Living under natural conditions of ocean acidification entails energy expenditure and oxidative stress in a mussel species.

We investigated the health conditions of the Mediterranean mussel Mytilus galloprovincialis recruited in the CO2 vents system of Castello Aragonese at Ischia Island (Mediterranean Sea). Individuals of M. galloprovincialis were sampled in three sites along the pH gradient (8.10, 7.7 and up to <7.4). Untargeted metabolomics and biochemical endpoints related to energetic metabolism, oxidative stress/damage, neurotoxicity and immune defense were analyzed. Corrosion of the valves occurred at low pH. A separation of the metabolome was observed along the pH gradient. Metabolites belonging to amino acids, nucleosides, lipids and organic osmolytes were significantly reduced in the organisms from the most acidified sites. The content of reactive oxygen species and the activity of glutathione peroxidase were reduced in organisms from the acidified sites compared to ambient pH, and no oxidative damage was induced. Overall results suggested the presence of an energy cost underpinning long-term survival in acidified conditions for this species.

Characterization of an undocumented CO2 hydrothermal vent system in the Mediterranean Sea: Implications for ocean acidification forecasting.

A previously undocumented shallow water hydrothermal field from Sicily (Southern Tyrrhenian Sea, Italy) is here described, based on a multidisciplinary investigation. The field, covering an area of nearly 8000 m2 and a depth from the surface to -5 m, was explored in June 2021 to characterise the main physico-chemical features of the water column, describe the bottom topography and features, and identify the main megabenthic and nektonic species. Twenty sites were investigated to characterise the carbonate system. Values of pH ranged between 7.84 and 8.04, ΩCa between 3.68 and 5.24 and ΩAr from 2.41 to 3.44. Geochemical analyses of hydrothermal gases revealed a dominance of CO2 (98.1%) together with small amounts of oxygen and reactive gases. Helium isotope ratios (R/Ra = 2.51) and δ13CCO2 suggest an inorganic origin of hydrothermal degassing of CO2 and the ascent of heat and deep-seated magmatic fluids to the surface. Visual census of fishes and megabenthos (mainly sessile organisms) allowed the identification of 64 species, four of which are protected by the SPA/BIO Protocol and two by the International Union for Conservation of Nature. The macroalgae Halopteris scoparia and Jania rubens and the sponge Sarcotragus sp. were the dominant taxa in the area, while among fishes Coris julis and Chromis chromis were the most abundant species. This preliminary investigation of San Giorgio vent field suggests that the site could be of interest and suitable for future experimental studies of ocean acidification.

Functional changes across marine habitats due to ocean acidification.

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

Effects of anthropogenic pressures on the seagrass Halophila stipulacea and its associated macrozoobenthic communities in the northern Gulf of Aqaba.

Halophila stipulacea is a tropical seagrass species, native to the Red Sea, Persian Gulf, and Indian Ocean, while invasive to the Mediterranean and Caribbean Seas. The benthic fauna assemblages associated with H. stipulacea in its native habitats and the potential effects of anthropogenic stressors on these assemblages remain unknown. We compared meadow characteristics, associated fauna assemblages and trophic niche structures of H. stipulacea from an impacted and a pristine site in the northern Red Sea. Seagrass cover and biomass were higher in the impacted site, however, the associated fauna community was more abundant and diverse in the pristine site. Both meadows showed comparable trophic niches based on stable isotope analysis. This study provides first insights into the macrozoobenthos associated with H. stipulacea in its native habitat and highlights the importance of better understanding the relationship between seagrasses and their associated biota and the potential effects of urbanization on this relationship.

Unraveling cellular and molecular mechanisms of acid stress tolerance and resistance in marine species: New frontiers in the study of adaptation to ocean acidification.

Since the industrial revolution, fossil fuel combustion has led to a 30 %-increase of the atmospheric CO2 concentration, also increasing the ocean partial CO2 pressure. The consequent lowered surface seawater pH is termed ocean acidification (OA) and severely affects marine life on a global scale. Cellular and molecular responses of marine species to lowered seawater pH have been studied but information on the mechanisms driving the tolerance of adapted species to comparatively low seawater pH is limited. Such information may be obtained from species inhabiting sites with naturally low water pH that have evolved remarkable abilities to tolerate such conditions. This review gathers information on current knowledge about species naturally facing low water pH conditions and on cellular and molecular adaptive mechanisms enabling the species to survive under, and even benefit from, adverse pH conditions. Evidences derived from case studies on naturally acidified systems and on resistance mechanisms will guide predictions on the consequences of future adverse OA scenarios for marine biodiversity.

Mediterranean Sea shelters for the gold coral Savalia savaglia (Bertoloni, 1819): An assessment of potential distribution of a rare parasitic species.

Savalia savaglia is an ecosystem engineer listed as Near-Threatened by the IUCN, even though effective management and proper monitoring efforts to assess its distribution is still lacking. The record of large, long-established colonies can indicate the occurrence of areas with limited human local pressure. These areas may be considered as proxies for the creation of baselines of reference useful to design restoration strategies. The aim of this work was to update the distribution of S. savaglia Mediterranean populations to develop an Ecological Niche Model, highlighting potential areas for future monitoring programs. Occurrence data were collected and harmonized into a single dataset using the scientific literature and validated observations to feed a presence-only MaxEnt model, obtaining a basin-level potential distribution of the species. The results of our study can support decision-makers in marine spatial planning measures including the preservation of mesophotic environments and prioritizing areas for conservation.

Acclimation to low pH does not affect the thermal tolerance of Arbacia lixula progeny.

As the ocean warms, the thermal tolerance of marine invertebrates is key to determining their distributional change, where acclimation to low pH may impact the thermal range of optimal development. We compared thermal tolerance of progeny from a low pH-acclimated sea urchin (Arbacia lixula) population from the CO2 vents of Ischia (Italy) and a nearby population living at ambient pH. The percentages of normally developing gastrulae and two-armed larvae were determined across 10 temperatures representing present and future temperature conditions (16-34°C). Vent-acclimated sea urchins showed a greater percentage of normal development at 24 h, with a larger optimal developmental temperature range than control sea urchins (12.3°C versus 5.4°C range, respectively). At 48 h, upper lethal temperatures for 50% survival with respect to ambient temperatures were similar between control (+6.8°C) and vent (+6.2°C) populations. Thus, acclimation to low pH did not impact the broad thermal tolerance of A. lixula progeny. With A. lixula's barrens-forming abilities, its wide thermotolerance and its capacity to acclimate to low pH, this species will continue to be an important ecological engineer in Mediterranean macroalgal ecosystems in a changing ocean.

Surviving in a changing ocean. Tolerance to acidification might affect the susceptibility of polychaetes to chemical contamination.

This study aimed to assess the combined effects of ocean acidification (OA) and pollution to the polychaete Syllis prolifera inhabiting the CO2 vent system of the Castello Aragonese (Ischia Island, Italy). We investigated the basal activities of antioxidant enzymes in organisms from the acidified site and from an ambient-pH control site in two different periods of the year. Results showed a limited influence of acidified conditions on the functionality of the antioxidant system. We then investigated the responsiveness of individuals living inside the CO2 vent compared to those from the control to face exposure to acetone and copper. Results highlighted a higher susceptibility of organisms from the vent to acetone and a different response of antioxidant enzymes in individuals from the two sites. Conversely, a higher tolerance to copper was observed in polychaetes from the acidified-site with respect to controls, but any significant oxidative stress was induced at sublethal concentrations.

Resilient consumers accelerate the plant decomposition in a naturally acidified seagrass ecosystem.

Anthropogenic stressors are predicted to alter biodiversity and ecosystem functioning worldwide. However, scaling up from species to ecosystem responses poses a challenge, as species and functional groups can exhibit different capacities to adapt, acclimate, and compensate under changing environments. We used a naturally acidified seagrass ecosystem (the endemic Mediterranean Posidonia oceanica) as a model system to examine how ocean acidification (OA) modifies the community structure and functioning of plant detritivores, which play vital roles in the coastal nutrient cycling and food web dynamics. In seagrass beds associated with volcanic CO2 vents (Ischia, Italy), we quantified the effects of OA on seagrass decomposition by deploying litterbags in three distinct pH zones (i.e., ambient, low, extreme low pH), which differed in the mean and variability of seawater pH. We replicated the study in two discrete vents for 117 days (litterbags sampled on day 5, 10, 28, 55, and 117). Acidification reduced seagrass detritivore richness and diversity through the loss of less abundant, pH-sensitive species but increased the abundance of the dominant detritivore (amphipod Gammarella fucicola). Such compensatory shifts in species abundance caused more than a threefold increase in the total detritivore abundance in lower pH zones. These community changes were associated with increased consumption (52%-112%) and decay of seagrass detritus (up to 67% faster decomposition rate for the slow-decaying, refractory detrital pool) under acidification. Seagrass detritus deployed in acidified zones showed increased N content and decreased C:N ratio, indicating that altered microbial activities under OA may have affected the decay process. The findings suggest that OA could restructure consumer assemblages and modify plant decomposition in blue carbon ecosystems, which may have important implications for carbon sequestration, nutrient recycling, and trophic transfer. Our study highlights the importance of within-community response variability and compensatory processes in modulating ecosystem functions under extreme global change scenarios.

The Nereid on the rise: Platynereis as a model system.

The Nereid Platynereis dumerilii (Audouin and Milne Edwards (Annales des Sciences Naturelles 1:195-269, 1833) is a marine annelid that belongs to the Nereididae, a family of errant polychaete worms. The Nereid shows a pelago-benthic life cycle: as a general characteristic for the superphylum of Lophotrochozoa/Spiralia, it has spirally cleaving embryos developing into swimming trochophore larvae. The larvae then metamorphose into benthic worms living in self-spun tubes on macroalgae. Platynereis is used as a model for genetics, regeneration, reproduction biology, development, evolution, chronobiology, neurobiology, ecology, ecotoxicology, and most recently also for connectomics and single-cell genomics. Research on the Nereid started with studies on eye development and spiralian embryogenesis in the nineteenth and early twentieth centuries. Transitioning into the molecular era, Platynereis research focused on posterior growth and regeneration, neuroendocrinology, circadian and lunar cycles, fertilization, and oocyte maturation. Other work covered segmentation, photoreceptors and other sensory cells, nephridia, and population dynamics. Most recently, the unique advantages of the Nereid young worm for whole-body volume electron microscopy and single-cell sequencing became apparent, enabling the tracing of all neurons in its rope-ladder-like central nervous system, and the construction of multimodal cellular atlases. Here, we provide an overview of current topics and methodologies for P. dumerilii, with the aim of stimulating further interest into our unique model and expanding the active and vibrant Platynereis community.

The Pandora's box: Morphological diversity within the genus emAmphiglena/em Claparède, 1864 (Sabellidae, Annelida) in the Mediterranean Sea, with description of nine new species.

We report the description of nine new taxa of sabellid polychaetes belonging to the genus Amphiglena, of which diversity in the Mediterranean Sea has been widely underestimated. Examined material derived from both new collections along the Italian coast, including four CO2 vents/hydrothermal systems, and from a re-examination of older material previously attributed to A. mediterranera (Leydig, 1851) which was so far the only species of the genus reported for the Mediterranean area. The analysis revealed the presence of different taxa also consistent with a previous molecular analysis conducted on material from the Gulf of Naples and the Salento coast (Ionian Sea). This led to an increase in the number of species in the genus and to highlight the occurrence in the Mediterranean Sea of a high diversity within the genus. A key to the Mediterranean Sea species of Amphiglena is also provided. Some taxa, however, remain for the moment undescribed due to the poor preservation of the old material, and the lack of the type material for this taxon. A major revision of all the Mediterranean material previously attributed to A. mediterranea from both morphological and molecular points of view is needed.

Boosted fish abundance associated with Posidonia oceanica meadows in temperate shallow CO2 vents.

Ocean acidification (OA) may induce major shifts in the structure and function of coastal marine ecosystems. Studies in volcanic CO2 vents, where seawater is naturally acidified, have reported an overall simplification of fish assemblages structure, while some primary producers are likely to increase their biomass under elevated concentration of CO2. Here we used temperate shallow CO2 vents located around the coast of Ischia island (Italy) to assess the effects of OA on necto-benthic fish assemblages associated with the foundation seagrass species Posidonia oceanica in the Mediterranean Sea. We compared P. oceanica meadow structure, its epiphytic community and the associated fish assemblage structure and diversity at vents with low pH sites and reference sites with ambient pH using underwater visual census strip transects, in two seasons (fall 2018 and summer 2019). Data were analysed using both univariate and multivariate statistical techniques. Results showed greater P. oceanica habitat complexity (i.e. shoot density) and lower abundance of epiphytic calcareous species (e.g. coralline algae) at the vents than reference sites. Total abundance of adult and juvenile fish was higher at vents than reference sites, while no differences were found for species richness and composition. Overall, the herbivore Sarpa salpa stands out among the species contributing the most to dissimilarity between vents and reference sites, showing higher abundances under OA conditions. This pattern could be explained by the combined effect of a positive response to the higher structural meadows complexity and the greater seagrasses palatability/nutritional value occurring at the vents, which may help herbivores to withstand the higher energetic cost to live under high pCO2/low pH conditions. Our results indicate that necto-benthic fish assemblages associated with the Mediterranean P. oceanica ecosystem may cope with OA under the CO2 emission scenarios forecasted for the end of this century.

Ocean acidification causes variable trait-shifts in a coral species.

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.

Habitat and benthic diversity in the bay of Bagnoli and surrounding areas (Gulf of Naples, Italy): A historical baseline for environmental restoration.

The aim of the study is to provide a synthesis on the biodiversity of zoobenthic species and benthic habitat distribution of Site of National Interest (SNI) of Bagnoli-Coroglio (Gulf of Naples, Tyrrhenian Sea, Italy), which represents one of the priority areas selected at National level for habitat restoration, and is located within the Gulf of Pozzuoli (a large Bay at the northern part of the Gulf of Naples). The work provides a dataset, covering the entire Gulf of Pozzuoli, and obtained consulting several sources of information, from historical to recent publications, and grey literature, aimed at the production of a check-list of species, and reconstruction map of the main marine habitats, in order to achieve a synoptic overview of the historical and recent zoobenthic fauna, as well as distribution of habitats, in order to serve as reference point for any future restoration plan in the area. Relevant information regarding the study area was found in 67 out of more than 250 sources consulted. Overall, 813 species of benthic organisms were recorded, summing up a total of 1006 records. Among them, 148 species were reported in the pre-industrial period (prior to 1911), 361 species during the industrial period (from 1911 to 1991), and 381 species in the course of the post-industrial period (from 1992 up to the present day). No differences in biodiversity or distribution of individual species were directly attributable to the industrial activities in the study area. Such a finding is possibly due to different sampling effort among periods and lack of quantitative data for the majority of the recorded taxa. A mosaic of various habitat and biocoenoses were documented in the zones (shallow and deep hard bottoms, soft-bottoms with different sediment types, seagrass meadows). An overall reduction of the cover and a higher habitat fragmentation was documented for seagrass meadows (mainly Posidonia oceanica) over time. Given that regression of this seagrass species is common in a much wider extent than that covered by the study area, the trend here observed is probably due to multiple impacts from different human activities, including the industrial one at the Bagnoli SNI. The present study highlights that the SNI area is placed in a wider area representing a mosaic of different habitat types, which can provide donor populations of both habitat formers (sponges, gorgonians, scleractinians, bryozoans and mollusks), and seagrasses. These organisms are potentially relevant in implementing restoration measures aimed to improve the ecological status of this post-industrial area.

Patterns in microbiome composition differ with ocean acidification in anatomic compartments of the Mediterranean coral Astroides calycularis living at CO2 vents.

Coral microbiomes, the complex microbial communities associated with the different anatomic compartments of the coral, provide important functions for the host's survival, such as nutrient cycling at the host's surface, prevention of pathogens colonization, and promotion of nutrient uptake. Microbiomes are generally referred to as plastic entities, able to adapt their composition and functionality in response to environmental change, with a possible impact on coral acclimatization to phenomena related to climate change, such as ocean acidification. Ocean sites characterized by natural gradients of pCO2 provide models for investigating the ability of marine organisms to acclimatize to decreasing seawater pH. Here we compared the microbiome of the temperate, shallow water, non-symbiotic solitary coral Astroides calycularis that naturally lives at a volcanic CO2 vent in Ischia Island (Naples, Italy), with that of corals living in non-acidified sites at the same island. Bacterial DNA associated with the different anatomic compartments (mucus, tissue and skeleton) of A. calycularis was differentially extracted and a total of 68 samples were analyzed by 16S rRNA gene sequencing. In terms of phylogenetic composition, the microbiomes associated with the different coral anatomic compartments were different from each other and from the microbial communities of the surrounding seawater. Of all the anatomic compartments, the mucus-associated microbiome differed the most between the control and acidified sites. The differences detected in the microbial communities associated to the three anatomic compartments included a general increase in subdominant bacterial groups, some of which are known to be involved in different stages of the nitrogen cycle, such as potential nitrogen fixing bacteria and bacteria able to degrade organic nitrogen. Our data therefore suggests a potential increase of nitrogen fixation and recycling in A. calycularis living close to the CO2 vent system.

Responses of sea urchin larvae to field and laboratory acidification.

Understanding the extent to which laboratory findings of low pH on marine organisms can be extrapolated to the natural environment is key toward making better projections about the impacts of global change on marine ecosystems. We simultaneously exposed larvae of the sea urchin Arbacia lixula to ocean acidification in laboratory and natural CO2 vents and assessed the arm growth response as a proxy of net calcification. Populations of embryos were simultaneously placed at both control and volcanic CO2 vent sites in Ischia (Italy), with a parallel group maintained in the laboratory in control and low pH treatments corresponding to the mean pH levels of the field sites. As expected, larvae grown at constant low pH (pHT 7.8) in the laboratory exhibited reduced arm growth, but counter to expectations, the larvae that developed at the low pH vent site (pHT 7.33-7.99) had the longest arms. The larvae at the control field site (pHT 7.87-7.99) grew at a similar rate to laboratory controls. Salinity, temperature, oxygen and flow regimes were comparable between control and vent sites; however, chlorophyll a levels and particulate organic carbon were higher at the vent site than at the control field site. This increased food availability may have modulated the effects of low pH, creating an opposite calcification response in the laboratory from that in the field. Divergent responses of the same larval populations developing in laboratory and field environments show the importance of considering larval phenotypic plasticity and the complex interactions among decreased pH, food availability and larval responses.

Resistance of seagrass habitats to ocean acidification via altered interactions in a tri-trophic chain.

Despite the wide knowledge about prevalent effects of ocean acidification on single species, the consequences on species interactions that may promote or prevent habitat shifts are still poorly understood. Using natural CO2 vents, we investigated changes in a key tri-trophic chain embedded within all its natural complexity in seagrass systems. We found that seagrass habitats remain stable at vents despite the changes in their tri-trophic components. Under high pCO2, the feeding of a key herbivore (sea urchin) on a less palatable seagrass and its associated epiphytes decreased, whereas the feeding on higher-palatable green algae increased. We also observed a doubled density of a predatory wrasse under acidified conditions. Bottom-up CO2 effects interact with top-down control by predators to maintain the abundance of sea urchin populations under ambient and acidified conditions. The weakened urchin herbivory on a seagrass that was subjected to an intense fish herbivory at vents compensates the overall herbivory pressure on the habitat-forming seagrass. Overall plasticity of the studied system components may contribute to prevent habitat loss and to stabilize the system under acidified conditions. Thus, preserving the network of species interactions in seagrass ecosystems may help to minimize the impacts of ocean acidification in near-future oceans.

Living in future ocean acidification, physiological adaptive responses of the immune system of sea urchins resident at a CO2 vent system.

The effects of ocean acidification, a major anthropogenic impact on marine life, have been mainly investigated in laboratory/mesocosm experiments. We used the CO2 vents at Ischia as a natural laboratory to study the long-term effects of ocean acidification on the sea urchin Paracentrotus lividus population resident in low-pH (7.8 ±â€¯0.2) compared to that at two control sites (pH 8.02 ±â€¯0.00; 8.02 ±â€¯0.01). The novelty of the present study is the analysis of the sea urchin immune cells, the sentinels of environmental stress responses, by a wide-ranging approach, including cell morphology, biochemistry and proteomics. Immune cell proteomics showed that 311 proteins were differentially expressed in urchins across sites with a general shift towards antioxidant processes in the vent urchins. The vent urchin immune cells showed higher levels of total antioxidant capacity, up-regulation of phagosome and microsomal proteins, enzymes of ammonium metabolism, amino-acid degradation, and modulation of carbon metabolism proteins. Lipid-hydroperoxides and nitric oxide levels were not different in urchins from the different sites. No differences in the coelomic fluid pH, immune cell composition, animal respiration, nitrogen excretion and skeletal mineralogy were observed. Our results reveal the phenotypic plasticity of the immune system of sea urchins adapted to life at vent site, under conditions commensurate with near-future ocean acidification projections.

Effects of short-term and long-term exposure to ocean acidification on carbonic anhydrase activity and morphometric characteristics in the invasive polychaete Branchiomma boholense (Annelida: Sabellidae): A case-study from a CO2 vent system.

The aim of this study was to test the effects of short- and long-term exposure to high pCO2 on the invasive polychaete Branchiomma boholense (Grube, 1878), (Sabellidae), through the implementation of a transplant experiment at the CO2 vents of the Castello Aragonese at the island of Ischia (Italy). Analysis of carbonic anhydrase (CA) activity, protein tissue content and morphometric characteristics were performed on transplanted individuals (short-term exposure) as well as on specimens resident to both normal and low pH/high pCO2 environments (long-term exposure). Results obtained on transplanted worms showed no significant differences in CA activity between individuals exposed to control and acidified conditions, while a decrease in weight was observed under short-term acclimatization to both control and low pH, although at low pH the decrease was more pronounced (∼20%). As regard individuals living under chronic exposure to high pCO2, the morphometric results revealed a significantly lower (70%) wet weight of specimens from the vents with respect to animals living in high pH/low pCO2 areas. Moreover, individuals living in the Castello vents showed doubled values of enzymatic activity and a significantly higher (50%) protein tissue content compared to specimens native from normal pH/low pCO2. The results of this study demonstrated that B. boholense is inclined to maintain a great homeostatic capacity when exposed to low pH, although likely at the energetic expense of other physiological processes such as growth, especially under chronic exposure to high pCO2.

Functional biodiversity loss along natural CO2 gradients.

The effects of environmental change on biodiversity are still poorly understood. In particular, the consequences of shifts in species composition for marine ecosystem function are largely unknown. Here we assess the loss of functional diversity, i.e. the range of species biological traits, in benthic marine communities exposed to ocean acidification (OA) by using natural CO2 vent systems. We found that functional richness is greatly reduced with acidification, and that functional loss is more pronounced than the corresponding decrease in taxonomic diversity. In acidified conditions, most organisms accounted for a few functional entities (i.e. unique combination of functional traits), resulting in low functional redundancy. These results suggest that functional richness is not buffered by functional redundancy under OA, even in highly diverse assemblages, such as rocky benthic communities.