Total alkalinity (TA) measurements in seawater are crucial for characterizing and monitoring the oceanic carbonate system. While international best practices and guidelines exist, the field still lacks widely available traceable reference materials and a well-established uncertainty budget of the measurement method. In this study, we applied key metrological principles – development of reference materials, inter-laboratory comparison and uncertainty quantification – to TA measurements. We developed two reference materials, including an artificial material with a rigorously characterized reference value and an associated uncertainty budget, being traceable to the International System of units (SI). These materials were tested in an inter-laboratory comparison involving five laboratories and demonstrated the applicability of the reference materials developed for quality control. Additionally, we established an uncertainty budget for the TA measurement method using two metrological approaches. The resulting expanded uncertainty was 5 µmol kg−1 (with a coverage factor k=2) in TA, approaching the 4 µmol kg−1 target set by the Global Ocean Acidification Observing Network for climate monitoring. These findings mark a significant step toward improving the quality and comparability of TA measurements, thereby strengthening long-term ocean carbonate system monitoring.

Seagrass meadows are natural carbon sinks, yet the effect of ocean acidification on their carbon burial capacity remains poorly understood. Here we investigated natural carbon dioxide vents in Ischia, Italy to assess how seawater pH influences carbon burial in an area dominated by the seagrass Posidonia oceanica . Organic carbon burial rates (mean ± standard error) between 1954 – 2021 were low under ambient conditions (1.5 ± 0.5 g m -2 yr -1 ) but increased sharply under acidified conditions (7 ± 1 g m -2 yr -1 ), reaching sevenfold higher values under extreme acidification (10 ± 3 g m -2 yr -1 ). Stable isotopes suggest that these patterns reflect changes in the relative contribution of seagrass, macroalgae, and epiphytes to buried carbon. These findings reveal that ocean acidification can substantially alter coastal carbon cycling, potentially through shifts in community composition, with important implications for understanding past and future feedbacks between seagrass ecosystems and the marine carbon cycle.

Ocean acidification (OA) driven by increasing atmospheric CO₂ is altering marine biodiversity. However, impacts of OA on ecosystem functioning at the community level, including calcification, primary production, and nutrient uptake, remain largely unknown. Here, we conducted community transplant experiments at natural CO2 vents to assess how declining pH affects marine community species composition, biomass, and key ecosystem processes over time. Our results indicate that community shifts caused by declining pH lead to decreased biomass and calcification rates, while photosynthesis and nutrient uptake rates increased. By leveraging OA field model systems and in situ measurements of ecosystem functioning, this study provides critical insights into how OA-induced biodiversity loss reshapes the structure and functioning of temperate marine coastal ecosystems.

Ocean acidification (OA) driven by increasing atmospheric CO 2 is altering marine biodiversity. However, impacts of OA on ecosystem functioning at the community level, including calcification, primary production and nutrient uptake, remain largely unknown. Here, we conducted community transplant experiments at natural CO 2 vents to assess how declining pH affects marine community species composition, biomass, and key ecosystem processes over time. Our results indicate that community shifts caused by declining pH lead to decreased biomass and calcification rates, while photosynthesis and nutrient uptake rates increased. By leveraging OA field model systems and in situ measurements of ecosystem functioning, this study provides critical insights into how OA-induced biodiversity loss reshapes the structure and functioning of temperate marine coastal ecosystems.

Abstract The Arctic region is experiencing rapid warming and an increase in the frequency and intensity of marine heatwaves (MHWs), posing an increasing risk to coastal ecosystems. Kelp communities in the Arctic are vital habitats that support biodiversity and resilience but are particularly vulnerable to increasing temperature anomalies. In this study, we conducted a mesocosm experiment over 23 days in summer to assess the effects of warming and MHWs on three kelp species ( Saccharina latissima , Laminaria digitata and Alaria esculenta ) and associated community members: coralline algae, snails, sea urchins and mussels. Three warming scenarios were tested, all of which were offset from a naturally variable control. The four conditions were a control (i.e. ambient seawater) which ranged from ~7.5 to 10°C (naturally variable), a constant high‐temperature treatment (+1.8°C from the control), a treatment simulating a single 13‐day heatwave with a maximum offset temperature of +2.8°C from the control, and a treatment simulating two consecutive 5‐day MHWs with a maximum offset of +3.9°C. For the three kelp species, net photosynthetic rate, maximum quantum yield, chlorophyll a content, carbon to nitrogen ratio and growth rate were investigated. For the associated species, only growth rates were evaluated. Laminaria digitata showed no significant changes in any of the measured parameters, suggesting a robust tolerance to heat stress. Conversely, the maximum quantum yield for S. latissima decreased in the two heatwaves treatment. In addition, A. esculenta displayed a decrease in chlorophyll a content and net photosynthesis over time in the two heatwaves treatment. The growth rates of coralline algae, mussels, snails and sea urchins were not affected by the treatments. Synthesis : These results suggest that Arctic kelp communities are fairly tolerant to warming and short‐term MHWs, despite the few minor effects on photosynthesis by some of the kelp species. We note that the future response of kelp communities must account for the potential indirect effects of climate change, such as predation and competition, along with any consequential effects of warming anomalies, as those presented here.

Total alkalinity (AT) and dissolved inorganic carbon (CT) in the oceans are important properties to understand the ocean carbon cycle and its link with global change (ocean carbon sinks and sources, ocean acidification) and ultimately to find carbon-based solutions or mitigation procedures (marine carbon removal). We present an extended database (SNAPO-CO2; Metzl et al., 2024c) with 24 700 new additional data for the period 2002 to 2023. The full database now includes more than 67 000 AT and CT observations along with basic ancillary data (time and space location, depth, temperature, and salinity) in various oceanic regions obtained since 1993 mainly in the framework of French research projects. This includes both surface and water columns data acquired in open oceans, coastal zones, rivers, the Mediterranean Sea, and either from time series stations or punctual cruises. Most AT and CT data in this synthesis were measured from discrete samples using the same closed-cell potentiometric titration calibrated with certified reference material, with an overall accuracy of ±4 µmol kg−1 for both AT and CT. The same technique was used on board for underway measurements during cruises conducted in the southern Indian and Southern oceans. The AT and CT data from these cruises are also added to this synthesis. The data are provided in one dataset for the global ocean (https://doi.org/10.17882/102337, Metzl et al., 2024c) that offers a direct use for regional or global purposes, e.g., AT–salinity relationships, long-term CT estimates, constraint and validation of diagnostics CT and AT reconstructed fields, ocean carbon and coupled climate–carbon models simulations, and data derived from Biogeochemical Argo (BGC-Argo) floats. These data can also be used to calculate pH, fugacity of CO2 (fCO2), and other carbon system properties to derive ocean acidification rates or air–sea CO2 fluxes.

Global environmental change drives diversity loss and shifts in community structure. A key challenge is to better understand the impacts on ecosystem function and to connect species and trait diversity of assemblages with ecosystem properties that are in turn linked to ecosystem functioning. Here we quantify shifts in species composition and trait diversity associated with ocean acidification (OA) by using field measurements at marine CO2 vent systems spanning four reef habitats across different depths in a temperate coastal ecosystem. We find that both species and trait diversity decreased, and that ecosystem properties (understood as the interplay between species, traits, and ecosystem function) shifted with acidification. Furthermore, shifts in trait categories such as autotrophs, filter feeders, herbivores, and habitat-forming species were habitat-specific, indicating that OA may produce divergent responses across habitats and depths. Combined, these findings reveal the importance of connecting species and trait diversity of marine benthic habitats with key ecosystem properties to anticipate the impacts of global environmental change. Our results also generate new insights on the predicted general and habitat-specific ecological consequences of OA.

Since 2004, the Service facility SNAPO-CO2 (Service National d’Analyse des Paramètres Océaniques du CO2) housed by the LOCEAN laboratory (Paris, France) has been in charge for the analysis of Total Alkalinity (AT) and Total dissolved inorganic carbon (CT) of seawater samples on a series of cruises or ships of opportunity conducted in different regions in the frame of French projects. More than 44000 observations are synthetized in this work. Sampling was performed either from CTD-Rosette casts (Niskin bottles) or collected from the ship’s seawater supply (intake at about 5m depth). After completion of each cruise, discrete samples were returned back at LOCEAN laboratory and stored in a dark room at 4 °C before analysis generally within 2-3 months after sampling (sometimes within a week). AT and CT were analyzed simultaneously by potentiometric titration using a closed cell (Edmond, 1970). Certified Reference Materials (CRMs) provided by Pr. A. Dickson (Scripps Institution of Oceanography, San Diego, USA) were used to calibrate the measurements. The same instrumentation was used for underway measurements during OISO cruises (https://doi.org/10.18142/228) and OISO AT-CT data for 1998-2018 in the South Indian Ocean added in this synthesis. The synthesis is organized in two files (one for Global ocean and the Coastal Zones, one for the Mediterranean Sea) with the same format: Cruise name, Ship name, day, month, year, hour, minute, second, latitude, longitude, depth, AT (µmol/kg), Flag-AT, CT (µmol/kg), Flag-CT, Temperature (°C), Flag-Temp, Salinity (PSU), Flag-Salinity, nsample/cruise, nsample on file, sampling method.

Arctic fjords are considered to be one of the ecosystems changing most rapidly in response to climate change. In the Svalbard archipelago, fjords are experiencing a shift in environmental conditions due to the Atlantification of Arctic waters and the retreat of sea-terminating glaciers. These environmental changes are predicted to facilitate expansion of large, brown macroalgae, into new ice-free regions. The potential resilience of macroalgal benthic communities in these fjord systems will depend on their response to combined pressures from freshening due to glacial melt, exposure to warmer waters, and increased turbidity from meltwater runoff which reduces light penetration. Current predictions, however, have a limited ability to elucidate the future impacts of multiple-drivers on macroalgal communities with respect to ecosystem function and biogeochemical cycling in Arctic fjords. To assess the impact of these combined future environmental changes on benthic productivity and resilience, we conducted a two-month mesocosm experiment exposing mixed kelp communities to three future conditions comprising increased temperature (+ 3.3 and + 5.3°C), seawater freshening by ∼ 3.0 and ∼ 5.0 units (i.e., salinity of 30 and 28, respectively), and decreased photosynthetically active radiation (PAR, - 25 and - 40 %). Exposure to these combined treatments resulted in non-significant differences in short-term productivity, and a tolerance of the photosynthetic capacity across the treatment conditions. We present the first robust estimates of mixed kelp community production in Kongsfjorden and place a median compensation irradiance of ∼12.5 mmol photons m$^−$$^2$ h$^−$$^1$ as the threshold for positive net community productivity. These results are discussed in the context of ecosystem productivity and biological tolerance of kelp communities in future Arctic fjord systems.

Since the beginning of the industrial revolution, atmospheric carbon dioxide (CO2) concentrations have risen steadily and have induced a decrease of the averaged surface ocean pH by 0.1 units, corresponding to an increase in ocean acidity of about 30 %. In addition to ocean warming, ocean acidification poses a tremendous challenge to some marine organisms, especially calcifiers. The need for long-term oceanic observations of pH and temperature is a key element to assess the vulnerability of marine communities and ecosystems to these pressures. Nearshore productive environments, where a large majority of shellfish farming activities are conducted, are known to present pH levels as well as amplitudes of daily and seasonal variations that are much larger than those observed in the open ocean. Yet, to date, there are very few coastal observation sites where these parameters are measured simultaneously and at high frequency. To bridge this gap, an observation network was initiated in 2021 in the framework of the CocoriCO2 project. Six sites were selected along the French Atlantic and Mediterranean coastlines based on their importance in terms of shellfish production and the presence of high- and low-frequency monitoring activities. At each site, autonomous pH sensors were deployed, both inside and outside shellfish production areas, next to high-frequency CTD (conductivity–temperature–depth) probes operated through two operating monitoring networks. pH sensors were set to an acquisition rate of 15 min, and discrete seawater samples were collected biweekly in order to control the quality of pH data (laboratory spectrophotometric measurements) as well as to measure total alkalinity and dissolved inorganic carbon concentrations for full characterization of the carbonate system. While this network has been up and running for more than 2 years, the acquired dataset has already revealed important differences in terms of pH variations between monitored sites related to the influence of diverse processes (freshwater inputs, tides, temperature, biological processes). Data are available at https://doi.org/10.17882/96982 (Petton et al., 2023a).

Abstract. The rapid environmental changes in aquatic systems as a result of anthropogenic forcings are creating a multitude of challenging conditions for organisms and communities. The need to better understand the interaction of environmental stressors now, and in the future, is fundamental to determining the response of ecosystems to these perturbations. This work describes an automated ex situ mesocosm perturbation system that can manipulate several variables of aquatic media in a controlled setting. This perturbation system was deployed in Kongsfjorden (Svalbard); within this system, ambient water from the fjord was heated and mixed with freshwater in a multifactorial design to investigate the response of mixed-kelp communities in mesocosms to projected future Arctic conditions. The system employed an automated dynamic offset scenario in which a nominal temperature increase was programmed as a set value above real-time ambient conditions in order to simulate future warming. A freshening component was applied in a similar manner: a decrease in salinity was coupled to track the temperature offset based on a temperature–salinity relationship in the fjord. The system functioned as an automated mixing manifold that adjusted flow rates of warmed and chilled ambient seawater, with unmanipulated ambient seawater and freshwater delivered as a single source of mixed media to individual mesocosms. These conditions were maintained via continuously measured temperature and salinity in 12 mesocosms (1 control and 3 treatments, all in triplicate) for 54 d. System regulation was robust, as median deviations from nominal conditions were < 0.15 for both temperature (∘C) and salinity across the three replicates per treatment. Regulation further improved during a second deployment that mimicked three marine heat wave scenarios in which a dynamic temperature regulation held median deviations to < 0.036 ∘C from the nominal value for all treatment conditions and replicates. This perturbation system has the potential to be implemented across a wide range of conditions to test single or multi-stressor drivers (e.g., increased temperature, freshening, and high CO2) while maintaining natural variability. The automated and independent control for each experimental unit (if desired) provides a large breadth of versatility with respect to experimental design.

Abstract. Total alkalinity (AT) and dissolved inorganic carbon (CT) in the oceans are important properties with respect to understanding the ocean carbon cycle and its link to global change (ocean carbon sinks and sources, ocean acidification) and ultimately finding carbon-based solutions or mitigation procedures (marine carbon removal). We present a database of more than 44 400 AT and CT observations along with basic ancillary data (spatiotemporal location, depth, temperature and salinity) from various ocean regions obtained, mainly in the framework of French projects, since 1993. This includes both surface and water column data acquired in the open ocean, coastal zones and in the Mediterranean Sea and either from time series or dedicated one-off cruises. Most AT and CT data in this synthesis were measured from discrete samples using the same closed-cell potentiometric titration calibrated with Certified Reference Material, with an overall accuracy of ±4 µmol kg−1 for both AT and CT. The data are provided in two separate datasets – for the Global Ocean and the Mediterranean Sea (https://doi.org/10.17882/95414, Metzl et al., 2023), respectively – that offer a direct use for regional or global purposes, e.g., AT–salinity relationships, long-term CT estimates, and constraint and validation of diagnostic CT and AT reconstructed fields or ocean carbon and coupled climate–carbon models simulations as well as data derived from Biogeochemical-Argo (BGC-Argo) floats. When associated with other properties, these data can also be used to calculate pH, the fugacity of CO2 (fCO2) and other carbon system properties to derive ocean acidification rates or air–sea CO2 fluxes.

Abstract Fjord systems in the Norwegian Arctic are experiencing an increasing frequency and magnitude of marine heatwaves. These episodic heat stress events can have varying degrees of acute impacts on primary production and nutrient uptake of mixed kelp communities, as well as modifying the biogeochemical cycling in nearshore systems where vast areas of kelp create structural habitat. To assess the impact of future marine heatwaves on kelp communities, we conducted a 23 day mesocosm experiment exposing mixed kelp communities to warming and heatwave scenarios projected for the year 2100. Three treatments were considered: a constant warming (+1.8°C from the control), a medium magnitude and long duration heatwave event (+2.8°C from the control for 13 days), and two short‐term, more intense, heatwaves(5 day long scenarios with temperature peaks at +3.9°C from the control). The results show that both marine heatwave treatments reduced net community production, whereas the constant warm temperature treatment displayed no difference from the control. The long marine heatwave scenario resulted in reduced accumulated net community production, indicating that prolonged exposure had a greater severity than two high magnitude, short‐term heatwave events. We estimated an 11°C temperature threshold at which negative effects to primary production appeared present. We highlight that marine heatwaves can induce sublethal effects on kelp communities by depressing net community production. These results are placed in the context of potential physiological resilience of kelp communities and implications of reduced net community production to future Arctic fjord environmental conditions.

Abstract. The Arctic is projected to warm by 2 to 5 °C by the end of the century. Warming causes melting of glaciers, shrinking of the areas covered by sea ice, and increased terrestrial runoff from snowfields and permafrost thawing. Warming, decreasing coastal underwater irradiance, and lower salinity are potentially threatening polar marine organisms, including kelps, that are key species of hard-bottom shallow communities. The present study investigates the physiological responses of four kelp species (Alaria esculenta, Laminaria digitata, Saccharina latissima, and Hedophyllum nigripes) to these environmental changes through a perturbation experiment in ex situ mesocosms. Kelps were exposed for 6 weeks to four experimental treatments: an unmanipulated control; a warming condition under the CO2 emission scenario SSP5-8.5; and two multifactorial conditions combining warming, low salinity, and low irradiance reproducing the future coastal Arctic exposed to terrestrial runoff under two CO2 emission scenarios (SSP2-4.5 and SSP5-8.5). The physiological effects on A. esculenta, L. digitata, and S. latissima were investigated, and gene expression patterns of S. latissima and H. nigripes were analyzed. Across all species and experimental treatments, growth rates were similar, underlying the acclimation potential of these species to future Arctic conditions. Specimens of A. esculenta increased their chlorophyll a content when exposed to low irradiance conditions, suggesting that they may be resilient to an increase in glacier and river runoff with the potential to become more dominant at greater depths. S. latissima showed a lower carbon : nitrogen (C : N) ratio under the SSP5-8.5 multifactorial conditions' treatment, suggesting tolerance to coastal erosion and permafrost thawing. In contrast, L. digitata showed no response to the conditions tested on any of the investigated physiological parameters. The down-regulation of genes coding for heat-shock proteins in H. nigripes and S. latissima underscores their ability to acclimate to heat stress, which portrays temperature as a key influencing factor. Based on these results, it is expected that kelp communities will undergo changes in species composition that will vary at local scale as a function of the changes in environmental drivers.

The objective of the TONGA oceanographic expedition was to study the control of productivity and carbon sequestration by micronutrients of shallow hydrothermal origin in the Western Tropical South Pacific (WTSP) Ocean. The 37-day oceanographic survey took place on board the R/V L’Atalante in 2019 between Oct. 31 to Dec. 6 (Nouméa-Nouméa). Over a large area of the WTSP the team acquired numerous results on both the entire water column (up to the sediment) and the atmosphere. Specific task are represented on figure 1: (task 1) to characterize chemically and optically shallow hydrothermal fluids and to compare the source from below (shallow hydrothermal fluids) with the source from above (atmospheric deposition); (task 2) to quantify the dynamical dispersion of the fluids at small and regional scale; (task 3) to investigate the impact of the shallow hydrothermal sources on the biological activity and diversity, and the feedback to the atmosphere via the oceanic emissions of primary and secondary aerosols. (Task 4) to communicate about the campaign (see for example our Tweeter account (https://twitter.com/tongaproject) and the movie (26’) both in French (https://www.youtube.com/watch?v=e5kAd0i6Dck) and English (https://www.youtube.com/watch?v=UeABf-cVR-k). A long west to east (up to the blue waters of the gyre) transect allowed to characterize the different biogeochemical provinces crossed and a focus in the region of the Lau Basin allowed to investigate the impact of shallow hydrothermal sources. A series of short and long stations allowed to fully characterize the stocks and the fluxes in the different provinces. Short-term (up to 10 days) processes studies have been conducted (drifting moorings and minicosms experiments). Part of these results will feed into important modeling work. A fixed mooring line launched at the end of the campaign and recovered in Nov. 2020 as well as the 7 ARGO floats and 20 drifting buoys that were dropped during the campaign provide a broader temporal context of the acquisitions done during the campaign. An important focus of the campaign was the trace metal characterization of the entire water column. For this, TONGA has been labeled by the international program GEOTRACES (https://www.geotraces.org/). The impact on biological communities of fluids is supported by the international IMBER program (https://imber.info/). The TONGA project is also part of the LEFE program (funding by LEFE-CYBER and LEFE-GMMC), the ANR (Appel à projets génériques) and the Fondation A-MIDeX of the Aix-Marseille Université.

Abstract. The rapid environmental changes in aquatic systems as a result of anthropogenic forcings are creating a multitude of challenging conditions for organisms and communities. The need to better understand the interaction of environmental stressors now, and in the future, is fundamental to determining the response of ecosystems to these perturbations. This work describes an in situ mesocosm perturbation system that can manipulate aquatic media in a controlled setting on land. The employed system manipulated ambient water from Kongsfjorden, (Svalbard) by increasing temperature and freshening the seawater to investigate the response of mixed kelp communities to projected future Arctic conditions. This system manipulated temperature and salinity in real-time as an offset from incoming ambient seawater to conditions simulating future Arctic fjords. The system adjusted flow rates and mixing regimes of chilled, heated, ambient seawater, and freshwater, based on continuously measured conditions in a total of 12 mesocosms (1 ambient-control and 3 treatments, all in triplicates) for 54 days. System regulation was robust as median deviations from setpoint conditions were < 0.15 for both temperature (°C) and salinity across the 3 replicates per treatment. The implementation of this system has a wide range of versatility and can be deployed in a range of conditions to test single or multi-stressor conditions while maintaining natural variability.

Abstract. The Arctic Ocean is subject to high rates of ocean warming and acidification, with critical implications for marine organisms as well as ecosystems and the services they provide. Carbonate system data in the Arctic realm are spotty in space and time, and, until recently, there was no time-series station measuring the carbonate chemistry at high frequency in this region, particularly in coastal waters. We report here on the first high-frequency (1 h), multi-year (5 years) dataset of salinity, temperature, CO2 partial pressure (pCO2) and pH at a coastal site (bottom depth of 12 m) in a high-Arctic fjord (Kongsfjorden, Svalbard). Discrete measurements of dissolved inorganic carbon and total alkalinity were also performed. We show that (1) the choice of formulations for calculating the dissociation constants of the carbonic acid remains unsettled for polar waters, (2) the water column is generally somewhat stratified despite the shallow depth, (3) the saturation state of calcium carbonate is subject to large seasonal changes but never reaches undersaturation (Ωa ranges between 1.4 and 3.0) and (4) pCO2 is lower than atmospheric CO2 at all seasons, making this site a sink for atmospheric CO2 (−9 to −16.8 molCO2m-2yr-1, depending on the parameterisation of the gas transfer velocity). Data are available on PANGAEA: https://doi.org/10.1594/PANGAEA.960131 (Gattuso et al., 2023a).

Reference materials, as well as inter-laboratory comparisons (ILCs), are two essential metrology principles allowing the validation and evaluation of a measurement method. This paper describes the use of a reference material for seawater pH(T) measurements in an inter-laboratory comparison carried out with the French coastal observation service SOMLIT, and laboratories within the project CocoriCO(2). The interpretation of the ILC results demonstrated the capability of the spectrophotometric measurement method to provide compatible results with the reference value. The ILC data also allowed the estimation of an uncertainty budget of the spectrophotometric pH(T) measurement method following the ISO standard 21748 guidelines. The estimated standard uncertainty of the method is of 0.0046 in terms of pH(T). This level of uncertainty doesn't meet the requirements of the "climate" quality objective of the Global Ocean Acidification Observing Network for the monitoring of ocean acidification. However, some recommendations that may allow reducing this uncertainty are proposed in the paper.

Marine forests are shrinking globally due to several anthropogenic impacts including climate change. Forest-forming macroalgae, such as Cystoseira s.l. species, can be particularly sensitive to environmental conditions (e.g. temperature increase, pollution or sedimentation), especially during early life stages. However, not much is known about their response to the interactive effects of ocean warming (OW) and acidification (OA). These drivers can also affect the performance and survival of crustose coralline algae, which are associated understory species likely playing a role in the recruitment of later successional species such as forest-forming macroalgae. We tested the interactive effects of elevated temperature, low pH and species facilitation on the recruitment of Cystoseira compressa. We demonstrate that the interactive effects of OW and OA negatively affect the recruitment of C. compressa and its associated coralline algae Neogoniolithon brassica-florida. The density of recruits was lower under the combinations OW and OA, while the size was negatively affected by the temperature increase but positively affected by the low pH. The results from this study show that the interactive effects of climate change and the presence of crustose coralline algae can have a negative impact on the recruitment of Cystoseira s.l. species. While new restoration techniques recently opened the door to marine forest restoration, our results show that the interactions of multiple drivers and species interactions have to be considered to achieve long-term population sustainability.

Abstract. Although atmospheric dust fluxes from arid as well as human-impacted areas represent a significant source of nutrients to surface waters of the Mediterranean Sea, studies focusing on the evolution of the metabolic balance of the plankton community following a dust deposition event are scarce, and none were conducted in the context of projected future levels of temperature and pH. Moreover, most of the experiments took place in coastal areas. In the framework of the PEACETIME project, three dust-addition perturbation experiments were conducted in 300 L tanks filled with surface seawater collected in the Tyrrhenian Sea (TYR), Ionian Sea (ION) and Algerian basin (FAST) on board the R/V Pourquoi Pas? in late spring 2017. For each experiment, six tanks were used to follow the evolution of chemical and biological stocks, biological activity and particle export. The impacts of a dust deposition event simulated at their surface were followed under present environmental conditions and under a realistic climate change scenario for 2100 (ca. +3 ∘C and −0.3 pH units). The tested waters were all typical of stratified oligotrophic conditions encountered in the open Mediterranean Sea at this period of the year, with low rates of primary production and a metabolic balance towards net heterotrophy. The release of nutrients after dust seeding had very contrasting impacts on the metabolism of the communities, depending on the station investigated. At TYR, the release of new nutrients was followed by a negative impact on both particulate and dissolved 14C-based production rates, while heterotrophic bacterial production strongly increased, driving the community to an even more heterotrophic state. At ION and FAST, the efficiency of organic matter export due to mineral/organic aggregation processes was lower than at TYR and likely related to a lower quantity/age of dissolved organic matter present at the time of the seeding and a smaller production of DOM following dust addition. This was also reflected by lower initial concentrations in transparent exopolymer particles (TEPs) and a lower increase in TEP concentrations following the dust addition, as compared to TYR. At ION and FAST, both the autotrophic and heterotrophic community benefited from dust addition, with a stronger relative increase in autotrophic processes observed at FAST. Our study showed that the potential positive impact of dust deposition on primary production depends on the initial composition and metabolic state of the investigated community. This impact is constrained by the quantity of nutrients added in order to sustain both the fast response of heterotrophic prokaryotes and the delayed one of primary producers. Finally, under future environmental conditions, heterotrophic metabolism was overall more impacted than primary production, with the consequence that all integrated net community production rates decreased with no detectable impact on carbon export, therefore reducing the capacity of surface waters to sequester anthropogenic CO2.

In low-nutrient low-chlorophyll areas, such as the Mediterranean Sea, atmospheric fluxes represent a considerable external source of nutrients likely supporting primary production, especially during periods of stratification. These areas are expected to expand in the future due to lower nutrient supply from sub-surface waters caused by climate-driven enhanced stratification, likely further increasing the role of atmospheric deposition as a source of new nutrients to surface waters. Whether plankton communities will react differently to dust deposition in a warmer and acidified environment remains; however, an open question. The potential impact of dust deposition both in present and future climate conditions was investigated in three perturbation experiments in the open Mediterranean Sea. Climate reactors (300 L) were filled with surface water collected in the Tyrrhenian Sea, Ionian Sea and in the Algerian basin during a cruise conducted in the frame of the PEACETIME project in May–June 2017. The experiments comprised two unmodified control tanks, two tanks enriched with a Saharan dust analogue and two tanks enriched with the dust analogue and maintained under warmer (+3 ∘C) and acidified (−0.3 pH unit) conditions. Samples for the analysis of an extensive number of biogeochemical parameters and processes were taken over the duration (3–4 d) of the experiments. Dust addition led to a rapid release of nitrate and phosphate, however, nitrate inputs were much higher than phosphate. Our results showed that the impacts of Saharan dust deposition in three different basins of the open northwestern Mediterranean Sea are at least as strong as those observed previously, all performed in coastal waters. The effects of dust deposition on biological stocks were different for the three investigated stations and could not be attributed to differences in their degree of oligotrophy but rather to the initial metabolic state of the community. Ocean acidification and warming did not drastically modify the composition of the autotrophic assemblage, with all groups positively impacted by warming and acidification. Although autotrophic biomass was more positively impacted than heterotrophic biomass under future environmental conditions, a stronger impact of warming and acidification on mineralization processes suggests a decreased capacity of Mediterranean surface plankton communities to sequester atmospheric CO2 following the deposition of atmospheric particles.

Cabled coastal observatories are often seen as future-oriented marine technology that enables science to conduct observational and experimental studies under water year-round, independent of physical accessibility to the target area. Additionally, the availability of (unrestricted) electricity and an Internet connection under water allows the operation of complex experimental setups and sensor systems for longer periods of time, thus creating a kind of laboratory beneath the water. After successful operation for several decades in the terrestrial and atmospheric research field, remote controlled observatory technology finally also enables marine scientists to take advantage of the rapidly developing communication technology. The continuous operation of two cabled observatories in the southern North Sea and off the Svalbard coast since 2012 shows that even highly complex sensor systems, such as stereo-optical cameras, video plankton recorders or systems for measuring the marine carbonate system, can be successfully operated remotely year-round facilitating continuous scientific access to areas that are difficult to reach, such as the polar seas or the North Sea. Experience also shows, however, that the challenges of operating a cabled coastal observatory go far beyond the provision of electricity and network connection under water. In this manuscript, the essential developmental stages of the "COSYNA Shallow Water Underwater Node" system are presented, and the difficulties and solutions that have arisen in the course of operation since 2012 are addressed with regard to technical, organizational and scientific aspects.

Coastal time series of ocean carbonate chemistry are critical for understanding how global anthropogenic change manifests in near-shore ecosystems. Yet, they are few and have low temporal resolution. At the time series station Point B in the northwestern Mediterranean Sea, seawater was sampled weekly from 2007 through 2015, at 1 and 50 m, and analyzed for total dissolved inorganic carbon (C T) and total alkalinity (A T). Parameters of the carbonate system such as pH (pH T , total hydrogen ion scale) were calculated and a deconvolution analysis was performed to identify drivers of change. The rate of surface ocean acidification was −0.0028 ± 0.0003 units pH T yr −1. This rate is larger than previously identified open-ocean trends due to rapid warming that occurred over the study period (0.072 ± 0.022 • C yr −1). The total pH T change over the study period was of similar magnitude as the diel pH T variability at this site. The acidification trend can be attributed to atmospheric carbon dioxide (CO 2) forcing (59 %, 2.08 ± 0.01 ppm CO 2 yr −1) and warming (41 %). Similar trends were observed at 50 m but rates were generally slower. At 1 m depth, the increase in atmospheric CO 2 accounted for approximately 40 % of the observed increase in C T (2.97 ± 0.20 µmol kg −1 yr −1). The remaining increase in C T may have been driven by the same unidentified process that caused an increase in A T (2.08 ± 0.19 µmol kg −1 yr −1). Based on the analysis of monthly trends, synchronous increases in C T and A T were fastest in the spring–summer transition. The driving process of the interannual increase in A T has a seasonal and shallow component, which may indicate riverine or groundwater influence. This study exemplifies the importance of understanding changes in coastal carbonate chemistry through the lens of biogeochemical cycling at the land–sea interface. This is the first coastal acidification time series providing multi-year data at high temporal resolution. The data confirm rapid warming in the Mediterranean Sea and demonstrate coastal acidification with a synchronous increase in total alkalinity.

There is a growing international interest in studying the effects of ocean acidification on plankton communities that play a major role in the global carbon cycle and in the consumption of atmospheric CO2 via the so-called biological pump. Recently, several mesocosm experiments reported on the effect of ocean acidification on marine plankton communities, although the majority were performed in eutrophic conditions or following nutrient addition. The objective of the present study was to perform two mesocosm experiments in the oligo-to meso-trophic Northwestern Mediterranean Sea during two seasons with contrasting environmental conditions: in summer 2012 in the Bay of Calvi (Corsica, France) and in winter 2013 in the Bay of Villefranche (France). This paper describes the objectives of these experiments, the study sites, the experimental set-up and the environmental and experimental conditions during the two experiments. The 20-day experiment in the Bay of Calvi was undoubtedly representative of summer conditions in the Northwestern Mediterranean Sea with low nutrient and chlorophyll a concentrations, warm waters and high surface solar irradiance. In contrast, the winter experiment, which was reduced to 12 days because of bad weather conditions, failed to reproduce the mesotrophic conditions typical of the wintertime in this area. Indeed, a rapid increase in phytoplankton biomass during the acidification phase led to a strong decrease in nitrate concentrations and an unrealistic N and P co-limitation at this period of the year. An overview of the 11 papers related to this study and published in this special issue is provided.

Alterations in seagrass epiphytic communities are expected under future ocean acidification conditions, yet this hypothesis has been little tested in situ. A Free Ocean Carbon Dioxide Enrichment system was used to lower pH by a ~0.3 unit offset within a partially enclosed portion (1.7 m3) of a Posidonia oceanica meadow (11 m depth) between June 21 and November 3, 2014. Leaf epiphytic community composition (% cover) and bulk epiphytic mineralogy were compared every 4 weeks within three treatments, located in the same meadow: a pH-manipulated (experimental enclosure) and a control enclosure, as well as a nearby ambient area. Percent coverage of invertebrate calcifiers and crustose coralline algae (CCA) did not appear to be affected by the lowered pH. Furthermore, fleshy algae did not proliferate at lowered pH. Only Foraminifera, which covered less than 3% of leaf surfaces, declined in manner consistent with ocean acidification predictions. Bulk epiphytic magnesium carbonate composition was similar between treatments and percentage of magnesium appeared to increase from summer to autumn. CCA did not exhibit any visible skeleton dissolution or mineral alteration at lowered pH and carbonate saturation state. Negative impacts from ocean acidification on P. oceanica epiphytic communities were smaller than expected. Epiphytic calcifiers were possibly protected from the pH treatment due to host plant photosynthesis inside the enclosure where water flow is slowed. The more positive outcome than expected suggests that calcareous members of epiphytic communities may find refuge in some conditions and be resilient to environmentally relevant changes in carbonate chemistry.

Alterations to colonization or early post-settlement stages may cause the reorganization of communities under future ocean acidification conditions. Yet, this hypothesis has been little tested by in situ pH manipulation. A Free Ocean Carbon Dioxide Enrichment (FOCE) system was used to lower pH by a ~ 0.3 unit offset within a partially enclosed portion (1.7 m3) of a Posidonia oceanica seagrass meadow (11 m depth) between 21 June and 3 November 2014. Epibiont colonization and early post settlement stages were assessed within the FOCE setup, as part of the larger community-level study, to better understand the outcome for a multispecies assemblage and the ecological processes that result in reported community shifts under altered carbonate chemistry. Two types of artificial collectors (tiles and scourers) were placed within three treatments: a pH-manipulated enclosure, an un-manipulated control enclosure, and an open plot in the ambient meadow. Tiles and scourers were collected after one to four months. Additionally, to see whether the outcome differed for communities in a later successional stage, previously settled scourer-collectors were also placed in the same three treatments. Enclosures acted to reduce settlement and migrant colonization. Scourers deployed for one to four months within the open-plot contained a community assemblage that could be distinguished from the assemblages within the enclosures. However, a comparison of enclosure assemblages on tiles showed evidence of a pH effect. There was lowered coverage of crustose coralline algae and fewer calcareous tube-forming polychaetes (Spirorbis sp. and Spirobranchus sp.) on tiles placed in the pH-manipulated enclosure compared to the un-manipulated enclosure. For assemblages in scourer collectors, shared and common taxa, in all treatments, were invertebrate polychaetes Psamathe fusca, Sphaerosyllis sp., Chrysopetalum sp., arthropods Harpacticoida, and Amphipoda, and the juvenile bivalve Lyonsia sp. Similar organism composition and abundance, as well as taxonomic richness and evenness, were found in scourers from both enclosures. Pre-settled scourers contained greater numbers of individuals and more calcified members, but the assemblage, as well as the growth rate of a juvenile bivalve Lyonsia sp., appeared unaffected by a two-month exposure to lowered pH and calcium carbonate saturation state. Results from this case study support the hypothesis that early stages of specific calcifiers (crustose coralline algae and calcareous tube-forming polychaetes) are sensitive to near future ocean acidification conditions yet suggest that negative effects on sessile micro-invertebrate assemblages will be minimal.

Seagrass is expected to benefit from increased carbon availability under future ocean acidification. This hypothesis has been little tested by in situ manipulation. To test for ocean acidification effects on seagrass meadows under controlled CO2/pH conditions, we used a Free Ocean Carbon Dioxide Enrichment (FOCE) system which allows for the manipulation of pH as continuous offset from ambient. It was deployed in a Posidonia oceanica meadow at 11 m depth in the Northwestern Mediterranean Sea. It consisted of two benthic enclosures, an experimental and a control unit both 1.7 m3, and an additional reference plot in the ambient environment (2 m2) to account for structural artifacts. The meadow was monitored from April to November 2014. The pH of the experimental enclosure was lowered by 0.26 pH units for the second half of the 8-month study. The greatest magnitude of change in P. oceanica leaf biometrics, photosynthesis, and leaf growth accompanied seasonal changes recorded in the environment and values were similar between the two enclosures. Leaf thickness may change in response to lower pH but this requires further testing. Results are congruent with other short-term and natural studies that have investigated the response of P. oceanica over a wide range of pH. They suggest any benefit from ocean acidification, over the next century (at a pH of ∼ 7.7 on the total scale), on Posidonia physiology and growth may be minimal and difficult to detect without increased replication or longer experimental duration. The limited stimulation, which did not surpass any enclosure or seasonal effect, casts doubts on speculations that elevated CO2 would confer resistance to thermal stress and increase the buffering capacity of meadows.

The energetically costly transition from free-swimming larvae to benthic life stage and maintenance of a calcareous structure can make calcifying marine invertebrates vulnerable to ocean acidification. The first goal of this study was to evaluate the impacts of ocean acidification on calcified tube growth for two Serpulidae polychaete worms. Spirorbis sp. and Spirobranchus triqueter were collected at 11 m depth from the Northwest Mediterranean Sea and maintained for 30 and 90 d, at three mean pHT levels (total scale) of 8.1 (ambient), 7.7, and 7.4. Moderately decreased tube elongation rates were observed in both species at a pHT of 7.7 while severe reductions occurred at pHT 7.4. There was visual evidence of dissolution and tubes were more fragile at lower pH but, fragility was not attributed to changes in fracture toughness. Instead, it appeared to be due to the presence of larger alveoli covered in a thinner calcareous layer. The second objective of the study was to test for effects in offspring development of the species S. triqueter. Spawning was induced, and offspring were reared in the same pH conditions the parents experienced. Trochophore size was reduced at the lowest pH level but settlement success was similar across pH conditions. Post-settlement tube growth was most affected. At 38 d post-settlement, juvenile tubes at pHT of 7.7 and 7.4 were half the size of those at pHT 8.1. Results suggest future carbonate chemistry will negatively affect initiation and persistence of both biofouling and epiphytic polychaete tube worms.

Temperate marine rocky habitats may be alternatively characterized by well vegetated macroalgal assemblages or barren grounds, as a consequence of direct and indirect human impacts (e.g. overfishing) and grazing pressure by herbivorous organisms. In future scenarios of ocean acidification, calcifying organisms are expected to be less competitive: among these two key elements of the rocky subtidal food web, coralline algae and sea urchins. In order to highlight how the effects of increased pCO 2 on individual calcifying species will be exacerbated by interactions with other trophic levels, we performed an experiment simultaneously testing ocean acidification effects on primary producers (calcifying and non-calcifying algae) and their grazers (sea urchins). Artificial communities, composed by juveniles of the sea urchin Paracentrotus lividus and calcifying (Corallina elongata) and non-calcifying (Cystoseira amentacea var stricta, Dictyota dichotoma) macroalgae, were subjected to pCO 2 levels of 390, 550, 750 and 1000 matm in the laboratory. Our study highlighted a direct pCO 2 effect on coralline algae and on sea urchin defense from predation (test robustness). There was no direct effect on the non-calcifying macroalgae. More interestingly, we highlighted diet-mediated effects on test robustness and on the Aristotle's lantern size. In a future scenario of ocean acidification a decrease of sea urchins' density is expected, due to lower defense from predation, as a direct consequence of pH decrease, and to a reduced availability of calcifying macroalgae, important component of urchins' diet. The effects of ocean acidification may therefore be contrasting on well vegetated macroalgal assemblages and barren grounds: in the absence of other human impacts, a decrease of biodiversity can be predicted in vegetated macroalgal assemblages, whereas a lower density of sea urchin could help the recovery of shallow subtidal rocky areas affected by overfishing from barren grounds to assemblages dominated by fleshy macroalgae.

The effect of ocean acidification on the balance between gross community production (GCP) and community respiration (CR) (i.e., net community production, NCP) of plankton communities was investigated in summer 2010 in Kongsfjorden, west of Svalbard. Surface water, which was characterized by low concentrations of dissolved inorganic nutrients and chlorophyll a (a proxy of phytoplankton biomass), was enclosed in nine mesocosms and subjected to eight pCO(2) levels (two replicated controls and seven enhanced pCO(2) treatments) for one month. Nutrients were added to all mesocosms on day 13 of the experiment, and thereafter increase of chlorophyll a was provoked in all mesocosms. No clear trend in response to increasing pCO(2) was found in the daily values of NCP, CR, and GCP. For further analysis, these parameters were cumulated for the following three periods: phase 1 - end of CO2 manipulation until nutrient addition (t4 to t13); phase 2 - nutrient addition until the second chlorophyll a minimum (t14 to t21); phase 3 - the second chlorophyll a minimum until the end of this study (t22 to t28). A significant response was detected as a decrease of NCP with increasing pCO(2) during phase 3. CR was relatively stable throughout the experiment in all mesocosms. As a result, the cumulative GCP significantly decreased with increasing pCO(2) during phase 3. After the nutrient addition, the ratios of cumulative NCP to cumulative consumption of NO3 and PO4 showed a significant decrease during phase 3 with increasing pCO(2). The results suggest that elevated pCO(2) influenced cumulative NCP and stoichiometric C and nutrient coupling of the plankton community in a high-latitude fjord only for a limited period. However provided that there were some differences or weak correlations between NCP data based on different methods in the same experiment, this conclusion should be taken with caution.

Pteropods are planktonic mollusks that play an important role in the food web of various ecosystems, particularly at high latitudes. Because they produce an aragonitic shell, ptero pods are expected to be very sensitive to ocean acidification driven by anthropogenic CO2 emissions. The effect of ocean acidification was investigated using juveniles of the Arctic pteropod Limacina helicina from the Canada Basin of the Arctic Ocean. The animals were maintained in 3 controlled pH conditions (total scale pH [pH(T)] approximate to 8.05, 7.90 or 7.75) for 8 d, and their mortality and the linear extension of their shell were monitored. The pH did not impact the mortality rate, but the linear extension of the shell decreased as a function of declining pH. Surprisingly, the ptero pods were still able to extend their shell at an aragonite saturation state as low as 0.6. Nevertheless, dissolution marks were visible on the whole shell, indicating that calcium carbonate dissolution had also occurred, casting doubts on the ability of the pteropods to maintain a positive balance between precipitation and dissolution of calcium carbonate under corrosive conditions.

Larvae of the Mediterranean pteropod Cavolinia inflexa were maintained at controlled pH(T) values of 8.1, 7.82 and 7.51, equivalent, respectively, to pCO(2) levels of 380, 857 and 1,713 mu atm. At pH(T) 7.82, larvae exhibited malformations and lower shell growth, compared to the control condition. At pH(T) 7.51, the larvae did not make shells but were viable and showed a normal development. However, smaller shells or no shells will have both ecological (food web) and biogeochemical (export of carbon and carbonate) consequences. These results suggest that pteropod larvae, as well as the species dependent upon them or upon adults as a food resource, might be significantly impacted by ocean acidification.