reproductive output and subsequently C. helgolandicus population size. Egg production rates (eggs female−1 day−1) were maximal in the spring to early-summer period of diatom blooms and high ciliate abundance, rather than during the equally large autumn blooms of autotrophic dinoflagellates. Egg hatch success was lower in spring however, with a greater proportion of naupliar deformities then also. Both the timing and the mean summer abundance of C. helgolandicus (CI–CVI) reflected those of spring total reproductive output. However this relationship was driven by inter-annual variability in female abundance and not that of egg production per female, which ranged only two-fold. Winter abundance of C. helgolandicus at L4 was much more variable than abundance in other seasons, and reflected conditions from the previous growing season. However, these low winter abundances had no clear carry-over signal to the following season’s population size. Overall, the C. helgolandicus population appears to be surprisingly resilient at this dynamic, inshore site, showing no long-term phenology shift and only a four-fold variation in mean abundance between years. This dampening effect may reflect a series of mortality sources, associated with the timing of stratification in the early part of the season, likely affecting egg sinking and loss, plus intense, density-dependent mortality of early stages in mid-summer likely through predation.

Questioning the role of phenology shifts and trophic mismatching in a planktonic food web

In a warming climate, differential shifts in the seasonal timing of predators and prey have been suggested to lead to trophic ‘‘mismatches’’ that decouple primary, secondary and tertiary production. We tested this hypothesis using a 25-year time-series of weekly sampling at the Plymouth L4 site, comparing 57 plankton taxa spanning 4 trophic levels. During warm years, there was a weak tendency for earlier timings of spring taxa and later timings of autumn taxa. While this is in line with many previous findings, numerous exceptions existed and only a few taxa (e.g. Gyrodinium spp., Pseudocalanus elongatus, and Acartia clausi) showed consistent, strong evidence for temperature-related timing shifts, revealed by all 4 of the timing indices that we used. Also, the calculated offsets in timing i.e. ‘‘mismatches’’) between predator and prey were no greater in extreme warm or cold years than during more average years. Further, the magnitude of these offsets had no effect on the ‘‘success’’ of the predator, in terms of their annual mean abundance or egg production rates. Instead numerous other factors override, including: inter-annual variability in food quantity, high food baseline levels, turnover rates and prolonged seasonal availability, allowing extended periods of production. Furthermore many taxa, notably meroplankton, increased well before the spring bloom. While theoretically a chronic mismatch, this likely reflects trade-offs for example in predation avoidance. Various gelatinous taxa (Phaeocystis, Noctiluca, ctenophores, appendicularians, medusae) may have reduced these predation constraints, with variable, explosive population outbursts likely responding to improved conditions. The match–mismatch hypothesis may apply for highly seasonal, pulsed systems or specialist feeders, but we suggest that the concept is being over-extended to other marine systems where multiple factors compensate.

Life-cycle modification in open oceans accounts for genome variability in a cosmopolitan phytoplankton

Emiliania huxleyi is the most abundant calcifying plankton in modern oceans with substantial intraspecific genome variability and a biphasic life cycle involving sexual alternation between calcified 2N and flagellated 1N cells. We show that high genome content variability in Emiliania relates to erosion of 1N-specific genes and loss of the ability to form flagellated cells. Analysis of 185 E. huxleyi strains isolated from world oceans suggests that loss of flagella occurred independently in lineages inhabiting oligotrophic open oceans over short evolutionary timescales. This environmentally linked physiogenomic change suggests life cycling is not advantageous in very large/diluted populations experiencing low biotic pressure and low ecological variability. Gene loss did not appear to reflect pressure for genome streamlining in oligotrophic oceans as previously observed in picoplankton. Life-cycle modifications might be common in plankton and cause major functional variability to be hidden from traditional taxonomic or molecular markers.

Life-cycle modification in open oceans accounts for genome variability in a cosmopolitan phytoplankton

Emiliania huxleyi is the most abundant calcifying plankton in modern oceans with substantial intraspecific genome variability and a biphasic life cycle involving sexual alternation between calcified 2N and flagellated 1N cells. We show that high genome content variability in Emiliania relates to erosion of 1N-specific genes and loss of the ability to form flagellated cells. Analysis of 185 E. huxleyi strains isolated from world oceans suggests that loss of flagella occurred independently in lineages inhabiting oligotrophic open oceans over short evolutionary timescales. This environmentally linked physiogenomic change suggests life cycling is not advantageous in very large/diluted populations experiencing low biotic pressure and low ecological variability. Gene loss did not appear to reflect pressure for genome streamlining in oligotrophic oceans as previously observed in picoplankton. Life-cycle modifications might be common in plankton and cause major functional variability to be hidden from traditional taxonomic or molecular markers.

DISCOVERY 2010: Spatial and temporal variability in a dynamic polar ecosystem

The Scotia Sea has been a focus of biological- and physical oceanographic study since the Discovery expeditions in the early 1900s. It is a physically energetic region with some of the highest levels of productivity in the Southern Ocean. It is also a region within which there have been greater than average levels of change in upper water column temperature. We describe the results of three cruises transecting the central Scotia Sea from south to north in consecutive years and covering spring, summer and autumn periods. We also report on some community level syntheses using both current-day and historical data from this region. A wide range of parameters were measured during the field campaigns, covering the physical oceanography of the region, air–sea CO2 fluxes, macro- and micronutrient concentrations, the composition and biomass of the nano-, micro- and mesoplankton communities, and the distribution and biomass of Antarctic krill and mesopelagic fish. Process studies examined the effect of iron-stress on the physiology of primary producers, reproduction and egestion in Antarctic krill and the transfer of stable isotopes between trophic layers, from primary consumers up to birds and seals. Community level syntheses included an examination of the biomass-spectra, food-web modelling, spatial analysis of multiple trophic layers and historical species distributions. The spatial analyses in particular identified two distinct community types: a northern warmer water community and a southern cold community, their boundary being broadly consistent with the position of the Southern Antarctic Circumpolar Current Front (SACCF). Temperature and ice cover appeared to be the dominant, over-riding factors in driving this pattern. Extensive phytoplankton blooms were a major feature of the surveys, and were persistent in areas such as South Georgia. In situ and bioassay measurements emphasised the important role of iron inputs as facilitators of these blooms. Based on seasonal DIC deficits, the South Georgia bloom was found to contain the strongest seasonal carbon uptake in the ice-free zone of the Southern Ocean. The surveys also encountered low-production, iron-limited regions, a situation more typical of the wider Southern Ocean. The response of primary and secondary consumers to spatial and temporal heterogeneity in production was complex. Many of the life-cycles of small pelagic organisms showed a close coupling to the seasonal cycle of food availability. For instance, Antarctic krill showed a dependence on early, non-ice-associated blooms to facilitate early reproduction. Strategies to buffer against environmental variability were also examined, such as the prevalence of multiyear life-cycles and variability in energy storage levels. Such traits were seen to influence the way in which Scotia Sea communities were structured, with biomass levels in the larger size classes being higher than in other ocean regions. Seasonal development also altered trophic function, with the trophic level of higher predators increasing through the course of the year as additional predator-prey interactions emerged in the lower trophic levels. Finally, our studies re-emphasised the role that the simple phytoplankton-krill-higher predator food chain plays in this Southern Ocean region, particularly south of the SACCF. To the north, alternative food chains, such as those involving copepods, macrozooplankton and mesopelagic fish, were increasingly important. Continued ocean warming in this region is likely to increase the prevalence of such alternative such food chains with Antarctic krill predicted to move southwards.

Coccolithophore surface distributions in the North Atlantic and their modulation of the air-sea flux of CO2 from 10 years of satellite Earth observation data

Coccolithophores are the primary oceanic phytoplankton responsible for the production of calcium carbonate (CaCO3). These climatically important plankton play a key role in the oceanic carbon cycle as a major contributor of carbon to the open ocean carbonate pump (similar to 50 %) and their calcification can affect the atmosphere-to-ocean (air-sea) uptake of carbon dioxide (CO2) through increasing the seawater partial pressure of CO2 (pCO(2)). Here we document variations in the areal extent of surface blooms of the globally important coccolithophore, Emiliania huxleyi, in the North Atlantic over a 10-year period (1998-2007), using Earth observation data from the Sea-viewing Wide Field-of-view Sensor (SeaWiFS). We calculate the annual mean sea surface areal coverage of E. huxleyi in the North Atlantic to be 474 000 +/- 104 000 km(2), which results in a net CaCO3 carbon (CaCO3-C) production of 0.14-1.71 Tg CaCO3-C per year. However, this surface coverage (and, thus, net production) can fluctuate inter-annually by -54/+81% about the mean value and is strongly correlated with the El Nino/Southern Oscillation (ENSO) climate oscillation index (r = 0.75, p < 0.02). Our analysis evaluates the spatial extent over which the E. huxleyi blooms in the North Atlantic can increase the pCO(2) and, thus, decrease the localised air-sea flux of atmospheric CO2. In regions where the blooms are prevalent, the average reduction in the monthly air-sea CO2 flux can reach 55%. The maximum reduction of the monthly air-sea CO2 flux in the time series is 155 %. This work suggests that the high variability, frequency and distribution of these calcifying plankton and their impact on pCO(2) should be considered if we are to fully understand the variability of the North Atlantic air-to-sea flux of CO2. We estimate that these blooms can reduce the annual N. Atlantic net sink atmospheric CO2 by between 3-28 %.

Seasonality and interannual variability of the European Slope Current from 20 years of altimeter data compared with in situ measurements

The European Slope Current (SC) is a major section of the warm poleward flow from the Atlantic to the Arctic, which also moderates the exchange of heat, salt, nutrients and carbon between the deep ocean and the European shelf seas. The mean structure of the geostrophic flow, seasonality, interannual variability and long-term trend of SC are appraised with an unprecedented continuous 20-year satellite altimeter dataset. Comparisons with long term in situ data showed a maximum correlation of r2=0.51 between altimeter and Acoustic Doppler Current Profilers (ADCP), with similar results for drogued buoy data. Mean geostrophic currents were appraised more comprehensively than previous attempts, and the paths of 4 branches of the North Atlantic Current (NAC) and positions of 5 eddies in the region were derived quantitatively. A consistent seasonal cycle in the flow of the SC was found at all 8 sections along the European shelf slope, with maximum poleward flow in the winter and minimum in the summer. The seasonal difference in the altimetry current speed amounted to ~8-10 cm s-1 at the northern sections, but only ~5 cm s-1 on the Bay of Biscay slopes. This extended altimeter dataset indicates significant regional and seasonal variations, and has revealed new insights into the interannual variability of the SC. It is shown that there is a peak poleward flow at most positions along a ~2000 km stretch of the continental slope from Portugal to Scotland during 1995-1997, but this did not clearly relate to the extreme negative North Atlantic Oscillation (NAO) in the winter of 1995-1996. The speed of the SC exhibited a long term decreasing trend of ~1% per year. By contrast the NAC showed no significant trend over the 20-year period. Major changes in the NAC occurred three times, and these changes followed decreases in the NAO index.

Long-term variability of the siphonophores Muggiaea atlantica and M. kochi in the Western English Channel

We investigated long-term variability of the calycophoran siphonophores Muggiaea atlantica and Muggiaea kochi in the Western English Channel (WEC) between 1930 and 2011. Our aims were to describe long-term changes in abundance and temporal distribution in relation to local environmental dynamics. In order to better understand mechanisms that regulate the species’ populations, we identified periods that were characteristic of in situ population growth and the environmental optima associated with these events. Our results show that between 1930 and the 1960s both M. atlantica and M. kochi were transient components of the WEC ecosystem. In the late 1960s M. atlantica, successfully established a resident population in the WEC, while the occurrence of M. kochi became increasingly sporadic. Once established as a resident species, the seasonal abundance and distribution of M. atlantica increased. Analysis of environmental conditions associated with in situ population growth revealed that temperature and prey were key determinants of the seasonal distribution and abundance of M. atlantica. Salinity was shown to have an indirect effect, likely representing a proxy for water circulation in the WEC. Anomalies in the seasonal cycle of salinity, indicating deviation from the usual circulation pattern in the WEC, were negatively associated with in situ growth, suggesting dispersal of the locally developing M. atlantica population. However, our findings identified complexity in the relationship between characteristics of the environment and M. atlantica variability. The transition from a period of transiency (1930–1968) to residency (1969–2011) was tentatively attributed to structural changes in the WEC ecosystem that occurred under the forcing of wider-scale hydroclimatic changes.

Spatio-temporal characteristics of Agulhas leakage: a model inter-comparison study

Investigating the variability of Agulhas leakage, the volume transport of water from the Indian Ocean to the South Atlantic Ocean, is highly relevant due to its potential contribution to the Atlantic Meridional Overturning Circulation as well as the global circulation of heat and salt and hence global climate. Quantifying Agulhas leakage is challenging due to the non-linear nature of this process; current observations are insufficient to estimate its variability and ocean models all have biases in this region, even at high resolution . An Eulerian threshold integration method is developed to examine the mechanisms of Agulhas leakage variability in six ocean model simulations of varying resolution. This intercomparison, based on the circulation and thermo- haline structure at the Good Hope line, a transect to the south west of the southern tip of Africa, is used to identify features that are robust regardless of the model used and takes into account the thermohaline biases of each model. When determined by a passive tracer method, 60 % of the magnitude of Agulhas leakage is captured and more than 80 % of its temporal fluctuations, suggesting that the method is appropriate for investigating the variability of Agulhas leakage. In all simulations but one, the major driver of variability is associated with mesoscale features passing through the section. High resolution (<1/10 deg.) hindcast models agree on the temporal (2–4 cycles per year) and spatial (300–500 km) scales of these features corresponding to observed Agulhas Rings. Coarser resolution models (<1/4 deg.) reproduce similar time scale of variability of Agulhas leakage in spite of their difficulties in representing the Agulhas rings properties. A coarser resolution climate model (2 deg.) does not resolve the spatio-temporal mechanism of variability of Agulhas leakage. Hence it is expected to underestimate the contribution of Agulhas Current System to climate variability.

Spatio-temporal characteristics of Agulhas leakage: a model inter-comparison study

Investigating the variability of Agulhas leakage, the volume transport of water from the Indian Ocean to the South Atlantic Ocean, is highly relevant due to its potential contribution to the Atlantic Meridional Overturning Circulation as well as the global circulation of heat and salt and hence global climate. Quantifying Agulhas leakage is challenging due to the non-linear nature of this process; current observations are insufficient to estimate its variability and ocean models all have biases in this region, even at high resolution . An Eulerian threshold integration method is developed to examine the mechanisms of Agulhas leakage variability in six ocean model simulations of varying resolution. This intercomparison, based on the circulation and thermo- haline structure at the Good Hope line, a transect to the south west of the southern tip of Africa, is used to identify features that are robust regardless of the model used and takes into account the thermohaline biases of each model. When determined by a passive tracer method, 60 % of the magnitude of Agulhas leakage is captured and more than 80 % of its temporal fluctuations, suggesting that the method is appropriate for investigating the variability of Agulhas leakage. In all simulations but one, the major driver of variability is associated with mesoscale features passing through the section. High resolution (<1/10 deg.) hindcast models agree on the temporal (2–4 cycles per year) and spatial (300–500 km) scales of these features corresponding to observed Agulhas Rings. Coarser resolution models (<1/4 deg.) reproduce similar time scale of variability of Agulhas leakage in spite of their difficulties in representing the Agulhas rings properties. A coarser resolution climate model (2 deg.) does not resolve the spatio-temporal mechanism of variability of Agulhas leakage. Hence it is expected to underestimate the contribution of Agulhas Current System to climate variability.

Is the Russell Cycle a true cycle? Multidecadal zooplankton and climate trends in the western English Channel

The Russell Cycle is one of the classical examples of climate influence on biological oceanography, represented as shifts in the marine plankton over several decades with warm and cool conditions. While the time-series data associated with the phenomenon indicate cyclical patterns, the question remains whether or not the Russell Cycle should be considered a “true cycle”. Zooplankton time-series data from 1924 to 2011 from the western English Channel were analysed with principal component (PC), correlation and spectral analyses to determine the dominant trends, and cyclic frequencies of the Russell Cycle indicators in relation to long-term hydroclimatic indices. PC1 accounted for 37.4% of the variability in the zooplankton data with the main contributions from non-clupeid fish larvae, southwestern zooplankton, and overall zooplankton biovolume. For PC2 (14.6% of data variance), the dominant groups were northern fish larvae, non-sardine eggs, and southern fish larvae. Sardine eggs were the major contributors to PC3 (representing 12.1% of data variance). No significant correlations were observed between the above three components and climate indices: Atlantic Multidecadal Oscillation, North Atlantic Oscillation, and local seawater temperature. Significant 44- and 29-year frequencies were observed for PC3, but the physical mechanisms driving the cycles are unclear. Harmonic analysis did not reveal any significant frequencies in the physical variables or in PCs 1 and 2. To a large extent, this is due to the dominant cycles in all datasets generally being long term (>50 years or so) and not readily resolved in the examined time frame of 88 years, hence restricting the ability to draw firm conclusions on the multidecadal relationship between zooplankton community dynamics in the western English Channel and environmental indices. Thus, the zooplankton time-series often associated and represented as the Russell Cycle cannot be concluded as being truly cyclical.

Is the Russell Cycle a true cycle? Multidecadal zooplankton and climate trends in the western English Channel

The Russell Cycle is one of the classical examples of climate influence on biological oceanography, represented as shifts in the marine plankton over several decades with warm and cool conditions. While the time-series data associated with the phenomenon indicate cyclical patterns, the question remains whether or not the Russell Cycle should be considered a “true cycle”. Zooplankton time-series data from 1924 to 2011 from the western English Channel were analysed with principal component (PC), correlation and spectral analyses to determine the dominant trends, and cyclic frequencies of the Russell Cycle indicators in relation to long-term hydroclimatic indices. PC1 accounted for 37.4% of the variability in the zooplankton data with the main contributions from non-clupeid fish larvae, southwestern zooplankton, and overall zooplankton biovolume. For PC2 (14.6% of data variance), the dominant groups were northern fish larvae, non-sardine eggs, and southern fish larvae. Sardine eggs were the major contributors to PC3 (representing 12.1% of data variance). No significant correlations were observed between the above three components and climate indices: Atlantic Multidecadal Oscillation, North Atlantic Oscillation, and local seawater temperature. Significant 44- and 29-year frequencies were observed for PC3, but the physical mechanisms driving the cycles are unclear. Harmonic analysis did not reveal any significant frequencies in the physical variables or in PCs 1 and 2. To a large extent, this is due to the dominant cycles in all datasets generally being long term (>50 years or so) and not readily resolved in the examined time frame of 88 years, hence restricting the ability to draw firm conclusions on the multidecadal relationship between zooplankton community dynamics in the western English Channel and environmental indices. Thus, the zooplankton time-series often associated and represented as the Russell Cycle cannot be concluded as being truly cyclical.

Linking environmental variables with regional- scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and >1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide.

Linking environmental variables with regional- scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and >1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide.