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.

Temporal variability in total, micro- and nano-phytoplankton primary production at a coastal site in the Western English Channel

Primary productivity and subsequent carbon cycling in the coastal zone have a significant impact on the global carbon budget. It is currently unclear how anthropogenic activity could alter these budgets but long term coastal time series of hydrological, biogeochemical and biological measurements represent a key means to better understand past drivers, and hence to predicting future seasonal and inter-annual variability in carbon fixation in coastal ecosystems. An 8-year time series of primary production from 2003 to 2010, estimated using a recently developed absorption-based algorithm, was used to determine the nature and extent of change in primary production at a coastal station (L4) in the Western English Channel (WEC). Analysis of the seasonal and inter-annual variability in production demonstrated that on average, nano- and pico-phytoplankton account for 48% of the total carbon fixation and micro-phytoplankton for 52%. A recent decline in the primary production of nano- and pico-phytoplankton from 2005 to 2010 was observed, corresponding with a decrease in winter nutrient concentrations and a decrease in the biomass of Phaeocystis sp. Micro-phytoplankton primary production (PPM) remained relatively constant over the time series and was enhanced in summer during periods of high precipitation. Increases in sea surface temperature, and decreases in wind speeds and salinity were associated with later spring maxima in PPM. Together these trends indicate that predicted increases in temperature and decrease in wind speeds in future would drive later spring production whilst predicted increases in precipitation would also continue these blooms throughout the summer at this site.

The global distribution of phytoplankton size spectrum and size classes from their light-absorption spectra derived from satellite data

The absorption spectra of phytoplankton in the visible domain hold implicit information on the phytoplankton community structure. Here we use this information to retrieve quantitative information on phytoplankton size structure by developing a novel method to compute the exponent of an assumed power-law for their particle-size spectrum. This quantity, in combination with total chlorophyll-a concentration, can be used to estimate the fractional concentration of chlorophyll in any arbitrarily-defined size class of phytoplankton. We further define and derive expressions for two distinct measures of cell size of mixed. populations, namely, the average spherical diameter of a bio-optically equivalent homogeneous population of cells of equal size, and the average equivalent spherical diameter of a population of cells that follow a power-law particle-size distribution. The method relies on measurements of two quantities of a phytoplankton sample: the concentration of chlorophyll-a, which is an operational index of phytoplankton biomass, and the total absorption coefficient of phytoplankton in the red peak of visible spectrum at 676 nm. A sensitivity analysis confirms that the relative errors in the estimates of the exponent of particle size spectra are reasonably low. The exponents of phytoplankton size spectra, estimated for a large set of in situ data from a variety of oceanic environments (similar to 2400 samples), are within a reasonable range; and the estimated fractions of chlorophyll in pico-, nano- and micro-phytoplankton are generally consistent with those obtained by an independent, indirect method based on diagnostic pigments determined using high-performance liquid chromatography. The estimates of cell size for in situ samples dominated by different phytoplankton types (diatoms, prymnesiophytes, Prochlorococcus, other cyanobacteria and green algae) yield nominal sizes consistent with the taxonomic classification. To estimate the same quantities from satellite-derived ocean-colour data, we combine our method with algorithms for obtaining inherent optical properties from remote sensing. The spatial distribution of the size-spectrum exponent and the chlorophyll fractions of pico-, nano- and micro-phytoplankton estimated from satellite remote sensing are in agreement with the current understanding of the biogeography of phytoplankton functional types in the global oceans. This study contributes to our understanding of the distribution and time evolution of phytoplankton size structure in the global oceans.

Role of light and photophysiological properties on phytoplankton succession during the spring bloom in the north-western Mediterranean Sea

This study aimed to determine the role of light on the succession of the phytoplankton community during the spring bloom in the northwestern Mediterranean Sea. To this end, three successive Lagrangian experiments were carried out between March and April 2003. The three experiments correspond to distinct phases of the bloom development (pre-bloom, bloom peak and post-bloom, respectively) and therefore to different trophic conditions. Phytoplankton (sampled on a daily scale) was grouped in size-based classes (pico and nano+micro) each of them were characterised in terms of chemotaxonomic composition, primary production and photophysiological properties. The phytoplankton community evolved with time changing in both size-class dominance and specie/group dominance within each size class. The bloom peak was characterised by highly dynamic condition (i.e. vertical mixing) and by the dominance of both small (pico) and large (nano and micro) diatoms, as a result of their capacity to photoacclimate to changing light regimes (‘physiological plasticity’). Concluding, we suggest that the physiological adaptation to light is the main factor driving the succession of the phytoplankton community during the first phases of the bloom (until the onset of thermal stratification) in the western Mediterranean Sea.

Interactions between multiple large macrofauna species and nematode communities — Mechanisms for indirect impacts of trawling disturbance

Highlights •We exposed meiofauna to 7 different large macrofauna species at high and low densities. •Macrofauna presence altered nematode community structure and reduced their abundance. •Macrofauna species had similar effects by reducing the few dominant nematode species. •Meio–macrofauna resource competition and spatial segregation are the main drivers. •Trawling effects on macrofauna affect nematode communities indirectly. Diverse assemblages of infauna in sediments provide important physical and biogeochemical services, but are under increasing pressure by anthropogenic activities, such as benthic trawling. It is known that trawling disturbance has a substantial effect on the larger benthic fauna, with reductions in density and diversity, and changes in community structure, benthic biomass, production, and bioturbation and biogeochemical processes. Largely unknown, however, are the mechanisms by which the trawling impacts on the large benthic macro- and megafauna may influence the smaller meiofauna. To investigate this, a mesocosm experiment was conducted whereby benthic nematode communities from a non-trawled area were exposed to three different densities (absent, low, normal) of 7 large (> 10 mm) naturally co-occurring, bioturbating species which are potentially vulnerable to trawling disturbance. The results showed that total abundances of nematodes were lower if these large macrofauna species were present, but no clear nematode abundance effects could be assigned to the macrofauna density differences. Nematode community structure changed in response to macrofauna presence and density, mainly as a result of the reduced abundance of a few dominant nematode species. Any detectable effects seemed similar for nearly all macrofauna species treatments, supporting the idea that there may be a general indirect, macrofauna-mediated trawling impact on nematode communities. Explanations for these results may be, firstly, competition for food resources, resulting in spatial segregation of the meio- and macrobenthic components. Secondly, different densities of large macrofauna organisms may affect the nematode community structure through different intensities of bioturbatory disturbance or resource competition. These results suggest that removal or reduced densities of larger macrofauna species as a result of trawling disturbance may lead to increased nematode abundance and hints at the validity of interference competition between large macrofauna organisms and the smaller meiofauna, and the energy equivalence hypothesis, where a trade-off is observed between groups of organisms that are dependent on a common source of energy.

Absorption-based algorithm of primary production for total and size-fractionated phytoplankton in coastal waters

Most satellite models of production have been designed and calibrated for use in the open ocean. Coastal waters are optically more complex, and the use of chlorophyll a (chl a) as a first-order predictor of primary production may lead to substantial errors due to significant quantities of coloured dissolved organic matter (CDOM) and total suspended material (TSM) within the first optical depth. We demonstrate the use of phytoplankton absorption as a proxy to estimate primary production in the coastal waters of the North Sea and Western English Channel for both total, micro- and nano+pico-phytoplankton production. The method is implemented to extrapolate the absorption coefficient of phytoplankton and production at the sea surface to depth to give integrated fields of total and micro- and nano+pico-phytoplankton primary production using the peak in absorption coefficient at red wavelengths. The model is accurate to 8% in the Western English Channel and 22% in this region and the North Sea. By comparison, the accuracy of similar chl a based production models was >250%. The applicability of the method to autonomous optical sensors and remotely sensed aircraft data in both coastal and estuarine environments is discussed.

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

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

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

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