Temperate Shelf Sea Research Articles

Rain triggers seasonal stratification in a temperate shelf sea

The North Atlantic Storm Track acts as a conveyor belt for extratropical cyclones that frequently deliver high winds and rainfall to northwest European shelf seas. Storms are primarily considered detrimental to shelf sea stratification due to wind-driven mixing countering thermal buoyancy, but their impact on shelf scale stratification cycles remains poorly understood. Here, we show that storms trigger stratification through enhanced surface buoyancy from rainfall. A multidecadal model confirms that rainfall contributed to triggering seasonal stratification 88% of the time from 1982 to 2015. Stratification could be further modulated by large-scale climate oscillations, such as the Atlantic Multidecadal Variability (AMV), with stratification onset dates being twice as variable during a positive AMV phase than a negative one. Further insights into how changing storm activity will impact shelf seas are discussed beyond the current view of increasing wind-driven mixing, with significant implications for marine productivity and ecosystem function.

Different flavours of oxygen help quantify seasonal variations of the biological carbon pump in the Celtic Sea

Shelf seas represent only 10% of the World’s Ocean by area but support up to 30% of its primary production. There are few measurements of biological production at high spatial and temporal resolution in these physically and biologically dynamic systems. Here, we use dissolved oxygen to-argon (O2/Ar) ratios and oxygen triple isotopes in O2 (16O, 17O, 18O) to estimate net community production, N(O2/Ar), and gross O2 production, G(17O), in summer and autumn 2014 and spring and summer 2015 in the Celtic Sea, as part of the UK Shelf-Sea Biogeochemistry Programme. Surface O2/Ar concentration ratios were measured continuously using a shipboard membrane inlet mass spectrometer. Additional depth profiles of O2/Ar concentration ratios, δ(17O) and δ(18O) were measured in discrete water samples from hydrocasts. The data were combined with wind-speed based gas exchange parameterisations to calculate biological air-sea oxygen fluxes. These fluxes were corrected for diapycnal diffusion, entrainment, production below the mixed layer, and changes over time to derive N(O2/Ar) and G(17O). The Celtic Sea showed the highest G(17O) in summer 2014 (825 mmol m–2 d–1) and lowest during autumn 2014 (153 mmol m–2 d–1). N(O2/Ar) was highest in spring 2015 (43 mmol m–2 d–1), followed by summer 2014 (42 mmol m–2 d–1), with a minimum in autumn 2014 (–24 mmol m–2 d–1). Dividing the survey region into three hydrographically distinct areas (Celtic Deep, Central Celtic Sea and Shelf Edge), we found that Celtic Deep and Shelf Edge had higher N(O2/Ar) in summer (71 and 63 mmol m–2 d–1, respectively) than in spring (49 and 22 mmol m–2 d–1). This study shows regional differences in the metabolic balance within the same season, as well as higher net community production in summer than in spring in some areas and years. The seasonal patterns in biological production rates and the export efficiency (f-ratio) identified the importance of biology for supporting the Celtic Sea’s ability to act as a net CO2 sink. Our measurements thus help improve our understanding of the biological carbon pump in temperate shelf seas.

Ocean circulation and biological processes drive seasonal variations of dissolved Al, Cd, Ni, Cu, and Zn on the Northeast Atlantic continental margin

Nutrients and nutrient-like dissolved trace metals (dTMs) are essential for the functioning of marine organisms and therefore form an important part of ocean biogeochemical cycles. Here, we report on the seasonal distributions of dissolved zinc (dZn), nickel (dNi), copper (dCu), cadmium (dCd), aluminum (dAl), and nutrients on the Northeast Atlantic continental margin (Celtic Sea), which is representative for temperate shelf seas globally. Variations in surface water dTM and nutrient concentrations were mainly regulated by seasonal changes in biological processes. The stoichiometry of dTMs (especially for dCu and dZn) and nutrients on the continental shelf was additionally affected by fluvial inputs. Nutrients and dTMs at depth on the continental slope were determined by water mass mixing driven by ocean circulation, without an important role for local remineralization processes. The Mediterranean Outflow Waters are especially important for delivering Mediterranean-sourced dTMs to the Northeast Atlantic Ocean and drive dTM:nutrient kinks at a depth of ~1000 m. These results highlight the importance of riverine inputs, seasonality of primary production and ocean circulation on the distributions of nutrients and nutrient-like dTMs in temperate continental margin seas. Future climate related changes in the forcing factors may impact the availability of nutrients and dTMs to marine organisms in highly productive continental shelf regions and consequently the regional carbon cycle.

Benthic ecosystem functioning under climate change: modelling the bioturbation potential for benthic key species in the southern North Sea.

Climate change affects the marine environment on many levels with profound consequences for numerous biological, chemical, and physical processes. Benthic bioturbation is one of the most relevant and significant processes for benthic-pelagic coupling and biogeochemical fluxes in marine sediments, such as the uptake, transport, and remineralisation of organic carbon. However, only little is known about how climate change affects the distribution and intensity of benthic bioturbation of a shallow temperate shelf sea system such as the southern North Sea. In this study, we modelled and projected changes in bioturbation potential (BPp) under a continuous global warming scenario for seven southern North Sea key bioturbators: Abra alba, Amphiura filiformis, Callianassa subterranea, Echinocardium cordatum, Goniada maculata, Nephtys hombergii, and Nucula nitidosa. Spatial changes in species bioturbation intensity are simulated for the years 2050 and 2099 based on one species distribution model per species driven by bottom temperature and salinity changes using the IPCC SRES scenario A1B. Local mean bottom temperature was projected to increase between 0.15 and 5.4 °C, while mean bottom salinity was projected to moderately decrease by 1.7. Our results show that the considered benthic species are strongly influenced by the temperature increase. Although the total BP remained rather constant in the southern North Sea, the BPp for four out of seven species was projected to increase, mainly due to a simultaneous northward range expansion, while the BPp in the core area of the southern North Sea declined for the same species. Bioturbation of the most important species, Amphiura filiformis and Echinocardium cordatum, showed no substantial change in the spatial distribution, but over time. The BPp of E. cordatum remained almost constant until 2099, while the BPp of A. filiformis decreased by 41%. The northward expansion of some species and the decline of most species in the south led to a change of relative contribution to bioturbation in the southern North Sea. These results indicate that some of the selected key bioturbators in the southern North Sea might partly compensate the decrease in bioturbation by others. But especially in the depositional areas where bioturbation plays a specifically important role for ecosystem functioning, bioturbation potential declined until 2099, which might affect the biochemical cycling in sediments of some areas of the southern North Sea.

Seasonal nutrient co-limitation in a temperate shelf sea: A modelling approach

Nutrient limitation on phytoplankton growth plays a critical role in ocean productivity, the functioning of marine ecosystems, and the ocean carbon cycle. In the Celtic Sea, a temperate shelf sea, many studies have shown the importance of nitrate on phytoplankton growth focusing on the seasonal cycle of nitrate and feedbacks with the physical environment; but only recently has it been demonstrated, through discrete measurements, that dissolved iron also plays an important role in the ecosystem of the region. A well-established one-dimensional model has been developed to analyse the nutrient co-limitation between dissolved iron and dissolved inorganic nitrogen in the Celtic Sea. This model allows us to study the full seasonal cycle and inter-annual variability of these two nutrients. Simulations show that dissolved iron is an important nutrient for the development of the spring bloom, while nitrate plays a more important role during the summer season. Sensitivity analyses show that these results are robust when varying the nutrient-related parameters; the largest variability observed for primary production was observed when varying the nutrient sediment flux rates for dissolved iron and nitrate while less impact on phytoplankton production occurs when changing the half saturation constants. Here, we demonstrate that dissolved iron is an important nutrient for the development of the spring bloom and it should not be neglected as a state variable when modelling the Celtic Sea or other temperate shelf seas.

Climatic Controls on the Spring Phytoplankton Growing Season in a Temperate Shelf Sea

AbstractThe Northwest European Shelf is positioned directly beneath the North Atlantic Storm Track, within which the frequency and intensity of transient storms are modulated by large‐scale climatic oscillations. In temperate shelf seas, the impact of storms on the physical environment has received considerable attention, but the effect on biogeochemistry is less studied. Here, we use output from a multidecadal (1982–2015) coupled physical‐biogeochemical model supported by observations from ocean gliders to investigate phytoplankton growth throughout the winter‐spring transition. We define two separate phytoplankton growth events: the spring bloom, defined as the exponential growth following seasonal stratification, and the prebloom, occurring before stratification, and accounting for up to 22% of the total spring growth. Our results support the paradigm that light is a first‐order control, with the spring bloom initiating up to 22 days after stratification onset should light levels be too low to trigger the bloom. The prebloom is heavily influenced by the phase of the Atlantic Multidecadal Oscillation (AMO), demonstrated by an acceleration in the rate of increase of total chlorophyll concentrations (±90% confidence limit) from 7.6 ± 2.8 mg m−2 d−1 (during a positive AMO) to 13.1 ± 4.3 mg m−2 d−1 (negative AMO), due to modulation of periods of ephemeral stratification that occur between successive storms. We propose that phytoplankton growth in prebloom events might help buffer the lag between phytoplankton supply and larval recruitment, particularly during years when the spring bloom is delayed.

Variability in structure and carbon content of plankton communities in autumn in the waters south-west of the UK

The size, type and abundance of planktonic organisms influence the efficiency with which carbon is transferred through the lower trophic levels, ultimately affecting dynamics at the higher trophic levels of the marine food web. In temperate shelf sea, such as the waters south-west of the UK, the plankton growing season span from early spring to autumn. While the plankton spring bloom has been extensively studied, the end of the growing season in September-October has received less attention, despite its potential importance for autumn-spawning fish and their larval stage survival. In this study we investigated the variability of the structure and carbon content of the planktonic communities in the waters south-west of the UK in October 2013 and 2014, discussing potential implications of these changes to small pelagic fish and higher trophic levels. Microphytoplankton (20–200 µm) dominated the plankton community in terms of carbon in 2013, while nanophytoplankton (<20 µm) in 2014. Ciliates, Copepoda, Decapoda and Cnidaria represented the highest proportion of carbon in the zooplankton component in both years, although ciliates and Copepoda biomass was higher in 2014. Environmental conditions were linked to these changes and were significant in describing the carbon content of the plankton groups. In particular, silicate concentration appeared to be a key variable, affecting diatom/plankton dynamics at the end of the growing season. Other important environmental variables associated with the structure of plankton groups were salinity, sea surface temperature, chlorophyll-a, difference between sea surface temperature and bottom temperature, phosphate and nitrogen concentrations. Although the composition and carbon distribution of the plankton community were different in the two years, cluster and Random Forest analyses showed similarities in the clusters of stations identified, defining an area of higher plankton carbon along the south coast of Cornwall, and an area of lower carbon in the Bristol Channel in both years. Presence of suitable prey for planktivorous small pelagic fish (e.g. Paracalanus and Pseudocalanus), particularly in 2014, provided supporting evidence of the importance of this sea area as a foraging and nursery ground for sardines and other small pelagic fish, as well as for their predators.

Simultaneous assessment of oxygen- and nitrate-based net community production in a temperate shelf sea from a single ocean glider

Abstract. The continental shelf seas are important at a global scale for ecosystem services. These highly dynamic regions are under a wide range of stresses, and as such future management requires appropriate monitoring measures. A key metric to understanding and predicting future change are the rates of biological production. We present here the use of an autonomous underwater glider with an oxygen (O2) and a wet-chemical microfluidic total oxidised nitrogen (NOx-=NO3-+NO2-) sensor during a spring bloom as part of a 2019 pilot autonomous shelf sea monitoring study. We find exceptionally high rates of net community production using both O2 and NOx- water column inventory changes, corrected for air–sea gas exchange in case of O2. We compare these rates with 2007 and 2008 mooring observations finding similar rates of NOx- consumption. With these complementary methods we determine the O2:N amount ratio of the newly produced organic matter (7.8 ± 0.4) and the overall O2:N ratio for the total water column (5.7 ± 0.4). The former is close to the canonical Redfield O2:N ratio of 8.6 ± 1.0, whereas the latter may be explained by a combination of new organic matter production and preferential remineralisation of more reduced organic matter at a higher O2:N ratio below the euphotic zone.

Intermittent Intense Thermocline Shear Associated With Wind‐Forced Near‐Inertial Internal Waves in a Summer Stratified Temperate Shelf Sea

AbstractThermocline shear (i.e., velocity shear in the thermocline) lies at the heart of stratified shelf sea systems, regulating the vertical mixing and transport of mass and biogeochemical constituents. Based on year‐long moored ADCP measurements in the southern Yellow Sea (YS), the baroclinic kinetic energy and thermocline shear are investigated in this study. The rotary spectra analysis of the velocity measurements and numerical simulations with the slab model demonstrate that the wind‐forced near‐inertial internal waves (NIWs) contribute most of the baroclinic kinetic energy (50%) and thermocline shear (55%) in this stratified temperate shelf sea. The NIWs are mostly in the first two modes and are frequently generated during the warm season, associated with the presence of strong stratification and disturbances from extratropical cyclones. Most interestingly, the observed thermocline shear shows intermittent enhanced near‐inertial shear in the YS for the first time. Further analysis reveals that the thermocline shear is in the form of a clockwise rotating vector with the shear maxima (hereafter the shear spike) occurring at an angle of ∼90° to the right of the wind vector. These shear spikes, which prevail in warm seasons, are further demonstrated to be related to the wind‐shear‐alignment mechanism through comparing with a theoretical shear production model. Given the prevalence of shear spikes in warm seasons, this process is believed to have important implications in inducing thermocline mixing, which may further modulate nutrients cycling and help the maintenance of primary productivity in seasonally stratified shelf seas.

Low Contribution of the Fast‐Sinking Particle Fraction to Total Plankton Metabolism in a Temperate Shelf Sea

AbstractTemperate shelf seas are productive areas with the potential to export high quantities of particulate organic carbon (POC), as sinking particles, to the sediments or off‐shelf to the open ocean. The amount of carbon which can be exported depends partly on the amount of POC produced and on the remineralization processes occurring on the sinking material. Here, we assessed the relative seasonal importance of microbial respiration and bacterial production associated with suspended, slow‐ and fast‐sinking particle fractions. The three fractions were collected in the Celtic Sea above and below the seasonal thermocline in November 2014, April and July 2015 using Marine Snow Catchers. The slow‐sinking fraction had higher microbial respiration and bacterial production rates than the fast‐sinking fractions, and these two fractions sustained rates of microbial respiration and bacterial production between 1 and 3 orders of magnitude lower than the suspended fraction. This low contribution of the slow‐ and fast‐sinking fractions was consistent with their low contribution to the POC concentration at the two depths sampled. The POC‐specific respiration rates associated with the slow‐ and fast‐sinking fractions were low (median 0.17 and 0.08 d−1, respectively), indicating low‐sinking particle degradation. Our results indicate that ∼5% of the POC in surface waters can be exported below the thermocline.

Intermittent intense thermocline shear associated with wind-forced near-inertial internal waves in a summer stratified temperate shelf sea

Intermittent intense thermocline shear associated with wind-forced near-inertial internal waves in a summer stratified temperate shelf sea

Control of a phytoplankton bloom by wind‐driven vertical mixing and light availability

AbstractThe balance of physical and biological processes governing phytoplankton growth rates and the accumulation of biomass is widely debated in the literature, notably during the winter–spring transition. Here we show, in a temperate shelf sea that variability in the depth of the actively mixing surface layer is the leading order control. During a 2‐week period preceding the peak of the spring bloom we observe two distinct regimes; first, growth within the euphotic zone during the day and re‐distribution of new biomass to the seasonal pycnocline at night by convective mixing; then, more rapid biomass accumulation trapped within a shallower, wind‐driven actively mixing layer that was decoupled from the pycnocline below. Our observations of the bloom in the Celtic Sea, Northwest European Shelf, were made using ocean gliders and include measurements of the dissipation of turbulent kinetic energy. A 1‐D phytoplankton growth model driven by our measurements of dissipation and incident irradiance replicates the observed bloom and reinforces the conclusion that physical processes that mediate light availability were key. Day‐to‐day variability in cloud cover and the ability of phytoplankton to acclimate to their light environment were also important factors in determining growth rates, and the timing of the biomass peak. Our results emphasize the need for accurate turbulent mixing parameterizations in coupled hydrodynamic‐ecosystem models. Our findings are applicable to any region where wind‐driven mixing can modify nutrient and light availability, especially across subpolar shelves in the northern hemisphere where light rather than nutrients is typically the limiting factor on phytoplankton growth.

Interannual variability of gelatinous mesozooplankton in a temperate shelf sea: greater abundance coincides with cooler sea surface temperatures

AbstractAlthough gelatinous zooplankton are an important component of marine ecosystems, gelatinous mesozooplankton that are &amp;lt;2 cm are underrepresented in monitoring programmes. Here, the interannual variability of gelatinous mesozooplankton abundance and diversity was estimated from 167 zooplankton net samples that were collected in the Celtic Sea during seven fisheries surveys between 2007 and 2019 and analysed alongside environmental parameters. Compositional changes occurred interannually, including an overturn in the abundance ratio of two siphonophores (Muggiaea atlantica and Agalma elegans). Analysis of annual mean gelatinous abundance revealed no linear trend over time (Spearman, r = −0.09, p = 0.287); however, the interannual abundance varied by a factor of 33 (minimum mean abundance in 2013 = 7.36 ± 4.86 individuals m−3; maximum in 2017 = 244.82 ± 84.59 individuals m−3). Holoplanktonic taxa dominated the abundance of the gelatinous community (93.27%) and their abundance was negatively associated with summer sea surface temperature (represented by the 16°C isotherm in July), and the Eastern Atlantic Pattern index 3 months prior (April). Our data suggest that gelatinous mesozooplankton in the Celtic Sea may become less abundant with further ocean warming, and further highlight the need to monitor gelatinous mesozooplankton with a high taxonomic resolution moving forward.

Long-term changes in ecological functioning of temperate shelf sea benthic communities

The functioning of ecosystems is decisively dependent on the composition and distribution of the functional traits of the constituent species. We used trait analysis to represent aspects of marine benthic functioning, using a 20-year time-series (1992–2011) on macrozoobenthos collected annually at four monitoring sites in the southern North Sea. Temporal patterns of trait composition and taxonomic composition were compared to test whether they exhibited similar or contrasting responses to environmental change. Temporal changes in trait composition were more similar among monitoring sites than the changes in the taxonomic composition suggesting that trait compositions converge towards a common structure sculpted by common environmental drivers. The relationship between species richness and functional diversity displayed a power-shaped curve with a shallow slope, implying considerable functional redundancy among species. The temporal trends in functional diversity were relatively stable as compared to taxonomic diversity, with only two irregularities coinciding with exceptionally cold winter events in the North Sea in 1995 and 2009. Following the temporary changes in ecological functioning, the functional diversity returned to previous levels within one year. The rapid functional recovery of the benthic infauna confirms the self-organizing ability of the ecosystem in response to stressors and may be attributed to the high functional redundancy in the temperate shelf sea system of the North Sea.

Constraining the response of phytoplankton to zooplankton grazing and photo-acclimation in a temperate shelf sea with a 1-D model – towards S2P3 v8.0

Abstract. An established one-dimensional Shelf Sea Physics and Primary Production (S2P3) model has been developed into three different new models: S2P3-NPZ which includes a nutrient–phytoplankton–zooplankton (NPZ) framework, where the grazing rate is no longer fixed but instead varies over time depending on different functions chosen to represent the predator–prey relationship between zooplankton and phytoplankton; S2P3-Photoacclim which includes a representation of the process of photo-acclimation and flexible stoichiometry in phytoplankton; and S2P3 v8.0 which combines the NPZ framework and the variable stoichiometry of phytoplankton at the same time. These model formulations are compared to buoy and conductivity–temperature–depth (CTD) observations, as well as zooplankton biomass and in situ phytoplankton physiological parameters obtained in the central Celtic Sea (CCS). Models were calibrated by comparison to observations of the timing and magnitude of the spring phytoplankton bloom, magnitude of the spring zooplankton bloom, and phytoplankton physiological parameters obtained throughout the water column. A sensitivity study was also performed for each model to understand the effects of individual parameters on model dynamics. Results demonstrate that better agreement with biological observations can be obtained through the addition of representations of photo-acclimation, flexible stoichiometry, and grazing provided these can be adequately constrained.

Observations of vertical mixing in autumn and its effect on the autumn phytoplankton bloom

This work examines the seasonal cycle of vertical density structure and its influence on primary production in a temperate shelf sea, with a particular focus on the breakdown of stratification in autumn. We do this by combining new, high resolution observations of water column structure, meteorological forcing, nitrate and chlorophyll fluorescence collected between March 2014 and July 2015 on the North West European Shelf.Our results challenge the generally accepted assumption that convection dominates over wind driven mixing resulting in seasonal breakdown of stratification. Furthermore we found, that vertical mixing in autumn not only transformed the vertical density structure but also the vertical structure of chlorophyll biomass and surface nutrients. The subsurface chlorophyll maximum was eroded and a vertically homogeneous profile of chlorophyll biomass established itself above the pycnocline. This increased mixing also led to replenishment of surface nitrate concentrations, which supported an autumn phytoplankton bloom. While the significance of phytoplankton blooms in autumn has previously not been well quantified, we argue that these can act as a significant contributor to the seasonal drawdown of carbon.

Estimation of Size-Fractionated Primary Production from Satellite Ocean Colour in UK Shelf Seas

Satellite ocean-colour based models of size-fractionated primary production (PP) have been developed for the oceans on a global level. Uncertainties exist as to whether these models are accurate for temperate Shelf seas. In this paper, an existing ocean-colour based PP model is tuned using a large in situ database of size-fractionated measurements from the Celtic Sea and Western English Channel of chlorophyll-a (Chl a) and the photosynthetic parameters, the maximum photosynthetic rate ( P m B ) and light limited slope ( α B ). Estimates of size fractionated PP over an annual cycle in the UK shelf seas are compared with the original model that was parameterised using in situ data from the open ocean and a climatology of in situ PP from 2009 to 2015. The Shelf Sea model captured the seasonal patterns in size-fractionated PP for micro- and picophytoplankton, and generally performed better than the original open ocean model, except for nanophytoplankton PP which was over-estimated. The overestimation in PP is in part due to errors in the parameterisation of the biomass profile during summer, stratified conditions. Compared to the climatology of in situ data, the shelf sea model performed better when phytoplankton biomass was high, but overestimated PP at low Chl a.

Seasonality in the cross-shelf physical structure of a temperate shelf sea and the implications for nitrate supply

We address a long-standing problem of how nutrients are transported from the shelf edge and from rivers to support regular, seasonal primary production in the interior of a wide, temperate, shelf sea. Cross-shelf sections of hydrography and nutrients, from a series of cruises between March 2014 and August 2015, along with time series of river discharge and river nutrient load are used to assess the seasonality of cross-shelf transports. Riverine nitrogen inputs are estimated to account for 30% of the nitrate available for the spring bloom on the inner shelf, and 10% in the mid- to outer-shelf. In the bottom layer in summer, high salinity, nutrient-rich waters are transported on-shelf as a result of wind-driven Ekman transport, cross-shelf pressure gradients and/or internal tidal wave Stoke’s drift. In the centre of the shelf this advection is responsible for 25% of the increase in bottom water nitrate seen between April and November 2014. The remaining nitrate increase suggests that about 50–62% of the nitrogen fixed into organic material during spring, summer and autumn phytoplankton growth is recycled in the bottom water over the 12 months between March 2014 and March 2015. In winter, when the water column is vertically mixed, there is a weak net off-shelf transport of about 1 m2 s−1, possibly driven by a reversal of the horizontal density gradient caused by excess cooling of shallower shelf waters. Overall, shelf nitrate concentrations are maintained by a combination of riverine supply, recycling of organic material, and summer on-shelf transports. We suggest that the main driver of inter-annual variability in pre-spring nitrate concentrations is variability in the depth of the winter mixed layer over the shelf slope.

Nitrate drawdown during a shelf sea spring bloom revealed using a novel microfluidic in situ chemical sensor deployed within an autonomous underwater glider

Here we describe, for the first time, the use of a miniaturized Lab-on-Chip (LoC) nutrient sensor deployed within an autonomous underwater vehicle (AUV; Kongsberg Seaglider) to collect high-resolution nitrate (nitrate + nitrite) data in a highly dynamic shelf sea environment. Seasonally stratified temperate shelf seas act as important carbon sinks, where primary production is controlled by the availability of nutrients such as nitrate. Spring phytoplankton blooms and sporadic mixing events can drastically modify the availability of nitrate on temporal scales from hours to days. Traditional sampling methods are unable to capture high frequency events that can be clearly observed using a wet-chemical microfluidic system deployed within a glider. We highlight firstly, an excellent agreement between the LoC and shipboard nitrate + nitrite measurements (r2 = 0.98 n = 11). Secondly, the LoC was able to observe a decrease in nitrate within the surface mixed layer from 5.74 μM (4th) to 1.42 μM (25th) during the onset of the spring bloom, whilst bottom layer concentrations remained constant (6.86 ± 0.16 μM), with an estimated analytical uncertaintly of <0.2 μM. Thirdly, the ability of an LoC sensor deployed within an AUV to accurately capture simultaneous biogeochemical and physical parameters at an enhanced resolution, on both spatial and temporal scales, improves our understanding of biogeochemical cycles within the dynamic temperate shelf sea environments.

Growth and mortality of coccolithophores during spring in a temperate Shelf Sea (Celtic Sea, April 2015)

Coccolithophores are key components of phytoplankton communities, exerting a critical impact on the global carbon cycle and the Earth’s climate through the production of coccoliths made of calcium carbonate (calcite) and bioactive gases. Microzooplankton grazing is an important mortality factor in coccolithophore blooms, however little is currently known regarding the mortality (or growth) rates within non-bloom populations. Measurements of coccolithophore calcite production (CP) and dilution experiments to determine microzooplankton (≤63 µm) grazing rates were made during a spring cruise (April 2015) at the Central Celtic Sea (CCS), shelf edge (CS2), and within an adjacent April bloom of the coccolithophore Emiliania huxleyi at station J2.CP at CCS ranged from 10.4 to 40.4 µmol C m−3 d−1 and peaked at the height of the spring phytoplankton bloom (peak chlorophyll-a concentrations ∼6 mg m−3). Cell normalised calcification rates declined from ∼1.7 to ∼0.2 pmol C cell−1 d−1, accompanied by a shift from a mixed coccolithophore species community to one dominated by the more lightly calcified species E. huxleyi and Calciopappus caudatus. At the CCS, coccolithophore abundance increased from 6 to 94 cells mL−1, with net growth rates ranging from 0.06 to 0.21 d−1 from the 4th to the 28th April. Estimates of intrinsic growth and grazing rates from dilution experiments, at the CCS ranged from 0.01 to 0.86 d−1 and from 0.01 to 1.32 d−1, respectively, which resulted in variable net growth rates during April. Microzooplankton grazers consumed 59 to >100% of daily calcite production at the CCS. Within the E. huxleyi bloom a maximum density of 1986 cells mL−1 was recorded, along with CP rates of 6000 µmol C m−3 d−1 and an intrinsic growth rate of 0.29 d−1, with ∼80% of daily calcite production being consumed.Our results show that microzooplankton can exert strong top-down control on both bloom and non-bloom coccolithophore populations, grazing over 60% of daily growth (and calcite production). The fate of consumed calcite is unclear, but may be lost either through dissolution in acidic food vacuoles, and subsequent release as CO2, or export to the seabed after incorporation into small faecal pellets. With such high microzooplankton-mediated mortality losses, the fate of grazed calcite is clearly a high priority research direction.