Papers by Stephen Widdicombe
Potential impact of CCS leakage on marine communities

Marine Environmental Research, 2017
The processes and patterns seen in coastal benthic communities can be strongly influenced by the ... more The processes and patterns seen in coastal benthic communities can be strongly influenced by the overlying pelagic environmental conditions. Integrating long-term biological and environmental data (both benthic and pelagic) can give insight into the specific relationships between key benthic functional groups and natural temporal changes in the marine environment. The identity and abundance of amphipod species found at Station L4 (Western English Channel) were tracked for 7 years (2008-2014), whilst simultaneously, annual changes in phytoplankton biomass, water temperature, salinity and chlorophyll a concentration were also characterized. The main species were persistent and showed little variability along the study period. Overall, however, there were significant changes in the structure of the whole community between sampling times, highlighting the importance of less numerically-dominant species in driving temporal variability. Surprisingly, the current study did not detect a significant influence of the phytoplankton biomass on benthic amphipod dynamics. On the other hand, there was a clear and constant correlation between bottom water temperatures and amphipod abundance. This pattern is different from that observed in other detritivorous species at L4 (e.g. polychaeta), highlighting the complexity of benthic-pelagic coupling and the high variability of the response to pelagic conditions among different groups. As a result of the biogeographic position of the Western English Channel, the key role of amphipods in benthic communities, the influence of the temperature in their populations dynamics, as well as the solid baseline provided here and in previous studies, the monitoring of long-term amphipod dynamics in the English Channel could be a valuable tool to evaluate the biological effect of climate change over marine benthic communities.

Marine pollution bulletin, Jan 8, 2016
Regulations pertaining to carbon dioxide capture with offshore storage (CCS) require an understan... more Regulations pertaining to carbon dioxide capture with offshore storage (CCS) require an understanding of the potential localised environmental impacts and demonstrably suitable monitoring practices. This study uses a marine ecosystem model to examine a comprehensive range of hypothetical CO2 leakage scenarios, quantifying both impact and recovery time within the benthic system. Whilst significant mortalities and long recovery times were projected for the larger and longer term scenarios, shorter-term or low level exposures lead to reduced projected impacts. This suggests that efficient monitoring and leak mitigation strategies, coupled with appropriate selection of storage sites can effectively limit concerns regarding localised environmental impacts from CCS. The feedbacks and interactions between physiological and ecological responses simulated reveal that benthic responses to CO2 leakage could be complex. This type of modelling investigation can aid the understanding of impact po...

ICES Journal of Marine Science, 2015
Ocean acidification, the result of increased dissolution of carbon dioxide (CO2) in seawater, is ... more Ocean acidification, the result of increased dissolution of carbon dioxide (CO2) in seawater, is a leading subject of current research. The effects of acidification on non-calcifying macroalgae are, however, still unclear. The current study reports two 1-month studies using two different macroalgae, the red alga Palmaria palmata (Rhodophyta) and the kelp Saccharina latissima (Phaeophyta), exposed to control (pHNBS = ∼8.04) and increased (pHNBS = ∼7.82) levels of CO2-induced seawater acidification. The impacts of both increased acidification and time of exposure on net primary production (NPP), respiration (R), dimethylsulphoniopropionate (DMSP) concentrations, and algal growth have been assessed. In P. palmata, although NPP significantly increased during the testing period, it significantly decreased with acidification, whereas R showed a significant decrease with acidification only. S. latissima significantly increased NPP with acidification but not with time, and significantly inc...

Marine Ecology Progress Series, 2014
The abundance of ammonia-oxidising bacterial (AOB) and ammonia-oxidising archaeal (AOA) (amoA) ge... more The abundance of ammonia-oxidising bacterial (AOB) and ammonia-oxidising archaeal (AOA) (amoA) genes and ammonia oxidation rates were compared bimonthly from July 2008 to May 2011 in 4 contrasting coastal sediments in the western English Channel. Despite a higher abundance of AOA amoA genes within all sediments and at all time-points, rates of ammonia oxidation correlated with AOB and not AOA amoA gene abundance. Sediment type was a major factor in determining both AOB amoA gene abundance and AOB community structure, possibly due to deeper oxygen penetration into the sandier sediments, increasing the area available for ammonia oxidation. Decreases in AOB amoA gene abundance were evident during summer and autumn, with maximum abundance and ammonia oxidation rates occurring in winter and early spring. PCR-DGGE of AOB amoA genes indicated that no seasonal changes to community composition occurred; however, a gradual movement in community composition occurred at 3 of the sites studied. The lack of correlation between AOA amoA gene abundance and ammonium oxidation rates, or any other environmental variable measured, may be related to the higher spatial variation amongst measurements, obscuring temporal trends, or the bimonthly sampling, which may have been too infrequent to capture temporal variability in the deposition of fresh organic matter. Alternatively, AOA may respond to changing substrate concentrations by an increase or decrease in transcript rather than gene abundance.
Progress in Oceanography, 2015
Highlights: 1) The response of a benthic bacterial community to a phytoplankton bloom was studied... more Highlights: 1) The response of a benthic bacterial community to a phytoplankton bloom was studied. 2) Phytoplankton material was heavily degraded only when it reached the sediment. 3) The abundance of benthic bacteria and chlorophyll were correlated. 4) Bacterial community composition shifted rapidly throughout the bloom. 5) This was due to an increase in relative abundance of members of the Flavobacteria.

ICES Journal of Marine Science, 2015
Increased atmospheric CO2 concentration is leading to changes in the carbonate chemistry and the ... more Increased atmospheric CO2 concentration is leading to changes in the carbonate chemistry and the temperature of the ocean. The impact of these processes on marine organisms will depend on their ability to cope with those changes, particularly the maintenance of calcium carbonate structures. Both a laboratory experiment (long-term exposure to decreased pH and increased temperature) and collections of individuals from natural environments characterized by low pH levels (individuals from intertidal pools and around a CO2 seep) were here coupled to comprehensively study the impact of near-future conditions of pH and temperature on the mechanical properties of the skeleton of the euechinoid sea urchin Paracentrotus lividus. To assess skeletal mechanical properties, we characterized the fracture force, Young's modulus, second moment of area, material nanohardness, and specific Young's modulus of sea urchin test plates. None of these parameters were significantly affected by low pH...
Queiros etal 2014 SupplementaryInformation
Modelling ocean acidification impacts: from biological experiments to economic assessments and social impacts

International Journal of Greenhouse Gas Control, 2015
The response of the benthic microbial community to a controlled sub-seabed CO 2 leak was assessed... more The response of the benthic microbial community to a controlled sub-seabed CO 2 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 CO 2 exposure, after 14 and 36 days of CO 2 release, and 6, 20 and 90 days after the CO 2 release had ended). Changes to the active community of microphytobenthos and bacteria were also assessed before, during and after CO 2 release. Increases in the abundance of microbial 16S rRNA were detected after 14 days of CO 2 release and at a distance of 25 m from the epicentre. CO 2 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 CO 2 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.

Biogeosciences, 2014
Free-ocean CO 2 enrichment (FOCE) systems are designed to assess the impact of ocean acidificatio... more Free-ocean CO 2 enrichment (FOCE) systems are designed to assess the impact of ocean acidification on biological communities in situ for extended periods of time (weeks to months). They overcome some of the drawbacks of laboratory experiments and field observations by enabling (1) precise control of CO 2 enrichment by monitoring pH as an offset of ambient pH, (2) consideration of indirect effects such as those mediated through interspecific relationships and food webs, and (3) relatively long experiments with intact communities. Bringing perturbation experiments from the laboratory to the field is, however, extremely challenging. The main goal of this paper is to provide guidelines on the general design, engineering, and sensor options required to conduct FOCE experiments. Another goal is to introduce xFOCE, a community-led initiative to promote awareness, provide resources for in situ perturbation experiments, and build a user community. Present and existing FOCE systems are briefly described and examples of data collected presented. Future developments are also addressed as it is anticipated that the next generation of FOCE systems will include, in addition to pH, options for oxygen and/or temperature control. FOCE systems should become an important experimental approach for projecting the future response of marine ecosystems to environmental change.

Global change biology, 2015
Understanding long-term, ecosystem-level impacts of climate change is challenging because experim... more 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 interdisciplinary 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 parameterize 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, u...

Journal of Experimental Biology, 2014
The distribution patterns of many species in the intertidal zone are partly determined by their a... more The distribution patterns of many species in the intertidal zone are partly determined by their ability to survive and recover from tidal emersion. During emersion most crustaceans experience gill-collapse impairing gas-exchange. Such collapse generates a state of hypoxemia and a hypercapnia-induced respiratory acidosis, leading to hyperlactaemia and metabolic acidosis. However, how such physiological responses to emersion are modified by prior exposure to elevated CO2 and temperature combinations, indicative of future climate change scenarios, is not known. We therefore investigated key physiological responses of velvet swimming crabs, Necora puber, kept for 14 days at one of four pCO2/temperature treatments (400 μatm/10 °C, 1000 μatm/10 °C, 400 μatm/15 °C, 1000 μatm/15 °C), to experimental emersion and recovery. Pre-exposure to elevated pCO2 and temperature increased pre-emersion bicarbonate ion concentrations [HCO3-], increasing resistance to short periods of emersion (90 min). H...

The ISME Journal, 2010
Bioturbation is a key process in coastal sediments, influencing microbially driven cycling of nut... more Bioturbation is a key process in coastal sediments, influencing microbially driven cycling of nutrients as well as the physical characteristics of the sediment. However, little is known about the distribution, diversity and function of the microbial communities that inhabit the burrows of infaunal macroorganisms. In this study, terminal-restriction fragment length polymorphism analysis was used to investigate variation in the structure of bacterial communities in sediment bioturbated by the burrowing shrimp Upogebia deltaura or Callianassa subterranea. Analyses of 229 sediment samples revealed significant differences between bacterial communities inhabiting shrimp burrows and those inhabiting ambient surface and subsurface sediments. Bacterial communities in burrows from both shrimp species were more similar to those in surface-ambient than subsurface-ambient sediment (R ¼ 0.258, Po0.001). The presence of shrimp was also associated with changes in bacterial community structure in surrounding surface sediment, when compared with sediments uninhabited by shrimp. Bacterial community structure varied with burrow depth, and also between individual burrows, suggesting that the shrimp's burrow construction, irrigation and maintenance behaviour affect the distribution of bacteria within shrimp burrows. Subsequent sequence analysis of bacterial 16S rRNA genes from surface sediments revealed differences in the relative abundance of bacterial taxa between shrimp-inhabited and uninhabited sediments; shrimp-inhabited sediment contained a higher proportion of proteobacterial sequences, including in particular a twofold increase in Gammaproteobacteria. Chao1 and ACE diversity estimates showed that taxon richness within surface bacterial communities in shrimp-inhabited sediment was at least threefold higher than that in uninhabited sediment. This study shows that bioturbation can result in significant structural and compositional changes in sediment bacterial communities, increasing bacterial diversity in surface sediments and resulting in distinct bacterial communities even at depth within the burrow. In an area of high macrofaunal abundance, this could lead to alterations in the microbial transformations of important nutrients at the sediment-water interface.

Philosophical Transactions of the Royal Society B: Biological Sciences, 2013
Ocean acidification (OA), caused by the dissolution of increasing concentrations of atmospheric c... more Ocean acidification (OA), caused by the dissolution of increasing concentrations of atmospheric carbon dioxide (CO 2 ) in seawater, is projected to cause significant changes to marine ecology and biogeochemistry. Potential impacts on the microbially driven cycling of nitrogen are of particular concern. Specifically, under seawater pH levels approximating future OA scenarios, rates of ammonia oxidation (the rate-limiting first step of the nitrification pathway) have been shown to dramatically decrease in seawater, but not in underlying sediments. However, no prior study has considered the interactive effects of microbial ammonia oxidation and macrofaunal bioturbation activity, which can enhance nitrogen transformation rates. Using experimental mesocosms, we investigated the responses to OA of ammonia oxidizing microorganisms inhabiting surface sediments and sediments within burrow walls of the mud shrimp Upogebia deltaura . Seawater was acidified to one of four target pH values (pH T...

Philosophical Transactions of the Royal Society B: Biological Sciences, 2013
Climate change may cause ecosystems to become trophically restructured as a result of primary pro... more Climate change may cause ecosystems to become trophically restructured as a result of primary producers and consumers responding differently to increasing CO 2 and temperature. This study used an integrative approach using a controlled microcosm experiment to investigate the combined effects of CO 2 and temperature on key components of the intertidal system in the UK, biofilms and their consumers ( Littorina littorea ). In addition, to identify whether pre-exposure to experimental conditions can alter experimental outcomes we explicitly tested for differential effects on L. littorea pre-exposed to experimental conditions for two weeks and five months. In contrast to predictions based on metabolic theory, the combination of elevated temperature and CO 2 over a five-week period caused a decrease in the amount of primary productivity consumed by grazers, while the abundance of biofilms increased. However, long-term pre-exposure to experimental conditions (five months) altered this effe...
Predicted levels of future ocean acidification and temperature rise could alter community structure and biodiversity in marine benthic communities
Oikos, 2011
... This community-based mesocosm study supports previ-ous suggestions, based on observations of ... more ... This community-based mesocosm study supports previ-ous suggestions, based on observations of direct physiological impacts, that ocean acidification induced changes in marine biodiversity will be driven by differential vulnerability within and between different taxonomical ...

Effects of anthropogenic seawater acidification on acid–base balance in the sea urchin Psammechinus miliaris
Marine Pollution Bulletin, 2007
The purple-tipped sea urchin, Psammechinus miliaris, was exposed to artificially acidified seawat... more The purple-tipped sea urchin, Psammechinus miliaris, was exposed to artificially acidified seawater treatments (pH(w) 6.16, 6.63 or 7.44) over a period of 8 days. Urchin mortality of 100% was observed at pH(w) 6.16 after 7 days and coincided with a pronounced hypercapnia in the coelomic fluid producing an irrecoverable acidosis. Coelomic fluid acid-base measures showed that an accumulation of CO(2) and a significant reduction in pH occurred in all treatments compared with controls. Bicarbonate buffering was employed in each case, reducing the resultant acidosis, but compensation was incomplete even under moderate environmental hypercapnia. Significant test dissolution was inferred from observable increases in the Mg(2+) concentration of the coelomic fluid under all pH treatments. We show that a chronic reduction of surface water pH to below 7.5 would be severely detrimental to the acid-base balance of this predominantly intertidal species; despite its ability to tolerate fluctuations in pCO(2) and pH in the rock pool environment. The absence of respiratory pigment (or any substantial protein in the coelomic fluid), a poor capacity for ionic regulation and dependency on a magnesium calcite test, make echinoids particularly vulnerable to anthropogenic acidification. Geological sequestration leaks may result in dramatic localised pH reductions, e.g. pH 5.8. P. miliaris is intolerant of pH 6.16 seawater and significant mortality is seen at pH 6.63.
The effect of chronic and acute low pH on the intracellular DMSP production and epithelial cell morphology of red coralline algae
Marine Biology Research, 2012
Abstract The release of dimethylsulphoniopropionate (DMSP) by marine algae has major impacts on t... more Abstract The release of dimethylsulphoniopropionate (DMSP) by marine algae has major impacts on the global sulphur cycle and may influence local climate through the formation of dimethylsulphide (DMS). However, the effect of global change on DMSP/DMS (DMS (P)) production by algae is not well understood. This study examined the effect of low pH on DMS (P) production and epithelial cell morphology of the free-living red coralline alga Lithothamnion glaciale. Three pH treatments were used in the 80-day experiment:(1) ...
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Papers by Stephen Widdicombe