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The worldwide decline of coral reefs has renewed interest in coral communities at the edge of environmental limits because they have the potential to serve as resilience hotspots and climate change refugia, and can provide insights into how coral reefs might function in future ocean conditions. These coral communities are often referred to as marginal or extreme but few definitions exist and usage of these terms has therefore been inconsistent. This creates significant challenges for categorising these often poorly studied communities and synthesising data across locations. Furthermore, this impedes our understanding of how coral communities can persist at the edge of their environmental limits and the lessons they provide for future coral reef survival. Here, we propose that marginal and extreme coral communities are related but distinct and provide a novel conceptual framework to redefine them. Specifically, we define coral reef extremeness solely based on environmental conditions (i.e., large deviations from optimal conditions in terms of mean and/or variance) and marginality solely based on ecological criteria (i.e., altered community composition and/or ecosystem functioning). This joint but independent assessment of environmental and ecological criteria is critical to avoid common pitfalls where coral communities existing outside the presumed optimal conditions for coral reef development are automatically considered inferior to coral reefs in more traditional settings. We further evaluate the differential potential of marginal and extreme coral communities to serve as natural laboratories, resilience hotspots and climate change refugia, and discuss strategies for their conservation and management as well as priorities for future research. Our new classification framework provides an important tool to improve our understanding of how corals can persist at the edge of their environmental limits and how we can leverage this knowledge to optimise strategies for coral reef conservation, restoration and management in a rapidly changing ocean.

Aquatic research is often recognized as the last real adventure with dedicated scientists sailing on famous research vessels to even remotest areas like the Arctic or Antarctica taking samples with highly sophisticated scientific equipment. Such cruises or even more, scientists in submarines or ROVs are eye-catchers and get a high level of attention in the scientific, public and political community. In contrast, there is a comparatively small group of scientists which indeed physically enter the aquatic ecosystem and do science there - the group of diving scientists. Most surprisingly, this small group of scientists is sometimes not well recognized in science. Very often, scientific divers are assumed to take up their hobby and have mainly fun under water instead of doing serious scientific work. We - the scientific diver community - are often confronted with statements like “oh, you can be outside and can dive the whole day long, I have to stand the entire day in the lab. Your work must be like holydays”. Most of these colleagues have never been out the entire day in a bulky drysuit, spending hours and hours on a small boat while the colleague is under water doing fine tuned work in the three-dimensional space having the air to survive in tanks on the back. Often these colleagues have never spend 6 or more hours outside in a steady swell or with outside temperatures below 0°C of above 30°C and 100% humidity - without a private toilet - on a 6 m long RIB-boat.In this talk, we present the well harmonized European standards for doing excellent science under water, the respective European and German national bodies for scientific diving, as well as the required and recommended occupational safety standards and procedures for a successful, safe and efficient scientific work under water. In contrast, we also stress some possible reasons why doing science under water as diving scientist is, most surprisingly, much less accepted and established in science as doing aquatic science from “outside”, e.g. from a ship floating at the surface.

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.

Understanding the drivers of net coral reef calcium carbonate production is increasingly important as ocean warming, acidification, and other anthropogenic stressors threaten the maintenance of coral reef structures and the services these ecosystems provide. Despite intense research effort on coral reef calcium carbonate production, the inclusion of a key reef forming/accreting calcifying group, the crustose coralline algae (CCA), remains challenging both from a theoretical and practical standpoint. While corals are typically the primary reef builders of today, ongoing declines in coral cover due to a range of environmental perturbations will likely increase the relative importance of CCA and other non-scleractinian calcifying taxa to coral reef carbonate production. Here, we demonstrate that CCA are important carbonate producers that, under certain conditions, can match or even exceed the contribution of corals to coral reef carbonate production. Despite their importance, CCA are often inaccurately recorded in benthic surveys or even entirely missing from coral reef carbonate budgets. We outline several recommendations to improve the inclusion of CCA into such carbonate budgets under the ongoing climate crisis.

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.

Este artículo contiene 18 páginas, 4 figuras., Climate change is causing an increase in the frequency and intensity of marine heatwaves (MHWs) and mass mortality events (MMEs) of marine organisms are one of their main ecological impacts. Here, we show that during the 2015–2019 period, the Mediterranean Sea has experienced exceptional thermal conditions resulting in the onset of five consecutive years of widespread MMEs across the basin. These MMEs affected thousands of kilometers of coastline from the surface to 45 m, across a range of marine habitats and taxa (50 taxa across 8 phyla). Significant relationships were found between the incidence of MMEs and the heat exposure associated with MHWs observed both at the surface and across depths. Our findings reveal that the Mediterranean Sea is experiencing an acceleration of the ecological impacts of MHWs which poses an unprecedented threat to its ecosystems' health and functioning. Overall, we show that increasing the resolution of empirical observation is critical to enhancing our ability to more effectively understand and manage the consequences of climate change., This paper was supported by Euromarine. Joaquim Garrabou acknowledges the funding by the “Severo Ochoa Centre of Excellence” (CEX2019-000928-S), the MCIU/AEI/FEDER [HEATMED; RTI2018-095346-B-I00], Interreg-Med Programme MPA-Engage (1MED15_3.2_M2_337), the European Union Horizon 2020 research and innovation programme (Futuremares SEP-210597628). Nuria Teixido acknowledges the French National Research Agency (4Oceans-MOPGA grant, ANR-17-MPGA-0001) and internal funds from the Stazione Zoologica Anton Dohrn. Gil Rilov was supported by the Israel Ministry of Environmental Protection. Hocein Bazairi, Free Spinosa, and Vasilis Gerovasileiou acknowledge the funding by the MAVA Fondation (MedKeyHabitats I Project) and the European Commission (Ecap-MED II Project; projects implemented by UNEP/MAP-RAC/SPA). Alfonso Ramos was supported the CIESM “Tropical Signals,” Stelios Katsanevakis and Maria Sini were supported by the Project “Coastal Environment Observatory and Risk Management in Island Regions AEGIS+” (MIS 5047038), implemented within the Operational Programme “Competitiveness, Entrepreneurship and Innovation” (NSRF 2014–2020), co-financed by the Hellenic Government (Ministry of Development and Investments) and the European Union (European Regional Development Fund, Cohesion Fund), Stelios Katsanevakis, Maria Sini and Vasilis Gerovasileiou acknowledge the support of the MARISCA Project, co-funded by 85% by the EEA GRANTS, 2009–2014, and 15% by the Public Investments Programme (PIP) of the Hellenic Republic. Ivan Guala and Daniele Grech thanks thank the support of the project “Pinna nobilis—ricerca per la sopravvivenza: un'iniziativa di Citizen Science per tracciare la mortalità di massa di Pinna nobilis in Sardegna” project and the L/7 grant (CUP 87G17000070002) funded by the Regione Autonoma Sardegna. Jean-Baptiste Ledoux was funded by an assistant researcher contract framework of the RD Unit—UID/Multi/04423/2019—Interdisciplinary Centre of Marine and Environmental Research—financed by the European Regional Development Fund (ERDF) through COMPETE2020—Operational Program for Competitiveness and Internationalization (POCI) and national funds through FCT/MCTES (PIDDAC). This research was supported by the Strategic Funding UIDB/04423/2020 and UIDP/04423/2020 through national funds provided by the FCT—Foundation for Science and Technology and European Regional Development Fund (ERDF), in the framework of the program PT2020. Bernat Hereu and Cristina Linares acknowledge the support of the long-term monitoring programme of the catalan Natural Parks, funded by the Departament de Territori i Sostenibilitat of the Generalitat de Catalunya. Cristina Linares acknowledges the support of the ICREA Academia programme. David Díaz acknowledges the support the research grant CTM2016-77027-R of the Programa Estatal de Investigación, Desarrollo e Innovación Orientada a los Retos de la Sociedad and Program of Marines Strategies of Spain funded by MITERD. Jamila Ben Soussi was partially funded by the Fondation Albert 2 Monaco (MIMOSA Project) and the Tropical Signals Program of CIESM. Giovanni Chimienti was supported by the Italian Ministry of Education, University and Research (PON 2014–2020, AIM 1807508–1, Linea 1), by the Ente Parco Nazionale del Gargano (Research agreement with CoNISMa N. 21/2018), and by the National Geographic Society (Grant EC-176R-18). Nathaniel Bensoussan acknowledges financial support from the European Commission through the programme “Caroline Herschell” in the context of the action “Developing Downstream applications and services on BIO-PHYsical characterization of the seascape for COASTal management” (BIOPHYCOAST). Monica Montefalcone and Annalissa Azzola collected some of their data on MMEs in the frame of the project “Mare Caldo” funded by Greenpeace Italy. Núria Marbà acknowledges financial support from the Spanish Ministries of Economy and Competitiveness (CTM2012-32603, CGL2015-71809-P) and Ministerio de Ciencia, Innovación y Universidades (RTI2018-095441-B-C21). Diego K. Kersting acknowledges support by the postdoctoral fellowship programme Beatriu de Pinós funded by the Secretary of Universities and Research (Government of Catalonia) and the Horizon 2020 programme of research and innovation of the European Union under the Marie Sklodowska-Curie grant agreement No 801370.

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 CO2 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 pHcf in their CF, but this characteristic was not linked with changes in abundance. Within less variable low pH seawater, corals with limited reductions in pHcf 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 pHcf homeostasis. Overall, although our findings indicate that there might be an association between ecological success and greater pHcf 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.

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.

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

Ocean acidification (OA) is a major threat to the persistence of biogenic reefs throughout the world's ocean. Coralline algae are comprised of high magnesium calcite and have long been considered one of the most susceptible taxa to the negative impacts of OA. We summarize these impacts and explore the causes of variability in coralline algal responses using a review/qualitative assessment of all relevant literature, meta-analysis, quantitative assessment of critical responses, and a discussion of physiological mechanisms and directions for future research. We find that most coralline algae experienced reduced abundance, calcification rates, recruitment rates, and declines in pH within the site of calcification in laboratory experiments simulating OA or at naturally elevated CO2 sites. There were no other consistent physiological responses of coralline algae to simulated OA (e.g., photo-physiology, mineralogy, and survival). Calcification/growth was the most frequently measured parameters in coralline algal OA research, and our meta-analyses revealed greater declines in seawater pH were associated with significant decreases in calcification in adults and similar but nonsignificant trends for juveniles. Adults from the family Mesophyllumaceae also tended to be more robust to OA, though there was insufficient data to test similar trends for juveniles. OA was the dominant driver in the majority of laboratory experiments where other local or global drivers were assessed. The interaction between OA and any other single driver was often additive, though factors that changed pH at the surface of coralline algae (light, water motion, epiphytes) acted antagonistically or synergistically with OA more than any other drivers. With advances in experimental design and methodological techniques, we now understand that the physiology of coralline algal calcification largely dictates their responses to OA. However, significant challenges still remain, including improving the geographic and life-history spread of research effort and a need for holistic assessments of physiology.

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, multi-generational 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 trade-off 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.

Significance The growth of coral reefs is threatened by the dual stressors of ocean warming and acidification. Despite a wealth of studies assessing the impacts of climate change on individual taxa, projections of their impacts on coral reef net carbonate production are limited. By projecting impacts across 233 different locations, we demonstrate that the majority of coral reefs will be unable to maintain positive net carbonate production globally by the year 2100 under representative concentration pathways RCP4.5 and 8.5, while even under RCP2.6, coral reefs will suffer reduced accretion rates. Our results provide quantitative projections of how different climate change stressors will influence whole ecosystem carbonate production across coral reefs in all major ocean basins., 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.

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 ( $${p}_{\mathrm{CO}_{2}}$$ ) 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 $${p}_{\mathrm{CO}_{2}}$$ by determining their chemistry in the CF during a year-long experiment. We found no evidence of acclimatization to elevated $${p}_{\mathrm{CO}_{2}}$$ across any of the tested taxa. The effects of increasing seawater $${p}_{\mathrm{CO}_{2}}$$ 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. Calcifying marine species are threatened by ocean acidification. A year-long study of calcifying fluid chemistry shows that reef species, both corals and calcifying macroalgal species, were not able to adapt to ocean acidification.

International audience; 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.

High pCO2 habitats and their populations provide an unparalleled opportunity to assess how species may survive under future ocean acidification conditions, and help to reveal the traits that confer tolerance. Here we utilize a unique CO2 vent system to study the effects of exposure to elevated pCO2 on trait-shifts observed throughout natural populations of Astroides calycularis, an azooxanthellate scleractinian coral endemic to the Mediterranean. Unexpected shifts in skeletal and growth patterns were found. Colonies shifted to a skeletal phenotype characterized by encrusting morphology, smaller size, reduced coenosarc tissue, fewer polyps, and less porous and denser skeletons at low pH. Interestingly, while individual polyps calcified more and extended faster at low pH, whole colonies found at low pH site calcified and extended their skeleton at the same rate as did those at ambient pH sites. Transcriptomic data revealed strong genetic differentiation among local populations of this warm water species whose distribution range is currently expanding northward. We found excess differentiation in the CO2 vent population for genes central to calcification, including genes for calcium management (calmodulin, calcium-binding proteins), pH regulation (V-type proton ATPase), and inorganic carbon regulation (carbonic anhydrase). Combined, our results demonstrate how coral populations can persist in high pCO2 environments, making this system a powerful candidate for investigating acclimatization and local adaptation of organisms to global environmental change.

Crustose coralline algae play a crucial role in the building of reefs in the photic zones of nearshore ecosystems globally, and are highly susceptible to ocean acidification1–3. Nevertheless, the extent to which ecologically important crustose coralline algae can gain tolerance to ocean acidification over multiple generations of exposure is unknown. We show that, while calcification of juvenile crustose coralline algae is initially highly sensitive to ocean acidification, after six generations of exposure the effects of ocean acidification disappear. A reciprocal transplant experiment conducted on the seventh generation, where half of all replicates were interchanged across treatments, confirmed that they had acquired tolerance to low pH and not simply to laboratory conditions. Neither exposure to greater pH variability, nor chemical conditions within the micro-scale calcifying fluid internally, appeared to play a role in fostering this capacity. Our results demonstrate that reef-accreting taxa can gain tolerance to ocean acidification over multiple generations of exposure, suggesting that some of these cosmopolitan species could maintain their critical ecological role in reef formation. Crustose coralline algae help build coral reef structures through calcification, a process threatened under ocean acidification. Juvenile algae were highly sensitive on initial exposure to ocean acidification, but continued exposure over six generations showed a gain of tolerance.

Reef-building corals are surrounded by complex microenvironments (i.e. concentration boundary layers) that partially isolate them from the ambient seawater. Although the presence of such concentration boundary layers (CBLs) could potentially play a role in mitigating the negative impacts of climate change stressors, their role is poorly understood. Furthermore, it is largely unknown how heat stress-induced bleaching affects O2 and pH dynamics across the CBLs of coral, particularly in branching species. We experimentally exposed the common coral species Acropora aspera to heat stress for 13 d and conducted a range of physiological and daytime microsensor measurements to determine the effects of bleaching on O2 and pH gradients across the CBL. Heat stress equivalent to 24 degree heating days (3.4 degree heating weeks) resulted in visible bleaching and significant declines in photochemical efficiency, photosynthesis rates and photosynthesis to respiration (P/R) ratios, whereas dark respiration and calcification rates were not impacted. As a consequence, bleached A. aspera had significantly lower (− 13%) surface O2 concentrations during the day than healthy corals, with concentrations being lower than that of the ambient seawater, thus resulting in O2 uptake from the seawater. Furthermore, we show here that Acropora, and potentially branching corals in general, have among the lowest surface pH elevation of all corals studied to date (0.041 units), which could contribute to their higher sensitivity to ocean acidification. Additionally, bleached A. aspera no longer elevated their surface pH above ambient seawater values and, therefore, had essentially no [H+] CBL. These findings demonstrate that heat stress-induced bleaching has negative effects on pH elevation and [H+] CBL thickness, which may increase the overall susceptibility of coral to the combined impacts of ocean acidification and warming.

The threat represented by ocean acidification (OA) for coral reefs has received considerable attention because of the sensitivity of calcifiers to changing seawater carbonate chemistry. However, most studies have focused on the organismic response of calcification to OA, and only a few have addressed community-level effects, or investigated parameters other than calcification, such as photosynthesis. Light (photosynthetically active radiation, PAR) is a driver of biological processes on coral reefs, and the possibility that these processes might be perturbed by OA has important implications for community function. Here we investigate how CO2 enrichment affects the relationships between PAR and community net O2 production (Pnet), and between PAR and community net calcification (Gnet), using experiments on three coral communities constructed to match (i) the back reef of Mo'orea, French Polynesia, (ii) the fore reef of Mo'orea, and (iii) the back reef of O'ahu, Hawaii. The results were used to test the hypothesis that OA affects the relationship between Pnet and Gnet. For the three communities tested, pCO2 did not affect the Pnet–PAR relationship, but it affected the intercept of the hyperbolic tangent curve fitting the Gnet–PAR relationship for both reef communities in Mo'orea (but not in O'ahu). For the three communities, the slopes of the linear relationships between Pnet and Gnet were not affected by OA, although the intercepts were depressed by the inhibitory effect of high pCO2 on Gnet. Our result indicates that OA can modify the balance between net calcification and net photosynthesis of reef communities by depressing community calcification, but without affecting community photosynthesis.

Turf algal assemblages are ubiquitous primary producers on coral reefs, but little is known about the response of this diverse group to ocean acidification (OA) across different temperatures. We tested the hypothesis that CO2 influences the functional response of epilithic and endolithic turf assemblages to increasing temperature. Replicate carbonate plugs covered by turf were collected from the reef and exposed to ambient and high pCO2 (1000 µatm) conditions for 3 weeks. Each pCO2 treatment was replicated across six temperatures (24.0–31.5 °C) that spanned the full seasonal temperature range on a fringing reef in Moorea, French Polynesia, and included one warming treatment (3 °C above daily average temperatures). Temperature and CO2 enrichment had complex, and sometimes interactive, effects on turf metabolism and growth. Photosynthetic and respiration rates were enhanced by increasing temperature, with an interactive effect of CO2 enrichment. Photosynthetic rates were amplified by high CO2 in the warmest temperatures, while the increase in respiration rates with temperature were enhanced under ambient CO2. Epilithic turf growth rates were not affected by temperature, but increased in response to CO2 enrichment. We found that CO2 and temperature interactively affected the endolithic assemblage, with the highest growth rates under CO2 enrichment, but only at the warmest temperatures. These results demonstrate how OA may influence algal physiology and growth across a range of ecologically relevant temperatures, and indicate that the effects of CO2 enrichment on coral-reef turf assemblages can be temperature dependent. The complex effects of CO2 enrichment and temperature across a suite of algal responses illustrates the importance of incorporating multiple stressors into global change experiments.

International audience; 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 poses a serious threat to marine calcifying organisms, yet experimental and field studies have found highly diverse responses among species and environments. Our understanding of the underlying drivers of differential responses to ocean acidification is currently limited by difficulties in directly observing and quantifying the mechanisms of bio‐calcification. Here, we present Raman spectroscopy techniques for characterizing the skeletal mineralogy and calcifying fluid chemistry of marine calcifying organisms such as corals, coralline algae, foraminifera, and fish (carbonate otoliths). First, our in vivo Raman technique is the ideal tool for investigating non‐classical mineralization pathways. This includes calcification by amorphous particle attachment, which has recently been controversially suggested as a mechanism by which corals resist the negative effects of ocean acidification. Second, high‐resolution ex vivo Raman mapping reveals complex banding structures in the mineralogy of marine calcifiers, and provides a tool to quantify calcification responses to environmental variability on various timescales from days to years. We describe the new insights into marine bio‐calcification that our techniques have already uncovered, and we consider the wide range of questions regarding calcifier responses to global change that can now be proposed and addressed with these new Raman spectroscopy tools., Newly developed in vivo and high‐resolution Raman spectroscopy analyses of marine calcifying organisms enable characterization of their mineralogy and calcification mechanisms. Insights from these new Raman spectroscopy tools are helping us understand the sensitivity of these important organisms to ocean warming and acidification.

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.

The effects of ocean acidification (OA) on coral reefs have been studied thoroughly with a focus on the response of calcification of corals and calcified algae. However, there are still large gaps in our knowledge of the effects of OA on photosynthesis and respiration of these organisms. Comparisons among species and determination of the functional relationships between pCO2 and either photosynthesis or respiration are difficult using previously published data, because experimental conditions typically vary widely between studies. Here, we tested the response of net photosynthesis, gross photosynthesis, dark respiration, and light-enhanced dark respiration (LEDR) of eight coral taxa and seven calcified alga taxa to six different pCO2 levels (from 280 to 2000 µatm). Organisms were maintained during 7–10 days incubations in identical conditions of light, temperature, and pCO2 to facilitate comparisons among species. Net photosynthesis was not affected by pCO2 in seven of eight corals or any of the algae; gross photosynthesis did not respond to pCO2 in six coral taxa and six algal taxa; dark respiration also was unaffected by pCO2 in six coral and six algae; and LEDR did not respond to pCO2 in any of the tested species. Overall, our results show that pCO2 levels up to 2000 µatm likely will not fertilize photosynthesis or modify respiration rates of most of the main calcifiers on the back reef of Moorea, French Polynesia.

‘Multiple drivers’ (also termed ‘multiple stressors’) is the term used to describe the cumulative effects of multiple environmental factors on organisms or ecosystems. Here, we consider ocean acidification as a multiple driver because many inorganic carbon parameters are changing simultaneously, including total dissolved inorganic carbon, CO2, HCO3–, CO32–, H+ and CaCO3 saturation state. With the rapid expansion of ocean acidification research has come a greater understanding of the complexity and intricacies of how these simultaneous changes to the seawater carbonate system are affecting marine life. We start by clarifying key terms used by chemists and biologists to describe the changing seawater inorganic carbon system. Then, using key groups of non-calcifying (fish, seaweeds, diatoms) and calcifying (coralline algae, coccolithophores, corals, molluscs) organisms, we consider how various physiological processes are affected by different components of the carbonate system.

Ocean acidification (OA) poses a severe threat to tropical coral reefs, yet much of what is know about these effects comes from individual corals and algae incubated in isolation under high pCO2. Studies of similar effects on coral reef communities are scarce. To investigate the response of coral reef communities to OA, we used large outdoor flumes in which communities composed of calcified algae, corals, and sediment were combined to match the percentage cover of benthic communities in the shallow back reef of Moorea, French Polynesia. Reef communities in the flumes were exposed to ambient (~ 400 μatm) and high pCO2 (~ 1300 μatm) for 8 weeks, and calcification rates measured for the constructed communities including the sediments. Community calcification was reduced by 59% under high pCO2, with sediment dissolution explaining ~ 50% of this decrease; net calcification of corals and calcified algae remained positive but was reduced by 29% under elevated pCO2. These results show that, despite the capacity of coral reef calcifiers to maintain positive net accretion of calcium carbonate under OA conditions, reef communities might transition to net dissolution as pCO2 increases, particularly at night, due to enhanced sediment dissolution.

International audience; 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.

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 geochemical techniques based on boron systematics and Raman spectroscopy, which together provide the first constraints on the sensitivity of coral calcifying fluid calcium concentrations ( ) to changing seawater pH. In response to simulated end-of-century pH conditions, Pocillopora damicornis increased to as much as 25% above that of seawater and maintained constant calcification rates. Conversely, Acropora youngei displayed less control over , and its calcification rates strongly declined at lower seawater pH. Although the role of in driving calcification has often been neglected, increasing 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.

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 (pHcf) was determined using δ11B systematics, and for both species pHcf declined slightly with seawater pH, with the decrease being more pronounced in P. damicornis. The dissolved inorganic carbon concentration at the site of calcification (DICcf) was estimated using geochemical proxies (B/Ca and δ11B) 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 (Ωcf) with OA was partially moderated by the DICcf increase. These results highlight that while pHcf, DICcf and Ωcf are important in the mineralization process, some corals are able to maintain their calcification rates despite shifts in their calcifying fluid carbonate chemistry.

The threat represented by ocean acidification (OA) for coral reef has received considerable attention because of the sensitivity of calcifiers to changing water carbonate chemistry. However most studies have focused on the organismic response of calcification to OA, and only a few have addressed community-level effects, or investigated parameters other than calcification, such as photosynthesis. Light (Photosynthetically Active Radiation, PAR) is a driver of biological processes on coral reefs, and the possibility that these processes might be perturbed by OA has important implications for community function. Here we investigate how CO2 enrichment affects the relationships between PAR and community net O2 production (Pnet), and between PAR and community net calcification (Gnet), using experiments on three coral communities constructed to match (i) the back reef of Moorea, French Polynesia, (ii) the fore reef of Moorea, and (iii) the reef flat of Oahu, Hawaii. The results were used to test the hypothesis that OA affects the relationship between Pnet and Gnet. For the three communities tested, pCO2 did not affect the Pnet-PAR relationship, but it affected the intercept of the hyperbolic tangent curve fitting the Gnet-PAR relationship for both reef communities in Moorea (but not in Oahu). For the three communities, the slopes of the linear relationships between Pnet and Gnet were not affected by OA, although the intercepts were depressed by the inhibitory effect of high pCO2 on Gnet. Our result indicates that OA can modify the balance between net calcification and net photosynthesis of reef communities by depressing community calcification, but without affecting community photosynthesis.

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.