Multiple environmental controls on phytoplankton growth strategies determine adaptive responses of the N : P ratio

The controls on the 'Redfield' N:P stoichiometry of marine phytoplankton and hence the N:P ratio of the deep ocean remain incompletely understood. Here, we use a model for phytoplankton ecophysiology and growth, based on functional traits and resource-allocation trade-offs, to show how environmental filtering, biotic interactions, and element cycling in a global ecosystem model determine phytoplankton biogeography, growth strategies and macromolecular composition. Emergent growth strategies capture major observed patterns in marine biomes. Using a new synthesis of experimental RNA and protein measurements to constrain per-ribosome translation rates, we determine a spatially variable lower limit on adaptive rRNA:protein allocation and hence on the relationship between the largest cellular P and N pools. Comparison with the lowest observed phytoplankton N:P ratios and N:P export fluxes in the Southern Ocean suggests that additional contributions from phospholipid and phosphorus storage compounds play a fundamental role in determining the marine biogeochemical cycling of these elements.

Evaluating the SEVIRI Fire Thermal Anomaly Detection Algorithm across the Central African Republic Using the MODIS Active Fire Product

Satellite-based remote sensing of active fires is the only practical way to consistently and continuously monitor diurnal fluctuations in biomass burning from regional, to continental, to global scales. Failure to understand, quantify, and communicate the performance of an active fire detection algorithm, however, can lead to improper interpretations of the spatiotemporal distribution of biomass burning, and flawed estimates of fuel consumption and trace gas and aerosol emissions. This work evaluates the performance of the Spinning Enhanced Visible and Infrared Imager (SEVIRI) Fire Thermal Anomaly (FTA) detection algorithm using seven months of active fire pixels detected by the Moderate Resolution Imaging Spectroradiometer (MODIS) across the Central African Republic (CAR). Results indicate that the omission rate of the SEVIRI FTA detection algorithm relative to MODIS varies spatially across the CAR, ranging from 25% in the south to 74% in the east. In the absence of confounding artifacts such as sunglint, uncertainties in the background thermal characterization, and cloud cover, the regional variation in SEVIRI's omission rate can be attributed to a coupling between SEVIRI's low spatial resolution detection bias (i.e., the inability to detect fires below a certain size and intensity) and a strong geographic gradient in active fire characteristics across the CAR. SEVIRI's commission rate relative to MODIS increases from 9% when evaluated near MODIS nadir to 53% near the MODIS scene edges, indicating that SEVIRI errors of commission at the MODIS scene edges may not be false alarms but rather true fires that MODIS failed to detect as a result of larger pixel sizes at extreme MODIS scan angles. Results from this work are expected to facilitate (i) future improvements to the SEVIRI FTA detection algorithm; (ii) the assimilation of the SEVIRI and MODIS active fire products; and (iii) the potential inclusion of SEVIRI into a network of geostationary sensors designed to achieve global diurnal active fire monitoring.

Mesoscale fronts as foraging habitats: composite front mapping reveals oceanographic drivers of habitat use for a pelagic seabird

The oceanographic drivers of marine vertebrate habitat use are poorly understood yet fundamental to our knowledge of marine ecosystem functioning. Here, we use composite front mapping and high-resolution GPS tracking to determine the significance of mesoscale oceanographic fronts as physical drivers of foraging habitat selection in northern gannets Morus bassanus. We tracked 66 breeding gannets from a Celtic Sea colony over 2 years and used residence time to identify area-restricted search (ARS) behaviour. Composite front maps identified thermal and chlorophyll-a mesoscale fronts at two different temporal scales—(i) contemporaneous fronts and (ii) seasonally persistent frontal zones. Using generalized additive models (GAMs), with generalized estimating equations (GEE-GAMs) to account for serial autocorrelation in tracking data, we found that gannets do not adjust their behaviour in response to contemporaneous fronts. However, ARS was more likely to occur within spatially predictable, seasonally persistent frontal zones (GAMs). Our results provide proof of concept that composite front mapping is a useful tool for studying the influence of oceanographic features on animal movements. Moreover, we highlight that frontal persistence is a crucial element of the formation of pelagic foraging hotspots for mobile marine vertebrates.

On the Front Line: frontal zones as priority at-sea conservation areas for mobile marine vertebrates

1.Identifying priority areas for marine vertebrate conservation is complex because species of conservation concern are highly mobile, inhabit dynamic habitats and are difficult to monitor. 2.Many marine vertebrates are known to associate with oceanographic fronts – physical interfaces at the transition between water masses – for foraging and migration, making them important candidate sites for conservation. Here, we review associations between marine vertebrates and fronts and how they vary with scale, regional oceanography and foraging ecology. 3.Accessibility, spatiotemporal predictability and relative productivity of front-associated foraging habitats are key aspects of their ecological importance. Predictable mesoscale (10s–100s km) regions of persistent frontal activity (‘frontal zones’) are particularly significant. 4.Frontal zones are hotspots of overlap between critical habitat and spatially explicit anthropogenic threats, such as the concentration of fisheries activity. As such, they represent tractable conservation units, in which to target measures for threat mitigation. 5.Front mapping via Earth observation (EO) remote sensing facilitates identification and monitoring of these hotspots of vulnerability. Seasonal or climatological products can locate biophysical hotspots, while near-real-time front mapping augments the suite of tools supporting spatially dynamic ocean management. 6.Synthesis and applications. Frontal zones are ecologically important for mobile marine vertebrates. We surmise that relative accessibility, predictability and productivity are key biophysical characteristics of ecologically significant frontal zones in contrasting oceanographic regions. Persistent frontal zones are potential priority conservation areas for multiple marine vertebrate taxa and are easily identifiable through front mapping via EO remote sensing. These insights are useful for marine spatial planning and marine biodiversity conservation, both within Exclusive Economic Zones and in the open oceans.

Family-level AMBI is valid for use in the north-eastern Atlantic but not for assessing the health of microtidal Australian estuaries

Analysis of benthic macroinvertebrate samples at a higher taxonomic level than species, e.g. family, potentially provides a more cost-effective protocol for environmental impact assessments and monitoring as it requires less time, funds and taxonomic expertise. Using the AMBI database, species ecological group scores are shown to be coherent within families. Faunal data from a wide range of environmental impact scenarios in the north-eastern Atlantic demonstrate that AMBI, calculated from mean values for families, exhibits a strong linear relationship with species-level AMBI, the correlation improving by using square-root transformed rather than untransformed abundances. In many regions of the world, however, the sensitivity of benthic macroinvertebrates to environmental perturbations is unknown, precluding the use of AMBI for environmental assessments. Yet the families are essentially the same as in the AMBI database. The utility of family-level AMBI is tested using data for four south-western Australian estuaries previously subjected to environmental quality assessments, but where only 17 species of the 144 taxa are included in the AMBI database. Although family-level AMBI scores reflect differences in environmental quality spatially and temporally within an estuary, they do not follow variations in environmental quality among estuaries. Indeed, south-western Australia estuaries are numerically dominated by families with high AMBI scores, probably due to the detrimental effects of natural accumulations of organic material in estuaries with long residence times. As taxonomic distinctness follows trends in environmental quality among estuaries, as well as temporally and spatially within a system, it provides an appropriate substitute for assessing the 'heath' of microtidal estuaries.

Marine baseline and monitoring strategies for carbon dioxide capture and storage (CCS)

The QICS controlled release experiment demonstrates that leaks of carbon dioxide (CO2) gas can be detected by monitoring acoustic, geochemical and biological parameters within a given marine system. However the natural complexity and variability of marine system responses to (artificial) leakage strongly suggests that there are no absolute indicators of leakage or impact that can unequivocally and universally be used for all potential future storage sites. We suggest a multivariate, hierarchical approach to monitoring, escalating from anomaly detection to attribution, quantification and then impact assessment, as required. Given the spatial heterogeneity of many marine ecosystems it is essential that environmental monitoring programmes are supported by a temporally (tidal, seasonal and annual) and spatially resolved baseline of data from which changes can be accurately identified. In this paper we outline and discuss the options for monitoring methodologies and identify the components of an appropriate baseline survey.

How many Coccolithovirus genotypes does it take to terminate an Emiliania huxleyi bloom?

Giant viruses are known to be significant mortality agents of phytoplankton, often being implicated in the terminations of large Emiliania huxleyi blooms. We have previously shown the high temporal variability of E. huxleyi-infecting coccolithoviruses (EhVs) within a Norwegian fjord mesocosm. In the current study we investigated EhV dynamics within a naturally-occurring E. huxleyi bloom in the Western English Channel. Using denaturing gradient gel electrophoresis and marker gene sequencing, we uncovered a spatially highly dynamic Coccolithovirus population that was associated with a genetically stable E. huxleyi population as revealed by the major capsid protein gene (mcp) and coccolith morphology motif (CMM), respectively. Coccolithoviruses within the bloom were found to be variable with depth and unique virus populations were detected at different stations sampled indicating a complex network of EhV-host infections. This ultimately will have significant implications to the internal structure and longevity of ecologically important E. huxleyi blooms.

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.

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.

Developments since 2005 in understanding potential environmental impacts of CO2 leakage from geological storage

This paper reviews research into the potential environmental impacts of leakage from geological storage of CO2 since the publication of the IPCC Special Report on Carbon Dioxide Capture and Storage in 2005. Possible impacts are considered on onshore (including drinking water aquifers) and offshore ecosystems. The review does not consider direct impacts on man or other land animals from elevated atmospheric CO2 levels. Improvements in our understanding of the potential impacts have come directly from CO2 storage research but have also benefitted from studies of ocean acidification and other impacts on aquifers and onshore near surface ecosystems. Research has included observations at natural CO2 sites, laboratory and field experiments and modelling. Studies to date suggest that the impacts from many lower level fault- or well-related leakage scenarios are likely to be limited spatially and temporarily and recovery may be rapid. The effects are often ameliorated by mixing and dispersion of the leakage and by buffering and other reactions; potentially harmful elements have rarely breached drinking water guidelines. Larger releases, with potentially higher impact, would be possible from open wells or major pipeline leaks but these are of lower probability and should be easier and quicker to detect and remediate.

Dynamics of air–sea CO2 fluxes in the northwestern European shelf based on voluntary observing ship and satellite observations

From January 2011 to December 2013, we constructed a comprehensive pCO2 data set based on voluntary observing ship (VOS) measurements in the western English Channel (WEC). We subsequently estimated surface pCO2 and air–sea CO2 fluxes in northwestern European continental shelf waters using multiple linear regressions (MLRs) from remotely sensed sea surface temperature (SST), chlorophyll a concentration (Chl a), wind speed (WND), photosynthetically active radiation (PAR) and modeled mixed layer depth (MLD). We developed specific MLRs for the seasonally stratified northern WEC (nWEC) and the permanently well-mixed southern WEC (sWEC) and calculated surface pCO2 with uncertainties of 17 and 16 μatm, respectively. We extrapolated the relationships obtained for the WEC based on the 2011–2013 data set (1) temporally over a decade and (2) spatially in the adjacent Celtic and Irish seas (CS and IS), two regions which exhibit hydrographical and biogeochemical characteristics similar to those of WEC waters. We validated these extrapolations with pCO2 data from the SOCAT and LDEO databases and obtained good agreement between modeled and observed data. On an annual scale, seasonally stratified systems acted as a sink of CO2 from the atmosphere of −0.6 ± 0.3, −0.9 ± 0.3 and −0.5 ± 0.3 mol C m−2 yr−1 in the northern Celtic Sea, southern Celtic sea and nWEC, respectively, whereas permanently well-mixed systems acted as source of CO2 to the atmosphere of 0.2 ± 0.2 and 0.3 ± 0.2 mol C m−2 yr−1 in the sWEC and IS, respectively. Air–sea CO2 fluxes showed important inter-annual variability resulting in significant differences in the intensity and/or direction of annual fluxes. We scaled the mean annual fluxes over these provinces for the last decade and obtained the first annual average uptake of −1.11 ± 0.32 Tg C yr−1 for this part of the northwestern European continental shelf. Our study showed that combining VOS data with satellite observations can be a powerful tool to estimate and extrapolate air–sea CO2 fluxes in sparsely sampled area.

Scaling up experimental ocean acidification and warming research: from individuals to the ecosystem

Understanding long-term, ecosystem-level impacts of climate change is challenging because experimental research frequently focuses on short-term, individual-level impacts in isolation. We address this shortcoming first through an inter-disciplinary ensemble of novel experimental techniques to investigate the impacts of 14-month exposure to ocean acidification and warming (OAW) on the physiology, activity, predatory behaviour and susceptibility to predation of an important marine gastropod (Nucella lapillus). We simultaneously estimated the potential impacts of these global drivers on N. lapillus population dynamics and dispersal parameters. We then used these data to parameterise a dynamic bioclimatic envelope model, to investigate the consequences of OAW on the distribution of the species in the wider NE Atlantic region by 2100. The model accounts also for changes in the distribution of resources, suitable habitat and environment simulated by finely resolved biogeochemical models, under three IPCC global emissions scenarios. The experiments showed that temperature had the greatest impact on individual level responses, while acidification has a similarly important role in the mediation of predatory behaviour and susceptibility to predators. Changes in Nucella predatory behaviour appeared to serve as a strategy to mitigate individual level impacts of acidification, but the development of this response may be limited in the presence of predators. The model projected significant large-scale changes in the distribution of Nucella by the year 2100 that were exacerbated by rising greenhouse gas emissions. These changes were spatially heterogeneous, as the degree of impact of OAW on the combination of responses considered by the model varied depending on local environmental conditions and resource availability. Such changes in macro-scale distributions cannot be predicted by investigating individual level impacts in isolation, or by considering climate stressors separately. Scaling up the results of experimental climate change research requires approaches that account for long-term, multi-scale responses to multiple stressors, in an ecosystem context.

Ocean-wide tracking of pelagic sharks reveals extent of overlap with longline fishing hotspots

Overfishing is arguably the greatest ecological threat facing the oceans, yet catches of many highly migratory fishes including oceanic sharks remain largely unregulated with poor monitoring and data reporting. Oceanic shark conservation is hampered by basic knowledge gaps about where sharks aggregate across population ranges and precisely where they overlap with fishers. Using satellite tracking data from six shark species across the North Atlantic, we show that pelagic sharks occupy predictable habitat ‘hotspots’ of high space use. Movement modelling showed sharks preferred habitats characterised by strong sea-surface-temperature gradients (fronts) over other available habitats. However, simultaneous Global Positioning System (GPS) tracking of the entire Spanish and Portuguese longline-vessel fishing fleets show an 80% overlap of fished areas with hotspots, potentially increasing shark susceptibility to fishing exploitation. Regions of high overlap between oceanic tagged sharks and longliners included the North Atlantic Current/Labrador Current convergence zone and the Mid-Atlantic Ridge south-west of the Azores. In these main regions, and sub-areas within them, shark/vessel co-occurrence was spatially and temporally persistent between years, highlighting how broadly the fishing exploitation efficiently ‘tracks’ oceanic sharks within their space-use hotspots year-round. Given this intense focus of longliners on shark hotspots our study argues the need for international catch limits for pelagic sharks and identifies a future role of combining fine-scale fish and vessel telemetry to inform the ocean-scale management of fisheries.

Spatio-temporal Bayesian network models with latent variables for revealing trophic dynamics and functional networks in fisheries ecology

Ecosystems consist of complex dynamic interactions among species and the environment, the understanding of which has implications for predicting the environmental response to changes in climate and biodiversity. However, with the recent adoption of more explorative tools, like Bayesian networks, in predictive ecology, few assumptions can be made about the data and complex, spatially varying interactions can be recovered from collected field data. In this study, we compare Bayesian network modelling approaches accounting for latent effects to reveal species dynamics for 7 geographically and temporally varied areas within the North Sea. We also apply structure learning techniques to identify functional relationships such as prey–predator between trophic groups of species that vary across space and time. We examine if the use of a general hidden variable can reflect overall changes in the trophic dynamics of each spatial system and whether the inclusion of a specific hidden variable can model unmeasured group of species. The general hidden variable appears to capture changes in the variance of different groups of species biomass. Models that include both general and specific hidden variables resulted in identifying similarity with the underlying food web dynamics and modelling spatial unmeasured effect. We predict the biomass of the trophic groups and find that predictive accuracy varies with the models' features and across the different spatial areas thus proposing a model that allows for spatial autocorrelation and two hidden variables. Our proposed model was able to produce novel insights on this ecosystem's dynamics and ecological interactions mainly because we account for the heterogeneous nature of the driving factors within each area and their changes over time. Our findings demonstrate that accounting for additional sources of variation, by combining structure learning from data and experts' knowledge in the model architecture, has the potential for gaining deeper insights into the structure and stability of ecosystems. Finally, we were able to discover meaningful functional networks that were spatially and temporally differentiated with the particular mechanisms varying from trophic associations through interactions with climate and commercial fisheries.

Chapter 11: Human activities and ecosystem service use: impacts and trade-offs

Ecosystem services provided by the marine environment are fundamental to human health and well-being. Despite this, many marine systems are being degraded to an extent that may reduce their capacity to provide these ecosystem services. The ecosystem approach is a strategy for the integrated management of land, water and living resources that promotes conservation and sustainable use in an equitable way (UN Convention on Biological Diversity, 2000). Its application to marine management and spatial planning has been proposed as a means of maintaining the economic and social value of the oceans, not only in the present but for generations to come. Characterising the susceptibility of services (and combinations of services) to particular human activities based on knowledge of impacts on biodiversity and ecosystem functioning (as described in preceding chapters) is a challenge for future management of the oceans. In this chapter, we highlight the existing, but limited knowledge of how ecosystem services may be impacted by different human activities. We discuss how impacts on one service can impact multiple services and explore how the impacts on services can vary both spatially and temporally and according to context. We focus particularly on the effects on ecosystem services of activities whose impacts on biodiversity and ecosystem functioning have already been considered in previous chapters. Some of these activities are associated with poor management of ecosystem benefits, for example, from provisioning services (aquaculture and fisheries), or with excessive input of wastes, fertilisers and contaminants into the system overburdening the waste treatment and assimilation services. Other impacts are associated with the construction of structures or use of space designed to generate benefits from environmental services such as the presence of water as a carrier for shipping, or sources of wind, wave and tidal power. We discuss the trade-offs that are made, consciously or otherwise, between different ecosystem services, which arise from human activities to optimise or manage specific ecosystem services.