Response of an Arctic Sediment Nitrogen Cycling Community to Increased CO2

Ocean acidification influences sediment/water nitrogen fluxes, possibly by impacting on the microbial process of ammonia oxidation. To investigate this further, undisturbed sediment cores collected from Ny Alesund harbour (Svalbard) were incubated with seawater adjusted to CO2 concentrations of 380, 540, 760, 1,120 and 3,000 μatm. DNA and RNA were extracted from the sediment surface after 14 days' exposure and the abundance of bacterial and archaeal ammonia oxidising (amoA) genes and transcripts quantified using quantitative polymerase chain reaction. While there was no change to the abundance of bacterial amoA genes, an increase to 760 μatm pCO2 reduced the abundance of bacterial amoA transcripts by 65 %, and this was accompanied by a shift in the composition of the active community. In contrast, archaeal amoA gene and transcript abundance both doubled at 3,000 μatm, with an increase in species richness also apparent. This suggests that ammonia oxidising bacteria and archaea in marine sediments have different pH optima, and the impact of elevated CO2 on N cycling may be dependent on the relative abundances of these two major microbial groups. Further evidence of a shift in the balance of key N cycling groups was also evident: the abundance of nirS-type denitrifier transcripts decreased alongside bacterial amoA transcripts, indicating that NO3 − produced by bacterial nitrification fuelled denitrification. An increase in the abundance of Planctomycete-specific 16S rRNA, the vastmajority of which grouped with known anammox bacteria, was also apparent at 3,000 μatm pCO2. This could indicate a possible shift from coupled nitrification–denitrification to anammox activity at elevated CO2.

Seasonal shelf-sea front mapping using satellite ocean colour and temperature to support development of a marine protected area network

Front detection and aggregation techniques were applied to 300m resolution MERIS satellite ocean colour data for the first time, to describe frequently occurring shelf-sea fronts near to the Scottish coast. Medium resolution (1km) thermal and colour data have previously been used to analyse the distribution of surface fronts, though these cannot capture smaller frontal zones or those in close proximity to the coast, particularly where the coastline is convoluted. Seasonal frequent front maps, derived from both chlorophyll and SST data, revealed a number of key frontal zones, a subset of which were based on new insights into the sediment and plankton dynamics provided exclusively by the higher-resolution chlorophyll fronts. The methodology is described for applying colour and thermal front data to the task of identifying zones of ecological importance that could assist the process of defining marine protected areas. Each key frontal zone is analysed to describe its spatial and temporal extent and variability, and possible mechanisms. It is hoped that these tools can provide guidance on the dynamic habitats of marine fauna towards aspects of marine spatial planning and conservation.

Macrofauna along the Sudanese Red Sea coast: potential effect of mangrove clearance on community and trophic structure

Mangroves along the Sudanese Red Sea coast are under constant anthropogenic pressure. To better understand the influence of mangrove clearance on the intertidal benthic community, we investigated the composition, biodiversity and standing stock of the macrofauna communities at high-, mid- and low-water levels in three contrasting habitats: a bare sand flat, a cleared mangrove and an intact mangrove. In addition, a community-wide metric approach based on taxon-specific carbon and nitrogen isotope values was used to compare the trophic structure between the three habitats. The habitats differed significantly in terms of macrofaunal standing stock, community composition and trophic structure. The high- and mid-water levels of the intact mangroves showed a distinct macrofaunal community characterized by elevated densities and biomass, largely governed by higher decapod and gastropod abundances. Diversity was similar for cleared and intact mangroves, but much lower for the bare sand flat. Community-wide metrics indicated highest trophic diversity and community niche breadth in the intact mangroves. Differences between the cleared and intact mangroves can be partly attributed to differences in sediment characteristics resulting from mangrove clearance. These results suggest a significant impact of mangrove clearance on the macrofaunal community and trophic structure. This study calls for further investigations and management actions to protect and restore these habitats, and ensure the survival of this ecologically valuable coastal ecosystem.

Scale-dependent foraging ecology of a marine top predator modelled using passive acoustic data

1.Understanding which environmental factors drive foraging preferences is critical for the development of effective management measures, but resource use patterns may emerge from processes that occur at different spatial and temporal scales. Direct observations of foraging are also especially challenging in marine predators, but passive acoustic techniques provide opportunities to study the behaviour of echolocating species over a range of scales. 2.We used an extensive passive acoustic data set to investigate the distribution and temporal dynamics of foraging in bottlenose dolphins using the Moray Firth (Scotland, UK). Echolocation buzzes were identified with a mixture model of detected echolocation inter-click intervals and used as a proxy of foraging activity. A robust modelling approach accounting for autocorrelation in the data was then used to evaluate which environmental factors were associated with the observed dynamics at two different spatial and temporal scales. 3.At a broad scale, foraging varied seasonally and was also affected by seabed slope and shelf-sea fronts. At a finer scale, we identified variation in seasonal use and local interactions with tidal processes. Foraging was best predicted at a daily scale, accounting for site specificity in the shape of the estimated relationships. 4.This study demonstrates how passive acoustic data can be used to understand foraging ecology in echolocating species and provides a robust analytical procedure for describing spatio-temporal patterns. Associations between foraging and environmental characteristics varied according to spatial and temporal scale, highlighting the need for a multi-scale approach. Our results indicate that dolphins respond to coarser scale temporal dynamics, but have a detailed understanding of finer-scale spatial distribution of resources.

Response of a floodplain fish community to river-floodplain connectivity: natural versus managed reconnection

To restore lateral connectivity in highly regulated river-floodplain systems, it has become necessary to implement localized, "managed" connection flows, made possible using floodplain irrigation infrastructure. These managed flows contrast with "natural", large-scale, overbank flood pulses. We compared the effects of a managed and a natural connection event on (i) the composition of the large-bodied fish community and (ii) the structure of an endangered catfish population of a large floodplain lake. The change in community composition following the managed connection was not greater than that exhibited between seasons or years during disconnection. By contrast, the change in fish community structure following the natural connection was much larger than that attributed to background, within-and between-year variability during disconnection. Catfish population structure only changed significantly following the natural flood. While the natural flood increased various population rates of native fishes, it also increased those of non-native carp, a pest species. To have a positive influence on native biodiversity, environmental flows may need to be delivered to floodplains in a way that simulates the properties of natural flood pulses. A challenge, however, will be managing river-floodplain connectivity to benefit native more than non-native species.

Challenges in integrative approaches to modelling the marine ecosystems of the North Atlantic: Physics to fish and coasts to ocean

It has long been recognised that there are strong interactions and feedbacks between climate, upper ocean biogeochemistry and marine food webs, and also that food web structure and phytoplankton community distribution are important determinants of variability in carbon production and export from the euphotic zone. Numerical models provide a vital tool to explore these interactions, given their capability to investigate multiple connected components of the system and the sensitivity to multiple drivers, including potential future conditions. A major driver for ecosystem model development is the demand for quantitative tools to support ecosystem-based management initiatives. The purpose of this paper is to review approaches to the modelling of marine ecosystems with a focus on the North Atlantic Ocean and its adjacent shelf seas, and to highlight the challenges they face and suggest ways forward. We consider the state of the art in simulating oceans and shelf sea physics, planktonic and higher trophic level ecosystems, and look towards building an integrative approach with these existing tools. We note how the different approaches have evolved historically and that many of the previous obstacles to harmonisation may no longer be present. We illustrate this with examples from the on-going and planned modelling effort in the Integrative Modelling Work Package of the EURO-BASIN programme.

Effects of increases in temperature and nutrients on phytoplankton community structure and photosynthesis in the Western English Channel

Anthropogenic climate change is exerting pressures on coastal ecosystems through increases in temperature, precipitation and ocean acidification. Phytoplankton community structure and photo-physiology are therefore adapting to these conditions. Changes in phytoplankton biomass and photosynthesis in relation to temperature and nutrient concentrations were assessed using a 14 year dataset from a coastal station in the Western English Channel (WEC). Dinoflagellate and coccolithophorid biomass exhibited a positive correlation with temperature, reaching the highest biomass at between 15 and 17°C. Diatoms showed a negative correlation with temperature, with highest biomass at 10°C. Chlorophyll a (chl a) normalised light-saturated photosynthetic rates (PBm) exhibited a hyperbolic response to increasing temperature, with an initial linear increase from 8 to 11°C, and reaching a plateau from 12°C. There was however no significant positive correlation between nutrients and phytoplankton biomass or PBm, which reflects the lag time between nutrient input and phytoplankton growth at this coastal site. The major phytoplankton groups that occurred at this site occupied distinct thermal niches, which in turn modified PBm. Increasing temperature, and higher water column stratification, was major factors in the initiation of dinoflagellates blooms at this site. Dinoflagellates blooms during summer also co-varied with silicate concentration, and acted as a tracer of dissolved inorganic nitrogen and phosphate from river run-off, which were subsequently reduced during these blooms. The data implies that increasing temperature and high river runoff during summer, will promote dinoflaglellates blooms in the WEC.

Effects of elevated CO2 and temperature on an intertidal meiobenthic community

In the near future, the marine environment is likely to be subjected to simultaneous increases in temperature and decreased pH. The potential effects of these changes on intertidal, meiofaunal assemblages were investigated using a mesocosm experiment. Artificial Substrate Units containing meiofauna from the extreme low intertidal zone were exposed for 60 days to eight experimental treatments (four replicates for each treatment) comprising four pH levels: 8.0 (ambient control), 7.7 & 7.3 (predicted changes associated with ocean acidification), and 6.7 (CO2 point-source leakage from geological storage), crossed with two temperatures: 12 °C (ambient control) and 16 °C (predicted). Community structure, measured using major meiofauna taxa was significantly affected by pH and temperature. Copepods and copepodites showed the greatest decline in abundance in response to low pH and elevated temperature. Nematodes increased in abundance in response to low pH and temperature rise, possibly caused by decreased predation and competition for food owing to the declining macrofauna density. Nematode species composition changed significantly between the different treatments, and was affected by both seawater acidification and warming. Estimated nematode species diversity, species evenness, and the maturity index, were substantially lower at 16 °C, whereas trophic diversity was slightly higher at 16 °C except at pH 6.7. This study has demonstrated that the combination of elevated levels of CO2 and ocean warming may have substantial effects on structural and functional characteristics of meiofaunal and nematode communities, and that single stressor experiments are unlikely to encompass the complexity of abiotic and biotic interactions. At the same time, ecological interactions may lead to complex community responses to pH and temperature changes in the interstitial environment

Satellite estimates of net community production indicate predominance of net autotrophy in the Atlantic Ocean

There is ongoing debate as to whether the oligotrophic ocean is predominantly net autotrophic and acts as a CO2 sink, or net heterotrophic and therefore acts as a CO2 source to the atmosphere. This quantification is challenging, both spatially and temporally, due to the sparseness of measurements. There has been a concerted effort to derive accurate estimates of phytoplankton photosynthesis and primary production from satellite data to fill these gaps; however there have been few satellite estimates of net community production (NCP). In this paper, we compare a number of empirical approaches to estimate NCP from satellite data with in vitro measurements of changes in dissolved O2 concentration at 295 stations in the N and S Atlantic Ocean (including the Antarctic), Greenland and Mediterranean Seas. Algorithms based on power laws between NCP and particulate organic carbon production (POC) derived from 14C uptake tend to overestimate NCP at negative values and underestimate at positive values. An algorithm that includes sea surface temperature (SST) in the power function of NCP and 14C POC has the lowest bias and root-mean square error compared with in vitro measured NCP and is the most accurate algorithm for the Atlantic Ocean. Nearly a 13 year time series of NCP was generated using this algorithm with SeaWiFS data to assess changes over time in different regions and in relation to climate variability. The North Atlantic subtropical and tropical Gyres (NATL) were predominantly net autotrophic from 1998 to 2010 except for boreal autumn/winter, suggesting that the northern hemisphere has remained a net sink for CO2 during this period. The South Atlantic subtropical Gyre (SATL) fluctuated from being net autotrophic in austral spring-summer, to net heterotrophic in austral autumn–winter. Recent decadal trends suggest that the SATL is becoming more of a CO2 source. Over the Atlantic basin, the percentage of satellite pixels with negative NCP was 27%, with the largest contributions from the NATL and SATL during boreal and austral autumn–winter, respectively. Variations in NCP in the northern and southern hemispheres were correlated with climate indices. Negative correlations between NCP and the multivariate ENSO index (MEI) occurred in the SATL, which explained up to 60% of the variability in NCP. Similarly there was a negative correlation between NCP and the North Atlantic Oscillation (NAO) in the Southern Sub-Tropical Convergence Zone (SSTC),which explained 90% of the variability. There were also positive correlations with NAO in the Canary Current Coastal Upwelling (CNRY) and Western Tropical Atlantic (WTRA)which explained 80% and 60% of the variability in each province, respectively. MEI and NAO seem to play a role in modifying phases of net autotrophy and heterotrophy in the Atlantic Ocean.

Assessing the sensitivity of seagrass bed biotopes to pressures associated with marine activities.

This project was commissioned to generate an improved understanding of the sensitivities of seagrass habitats to pressures associated with human activities in the marine environment - to provide an evidence base to facilitate and support management advice for Marine Protected Areas; development of UK marine monitoring and assessment, and conservation advice to offshore marine industries. Seagrass bed habitats are identified as a Priority Marine Feature (PMF) under the Marine (Scotland) Act 2010, they are also included on the OSPAR list of threatened and declining species and habitats, and are a Habitat of Principle Importance (HPI) under the Natural Environment and Rural Communities (NERC) Act 2006, in England and Wales. The purpose of this project was to produce sensitivity assessments with supporting evidence for the HPI, OSPAR and PMF seagrass/Zostera bed habitat definitions, clearly documenting the evidence behind the assessments and any differences between assessments. Nineteen pressures, falling in five categories - biological, hydrological, physical damage, physical loss, and pollution and other chemical changes - were assessed in this report. Assessments were based on the three British seagrasses Zostera marina, Z. noltei and Ruppia maritima. Z. marina var. angustifolia was considered to be a subspecies of Z. marina but it was specified where studies had considered it as a species in its own rights. Where possible other components of the community were investigated but the basis of the assessment focused on seagrass species. To develop each sensitivity assessment, the resistance and resilience of the key elements were assessed against the pressure benchmark using the available evidence. The benchmarks were designed to provide a ‘standard’ level of pressure against which to assess sensitivity. Overall, seagrass beds were highly sensitive to a number of human activities: • penetration or disturbance of the substratum below the surface; • habitat structure changes – removal of substratum; • physical change to another sediment type; • physical loss of habitat; • siltation rate changes including and smothering; and • changes in suspended solids. High sensitivity was recorded for pressures which directly impacted the factors that limit seagrass growth and health such as light availability. Physical pressures that caused mechanical modification of the sediment, and hence damage to roots and leaves, also resulted in high sensitivity. Seagrass beds were assessed as ‘not sensitive’ to microbial pathogens or ‘removal of target species’. These assessments were based on the benchmarks used. Z. marina is known to be sensitive to Labyrinthula zosterae but this was not included in the benchmark used. Similarly, ‘removal of target species’ addresses only the biological effects of removal and not the physical effects of the process used. For example, seagrass beds are probably not sensitive to the removal of scallops found within the bed but are highly sensitive to the effects of dredging for scallops, as assessed under the pressure penetration or disturbance of the substratum below the surface‘. This is also an example of a synergistic effect Assessing the sensitivity of seagrass bed biotopes to pressures associated with marine activities between pressures. Where possible, synergistic effects were highlighted but synergistic and cumulative effects are outside the scope off this study. The report found that no distinct differences in sensitivity exist between the HPI, PMF and OSPAR definitions. Individual biotopes do however have different sensitivities to pressures. These differences were determined by the species affected, the position of the habitat on the shore and the sediment type. For instance evidence showed that beds growing in soft and muddy sand were more vulnerable to physical damage than beds on harder, more compact substratum. Temporal effects can also influence the sensitivity of seagrass beds. On a seasonal time frame, physical damage to roots and leaves occurring in the reproductive season (summer months) will have a greater impact than damage in winter. On a daily basis, the tidal regime could accentuate or attenuate the effects of pressures depending on high and low tide. A variety of factors must therefore be taken into account in order to assess the sensitivity of a particular seagrass habitat at any location. No clear difference in resilience was established across the three seagrass definitions assessed in this report. The resilience of seagrass beds and the ability to recover from human induced pressures is a combination of the environmental conditions of the site, growth rates of the seagrass, the frequency and the intensity of the disturbance. This highlights the importance of considering the species affected as well as the ecology of the seagrass bed, the environmental conditions and the types and nature of activities giving rise to the pressure and the effects of that pressure. For example, pressures that result in sediment modification (e.g. pitting or erosion), sediment change or removal, prolong recovery. Therefore, the resilience of each biotope and habitat definitions is discussed for each pressure. Using a clearly documented, evidence based approach to create sensitivity assessments allows the assessment and any subsequent decision making or management plans to be readily communicated, transparent and justifiable. The assessments can be replicated and updated where new evidence becomes available ensuring the longevity of the sensitivity assessment tool. The evidence review has reduced the uncertainty around assessments previously undertaken in the MB0102 project (Tillin et al 2010) by assigning a single sensitivity score to the pressures as opposed to a range. Finally, as seagrass habitats may also contribute to ecosystem function and the delivery of ecosystem services, understanding the sensitivity of these biotopes may also support assessment and management in regard to these. Whatever objective measures are applied to data to assess sensitivity, the final sensitivity assessment is indicative. The evidence, the benchmarks, the confidence in the assessments and the limitations of the process, require a sense-check by experienced marine ecologists before the outcome is used in management decisions.

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.

Benthic megafauna and CO2 bubble dynamics observed by underwater photography during a controlled sub-seabed release of CO2

In 2012, a controlled sub-seabed release of carbon dioxide (CO2) was conducted in Ardmucknish Bay, a shallow (12 m) coastal bay on the west coast of Scotland. During the experiment, CO2 gas was released 12 m below the seabed for 37 days, causing significant disruption to sediment and water carbonate chemistry as the gas passed up through the sediment and into the overlying water. One of the aims of the study was to investigate how the impacts caused by leakage from geological CO2 Capture and Storage (CCS) could be detected and quantified in the context of natural heterogeneity and dynamics. To do this underwater photography was used to analyze (i) the benthic megafaunal response to the CO2 release and (ii) the dynamics of the CO2 bubble streams, emerging from the seabed into the overlying water column. The frequently observed megafauna species in the study area were Virgularia mirabilis (Cnidaria), Turritella communis (Mollusca), Asterias rubens (Echinodermata), Pagurus bernhardus (Crustacea), Liocarcinus depurator (Crustacea), and Gadus morhua (Osteichthyes). No discernable abnormal behavior was observed for these megafauna, in any of the zones investigated, during or after the CO2 release. Time-lapse photography revealed that the intensity and presence of the CO2 bubble plume was affected by the tides, with the most active bubbling seen at low tides and the larger hydrostatic pressure at high tide suppressing CO2 bubbling from the seabed.

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.