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.

Rapid response of the active microbial community to CO2 exposure from a controlled sub-seabed CO2 leak in Ardmucknish Bay (Oban, Scotland)

The response of the benthic microbial community to a controlled sub-seabed CO2 leak was assessed using quantitative PCR measurements of benthic bacterial, archaeal and cyanobacteria/chloroplast 16S rRNA genes. Samples were taken from four zones (epicentre; 25 m distant, 75 m distant and 450 m distant) during 6 time points (7 days before CO2 exposure, after 14 and 36 days of CO2 release, and 6, 20 and 90 days after the CO2 release had ended). Changes to the active community of microphytobenthos and bacteria were also assessed before, during and after CO2 release. Increases in the abundance of microbial 16S rRNA were detected after 14 days of CO2 release and at a distance of 25 m from the epicentre. CO2 related changes to the relative abundance of both major and minor bacterial taxa were detected: most notably an increase in the relative abundance of the Planctomycetacia after 14 days of CO2 release. Also evident was a decrease in the abundance of microbial 16S rRNA genes at the leak epicentre during the initial recovery phase: this coincided with the highest measurements of DIC within the sediment, but may be related to the release of potentially toxic metals at this time point.

Impact of sub-seabed CO2 leakage on macrobenthic community structure and diversity

A sub-seabed release of carbon dioxide (CO2) was conducted to assess the potential impacts of leakage from sub-seabed geological CO2 Capture and Storage CCS) on benthic macrofauna. CO2 gas was released 12 m below the seabed for 37 days, causing significant disruption to sediment carbonate chemistry. Regular macrofauna samples were collected from within the area of active CO2 leakage (Zone 1) and in three additional reference areas, 25 m, 75 m and 450 m from the centre of the leakage (Zones 2, 3 and 4 respectively). Macrofaunal community structure changed significantly in all zones during the study period. However, only the changes in Zone 1 were driven by the CO2 leakage with the changes in reference zones appearing to reflect natural seasonal succession and stochastic weather events. The impacts in Zone 1 occurred rapidly (within a few days), increased in severity through the duration of the leak, and continued to worsen after the leak had stopped. Considerable macrofaunal recovery was seen 18 days after the CO2 gas injection had stopped. In summary, small short-term CCS leakage events are likely to cause highly localised impacts on macrofaunal communities and there is the potential for rapid recovery to occur, depending on the characteristics of the communities and habitats impacted.

Can benthic community structure be used to predict the process of bioturbation in real ecosystems

Disentangling the roles of environmental change and natural environmental variability on biologically mediated ecosystem processes is paramount to predict future marine ecosystem functioning. Bioturbation, the biogenic mixing of sediments, has a regulating role in marine biogeochemical processes. However, our understanding of bioturbation as a community level process and of its environmental drivers is still limited by loose use of terminology, and a lack of consensus about what bioturbation is. To help resolve these challenges, this empirical study investigated the links between four different attributes of bioturbation (bioturbation depth, activity and distance, and biodiffusive transport); the ability of an index of bioturbation (BPc) to predict each of them; and their relation to seasonality, in a shallow coastal system – the Western Channel Observatory, UK. Bioturbation distance depended on changes in benthic community structure, while the other three attributes were more directly influenced by seasonality in food availability. In parallel, BPc successfully predicted bioturbation distance but not the other attributes of bioturbation. This study therefore highlights that community bioturbation results from this combination of processes responding to environmental variability at different time-scales. However, community level measurements of bioturbation across environmental variability are still scarce, and BPc is calculated using commonly available data on benthic community structure and the functional classification of invertebrates. Therefore, BPc could be used to support the growth of landscape scale bioturbation research, but future uses of the index need to consider which bioturbation attributes the index actually predicts. As BPc predicts bioturbation distance, estimated here using a random-walk model applicable to community settings, studies using either of the metrics should be directly comparable and contribute to a more integrated future for bioturbation research.

Looking backwards to look forwards: the role of natural history in temperate reef ecology

Temperate reefs are superb tractable systems for testing hypotheses in ecology and evolutionary biology. Accordingly there is a rich history of research stretching back over 100 years, which has made major contributions to general ecological and evolutionary theory as well as providing better understanding of how littoral systems work by linking pattern with process. A brief resumé of the history of temperate reef ecology is provided to celebrate this rich heritage. As a community, temperate reef ecologists generally do well designed experiments and test well formulated hypotheses. Increasingly large datasets are being collected, collated and subjected to complex meta-analyses and used for modelling. These datasets do not happen spontaneously – the burgeoning subject of macroecology is possible only because of the efforts of dedicated natural historians whether it be observing birds, butterflies, or barnacles. High-quality natural history and old-fashioned field craft enable surveys or experiments to be stratified (i.e. replicates are replicates and not a random bit of rock) and lead to the generation of more insightful hypotheses. Modern molecular approaches have led to the discovery of cryptic species and provided phylogeographical insights, but natural history is still required to identify species in the field. We advocate a blend of modern approaches with old school skills and a fondness for temperate reefs in all their splendour.

Ocean acidification has different effects on the production of DMS and DMSP measured in cultures of Emiliania huxleyi and a mesocosm study: a comparison of laboratory monocultures and community interactions

The human-induced rise in atmospheric carbon dioxide since the industrial revolution has led to increasing oceanic carbon uptake and changes in seawater carbonate chemistry, resulting in lowering of surface water pH. In this study we investigated the effect of increasing CO2 partial pressure (pCO2) on concentrations of volatile biogenic dimethylsulfide (DMS) and its precursor dimethylsulfoniopropionate (DMSP), through monoculture studies and community pCO2 perturbation. DMS is a climatically important gas produced by many marine algae: it transfers sulfur into the atmosphere and is a major influence on biogeochemical climate regulation through breakdown to sulfate and formation of subsequent cloud condensation nuclei (CCN). Overall, production of DMS and DMSP by the coccolithophore Emiliania huxleyi strain RCC1229 was unaffected by growth at 900 matm pCO2, but DMSP production normalised to cell volume was 12% lower at the higher pCO2 treatment. These cultures were compared with community DMS and DMSP production during an elevated pCO2 mesocosm experiment with the aim of studying E. huxleyi in the natural environment. Results contrasted with the culture experiments and showed reductions in community DMS and DMSP concentrations of up to 60 and 32% respectively at pCO2 up to 3000 matm, with changes attributed to poorer growth of DMSP-producing nanophytoplankton species, including E. huxleyi, and potentially increased microbial consumption of DMSand dissolvedDMSPat higher pCO2.DMSandDMSPproduction differences between culture and community likely arise from pH affecting the inter-species responses between microbial producers and consumers.

The combined effects of seasonal community succession and adaptive algal physiology on lipid profiles of coastal phytoplankton in the western English channel

Lipids are key constituents of marine phytoplankton, and some fatty acids (key constituents of lipids) are essential dietary components for secondary producers. However, in natural marine ecosystems the interactions of factors affecting seasonal phytoplankton lipid composition are still poorly understood. The aim of this study was to assess the roles of seasonal succession in phytoplankton community composition and nutrient concentrations, on the lipid composition of the phytoplankton community. Fatty acid and polar lipid composition in seston was measured in surface waters at the time series station L4, an inshore station in the Western English Channel, from January to December 2013. Redundancy analyses (RDA) were used to identify factors (abiotic and biotic) that explained the seasonal variability in phytoplankton lipids. RDA demonstrated that nutrients (namely nitrogen) explained the majority of variation in phytoplankton lipid composition, as well as a smaller explanatory contribution from changes in phytoplankton community composition. The physiological adaptations of the phytoplankton community to nutrient deplete conditions during the summer season when the water column was stratified, was further supported by changes in the polar lipid to phytoplankton biomass ratios (also modelled with RDA) and increases in the lipid to chlorophyll a ratios, which are both indicative of nutrient stress. However, the association of key fatty acid markers with phytoplankton groups e.g. 22:6 n-3 and dinoflagellate biomass (predominant in summer), meant there were no clear seasonal differences in the overall degree of fatty acid saturation, as might have been expected from typical nutrient stress on phytoplankton. Based on the use of polyunsaturated fatty acids (PUFA) as markers of ‘food quality’ for grazers, our results suggest that in this environment high food quality is maintained throughout summer, due to seasonal succession towards flagellated phytoplankton species able to maintain PUFA synthesis under surface layer nutrient depletion.

Abundance of a chlorophyll a precursor and the oxidation product hydroxychlorophyll a during seasonal phytoplankton community progression in the Western English Channel

This study presents the first in-situ measurements of the chlorophyll a oxidation product, hydroxychlorophyll a as well as the chlorophyll a precursor, chlorophyll aP276 conducted over an annual cycle. Chlorophyll a oxidation products, such as hydroxychlorophyll a may be associated with the decline of algal populations and can act as an initial step in the degradation of chlorophyll a into products which can be found in the geochemical record, important for studying past climate change events. Here, hydroxychlorophyll a and chlorophyll aP276 were measured at the long-term monitoring station L4, Western Channel Observatory (UK, www.westernchannelobservatory.org) over an annual cycle (2012). Weekly measurements of phytoplankton species composition and abundance enabled detailed analysis of possible sources of hydroxychlorophyll a. Dinoflagellates, 2 diatom species, the prymnesiophyte Phaeocystis spp. and the coccolithophorid Emiliania huxleyi were all associated with hydroxychlorophyll a occurrence. However, during alternate peaks in abundance of the diatoms, no association with hydroxychlorophyll a occurred, indicating that the oxidation of chlorophyll a was dependant not only on species but also on additional factors such as the mode of mortality, growth limiting factor (i.e. nutrient concentration) or phenotypic plasticity. Surface sediment samples contained 10 times more hydroxychlorophyll a (relative to chlorophyll a) than pelagic particulate samples, indicating that more chlorophyll a oxidation occurred during sedimentation or at the sediment–water interface, than in the pelagic environment. In addition, chlorophyll aP276 correlated with chl-a concentration, thus supporting its assignment as a chl-a precursor.

Dynamic responses of the benthic bacterial community at the Western English Channel observatory site L4 are driven by deposition of fresh phytodetritus

The impact of the seasonal deposition of phytoplankton and phytodetritus on surface sediment bacterial abundance and community composition was investigated at the Western English Channel site L4. Sediment and water samples were collected from January to September in 2012, increasing in frequency during periods of high water column phytoplankton abundance. Compared to the past two decades, the spring bloom in 2012 was both unusually long in duration and contained higher than average biomass. Within spring months, the phytoplankton bloom was well mixed through the water column and showed accumulations near the sea bed, as evidenced by flow cytometry measurements of nanoeukaryotes, water column chlorophyll a and the appearance of pelagic phytoplankton at the sediment. Measurements of chlorophyll and chlorophyll degradation products indicated phytoplankton material was heavily degraded after it reached the sediment surface: the nature of the chlorophyll degradation products (predominantly pheophorbide, pyropheophorbide and hydroxychlorophyllone) was indicative of grazing activity. The abundance of bacterial 16S rRNA genes g−1 sediment (used as a proxy for bacterial biomass) increased markedly with the onset of the phytoplankton bloom, and correlated with measurements of chlorophyll at the surface sediment. Together, this suggests that bacteria may have responded to nutrients released via grazing activity. In depth sequencing of the 16S rRNA genes indicated that the composition of the bacterial community shifted rapidly through-out the prolonged spring bloom period. This was primarily due to an increase in the relative sequence abundance of Flavobacteria.

Making modelling count - increasing the contribution of shelf-seas community and ecosystem models to policy development and management

Marine legislation is becoming more complex and marine ecosystem-based management is specified in national and regional legislative frameworks. Shelf-seas community and ecosystem models (hereafter termed ecosystem models) are central to the delivery of ecosystem-based management, but there is limited uptake and use of model products by decision makers in Europe and the UK in comparison with other countries. In this study, the challenges to the uptake and use of ecosystem models in support of marine environmental management are assessed using the UK capability as an example. The UK has a broad capability in marine ecosystem modelling, with at least 14 different models that support management, but few examples exist of ecosystem modelling that underpin policy or management decisions. To improve understanding of policy and management issues that can be addressed using ecosystem models, a workshop was convened that brought together advisors, assessors, biologists, social scientists, economists, modellers, statisticians, policy makers, and funders. Some policy requirements were identified that can be addressed without further model development including: attribution of environmental change to underlying drivers, integration of models and observations to develop more efficient monitoring programmes, assessment of indicator performance for different management goals, and the costs and benefit of legislation. Multi-model ensembles are being developed in cases where many models exist, but model structures are very diverse making a standardised approach of combining outputs a significant challenge, and there is a need for new methodologies for describing, analysing, and visualising uncertainties. A stronger link to social and economic systems is needed to increase the range of policy-related questions that can be addressed. It is also important to improve communication between policy and modelling communities so that there is a shared understanding of the strengths and limitations of ecosystem models.

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

Analyses of sublittoral macrobenthic community change in a marine nature reserve using similarity profiles (SIMPROF).

Sublittoral macrobenthic communities in the Skomer Marine Nature Reserve (SMNR), Pembrokeshire, Wales, were sampled at 10 stations in 1993, 1996, 1998, 2003, 2007 and 2009 using a Day grab and a 0.5 mm mesh. The time series is analysed using Similarities Profiles (SIMPROF) tests and associated methods. Q-mode analysis using clustering with Type 1 SIMPROF addresses multivariate structure among samples, showing that there is clear structure associated with differences among years. Inverse (r-mode) analysis using Type 2 SIMPROF decisively rejects a hypothesis that species are not associated with each other. Clustering of the variables (species) with Type 3 SIMPROF identifies groups of species which covary coherently through the time-series. The time-series is characterised by a dramatic decline in abundances and diversity between the 1993 and 1996 surveys. By 1998 there had been a shift in community composition from the 1993 situation, with different species dominating. Communities had recovered in terms of abundance and species richness, but different species dominated the community. No single factor could be identified which unequivocally explained the dramatic changes observed in the SMNR. Possible causes were the effects of dispersed oil and dispersants from the Sea Empress oil spill in February 1996 and the cessation of dredge-spoil disposal off St Anne’s Head in 1995, but the most likely cause was severe weather. With many species, and a demonstrable recovery from an impact, communities within the SMNR appear to be diverse and resilient. If attributable to natural storms, the changes observed here indicate that natural variability may be much more important than is generally taken into account in the design of monitoring programmes.