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 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.

Coral reefs form complex physical structures that can help to mitigate coastal flooding risk 1,2 . This function will be reduced by sea-level rise (SLR) and impaired reef growth caused by climate change and local anthropogenic stressors 3 . Water depths above reef surfaces are projected to increase as a result, but the magnitudes and timescales of this increase are poorly constrained, which limits modelling of coastal vulnerability 4,5 . Here we analyse fossil reef deposits to constrain links between reef ecology and growth potential across more than 400 tropical western Atlantic sites, and assess the magnitudes of resultant above-reef increases in water depth through to 2100 under various shared socioeconomic pathway (SSP) emission scenarios. Our analysis predicts that more than 70% of tropical western Atlantic reefs will transition into net erosional states by 2040, but that if warming exceeds 2 °C (SSP2–4.5 and higher), nearly all reefs (at least 99%) will be eroding by 2100. The divergent trajectories of reef growth and SLR will thus magnify the effects of SLR; increases in water depth of around 0.3–0.5 m above the present are projected under all warming scenarios by 2060, but depth increases of 0.7–1.2 m are predicted by 2100 under scenarios in which warming surpasses 2 °C. This would increase the risk of flooding along vulnerable reef-fronted coasts and modify nearshore hydrodynamics and ecosystems. Reef restoration offers one pathway back to higher reef growth 6,7 , but would dampen the effects of SLR in 2100 only by around 0.3–0.4 m, and only when combined with aggressive climate mitigation.

Unraveling the functional future of marine ecosystems amid global change poses a pressing challenge. This is particularly critical in the Mediterranean Sea, which is highly impacted by global and local drivers. Utilizing extensive mass mortality events (MMEs) datasets spanning from 1986 to 2020 across the Mediterranean Sea, we investigated the trait vulnerability of benthic species that suffered from MMEs induced by nine distinct mortality drivers. By analyzing changes in ten ecological traits across 389 benthic species—constituting an extensive compendium of Mediterranean ecological traits to date—we identified 228 functional entities (FEs), defined as groups of species sharing the same trait values. Our findings indicate that of these 55 FEs were impacted by MMEs, accentuating a heightened vulnerability within specific trait categories. Notably, more than half of the mortality records showed severe impacts on calcifying and larger species with slower growth which mostly account for tree-like and massive forms. Altogether, we highlight that 29 FEs suffered extreme mortality, leading to a maximum increase of 19.1% of the global trait volume vulnerability over 35 years. We also reveal that 10.8% of the trait volume may have been temporarily lost over the last five years, emphasizing the risk of a rapid ecological transformation in the Mediterranean Sea.

Crustose coralline algae (CCA) and their bacterial communities can emit chemical cues favoring coral larval settlement. Larvae of Eunicella singularis (white gorgonian) preferentially settle on CCA. We investigated the effect of two Mediterranean CCA holobionts, Macroblastum dendrospermum and Lithophyllum stictiforme, on E. singularis larvae settlement and their bacterial communities, even after warming and acidification treatments. We exposed CCA to temperature and pH expected for 2100 (SSP5-8.5), and to a marine heatwave event. Larval settlement increased 1.8-2.7 times in presence of CCA exposed to warming and acidification compared to non-exposed CCA. High abundance of bacteria belonging to the Pirellulaceae family was observed in all CCA, while a higher abundance of monosaccharides was found in exudates of exposed CCA. Based on CCA-related 16S rDNA metabarcoding and metabolomics results, we hypothesize that the enhanced larval settlement was driven by the Pirellulaceae breakdown and utilization of CCA polysaccharides, in combination with polysaccharides release through the CCA cell walls likely augmented by decalcification. Furthermore, CCA acted as sources of bacterial taxa that may establish and persist in adult E. singularis holobiont, independently of climate change effects. We conclude that CCA are key for E. singularis recruitment success, especially under future climate conditions, and microbiome development.

Abstract. 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.

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.

Crustose coralline algae (CCA) and their bacterial communities can emit chemical cues favoring coral larval settlement. Indeed, larvae of Eunicella singularis (white gorgonian) preferentially settle on CCA. Here, we investigated the effect of two Mediterranean CCA holobionts, Macroblastum dendrospermum and Lithophyllum stictiforme , on E. singularis larvae settlement and their bacterial communities, after warming and acidification treatments. We exposed CCA to temperature and pH expected for 2100 (SSP5‐8.5) and to a marine heatwave event. Larval settlement increased 1.8–2.7 times in the presence of CCA exposed to warming and acidification compared to non‐exposed CCA. High abundance of bacteria belonging to the Pirellulaceae family was observed in all CCA, while a higher abundance of monosaccharides was found in exudates of exposed CCA. Based on CCA‐related 16S rDNA metabarcoding and metabolomics results, we hypothesize that the enhanced larval settlement was driven by the Pirellulaceae breakdown and utilization of CCA polysaccharides, in combination with polysaccharide release through the CCA cell walls likely augmented by decalcification. Furthermore, CCA acted as sources of bacterial taxa that may establish and persist in the adult E. singularis holobiont, independently of climate change effects. We conclude that CCA are key for E. singularis recruitment success, especially under future climate conditions, and contribute to their microbiome development.

Human activities over the past 150 yr have led to significant carbon dioxide (CO$_2$) emissions, causing global warming and ocean acidification. Surface ocean temperature has risen by 0.93°C since 1850, with projections of an additional +1.42°C to 3.47°C by 2080–2099. Ocean acidification, driven by CO$_2$ absorption, has already lowered seawater pH by 0.1 units, affecting calcifying organisms, including shelled mollusks. Long‐term multigenerational studies on mollusk responses to both ocean acidification and warming, under realistic environmental conditions, are scarce. To address this knowledge gap, two mobile experimental units that can be deployed at the vicinity of shellfish farming areas were developed within the framework of the CocoriCO$_2$ project. The experimental systems were designed to manipulate temperature and pH as offsets from ambient conditions. The experimental units have shown their effectiveness in terms of controlling and maintaining pH and temperature to assess the multigenerational effects of ocean warming and acidification on benthic invertebrates. Finally, the developed experimental systems can be modified easily to provide an educated assessment of the impact of other relevant environmental changes such as deoxygenation and changes in salinity.

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 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. 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 Mediterranean Sea is a hotspot of global change, particularly exposed to ocean warming and the increasing occurrence of marine heatwaves (MHWs). However, experiments based on long-term temperature data from the field are scarce. Here, we investigate the response of the zooxanthellate coral Cladocora caespitosa and the azooxanthellate coral Astroides calycularis to future warming and MHWs based on 8 years of in situ data. Corals were maintained in the laboratory for five months under four temperature conditions: Warming (3.2°C above the in situ mean from 2012 to 2020), Heatwave (temperatures of 2018 with two heatwaves), Ambient ( in situ mean) and Cool (deeper water temperatures). Under the Warming treatment, some C. caespitosa colonies severely bleached and A. calycularis colonies presented necrosis. Cladocora caespitosa symbiosis was impaired by temperature with a decrease in the density of endosymbiotic algae and an increase in per cent whiteness in all the treatments except for the coolest. Recovery for both species was observed through different mechanisms such as regrowth of polyps of A. calycularis and recovery of pigmentation for C. caespitosa . These results suggest that A. calycularis and C. caespitosa may be resilient to heat stress and can recover from physiological stresses caused by heatwaves in the laboratory.

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.

Abstract. Ocean acidification (OA) is predicted to cause profound shifts in many marine ecosystems by impairing the ability of calcareous taxa to calcify and grow and by influencing the physiology of many others. In both calcifying and non-calcifying taxa, ocean acidification could further impair the ability of marine life to regulate internal pH and thus metabolic function and/or behaviour. Identifying tipping points at which these effects will occur for different taxa due to the direct impacts of ocean acidification on organism physiology is difficult because they have not adequately been determined for most taxa nor for ecosystems at higher levels. This is due to the presence of both resistant and sensitive species within most taxa. However, calcifying taxa such as coralline algae, corals, molluscs, and sea urchins appear to be most sensitive to ocean acidification. Conversely, non-calcareous seaweeds, seagrasses, diatoms, cephalopods, and fish tend to be more resistant or even benefit from the direct effects of ocean acidification, though the effects of ocean acidification are more subtle for these taxa. While physiological tipping points of the effects of ocean acidification either do not exist or are not well defined, their direct effects on organism physiology will have flow-on indirect effects. These indirect effects will cause ecological tipping points in the future through changes in competition, herbivory, and predation. Evidence for indirect effects and ecological change is mostly taken from benthic ecosystems in warm temperate–tropical locations in situ that have elevated CO2. Species abundances at these locations indicate a shift away from calcifying taxa and towards non-calcareous taxa at high-CO2 concentrations. For example, lower abundance of corals and coralline algae and higher covers of non-calcareous macroalgae, often turfing species, are often found at elevated CO2. However, there are some locations where only minor changes or no detectable changes occur. Where ecological tipping points do occur, it is usually at locations with naturally elevated mean pCO2 concentrations of 500 µatm or more, which also corresponds to just under that concentration where the direct physiological impacts of ocean acidification are detectable in the most sensitive taxa in laboratory research (coralline algae and corals). Collectively, the available data support the concern that ocean acidification will most likely cause ecological change in the near future in most benthic marine ecosystems, with tipping points in some ecosystems as low as 500 µatm pCO2. However, further research is required to more adequately quantify and model the extent of these impacts in order to accurately project future marine ecosystem tipping points under ocean acidification.

Arctic coastal ecosystems include benthic communities that hold an important role within the marine food chain. Kelps, fucoid species, and coralline algae dominate rocky habitats, offering food and shelter for various species. Kelps and fucoid species also aid in carbon sequestration, sediment stabilization, and erosion mitigation. In summer, the influx of freshwater from glacier and permafrost melt alters coastal waters conditions. The input of turbid freshwater influences underwater light, salinity, and substrate, impacting benthic organism distribution. This study investigates possible link between environmental conditions and benthic diversity through environmental DNA (eDNA). Six sites were monitored along Kongsfjorden (Svalbard,Norway) during the summer of 2021. Contrary to expectations, macroalgal distribution didn’t correlate with light, and suspension feeders showed no clear links with chlorophyll a or nutrient concentrations. Glacial influence may have contributed to higher benthic diversity. Predators’ presence, tied to glacier proximity, possibly explained this trend. However, further studies are needed to validate these observations and assumptions.

Abstract Projecting the effects of climate change on net reef calcium carbonate production is critical to understanding the future impacts on ecosystem function, but prior estimates have not included corals' natural adaptive capacity to such change. Here we estimate how the ability of symbionts to evolve tolerance to heat stress, or for coral hosts to shuffle to favourable symbionts, and their combination, may influence responses to the combined impacts of ocean warming and acidification under three representative concentration pathway (RCP) emissions scenarios (RCP2.6, RCP4.5 and RCP8.5). We show that symbiont evolution and shuffling, both individually and when combined, favours persistent positive net reef calcium carbonate production. However, our projections of future net calcium carbonate production (NCCP) under climate change vary both spatially and by RCP. For example, 19%–35% of modelled coral reefs are still projected to have net positive NCCP by 2050 if symbionts can evolve increased thermal tolerance, depending on the RCP. Without symbiont adaptive capacity, the number of coral reefs with positive NCCP drops to 9%–13% by 2050. Accounting for both symbiont evolution and shuffling, we project median positive NCPP of coral reefs will still occur under low greenhouse emissions (RCP2.6) in the Indian Ocean, and even under moderate emissions (RCP4.5) in the Pacific Ocean. However, adaptive capacity will be insufficient to halt the transition of coral reefs globally into erosion by 2050 under severe emissions scenarios (RCP8.5).

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.

Ocean warming is transforming the world's coral reefs, which are governed by the growth of marine calcifiers, most notably branching corals. Critical to skeletal growth is the corals' regulation of their internal chemistry to promote calcification. Here we investigate the effects of temperature and light on the calcifying fluid chemistry (using boron isotope systematics), calcification rates, metabolic rates and photo-physiology of Acropora nasuta during two mesocosm experiments simulating seasonal and static temperature and light regimes. Under the seasonal regime, coral calcification rates, calcifying fluid carbonate chemistry, photo-physiology and metabolic productivity responded to both changes in temperature and light. However, under static conditions the artificially prolonged exposure to summer temperatures resulted in heat stress and a heightened sensitivity to light. Our results indicate that temperature and light effects on coral physiology and calcification mechanisms are interactive and context-specific, making it essential to conduct realistic multi-variate dynamic experiments in order to predict how coral calcification will respond to ocean warming.

Abstract. The ability of coral populations to recover from disturbance depends on larval dispersion and recruitment. While ocean warming and acidification effects on adult corals are well documented, information on early life stages is comparatively scarce. Here, we investigate whether ocean warming and acidification can affect the larval and recruit development of the Mediterranean azooxanthellate coral Astroides calycularis. Larvae and recruits were raised for 9 months at ambient (23 ∘C) and warm (26 ∘C) temperatures and ambient (8.0) and low pH (7.7, on the total scale). The timing of the larval metamorphosis, growth of the recruit polyp by linear extension and budding, and skeletal characteristics of the 9-month-old polyps were monitored. Settlement and metamorphosis were more successful and hastened under a warm temperature. In contrast, low pH delayed the metamorphosis and affected the growth of the recruits by reducing the calcified area of attachment to the substrate as well as by diminishing the skeleton volume and the number of septa. However, skeleton density was higher under low pH and ambient temperature. The warm temperature and low-pH treatment had a negative impact on the survival, settlement, and growth of recruits. This study provides evidence of the threat represented by ocean warming and acidification for the larval recruitment and the growth of recruits of A. calycularis.

Ocean warming (OW) and acidification (OA) are two of the greatest global threats to the persistence of coral reefs. Calcifying reef taxa such as corals and coralline algae provide the essential substrate and habitat in tropical reefs but are at particular risk due to their susceptibility to both OW and OA. OW poses the greater threat to future reef growth and function, via its capacity to destabilise the productivity of both taxa, and to cause mass bleaching events and mortality of corals. Marine heatwaves are projected to increase in frequency, intensity, and duration over the coming decades, raising the question of whether coral reefs will be able to persist as functioning ecosystems and in what form. OA should not be overlooked, as its negative impacts on the calcification of reef-building corals and coralline algae will have consequences for global reef accretion. Given that OA can have negative impacts on the reproduction and early life stages of both coralline algae and corals, the interdependence of these taxa may result in negative feedbacks for reef replenishment. However, there is little evidence that OA causes coral bleaching or exacerbates the effects of OW on coral bleaching. Instead, there is some evidence that OA alters the photo-physiology of both taxa. Tropical coralline algal possess shorter generation times than corals, which could enable more rapid evolutionary responses. Future reefs will be dominated by taxa with shorter generation times and high plasticity, or those individuals inherently resistant and resilient to both marine heatwaves and OA.

The Arctic region faces a warming rate that is more than twice the global average. Seaice loss, increase in precipitation and freshwater discharge, changes in underwater light, and amplification of ocean acidification modify benthic habitats and the communities they host. Here we synthesize existing information on the impacts of climate change on the macroalgal communities of Arctic coasts. We review the shortand long-term changes in environmental characteristics of shallow hard-bottomed Arctic coasts, the floristics of Arctic macroalgae (description, distribution, life-cycle, adaptations), the responses of their biological and ecological processes to climate change, the resulting winning and losing species, and the effects on ecosystem functioning. The focus of this review is on fucoid species, kelps, and coralline algae which are key ecosystem engineers in hard-bottom shallow areas of the Arctic, providing food, substrate, shelter, and nursery ground for many species. Changes in seasonality, benthic functional diversity, food-web structure, and carbon cycle are already occurring and are reshaping Arctic benthic ecosystems. Shallow communities are projected to shift from invertebrate-to algal-dominated communities. Fucoid and several kelp species are expected to largely spread and dominate the area with possible extinctions of native species. A considerable amount of functional diversity could be lost impacting the processing of land-derived nutrients and organic matter and significantly altering trophic structure and energy flow up to the apex consumers. However, many factors are not well understood yet, making it difficult to appreciate the current situation and predict the future coastal Arctic ecosystem. Efforts must be made to improve knowledge in key regions with proper seasonal coverage, taking into account interactions between stressors and across species.

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.

Ocean acidification (OA) is a serious consequence of climate change with complex organism-to-ecosystem effects that have been observed through field observations but are mainly derived from experimental studies. Although OA trends and the resulting biological impacts are likely exacerbated in the semi-enclosed and highly populated Mediterranean Sea, some fundamental knowledge gaps still exist. These gaps are at tributed to both the uneven capacity for OA research that exists between Mediterranean countries, as well as to the subtle and long-term biological, physical and chemical interactions that define OA impacts. In this paper, we systematically analyzed the different aspects of OA research in the Mediterranean region based on two sources: the United Nation’s International Atomic Energy Agency’s (IAEA) Ocean Acidification International Coordination Center (OA-ICC) database, and an extensive survey. Our analysis shows that 1) there is an uneven geographic capacity in OA research, and illustrates that both the Algero-Provencal and Ionian sub-basins are currently the least studied Mediterranean areas, 2) the carbonate system is still poorly quantified in coastal zones, and long-term time-series are still sparse across the Mediterranean Sea, which is a challenge for studying its variability and assessing coastal OA trends, 3) the most studied groups of organisms are autotrophs (algae, phanerogams, phytoplankton), mollusks, and corals, while microbes, small mollusks (mainly pteropods), and sponges are among the least studied, 4) there is an overall paucity in socio-economic, paleontological, and modeling studies in the Mediterranean Sea, and 5) in spite of general resource availability and the agreement for improved and coordinated OA governance, there is a lack of consistent OA policies in the Mediterranean Sea. In addition to highlighting the current status, trends and gaps of OA research, this work also provides recommendations, based on both our literature assessment and a survey that targeted the Mediterranean OA scientific community. In light of the ongoing 2021-2030 United Nations Decade of Ocean Science for Sustainable Development, this work might provide a guideline to close gaps of knowledge in the Mediterranean OA research. Systematic Review Registration https://www.oceandecade.org

Abstract Coral reefs are iconic ecosystems with immense ecological, economic and cultural value, but globally their carbonate‐based skeletal construction is threatened by ocean acidification (OA). Identifying coral species that have specialised mechanisms to maintain high rates of calcification in the face of declining seawater pH is of paramount importance in predicting future species composition, and growth of coral reefs. Here, we studied multiple coral species from two distinct volcanic CO 2 seeps in Papua New Guinea to assess their capacity to control their calcifying fluid (CF) chemistry. Several coral species living under conditions of low mean seawater pH, but with either low or high variability in seawater pH, were examined and compared with those living in ‘normal’ (non‐seep) ambient seawater pH. We show that when mean seawater pH is low but highly variable, corals have a greater ability to maintain constant pH cf in their CF, but this characteristic was not linked with changes in abundance. Within less variable low pH seawater, corals with limited reductions in pH cf at the seep sites compared with controls tended to be more abundant at the seep site than at the control site. However, this finding was strongly influenced by a single species ( Montipora foliosa ), which was able to maintain complete pH cf homeostasis. Overall, although our findings indicate that there might be an association between ecological success and greater pH cf homeostasis, further research with additional species and at more sites with differing seawater pH regimes is required to solidify inferences regarding coral ecological success under future OA.

In corals, pH regulation of the extracellular calcifying medium (ECM) by the calcifying cell layer is a crucial step in the calcification process and is potentially important to influencing how corals respond to ocean acidification. Here, we analyzed the growing edge of the reef coral Stylophora pistillata to make the first characterization of the proton gradient across the coral calcifying epithelium. At seawater pH 8 we found that while the calcifying epithelium elevates pH in the ECM on its apical side above that of seawater, pH on its basal side in the mesoglea is markedly lower, highlighting that the calcifying cells are exposed to a microenvironment distinct from the external environment. Coral symbiont photosynthesis elevates pH in the mesoglea, but experimental ocean acidification and decreased seawater inorganic carbon concentration lead to large declines in mesoglea pH relative to the ECM, which is maintained relatively stable. Together, our results indicate that the coral calcifying epithelium is functionally polarized and that environmental variation impacts pH ECM regulation through its effects on the basal side of the calcifying cells.

The future of coral reef ecosystems is under threat because vital reef-accreting species such as coralline algae are highly susceptible to ocean acidification. Although ocean acidification is known to reduce coralline algal growth rates, its direct effects on the development of coralline algal reproductive structures (conceptacles) is largely unknown. Furthermore, the long-term, multigenerational response of coralline algae to ocean acidification is extremely understudied. Here, we investigate how mean pH, pH variability and the pH regime experienced in their natural habitat affect coralline algal conceptacle abundance and size across six generations of exposure. We show that second generation coralline algae exposed to ocean acidification treatments had conceptacle abundances 60% lower than those kept in present day conditions, suggesting that conceptacle development is initially highly sensitive to ocean acidification. However, this negative effect of ocean acidification on conceptacle abundance disappears after three generations of exposure. Moreover, we show that this transgenerational acclimation of conceptacle development is not facilitated by a tradeoff with reduced investment in growth, as higher conceptacle abundances are associated with crusts with faster growth rates. These results indicate that the potential reproductive output of coralline algae may be sustained under future ocean acidification.

Ocean acidification is perceived to be a major threat for many calcifying organisms, including scleractinian corals. Here we investigate (1) whether past exposure to low pH environments associated with CO2 vents could increase corals tolerance to low pH and (2) whether zooxanthellate corals are more tolerant to low pH than azooxanthellate corals. To test these hypotheses, two Mediterranean colonial corals Cladocora caespitosa (zooxanthellate) and Astroides calycularis (azooxanthellate) were collected from CO2 vents and reference sites and incubated in the laboratory under present-day (pH on the total scale, pHT 8.07) and low pH conditions (pHT 7.70). Rates of net calcification, dark respiration and photosynthesis were monitored during a six-month experiment. Monthly net calcification was assessed every 27 to 35 d using the buoyant weight technique, whereas light and dark net calcification was estimated using the alkalinity anomaly technique during 1 h incubations. Neither species showed any change in net calcification rates, respiration, and photosynthesis regardless of their environmental history, pH treatment and trophic strategy. Our results indicate that C. caespitosa and A. calycularis could tolerate future ocean acidification conditions for at least 6 months. These results will aid in predicting species' future responses to ocean acidification, and thus improve the management and conservation of Mediterranean corals.

Ocean warming and acidification threaten the future growth of coral reefs. This is because the calcifying coral reef taxa that construct the calcium carbonate frameworks and cement the reef together are highly sensitive to ocean warming and acidification. However, the global-scale effects of ocean warming and acidification on rates of coral reef net carbonate production remain poorly constrained despite a wealth of studies assessing their effects on the calcification of individual organisms. Here, we present global estimates of projected future changes in coral reef net carbonate production under ocean warming and acidification. We apply a meta-analysis of responses of coral reef taxa calcification and bioerosion rates to predicted changes in coral cover driven by climate change to estimate the net carbonate production rates of 183 reefs worldwide by 2050 and 2100. We forecast mean global reef net carbonate production under representative concentration pathways (RCP) 2.6, 4.5, and 8.5 will decline by 76, 149, and 156%, respectively, by 2100. While 63% of reefs are projected to continue to accrete by 2100 under RCP2.6, 94% will be eroding by 2050 under RCP8.5, and no reefs will continue to accrete at rates matching projected sea level rise under RCP4.5 or 8.5 by 2100. Projected reduced coral cover due to bleaching events predominately drives these declines rather than the direct physiological impacts of ocean warming and acidification on calcification or bioerosion. Presently degraded reefs were also more sensitive in our analysis. These findings highlight the low likelihood that the world's coral reefs will maintain their functional roles without near-term stabilization of atmospheric CO2 emissions.

Coralline algae are foundation species in many hard-bottom ecosystems acting as a settlement substrate, and binding together and even creating reefs in some locations. Ocean acidification is known to be a major threat to coralline algae. However, the effects of ocean warming are less certain. Here we bring multiple lines of evidence together to discuss the potential impacts of ocean warming on these ecologically crucial taxa. We use a meta-analysis of 40 responses within 14 different studies available which assessed the effects of increasing temperature on coralline algal calcification in laboratory experiments. We find a net negative impact of increasing temperature on coralline algal calcification at 5.2°C above ambient conditions. Conversely, negative effects are observed when temperature drops below 2.0°C from ambient conditions. We propose that some coralline algae will be more capable of both acclimatizing and locally adapting to increasing ocean temperatures over the coming decades. This is because many species possess short generation times, the ability to opportunistically rapidly utilize open space, and relatively high phenotypic plasticity. However, less resistant and resilient species will be those that are long-lived, those with long generation times, or with narrow thermal tolerances (e.g., tropical taxa living close to their thermal maxima). Additionally, ocean warming will occur simultaneously with ocean acidification, a potentially greater threat to coralline algae, which could also reduce any tolerance to ocean warming for many species. To maximize the potential to accurately determine how coralline algae will respond to future ocean warming and marine heatwaves, future research should use environmentally relevant temperature treatments, use appropriate acclimation times and follow best practices in experimental design.

Ocean acidification (OA) is a major threat to coral reefs, which are built by calcareous species. However, long-term assessments of the impacts of OA are scarce, limiting the understanding of the capacity of corals and coralline algae to acclimatize to high partial pressure of carbon dioxide (pCO2) levels. Species-specific sensitivities to OA are influenced by its impacts on chemistry within the calcifying fluid (CF). Here, we investigate the capacity of multiple coral and calcifying macroalgal species to acclimatize to elevated pCO2 by determining their chemistry in the CF during a year-long experiment. We found no evidence of acclimatization to elevated pCO2 across any of the tested taxa. The effects of increasing seawater pCO2 on the CF chemistry were rapid and persisted until the end of the experiment. Our results show that acclimatization of the CF chemistry does not occur within one year, which confirms the threat of OA for future reef accretion and ecological function.

Natural variability in pH in the diffusive boundary layer (DBL), the discrete layer of seawater between bulk seawater and the outer surface of organisms, could be an important factor determining the response of corals and coralline algae to ocean acidification (OA). Here, two corals with different morphologies and one coralline alga were maintained under two different regimes of flow velocities, pH, and light intensities in a 12 flumes experimental system for a period of 27 weeks. We used a combination of geochemical proxies, physiological and micro-probe measurements to assess how these treatments affected the conditions in the DBL and the response of organisms to OA. Overall, low flow velocity did not ameliorate the negative effect of low pH and therefore did not provide a refugia from OA. Flow velocity had species-specific effects with positive effects on calcification for two species. pH in the calcifying fluid (pHcf) was reduced by low flow in both corals at low light only. pHcf was significantly impacted by pH in the DBL for the two species capable of significantly modifying pH in the DBL. The dissolved inorganic carbon in the calcifying fluid (DICcf) was highest under low pH for the corals and low flow for the coralline, while the saturation state in the calcifying fluid and its proxy (FWHM) were generally not affected by the treatments. This study therefore demonstrates that the effects of OA will manifest most severely in a combination of lower light and lower flow habitats for sub-tropical coralline algae. These effects will also be greatest in lower flow habitats for some corals. Together with existing literature, these findings reinforce that the effects of OA are highly context dependent, and will differ greatly between habitats, and depending on species composition.

Ocean acidification is a threat to the continued accretion of coral reefs, though some undergo daily fluctuations in pH exceeding declines predicted by 2100. We test whether exposure to greater pH variability enhances resistance to ocean acidification for the coral Goniopora sp. and coralline alga Hydrolithon reinboldii from two sites: one with low pH variability (less than 0.15 units daily; Shell Island) and a site with high pH variability (up to 1.4 pH units daily; Tallon Island). We grew populations of both species for more than 100 days under a combination of differing pH variability (high/low) and means (ambient pH 8.05/ocean acidification pH 7.65). Calcification rates of Goniopora sp. were unaffected by the examined variables. Calcification rates of H. reinboldii were significantly faster in Tallon than in Shell Island individuals, and Tallon Island individuals calcified faster in the high variability pH 8.05 treatment compared with all others. Geochemical proxies for carbonate chemistry within the calcifying fluid (cf) of both species indicated that only mean seawater pH influenced pH(cf). pH treatments had no effect on proxies for Omega(cf). These limited responses to extreme pH treatments demonstrate that some calcifying taxa may be capable of maintaining constant rates of calcification under ocean acidification by actively modifying Omega(cf).

Ocean acidification threatens the persistence of biogenic calcium carbonate (CaCO 3) production on coral reefs. However, some coral genera show resistance to declines in seawater pH, potentially achieved by modulating the chemistry of the fluid where calcification occurs. We use two novel geochem-ical techniques based on boron systematics and Raman spectroscopy, which together provide the first constraints on the sensitivity of coral calcifying fluid calcium concentrations (½Ca 2þ cf) to changing seawater pH. In response to simulated end-of-century pH conditions, Pocillopora damicornis increased ½Ca 2þ cf to as much as 25% above that of seawater and maintained constant calcification rates. Conversely, Acropora youngei displayed less control over ½Ca 2þ cf , and its calcification rates strongly declined at lower seawater pH. Although the role of ½Ca 2þ cf in driving calcification has often been neglected, increasing ½Ca 2þ cf may be a key mechanism enabling more resistant corals to cope with ocean acidification and continue to build CaCO 3 skeletons in a high-CO 2 world.

Ocean acidification (OA) is a major threat to marine ecosystems, particularly coral reefs which are heavily reliant on calcareous species. OA decreases seawater pH and calcium carbonate saturation state (Ω), and increases the concentration of dissolved inorganic carbon (DIC). Intense scientific effort has attempted to determine the mechanisms via which ocean acidification (OA) influences calcification, led by early hypotheses that calcium carbonate saturation state (Ω) is the main driver. We grew corals and coralline algae for 8-21 weeks, under treatments where the seawater parameters Ω, pH, and DIC were manipulated to examine their differential effects on calcification rates and calcifying fluid chemistry (Ω cf , pH cf , and DIC cf). Here, using long duration experiments, we provide geochemical evidence that differing physiological controls on carbonate chemistry at the site of calcification, rather than seawater Ω, are the main determinants of calcification. We found that changes in seawater pH and DIC rather than Ω had the greatest effects on calcification and calcifying fluid chemistry, though the effects of seawater carbonate chemistry were limited. Our results demonstrate the capacity of organisms from taxa with vastly different calcification mechanisms to regulate their internal chemistry under extreme chemical conditions. These findings provide an explanation for the resistance of some species to OA, while also demonstrating how changes in seawater DIC and pH under OA influence calcification of key coral reef taxa.

Evaluating the factors responsible for differing species-specific sensitivities to declining seawater pH is central to understanding the mechanisms via which ocean acidification (OA) affects coral calcification. We report here the results of an experiment comparing the responses of the coral Acropora yongei and Pocillopora damicornis to differing pH levels (8.09, 7.81, and 7.63) over an 8-week period. Calcification of A. youngei was reduced by 35% at pH 7.63, while calcification of P. damicornis was unaffected. The pH in the calcifying fluid (pH cf) was determined using δ 11 B systematics, and for both species pH cf declined slightly with seawater pH, with the decrease being more pronounced in P. damicornis. The dissolved inorganic carbon concentration at the site of calcification (DIC cf) was estimated using geochemical proxies (B/Ca and δ 11 B) and found to be double that of seawater DIC, and increased in both species as seawater pH decreased. As a consequence, the decline of the saturation state at the site of calcification (Ω cf) with OA was partially moderated by the DIC cf increase. These results highlight that while pH cf , DIC cf and Ω cf are important in the mineralization process, some corals are able to maintain their calcification rates despite shifts in their calcifying fluid carbonate chemistry.

The impact of anthropogenic ocean acidification (OA) on marine ecosystems is a vital concern facing marine scientists and managers of ocean resources. Euthecosomatous pteropods (holoplanktonic gastropods) represent an excellent sentinel for indicating exposure to anthropogenic OA because of the sensitivity of their aragonite shells to the OA conditions less favorable for calcification. However, an integration of observations, experiments and modelling efforts is needed to make accurate predictions of how these organisms will respond to future changes to their environment. Our understanding of the underlying organismal biology and life history is far from complete and must be improved if we are to comprehend fully the responses of these organisms to the multitude of stressors in their environment beyond OA. This review considers the present state of research and understanding of euthecosomatous pteropod biology and ecology of these organisms and considers promising new laboratory methods, advances in instrumentation (such as molecular, trace elements, stable isotopes,

Purpose of Review We investigate whether regimes of greater daily variability in temperature or pH result in greater tolerance to ocean warming and acidification in key reef-building taxa (corals, coralline algae). Recent Findings Temperature and pH histories will likely influence responses to future warming and acidification. Past exposure of corals to increased temperature variability generally leads to greater thermotolerance. However, the effects of past pH variability are unclear. Variability in pH or temperature will likely modify responses during exposure to stressors, independent of environmental history. In the laboratory, pH variability often limited the effects of ocean acidification, but the effects of temperature variability on responses to warming were equivocal. Summary Environmental variability could alter responses of coral reef organisms to climate change. Determining how both environmental history as well as the direct impacts of environmental variability will interact with the effects of anthropogenic climate change should now be high priority.

Current research on coral reefs seeks to link the responses to anthropogenic stressors (such as global warming and ocean acidification [OA]) among differing functional levels of biological organization. While experimental studies have identified ex situ taxon-specific responses to OA and global warming, isolating and connecting these effects in situ at the community-level has proved difficult. The difficulties arise from the large number of naturally varying parameters affecting corals reefs, such as light intensity and seawater residence time that affect net community production and calcification. To control variation in seawater residence time and allow light intensity to vary naturally, experimental outer reef (17-m depth) benthic communities composed of calcified algae, corals, and reef pavement were constructed in large outdoor flumes in Mo'orea, French Polynesia. Net community production (P), net community calcification (G), the ratio of P/G (P/G(ratio)), and slope of P regressed on G (P/G(slope)) were calculated for the communities, and concurrently for the constituent members under the same temperature, light, and flow conditions. P and G, for both the communities and constituent members, were correlated positively with light intensity, whereas P/G(rati)o and P/G(siope) were unaffected by light intensity. P/G(ratios) and P/G(siopes) exhibited values that were specific to each community member. These results suggest that the P/G(ratio) and P/G(slope) may be unaffected by natural variability in light intensity and could serve as useful metrics to relate responses at the taxon and community level, which is an important step in assessing the effects of environmental changes on coral reefs.

In 2015/16, a marine heatwave associated with a record El Nino led to the third global mass bleaching event documented to date. This event impacted coral reefs around the world, including in Western Australia (WA), although WA reefs had largely escaped bleaching during previous strong El Nino years. Coral health surveys were conducted during the austral summer of 2016 in four bioregions along the WA coast (similar to 17 degrees of latitude), ranging from tropical to temperate locations. Here we report the first El Nino-related regional-scale mass bleaching event in WA. The heatwave primarily affected the macrotidal Kimberley region in northwest WA (similar to 16 degrees S), where 4.5-9.3 degree heating weeks (DHW) resulted in 56.6-80.6% bleaching, demonstrating that even heat-tolerant corals from naturally extreme, thermally variable reef environments are threatened by heatwaves. Some heat stress (2.4 DHW) and bleaching (<30%) also occurred at Rottnest Island (32 degrees 01'S), whereas coral communities at Ningaloo Reef (23 degrees 9'S) and Bremer Bay (34 degrees 25'S) were not impacted. The only other major mass bleaching in WA occurred during a strong La Nina event in 2010/11 and primarily affected reefs along the central-to-southern coast. This suggests that WA reefs are now at risk of severe bleaching during both El Nino and La Nina years.

The response of tropical corals and calcifying algae to ocean acidification (OA) and warming has received much attention in the past decade. However, most studies have evaluated the response of organisms to two or three temperature treatments, which does not allow the functional relationship between calcification and temperature under ambient and future pCO(2) to be determined. This study tested the hypothesis that the relationship between calcification and temperature is affected by OA in the coral Acropora pulchra and the calcified alga Lithophyllum kotschyanum. Pieces of each organism were incubated under five (24-30 A degrees C) or six (24-31.5 A degrees C) temperatures crossed with two pCO(2) levels (400 and 1000 mu atm), and calcification was assessed in trials conducted in the spring and summer. The response of coral calcification to temperature was a positive asymmetric parabola with a maximum at similar to 28 A degrees C under both pCO(2) levels and in both seasons; the effects of pCO(2) on calcification were largest at similar to 28 A degrees C and lowest in both cool and warm temperatures. In contrast, calcification of the alga at both levels of pCO(2) was unaffected by temperature in spring, but declined linearly with temperature in summer. This study demonstrates that the calcification response of coral reef organisms to the crossed effect of warming and OA is complex and cannot be fully assessed without using multiple temperature treatments that are ecologically relevant.

There are concerns about the future of coral reefs in the face of ocean acidification and warming, and although studies of these phenomena have advanced quickly, efforts have focused on pieces of the puzzle rather than integrating them to evaluate ecosystem-level effects. The field is now poised to begin this task, but there are information gaps that first must be overcome before progress can be made. Many of these gaps focus on calcification at the levels of cells, organisms, populations, communities, and ecosystem, and their closure will be made difficult by the complexity of the interdependent processes by which coral reefs respond to ocean acidification, with effects scaling from cells to ecosystems and from microns to kilometers. Existing ecological theories provide an important and largely untapped resource for overcoming these difficulties, and they offer great potential for integrating the effects of ocean acidification across scales on coral reefs.

The widespread distribution of pteropods, their role in ocean food webs and their sensitivity to ocean acidification and warming has renewed scientific interest in this group of zooplankton. Unfortunately, their fragile shell, sensitivity to handling, unknowns surrounding buoyancy regulation and poorly described feeding mechanisms make thecosome pteropods notoriously difficult to maintain in the laboratory. The resultant high mortality rates and unnatural behaviours may confound experimental findings. The high mortality rate also discourages the use of periods of acclimation to experimental conditions and precludes vital long-term studies. Here we summarize the current status of culture methodology to provide a comprehensive basis for future experimental work and culture system development.

Over the next century, elevated quantities of atmospheric CO2 are expected to penetrate into the oceans, causing a reduction in pH (-0.3/-0.4 pH unit in the surface ocean) and in the concentration of carbonate ions (so-called ocean acidification). Of growing concern are the impacts that this will have on marine and estuarine organisms and ecosystems. Marine shelled molluscs, which colonized a large latitudinal gradient and can be found from intertidal to deep-sea habitats, are economically and ecologically important species providing essential ecosystem services including habitat structure for benthic organisms, water purification and a food source for other organisms. The effects of ocean acidification on the growth and shell production by juvenile and adult shelled molluscs are variable among species and even within the same species, precluding the drawing of a general picture. This is, however, not the case for pteropods, with all species tested so far, being negatively impacted by ocean acidification. The blood of shelled molluscs may exhibit lower pH with consequences for several physiological processes (e.g. respiration, excretion, etc.) and, in some cases, increased mortality in the long term. While fertilization may remain unaffected by elevated pCO(2), embryonic and larval development will be highly sensitive with important reductions in size and decreased survival of larvae, increases in the number of abnormal larvae and an increase in the developmental time. There are big gaps in the current understanding of the biological consequences of an acidifying ocean on shelled molluscs. For instance, the natural variability of pH and the interactions of changes in the carbonate chemistry with changes in other environmental stressors such as increased temperature and changing salinity, the effects of species interactions, as well as the capacity of the organisms to acclimate and/or adapt to changing environmental conditions are poorly described.

Thecosome pteropods play a key role in the food web of various marine ecosystems and they calcify, secreting the unstable CaCO 3 mineral aragonite to form their shell material. Here, we have estimated the effect of ocean acidification on pteropod calcification by exploiting empirical relationships between their gross calcification rates (CaCO 3 precipitation) and aragonite saturation state Ω a , combined with model projections of future Ω a . These were corrected for modern model-data bias and taken over the depth range where pteropods are observed to migrate vertically. Results indicate large reductions in gross calcification at temperate and high latitudes. Over much of the Arctic, the pteropod Limacina helicina will become unable to precipitate CaCO 3 by the end of the century under the IPCC SRES A2 scenario. These results emphasize concerns over the future of shelled pteropods, particularly L. helicina in high latitudes. Shell-less L. helicina are not known to have ever existed nor would we expect them to survive. Declines of pteropod populations could drive dramatic ecological changes in the various pelagic ecosystems in which they play a critical role.

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.

Although shelled pteropods are expected to be particularly sensitive to ocean acidification, the few available studies have mostly focused on polar species and have not allowed determining which parameter of the carbonate system controls their calcification. Specimens of the temperate Mediterranean species Creseis acicula were maintained under seven different conditions of the carbonate chemistry, obtained by manipulating pH and total alkalinity, with the goal to disentangle the effects of the pH and the saturation state with respect to aragonite (Ω_a). Our results tend to show that respiration, excretion as well as rates of net and gross calcification were not directly affected by a decrease in pH but decreased significantly with a decrease in Ω_a. Due to the difficulties in maintaining pteropods in the laboratory and the important variability in their abundances in our study site, long-term acclimation as well as replication of the experiment was not possible. However, we strongly believe that these results represent an important step in the mechanistic understanding of the effect of ocean acidification on pteropods physiology.

Thecosome pteropods (pelagic mollusks) can play a key role in the food web of various marine ecosystems. They are a food source for zooplankton or higher predators such as fishes, whales and birds that is particularly important in high latitude areas. Since they harbor a highly soluble aragonitic shell, they could be very sensitive to ocean acidification driven by the increase of anthropogenic CO2 emissions. The effect of changes in the seawater chemistry was investigated on Limacina helicina, a key species of Arctic pelagic ecosystems. Individuals were kept in the laboratory under controlled pCO(2) levels of 280, 380, 550, 760 and 1020 mu atm and at control (0 degrees C) and elevated (4 degrees C) temperatures. The respiration rate was unaffected by pCO(2) at control temperature, but significantly increased as a function of the pCO(2) level at elevated temperature. pCO(2) had no effect on the gut clearance rate at either temperature. Precipitation of CaCO3, measured as the incorporation of Ca-45, significantly declined as a function of pCO(2) at both temperatures. The decrease in calcium carbonate precipitation was highly correlated to the aragonite saturation state. Even though this study demonstrates that pteropods are able to precipitate calcium carbonate at low aragonite saturation state, the results support the current concern for the future of Arctic pteropods, as the production of their shell appears to be very sensitive to decreased pH. A decline of pteropod populations would likely cause dramatic changes to various pelagic ecosystems.

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.

Abstract. Thecosome pteropods (shelled pelagic molluscs) can play an important role in the food web of various ecosystems and play a key role in the cycling of carbon and carbonate. Since they harbor an aragonitic shell, they could be very sensitive to ocean acidification driven by the increase of anthropogenic CO2 emissions. The impact of changes in the carbonate chemistry was investigated on Limacina helicina, a key species of Arctic ecosystems. Pteropods were kept in culture under controlled pH conditions corresponding to pCO2 levels of 350 and 760 μatm. Calcification was estimated using a fluorochrome and the radioisotope 45Ca. It exhibits a 28% decrease at the pH value expected for 2100 compared to the present pH value. This result supports the concern for the future of pteropods in a high-CO2 world, as well as of those species dependent upon them as a food resource. A decline of their populations would likely cause dramatic changes to the structure, function and services of polar ecosystems.