Description: We present measured and estimated seawater physiochemical parameters at CO2 vents and ambient pH sites along the coast of Ischia (Italy) across water depths from 1 m to 40 m. We characterized the physical and chemical parameters in four CO2 venting sites (vent 1 to vent 4) and reference sites with ambient pH with no venting activity (two reference sites for each CO2 vent). SeaFETTM Ocean pH sensors (Satlantic) were deployed to quantify variation in pH at the CO2 vents and their corresponding ambient pH sites at the same depths where benthic surveys were performed from May to October 2019. Before deployment, the SeaFETs were calibrated with ambient pH water. The mean offset between calibration samples and calibrated SeaFET pH was ± 0.006 units (n= 44 water samples), indicating a high-quality pH dataset. The pH sensors were deployed in the recently discovered CO2 vents (Vent 2, Vent 3, Vent 4) and one of the corresponding reference areas with no visible vent activity during the same period (ambient 2a, ambient 3a, ambient 4a). The pH and seawater physicochemical parameters for vent 1 and their corresponding ambient sites (site Castello Aragonese) are also included here. This data from the Castello Aragonese site was originally reported by Kroeker et al., 2011. Discrete water samples were collected using Niskin bottles at the vent and reference areas with ambient pH to measure: i) the carbonate system parameters during the pH sensor deployment, and ii) the dissolved inorganic nutrients. Salinity was measured using a CTD (CTD Sea-Bird Electronics SBE 19 Plus Seacat). Discrete water samples for total alkalinity (AT) were collected within 0.25 m of the pH sensors using standard operating protocols. Precision of the AT measurements of CRMs was 〈 2 μmol kg-1 from nominal values. AT and pHT along with in situ temperature and depth were used to determine the remaining carbonate system parameters for each sampling period using the R package seacarb v3.2.12 (Gattuso et al., 2023).

Description: Ocean acidification is predicted to impact all areas of the oceans and affect a diversity of marine organisms. However, the diversity of responses among species prevents clear predictions about the impact of acidification at the ecosystem level. Here, we used shallow water CO2 vents in the Mediterranean Sea as a model system to examine emergent ecosystem responses to ocean acidification in rocky reef communities. We assessed in situ benthic invertebrate communities in three distinct pH zones (ambient, low, and extreme low), which differed in both the mean and variability of seawater pH along a continuous gradient. We found fewer taxa, reduced taxonomic evenness, and lower biomass in the extreme low pH zones. However, the number of individuals did not differ among pH zones, suggesting that there is density compensation through population blooms of small acidification-tolerant taxa. Furthermore, the trophic structure of the invertebrate community shifted to fewer trophic groups and dominance by generalists in extreme low pH, suggesting that there may be a simplification of food webs with ocean acidification. Despite high variation in individual species' responses, our findings indicate that ocean acidification decreases the diversity, biomass, and trophic complexity of benthic marine communities. These results suggest that a loss of biodiversity and ecosystem function is expected under extreme acidification scenarios.

Description: We report on fine taxonomic and functional analyses of polychaetes associated with rocky reefs along a gradient of ocean acidification (OA) at the volcanic CO2 vent system off the Castello Aragonese (Ischia Island, Italy). Percent cover of algae and sessile invertebrates (a determinant of polychaete distribution) was classified into functional groups to disentangle the direct effects of low pH on polychaete abundance from the indirect effects of pH on habitat and other species associations. A total of 6459 polychaete specimens belonging to 83 taxa were collected. Polychaete species richness and abundance dramatically dropped under the extreme low pH conditions due to the disappearance of both calcifying and non-calcifying species. Differences in distribution patterns indicate that the decreasing pH modified the structure and biological traits of polychaete assemblages independent of changes in habitat. A detailed taxonomic analysis highlighted species-specific responses to OA, with closely related species having opposing responses to decreasing pH. This resulted in an increase in the abundance of filter feeders and herbivores with decreasing pH, while sessile polychaetes disappeared in the extreme low pH zones, and were replaced by discretely motile forms. Reproductive traits of the polychaete assemblages changed as well, with brooding species dominating the most acidified zones. The few taxa that were abundant in extreme low pH conditions showed high tolerance to OA (e.g. Amphiglena mediterranea, Syllis prolifera, Platynereis cf. dumerilii, Parafabricia mazzellae, Brifacia aragonensis), and are promising models for further studies on the responses of benthic organisms to the effects of reduced pH.

Description: Local adaptation can cause predator populations to vary in traits and their effects on prey, but few studies have tested whether divergent predator populations respond differently to acute environmental stressors. We tested how Nucella dogwhelks from 3 populations with natural exposure to distinct environmental regimes in the California Current System altered consumption of mussel prey (Mytilus californianus) in ambient (pH 8.0, 429 µatm partial pressure of CO2 [pCO2]) and acidified (pH 7.6, 1032 µatm pCO2) seawater. Overall, experimental acidification increased the variation in consumption time observed among populations. We found reduced consumption time for the population that experienced more frequent exposure to low pH conditions in nature but not for populations with less prior exposure. Exposure to acidification also altered the individual components of consumption time—search time and handling time—depending on source population. These results indicate that impaired predator performance is not a universal response to acidification, that predation responses to acute acidification can be population specific, and that individual population responses may relate to prior exposure. Our study highlights how population-specific responses to climate change can lead to differences in ecological effects that may restructure prey communities at local scales.

Description: The emergent responses of vulnerable species to global change can vary depending on the relative quality of resources available to support their productivity under increased stress, as well as the biotic interactions with other species that may alter their access to these resources. This research tested how seawater pCO2 may interact with seasonal light availability to affect the photosynthesis and calcification of high-latitude coralline algae, and whether the responses of these calcified macroalgae are modified by physical association with a non-calcified seaweed. Through an in situ approach, our study first investigated how current seasonal environmental variation affects the growth of the understory coralline algae Crusticorallina spp. and Bossiella orbigniana in Southeast Alaska's kelp forests. We then experimentally manipulated pH to simulate end-of-century acidification scenarios, light regime to simulate seasonal light availability at the benthos, and pairings of coralline algal species with and without a fleshy red alga to examine the interactive effects of these variables on coralline productivity and calcification. Our results indicate that: 1) coralline species may face net dissolution under projected future winter pH and carbonate saturation state conditions, 2) differences in seasonal light availability in productive, high-latitude waters may not be distinct enough to modify coralline algal net calcification, and 3) association with a non-calcified red alga does not alter the response of these coralline algal species to ocean acidification scenarios. This research highlights the necessity of incorporating locally informed scenarios of environmental variability and community interactions when predicting species' vulnerability to global change.

Description: Despite a growing interest in identifying tipping points in response to environmental change, our understanding of the ecological mechanisms underlying non-linear ecosystem dynamics is limited. Ecosystems governed by strong species interactions can provide important insight into how non-linear relationships between organisms and their environment propagate through ecosystems, and the potential for environmentally mediated species interactions to drive or protect against sudden ecosystem shifts. Here, we experimentally determine the functional relationships (i.e., the shapes of the relationships between predictor and response variables) of a seagrass assemblage with well-defined species interactions to ocean acidification (enrichment of CO2) in isolation and in combination with nutrient loading. We demonstrate that the effect of ocean acidification on grazer biomass (Phyllaplysia taylori and Idotea resecata) was quadratic, with the peak of grazer biomass at mid-pH levels. Algal grazing was negatively affected by nutrients, potentially due to low grazer affinity for macroalgae (Ulva intestinalis), as recruitment of both macroalgae and diatoms were favored in elevated nutrient conditions. This led to an exponential increase in macroalgal and epiphyte biomass with ocean acidification, regardless of nutrient concentration. When left unchecked algae can cause declines in seagrass productivity and persistence through shading and competition. Despite quadratic and exponential functional relationships to stressors that could cause a non-linear decrease in seagrass biomass, productivity of our model seagrass – the eelgrass (Zostera marina)- remained highly resilient to increasing acidification. These results suggest that important species interactions governing ecosystem dynamics may shift with environmental change, and ecosystem state may be decoupled from ecological responses at lower levels of organization.

Description: Percent cover of 215 benthic species was quantified using two approaches with scuba diving: i) visual census techniques in Shallow Reef (benthic surveys were performed from 0.5 to 3 m depth) and Reef habitats (benthic surveys at ~10 m depth) dominated by algae and ii) photographic surveys in Caves (benthic surveys at ~3 m depth) and Deep Reef habitats (benthic surveys at ~ 40 m depth) dominated by sessile invertebrate animals. For visual censuses, 12 quadrats (25x25 cm) were haphazardly placed at Vent 1 and Vent 3 and the two corresponding ambient pH sites (n = 3 sites per habitat, n = 36 quadrats for Shallow Reefs and Reefs, respectively). For photographic surveys, 48–54 and 24 photoquadrats (25x25 cm) were taken at Vent 2 and Vent 4 and the two corresponding reference sites with ambient pH for each habitat (n = 3 sites per habitat, n = 144 photographic quadrats for Caves and n = 72 photographic quadrats for Deep Reefs, respectively). Both types of quadrats (visual censuses in the field and photographs in the lab) were divided into a grid of 25 squares (5 cm x 5 cm each). Percentage cover was quantified by counting the number of squares filled in the grid by the species and expressing the final values as relative percentages.

Description: Anthropogenic emissions of carbon dioxide (CO2) are causing ocean acidification, lowering seawater aragonite (CaCO3) saturation state (Omega arag), with potentially substantial impacts on marine ecosystems over the 21st Century. Calcifying organisms have exhibited reduced calcification under lower saturation state conditions in aquaria. However, the in situ sensitivity of calcifying ecosystems to future ocean acidification remains unknown. Here we assess the community level sensitivity of calcification to local CO2-induced acidification caused by natural respiration in an unperturbed, biodiverse, temperate intertidal ecosystem. We find that on hourly timescales nighttime community calcification is strongly influenced by Omega arag, with greater net calcium carbonate dissolution under more acidic conditions. Daytime calcification however, is not detectably affected by Omega arag. If the short-term sensitivity of community calcification to Omega arag is representative of the long-term sensitivity to ocean acidification, nighttime dissolution in these intertidal ecosystems could more than double by 2050, with significant ecological and economic consequences.

Description: Ocean acidification is a pervasive threat to coral reef ecosystems, and our understanding of the ecological processes driving patterns in tropical benthic community development in conditions of acidification is limited. We deployed limestone recruitment tiles in low aragonite saturation (Omega arag) waters during an in-situ field experiment at Puerto Morelos, Mexico, and compared them to tiles placed in control zones over a 14-month investigation. The early stages of succession showed relatively little difference in coverage of calcifying organisms between the low Omega arag and control zones. However, after 14 months of development, tiles from the low Omega arag zones had up to 70% less cover of calcifying organisms coincident with 42% more fleshy algae than the controls. The percent cover of biofilm and turf algae was also significantly greater in the low Omega arag zones, while the number of key grazing taxa remained constant. We hypothesize that fleshy algae have a competitive edge over the primary calcified space holders, coralline algae, and that acidification leads to altered competitive dynamics between various taxa. We suggest that as acidification impacts reefs in the future, there will be a shift in community assemblages away from upright and crustose coralline algae toward more fleshy algae and turf, established in the early stages of succession.

Description: Acidification-induced changes in neurological function have been documented in several tropical marine fishes. Here, we investigate whether similar patterns of neurological impacts are observed in a temperate Pacific fish that naturally experiences regular and often large shifts in environmental pH/pCO2. In two laboratory experiments, we tested the effect of acidification, as well as pH/pCO2 variability, on gene expression in the brain tissue of a common temperate kelp forest/estuarine fish, Embiotoca jacksoni. Experiment 1 employed static pH treatments (target pH = 7.85/7.30), while Experiment 2 incorporated two variable treatments that oscillated around corresponding static treatments with the same mean (target pH = 7.85/7.70) in an eight-day cycle (amplitude ± 0.15). We found that patterns of global gene expression differed across pH level treatments. Additionally, we identified differential expression of specific genes and enrichment of specific gene sets (GSEA) in comparisons of static pH treatments and in comparisons of static and variable pH treatments of the same mean pH. Importantly, we found that pH/pCO2 variability decreased the number of differentially expressed genes detected between high and low pH treatments, and that inter-individual variability in gene expression was greater in variable treatments than static treatments. These results provide important confirmation of neurological impacts of acidification in a temperate fish species and, critically, that natural environmental variability may mediate the impacts of ocean acidification.