Southern Ocean biogeography and taxonomic resolution: what's in the name?

We examined the taxonomic resolution of zooplankton data required to identify ocean basin scale biogeographic zonation in the Southern Ocean. A 2,154 km transect was completed south of Australia. Sea surface temperature (SST) measured at 1 min intervals showed that seven physical zones were sampled. Zooplankton were collected at a spatial resolution of similar to 9.2 km with a continuous plankton recorder, identified to the highest possible taxonomic resolution and enumerated. Zooplankton assemblage similarity between samples was calculated using the Bray-Curtis index for the taxonomic levels of species, genus, family, order and class after first log(10)(x + 1) (LA) and then presence/absence (PA) transformation of abundance data. Although within and between zone sample similarity increased with decreasing taxonomic resolution, for both data transformations, cluster analysis demonstrated that the biogeographic separation of zones remained at all taxonomic levels when using LA data. ANOSIM confirmed this, detecting significant differences in zooplankton assemblage structure between all seven a priori determined physical zones for all taxonomic levels when using the LA data. In the case of the PA data for the complete data set, and both LA and PA data for a crustacean only data set, no significant differences were detected between zooplankton assemblages in the Polar frontal zone (PFZ) and inter-PFZ at any taxonomic level. Loss of information at resolutions below the species level, particularly in the PA data, prevented the separation of some zones. However, the majority of physical zones were biogeographically distinct from species level to class using both LA and PA transformations. Significant relationships between SST and zooplankton community structure, summarised as NMDS scores, at all taxonomic levels, for both LA and PA transformations, and complete and crustacean only data sets, highlighted the biogeographic relevance of low resolution taxonomic data. The retention of biogeographic information in low taxonomic resolution data shows that data sets collected with different taxonomic resolutions may be meaningfully merged for the post hoc generation of Southern Ocean time series.

Going Global: planning the next 80 years of the Continuous Plankton Recorder Survey

Going Global: planning the next 80 years of the Continuous Plankton Recorder Survey. Operated by the Sir Alister Hardy Foundation for Ocean Science (SAHFOS), the Continuous Plankton Recorder (CPR) survey is the world’s largest, sampling 4 ocean basins, and longest running (since 1931) plankton biodiversity monitoring programme. Having sampled enough miles to circumnavigate the globe over 200 times, the CPR database houses over 2.5 million entries, describing the distribution of 500 phytoplankton and zooplankton taxa. Routinely sampling in the Arctic, Atlantic, Pacific and Southern Oceans, the survey analyses 4000 samples yearly. Data collected from these samples are made freely available for bona fide scientific purposes. The CPR survey data is used to generate a better understanding of changes in the plankton and to date some 1000 papers have been published on plankton biodiversity. This year sees the 80th anniversary of the CPR survey and to celebrate and build upon this unique monitoring programme, SAHFOS intends to further develop its global plankton perspective. Work will be extended into the South Atlantic and Indian Ocean and an international partnership with complementary surveys in Australia, Canada, America, Japan and South Africa will be implemented. The Digital Object will describe the CPR survey using compilations made by Plymouth Art College and BBC film footage.

Lagrangian evolution of DMS during the Southern Ocean gas exchange experiment: The effects of vertical mixing and biological community shift

Concentrations of dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP) are highly variable in time and space. What is driving the variability in DMS(P), and can those variability be explained by physical processes and changes in the biological community? During the Southern Ocean Gas Exchange Experiment (SO GasEx) in the austral fall of 2008, two 3He/SF6 labeled patches were created in the surface water. SF6 and DMS were surveyed continuously in a Lagrangian framework, while direct measurements of air-sea exchange further constrained the gas budgets. Turbulent diffusivity at the base of the mixed layer was estimated from SF6 profiles and used to calculate the vertical fluxes of DMS and nutrients. Increasing mixed layer nutrient concentrations due to mixing were associated with a shift in the phytoplankton community structure, which in turned likely affected the sulfur dynamics on timescales of days. DMS concentration as well as air-sea DMS flux appeared to be decoupled from the DMSP concentration, possibly due to grazing and bacterial DMS production. Contrary to expectations, in an environment with high winds and modest productivity, physical processes (air-sea exchange, photochemistry, vertical mixing) only accounted for a small fraction of DMS loss from the surface water. Among the DMS sinks, inferred biological consumption most likely dominated during SO GasEx.

Strong responses of Southern Ocean phytoplankton communities to volcanic ash

Volcanic eruptions have been hypothesized as an iron supply mechanism for phytoplankton blooms; however, little direct evidence of stimulatory responses has been obtained in the field. Here we present the results of twenty-one 1–2 day bottle enrichment experiments from cruises in the South Atlantic and Southern Ocean which conclusively demonstrated a photophysiological and biomass stimulation of phytoplankton communities following supply of basaltic or rhyolitic volcanic ash. Furthermore, experiments in the Southern Ocean demonstrated significant phytoplankton community responses to volcanic ash supply in the absence of responses to addition of dissolved iron alone. At these sites, dissolved manganese concentrations were among the lowest ever measured in seawater, and we therefore suggest that the enhanced response to ash may have been a result of the relief of manganese (co)limitation. Our results imply that volcanic ash deposition events could trigger extensive phytoplankton blooms, potentially capable of significant impacts on regional carbon cycling.

Ecological Investigations with the Continuous Plankton Recorder: The Phytoplankton in the Southern North Sea. 1932-37.

1. The changes in the composition and distribution of the plankton of the southern North Sea have been investigated month by month, from June 1932 to December 1937; the present report deals with the phytoplankton. The survey was carried out by the Continuous Plankton Recorder, towed at a standard depth of 10 metres, by ships on regular steamship lines across the North Sea from Hull towards the Skagerrak, to Bremen and to Rotterdam, and later between London and Esbjerg. 2. The material and methods are described, together with a discussion on the validity of this type of survey and some comparison of its results with those obtained by other methods (pp. 76-86). 3. Particular attention has been paid to Rhizosolenia styliformis (pp. 92- 107), Biddulphia sinensis (pp. 108-115), Phaeocystis (pp. 149-153), and the Dinoflagellates (pp. 134-149); of these the first three are known to be of particular importance in relation to the herring fisheries. More generalised data are available for the principal diatoms other than R. styliformis and B. sinensis (pp. 116-134). 4. The main part of the work is an ecological study of the phytoplankton changes in time and space over the 5½ years. Each year is marked by some distinct variations in the abundance and the times of increase, maximum numbers and decline as recorded in the different forms. These variations in the annual cycles are compared on the different lines by a series of graphs arranged against a time scale of months, a set for each year being placed side by side (Plates I-XXI). More detailed studies by more frequent records were made in the autumns of 1934, 1935, 1936 and 1937 (cf. Figs. 3 and 4). The changes in spatial distribution are shown by a series of monthly maps arranged in a similar manner for each year (Plates XXII-LXIV). These intensive studies of the changes in time and space are also intended to form the basis for correlations with other features in the general ecology of the area (e. g. the zooplankton, hydrology, meteorology and fisheries) to be made in later publications. 5. Whilst each form has shown its own peculiar features, a trend towards a general increase in the phytoplankton as a whole has been observed during the period, although the years 1934 and 1936 have in some respects shown deviations and regressive features, and not all organisms have revealed the same trend. The possible relation of this gradual trend to other events observed in recent years in these and neighbouring waters is discussed (pp. 162-167). 6. The application of these results to the study of patchiness (pp. 154-158), inter-relationships in the plankton (pp. 159-160) and to water movements (pp. 160-162) is briefly discussed.

Ecological Investigations with the Continuous Plankton Recorder: The Copepoda of the Southern North Sea. 1932-37.

I. The monthly changes in the distribution and abundance of the Copepoda in the southern North Sea have been investigated from June 1932 to December 1937 by using the Continuous Plankton Recorder. This was towed at a standard depth of 10 metres by ships sailing on regular lines from Hull to Rotterdam, to Bremen and towards the Skagerrak, and later from London to Esbjerg. 2. The methods are described and those limitations which apply more particularly to the Copepoda are discussed (pp. 175 to 186 and 198 to 203). 3. The first part of the report deals with the Copepoda as a whole, i.e. the total population. The difference between the summer and winter distributions is stressed. The variations in numbers from year to year are found to be considerable and it is suggested that they are sufficiently large to be reflected in the success or failure of the broods of those fish which are at some period of their development dependent upon the Copepoda for food. 4. The second part deals with the data concerning the constituent species or groups of allied species ; a list of these is given on p. 197. 5. The group Paracalanus + Pseudocalanus was by far the most abundant and together with the genera Temora and Acartia was found to be responsible for most of the fluctuations in the population (pp. 205 to 208). 6. The distributions, seasonal and spatial, of the other common forms are described, with the exception of that of Oalantts finmarchicus which is to be the subject of a later report. 7. The recorder results are compared with the findings of the International Council survey from 1902 to 1908; some marked disagreements are discussed (pp. 227 to 232). 8. The appearance of the northern forms Oandacia armata and Metridia lucens during the winters of 1932-33, 1933-34 and 1937 are recorded (pp. 222 to 223) 9. A summarised account of the main seasonal changes in the area is given (pp. 232 to 234) and followed by a brief comparison of the 5½ years investigated.

Ecological Investigations with the Continuous Plankton Recorder: The Zooplankton (other than Copepoda and Young Fish) in the Southern North Sea, 1932-37.

I. The report describes the main monthly changes in the distribution and abundance of the zooplankton, other than Copepoda and young fish (dealt with in separate reports), over the southern part of the North Sea from 1932 to 1937. The work is part of the survey carried out by Continuous Plankton Recorders towed at a depth of 10 metres on regular steamship lines between England and the Continent. 2. The limitations to the sampling method are discussed, and it is shown to be unsuitable for recording Mysidacea and Euphausiacea on account of their marked diurnal variation due presumably to vertical migration; they are omitted from the report. 3. The changing distribution of Sagitta, Limacina, Clione, Lamellibranch larvae, Cladocera, Caprellid Amphipoda, Decapod larvae, Echinoderm larvae and Oikopleura are shown in a series of monthly charts while their seasonal fluctuations are compared in time-chart histograms. 4. The Alima larvae of Squilla are recorded on a few occasions in the regions where the Channel opens into the North Sea. 5. The distributional characteristics of the different forms, i.e. their tendencies to even or " patchy " production, are compared.

Dimethylsulfide gas transfer coefficients from algal blooms in the Southern Ocean

Air–sea dimethylsulfide (DMS) fluxes and bulk air–sea gradients were measured over the Southern Ocean in February–March 2012 during the Surface Ocean Aerosol Production (SOAP) study. The cruise encountered three distinct phytoplankton bloom regions, consisting of two blooms with moderate DMS levels, and a high biomass, dinoflagellate-dominated bloom with high seawater DMS levels (> 15 nM). Gas transfer coefficients were considerably scattered at wind speeds above 5 m/s. Bin averaging the data resulted in a linear relationship between wind speed and mean gas transfer velocity consistent with that previously observed. However, the wind-speed-binned gas transfer data distribution at all wind speeds is positively skewed. The flux and seawater DMS distributions were also positively skewed, which suggests that eddy covariance-derived gas transfer velocities are consistently influenced by additional, log-normal noise. A flux footprint analysis was conducted during a transect into the prevailing wind and through elevated DMS levels in the dinoflagellate bloom. Accounting for the temporal/spatial separation between flux and seawater concentration significantly reduces the scatter in computed transfer velocity. The SOAP gas transfer velocity data show no obvious modification of the gas transfer–wind speed relationship by biological activity or waves. This study highlights the challenges associated with eddy covariance gas transfer measurements in biologically active and heterogeneous bloom environments.

Trait-based representation of diatom functional diversity in a plankton functional type model of the eutrophied southern North Sea

We introduce a trait-based description of diatom functional diversity to an existing plankton functional type (PFT) model, implemented for the eutrophied coastal ecosystem in the Southern Bight of the North Sea. The trait-based description represents a continuum of diatom species, each characterized by a distinct cell volume, and includes size dependence of four diatom traits: the maximum growth rate, the half-saturation constants for nutrient uptake, the photosynthetic efficiency, and the relative affinity of copepods for diatoms. Through competition under seasonally varying forcing, the fitness of each diatom varies throughout time, and the outcome of competition results in a changing community structure. The predicted seasonal change in mean cell volume of the community is supported by field observations: smaller diatoms, which are more competitive in terms of resource acquisition, prevail during the first spring bloom, whereas the summer bloom is dominated by larger species which better resist grazing. The size-based model is used to determine the ecological niche of diatoms in the area and identifies a range of viable sizes that matches observations. The general trade-off between small, competitive diatoms and large, grazing-resistant species is a convenient framework to study patterns in diatom functional diversity. PFT models and trait-based approaches constitute promising complementary tools to study community structure in marine ecosystems.

High-resolution view of the spring bloom initiation and net community production in the Subantarctic Southern Ocean using glider data

In the Southern Ocean, there is increasing evidence that seasonal to subseasonal temporal scales, and meso- to submesoscales play an important role in understanding the sensitivity of ocean primary productivity to climate change. This drives the need for a high-resolution approach to re- solving biogeochemical processes. In this study, 5.5 months of continuous, high-resolution (3 h, 2 km horizontal resolution) glider data from spring to summer in the Atlantic Subantarctic Zone is used to investigate: (i) the mechanisms that drive bloom initiation and high growth rates in the region and (ii) the seasonal evolution of water column production and respiration. Bloom initiation dates were analysed in the context of upper ocean boundary layer physics highlighting sensitivities of different bloom detection methods to different environmental processes. Model results show that in early spring (September to mid-November) increased rates of net community production (NCP) are strongly affected by meso- to submesoscale features. In late spring/early summer (late-November to mid-December) seasonal shoaling of the mixed layer drives a more spatially homogenous bloom with maximum rates of NCP and chlorophyll biomass. A comparison of biomass accumulation rates with a study in the North Atlantic highlights the sensitivity of phytoplankton growth to fine-scale dynamics and emphasizes the need to sample the ocean at high resolution to accurately resolve phytoplankton phenology and improve our ability to estimate the sensitivity of the biological carbon pump to climate change.

Southern Annular Mode and westerly-wind-driven changes in Indian-Atlantic exchange mechanisms

The dynamical link between the Indian Ocean and Atlantic Meridional Overturning Circulation (AMOC) remains poorly understood. This partly arises from the complex Agulhas leakage, which occurs via rings, cyclones, and non-eddy flux. Hindcast simulations suggest that leakage has recently increased but have not decomposed this signal into its constituent mechanisms. Here these are isolated in a realistic ocean model. Increases in simulated leakage are attributed to stronger eddy and non-eddy-driven transports, and a strong warming and salinification, especially within Agulhas rings. Variability in both regimes is associated with strengthening Indian Ocean westerly winds, reflecting an increasingly positive Southern Annular Mode. While eddy and non-eddy flux signals are tied through turbulent eddy dissipation, the ratio between the two varies decadally. Consequently, while altimetry suggests a recent increase in retroflection turbulence and implied leakage, non-eddy flux may also play a significant role in modulating the leakage AMOC connection.

Role of zooplankton dynamics for Southern Ocean phytoplankton biomass and global biogeochemical cycles

Global ocean biogeochemistry models currently employed in climate change projections use highly simplified representations of pelagic food webs. These food webs do not necessarily include critical pathways by which ecosystems interact with ocean biogeochemistry and climate. Here we present a global biogeochemical model which incorporates ecosystem dynamics based on the representation of ten plankton functional types (PFTs); six types of phytoplankton, three types of zooplankton, and heterotrophic bacteria. We improved the representation of zooplankton dynamics in our model through (a) the explicit inclusion of large, slow-growing zooplankton, and (b) the introduction of trophic cascades among the three zooplankton types. We use the model to quantitatively assess the relative roles of iron vs. grazing in determining phytoplankton biomass in the Southern Ocean High Nutrient Low Chlorophyll (HNLC) region during summer. When model simulations do not represent crustacean macrozooplankton grazing, they systematically overestimate Southern Ocean chlorophyll biomass during the summer, even when there was no iron deposition from dust. When model simulations included the developments of the zooplankton component, the simulation of phytoplankton biomass improved and the high chlorophyll summer bias in the Southern Ocean HNLC region largely disappeared. Our model results suggest that the observed low phytoplankton biomass in the Southern Ocean during summer is primarily explained by the dynamics of the Southern Ocean zooplankton community rather than iron limitation. This result has implications for the representation of global biogeochemical cycles in models as zooplankton faecal pellets sink rapidly and partly control the carbon export to the intermediate and deep ocean.

Birth, life and death of a cyclonic eddy in the Southern Ocean

The ACC is a climatically relevant frontal structure of global importance that regularly develops instabilities which grow into meanders that eventually evolve into long-lived cyclonic eddies. These eddies exhibit sustain primary productivity that can last several months fuelled by local resupply of nutrients. During April-May 2015 we conducted an intensive field experiment in the Southern Ocean (SMILES) where we sampled and tracked an ACC meander as it developed into an eddy and later vanished some 90 days later. The meander and later eddy physical characteristics were observed with a combination of high resolution hydrography, ADCP and turbulence observations in addition to surface and depth resolved biogeochemical observations of nutrients and phytoplankton. The life and death of the eddy was subsequently tracked through ARGO, BIO-ARGO and remote sensing.

Depth-related gradients in community structure and relatedness of bivalves and isopods in the Southern Ocean.

Despite increased research over the last decade, diversity patterns in Antarctic deep-sea benthic taxa and their driving forces are only marginally known. Depth-related patterns of diversity and distribution of isopods and bivalves collected in the Atlantic sector of the Southern Ocean are analysed. The data, sampled by epibenthic sledge at 40 deep-sea stations from the upper continental slope to the hadal zone (774 – 6348 m) over a wide area of the Southern Ocean, comprises 619 species of isopods and 81 species of bivalves,. There were more species of isopods than bivalves in all samples, and species per station varied from 2 to 85 for isopods and from 0 to 18 for bivalves. Most species were rare, with 72% of isopod species restricted to one or two stations, and 45% of bivalves. Among less-rare species bivalves tended to have wider distributions than isopods. The species richness of isopods varied with depth, showing a weak unimodal curve with a peak at 2000 – 4000 m, while the richness of bivalves did not. Multivariate analyses indicate that there are two main assemblages in the Southern Ocean, one shallow and one deep. These overlap over a large depth-range (2000 – 4000 m). Comparing analyses based on the Sørensen resemblance measure (presence/absence) and Γ+ (presence/absence incorporating relatedness among species) indicates that rare species tend to have other closely related species within the same depth band. Analysis of relatedness among species indicates that the taxonomic variety of bivalves tends to decline at depth, whereas that of isopods is maintained. This, it is speculated, may indicate that the available energy at depth is insufficient to maintain a range of bivalve life-history strategies