Competition Drives Clumpy Species Coexistence in Estuarine Phytoplankton

Understanding the mechanisms that maintain biodiversity is a fundamental problem in ecology. Competition is thought to reduce diversity, but hundreds of microbial aquatic primary producers species coexist and compete for a few essential resources (e.g., nutrients and light). Here, we show that resource competition is a plausible mechanism for explaining clumpy distribution on individual species volume (a proxy for the niche) of estuarine phytoplankton communities ranging from North America to South America and Europe, supporting the Emergent Neutrality hypothesis. Furthermore, such a clumpy distribution was also observed throughout the Holocene in diatoms from a sediment core. A Lotka-Volterra competition model predicted position in the niche axis and functional affiliation of dominant species within and among clumps. Results support the coexistence of functionally equivalent species in ecosystems and indicate that resource competition may be a key process to shape the size structure of estuarine phytoplankton, which in turn drives ecosystem functioning.

Transregional linkages in the North-Eastern Atlantic - An ‘end-to-end ’ analysis of pelagic ecosystems

This review examines interregional linkages and gives an overview perspective on marine ecosystem functioning in the north-eastern Atlantic. It is based on three of the 'systems' considered by the European Network of Excellence for Ocean Ecosystems Analysis (EUR-OC EANS was established in 2004 under the European Framework VI funding programme to promote integration of marine ecological research within Europe), the Arctic and Nordic Seas, North Atlantic shelf seas and North Atlantic. The three systems share common open boundaries and the transport of water, heat, nutrients and particulates across these boundaries modifies local processes. Consistent with the EUR-OC EANS concept of 'end-to-end' analyses of marine food webs, the review takes an integrated approach linking ocean physics, lower trophic levels and working up the food web to top predators such as marine mammals. We begin with an overview of the regions focusing on the major physical patterns and their implications for the microbial community, phytoplankton, zooplankton, fish and top predators. Human-induced links between the regional systems are then considered and finally possible changes in the regional linkages over the next century are discussed. Because of the scale of potential impacts of climate change, this issue is considered in a separate section. The review demonstrates that the functioning of the ecosystems in each of the regions cannot be considered in isolation and the role of the atmosphere and ocean currents in linking the North Atlantic Ocean, North Atlantic shelf seas and the Arctic and Nordic Seas must be taken into account. Studying the North Atlantic and associated shelf seas as an integrated 'basin-scale' system will be a key challenge for the early twenty-first century. This requires a multinational approach that should lead to improved ecosystem-based approaches to conservation of natural resources, the maintenance of biodiversity, and a better understanding of the key role of the north-eastern Atlantic in the global carbon cycle.

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.

Assimilation of remotely-sensed optical properties to improve marine biogeochemistry modelling

In this paper we evaluate whether the assimilation of remotely-sensed optical data into a marine ecosystem model improves the simulation of biogeochemistry in a shelf sea. A localized Ensemble Kalman filter was used to assimilate weekly diffuse light attenuation coefficient data, Kd(443) from SeaWiFs, into an ecosystem model of the western English Channel. The spatial distributions of (unassimilated) surface chlorophyll from satellite, and a multivariate time series of eighteen biogeochemical and optical variables measured in situ at one long-term monitoring site were used to evaluate the system performance for the year 2006. Assimilation reduced the root mean square error and improved the correlation with the assimilated Kd(443) observations, for both the analysis and, to a lesser extent, the forecast estimates, when compared to the reference model simulation. Improvements in the simulation of (unassimilated) ocean colour chlorophyll were less evident, and in some parts of the Channel the simulation of this data deteriorated. The estimation errors for the (unassimilated) in situ data were reduced for most variables with some exceptions, e.g. dissolved nitrogen. Importantly, the assimilation adjusted the balance of ecosystem processes by shifting the simulated food web towards the microbial loop, thus improving the estimation of some properties, e.g. total particulate carbon. Assimilation of Kd(443) outperformed a comparative chlorophyll assimilation experiment, in both the estimation of ocean colour data and in the simulation of independent in situ data. These results are related to relatively low error in Kd(443) data, and because it is a bulk optical property of marine ecosystems. Assimilation of remotely-sensed optical properties is a promising approach to improve the simulation of biogeochemical and optical variables that are relevant for ecosystem functioning and climate change studies.

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.

Next Generation Sequencing Reveals the Hidden Diversity of Zooplankton Assemblages

Zooplankton play an important role in our oceans, in biogeochemical cycling and providing a food source for commercially important fish larvae. However, difficulties in correctly identifying zooplankton hinder our understanding of their roles in marine ecosystem functioning, and can prevent detection of long term changes in their community structure. The advent of massively parallel next generation sequencing technology allows DNA sequence data to be recovered directly from whole community samples. Here we assess the ability of such sequencing to quantify richness and diversity of a mixed zooplankton assemblage from a productive time series site in the Western English Channel. Methodology/Principle Findings Plankton net hauls (200 µm) were taken at the Western Channel Observatory station L4 in September 2010 and January 2011. These samples were analysed by microscopy and metagenetic analysis of the 18S nuclear small subunit ribosomal RNA gene using the 454 pyrosequencing platform. Following quality control a total of 419,041 sequences were obtained for all samples. The sequences clustered into 205 operational taxonomic units using a 97% similarity cut-off. Allocation of taxonomy by comparison with the National Centre for Biotechnology Information database identified 135 OTUs to species level, 11 to genus level and 1 to order, <2.5% of sequences were classified as unknowns. By comparison a skilled microscopic analyst was able to routinely enumerate only 58 taxonomic groups. Conclusions Metagenetics reveals a previously hidden taxonomic richness, especially for Copepoda and hard-to-identify meroplankton such as Bivalvia, Gastropoda and Polychaeta. It also reveals rare species and parasites. We conclude that Next Generation Sequencing of 18S amplicons is a powerful tool for elucidating the true diversity and species richness of zooplankton communities. While this approach allows for broad diversity assessments of plankton it may become increasingly attractive in future if sequence reference libraries of accurately identified individuals are better populated.

Impacts of ocean acidification in a warming Mediterranean Sea: An overview

Mediterranean Sea fisheries supply significant local and international markets, based largely on small pelagic fish, artisanal fisheries and aquaculture of finfish (mainly seabass and seabream) and shellfish (mussels and oysters). Fisheries and aquaculture contribute to the economy of countries bordering this sea and provide food and employment to coastal communities employing ca 600,000 people. Increasing temperatures and heat wave frequency are causing stress and mortality in marine organisms and ocean acidification is expected to worsen these effects, especially for bivalves and coralligenous systems. Recruitment and seed production present possible bottlenecks for shellfish aquaculture in the future since early life stages are vulnerable to acidification and warming. Although adult finfish seem able to withstand the projected increases in seawater CO2, degradation of seabed habitats and increases in harmful blooms of algae and jellyfish might adversely affect fish stocks. Ocean acidification should therefore be factored into fisheries and aquaculture management plans. Rising CO2 levels are expected to reduce coastal biodiversity, altering ecosystem functioning and possibly impacting tourism being the Mediterranean the world’s most visited region. We recommend that ocean acidification is monitored in key areas of the Mediterranean Sea, with regular assessments of the likely socio-economic impacts to build adaptive strategies for the Mediterranean countries concerned.

The Continuous Plankton Recorder survey: How can long-term phytoplankton datasets contribute to the assessment of Good Environmental Status?

Phytoplankton are crucial to marine ecosystem functioning and are important indicators of environmental change. Phytoplankton data are also essential for informing management and policy, particularly in supporting the new generation of marine legislative drivers, which take a holistic ecosystem approach to management. The Marine Strategy Framework Directive (MSFD) seeks to achieve Good Environmental Status (GES) of European seas through the implementation of such a management approach. This is a regional scale directive which recognises the importance of plankton communities in marine ecosystems; plankton data at the appropriate spatial, temporal and taxonomic scales are therefore required for implementation. The Continuous Plankton Recorder (CPR) survey is a multidecadal, North Atlantic basin scale programme which routinely records approximately 300 phytoplankton taxa. Because of these attributes, the survey plays a key role in the implementation of the MSFD and the assessment of GES in the Northeast Atlantic region. This paper addresses the role of the CPR's phytoplankton time-series in delivering GES through the development and informing of MSFD indicators, the setting of targets against a background of climate change and the provision of supporting information used to interpret change in non-plankton indicators. We also discuss CPR data in the context of other phytoplankton data types that may contribute to GES, as well as explore future possibilities for the use of new and innovative applications of CPR phytoplankton datasets in delivering GES. Efforts must be made to preserve long-term time series, such as the CPR, which supply vital ecological information used to informed evidence-based environmental policy.

The Continuous Plankton Recorder survey: How can long-term phytoplankton datasets contribute to the assessment of Good Environmental Status?

Phytoplankton are crucial to marine ecosystem functioning and are important indicators of environmental change. Phytoplankton data are also essential for informing management and policy, particularly in supporting the new generation of marine legislative drivers, which take a holistic ecosystem approach to management. The Marine Strategy Framework Directive (MSFD) seeks to achieve Good Environmental Status (GES) of European seas through the implementation of such a management approach. This is a regional scale directive which recognises the importance of plankton communities in marine ecosystems; plankton data at the appropriate spatial, temporal and taxonomic scales are therefore required for implementation. The Continuous Plankton Recorder (CPR) survey is a multidecadal, North Atlantic basin scale programme which routinely records approximately 300 phytoplankton taxa. Because of these attributes, the survey plays a key role in the implementation of the MSFD and the assessment of GES in the Northeast Atlantic region. This paper addresses the role of the CPR's phytoplankton time-series in delivering GES through the development and informing of MSFD indicators, the setting of targets against a background of climate change and the provision of supporting information used to interpret change in non-plankton indicators. We also discuss CPR data in the context of other phytoplankton data types that may contribute to GES, as well as explore future possibilities for the use of new and innovative applications of CPR phytoplankton datasets in delivering GES. Efforts must be made to preserve long-term time series, such as the CPR, which supply vital ecological information used to informed evidence-based environmental policy.

Linking environmental variables with regional- scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and >1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide.

Linking environmental variables with regional- scale variability in ecological structure and standing stock of carbon within UK kelp forests

Kelp forests represent some of the most productive and diverse habitats on Earth. Understanding drivers of ecological patterns at large spatial scales is critical for effective management and conservation of marine habitats. We surveyed kelp forests dominated by Laminaria hyperborea (Gunnerus) Foslie 1884 across 9° latitude and >1000 km of coastline and measured a number of physical parameters at multiple scales to link ecological structure and standing stock of carbon with environmental variables. Kelp density, biomass, morphology and age were generally greater in exposed sites within regions, highlighting the importance of wave exposure in structuring L. hyperborea populations. At the regional scale, wave-exposed kelp canopies in the cooler regions (the north and west of Scotland) were greater in biomass, height and age than in warmer regions (southwest Wales and England). The range and maximal values of estimated standing stock of carbon contained within kelp forests was greater than in historical studies, suggesting that this ecosystem property may have been previously undervalued. Kelp canopy density was positively correlated with large-scale wave fetch and fine-scale water motion, whereas kelp canopy biomass and the standing stock of carbon were positively correlated with large-scale wave fetch and light levels and negatively correlated with temperature. As light availability and summer temperature were important drivers of kelp forest biomass, effective management of human activities that may affect coastal water quality is necessary to maintain ecosystem functioning, while increased temperatures related to anthropogenic climate change may impact the structure of kelp forests and the ecosystem services they provide.

Consequences of a simulated rapid ocean acidification event for benthic ecosystem processes and functions

Whilst the biological consequences of long-term, gradual changes in acidity associated with the oceanic uptake of atmospheric carbon dioxide (CO2) are increasingly studied, the potential effects of rapid acidification associated with a failure of sub-seabed carbon storage infrastructure have received less attention. This study investigates the effects of severe short-term (8 days) exposure to acidified seawater on infaunal mediation of ecosystem processes (bioirrigation and sediment particle redistribution) and functioning (nutrient concentrations). Following acidification, individuals of Amphiura filiformis exhibited emergent behaviour typical of a stress response, which resulted in altered bioturbation, but limited changes in nutrient cycling. Under acidified conditions, A. filiformis moved to shallower depths within the sediment and the variability in occupancy depth reduced considerably. This study indicated that rapid acidification events may not be lethal to benthic invertebrates, but may result in behavioural changes that could have longer-term implications for species survival, ecosystem structure and functioning.

Impact of elevated levels of CO2 on animal mediated ecosystem function: The modification of sediment nutrient fluxes by burrowing urchins

A mesocosm experiment was conducted to quantify the relationships between the presence and body size of two burrowing heart urchins (Brissopsis lyrifera and Echinocardium cordatum) and rates of sediment nutrient flux. Furthermore, the impact of seawater acidification on these relationships was determined during this 40-day exposure experiment. Using carbon dioxide (CO2) gas, seawater was acidified to pHNBS 7.6, 7.2 or 6.8. Control treatments were maintained in natural seawater (pH8.0). Under normocapnic conditions, burrowing urchins were seen to reduce the sediment uptake of nitrite or nitrate whilst enhancing the release of silicate and phosphate. In acidified (hypercapnic) treatments, the biological control of biogeochemical cycles by urchins was significantly affected, probably through the combined impacts of high CO2 on nitrifying bacteria, benthic algae and urchin behaviour. This study highlights the importance of considering biological interactions when predicting the consequences of seawater acidification on ecosystem function.