ALBERT

Description: We present the first study of the effects of ocean acidification on settlement of benthic invertebrates and microfauna. Artificial collectors were placed for 1 month along pH gradients at CO2 vents off Ischia (Tyrrhenian Sea, Italy). Seventy-nine taxa were identified from six main taxonomic groups (foraminiferans, nematodes, polychaetes, molluscs, crustaceans and chaetognaths). Calcareous foraminiferans, serpulid polychaetes, gastropods and bivalves showed highly significant reductions in recruitment to the collectors as pCO2 rose from normal (336-341 ppm, pH 8.09-8.15) to high levels (886-5,148 ppm) causing acidified conditions near the vents (pH 7.08-7.79). Only the syllid polychaete Syllis prolifera had higher abundances at the most acidified station, although a wide range of polychaetes and small crustaceans was able to settle and survive under these conditions. A few taxa (Amphiglena mediterranea, Leptochelia dubia, Caprella acanthifera) were particularly abundant at stations acidified by intermediate amounts of CO2 (pH 7.41-7.99). These results show that increased levels of CO2 can profoundly affect the settlement of a wide range of benthic organisms.

Description: Ocean acidification (OA) is likely to exert selective pressure on natural populations. Our ability to predict which marine species will adapt to OA, and what underlies this adaptive potential, are of high conservation and resource management priority. Using a naturally low pH vent site in the Mediterranean Sea (Castello Aragonese, Ischia) mirroring projected future OA conditions, we carried out a reciprocal transplant experiment to investigate the relative importance of phenotypic plasticity and local adaptation in two populations of the sessile, calcifying polychaete /Simplaria /sp. (Annelida, Serpulidae, Spirorbinae): one residing in low pH and the other from a nearby ambient (i.e. high) pH site. We measured a suite of fitness related traits (i.e. survival, reproductive output, maturation, population growth) and tube growth rates in laboratory-bred F2 generation individuals from both populations reciprocally transplanted back into both ambient and low pH /in situ/ habitats. Both populations showed lower expression in all traits, but increased tube growth rates, when exposed to low pH compared to high pH conditions, regardless of their site of origin suggesting that local adaptation to low pH conditions has not occurred. We also found comparable levels of plasticity in the two populations investigated, suggesting no influence of long-term exposure to low pH on the ability of populations to adjust their phenotype. Despite high variation in trait values among sites and the relatively extreme conditions at sites close to the vents (pH 〈 7.36), response trends were consistent across traits. Hence, our data suggest that, for /Simplaria /and possibly other calcifiers, neither local adaptations nor sufficient phenotypic plasticity levels appear to suffice in order to compensate for the negative impacts of OA on long-term survival. Our work also underlines the utility of field experiments in natural environments subjected to high level of /p/CO_2 for elucidating the potential for adaptation to future scenarios of OA.

Description: There are serious concerns that ocean acidification will combine with the effects of global warming to cause major shifts in marine ecosystems, but there is a lack of field data on the combined ecological effects of these changes due to the difficulty of creating large-scale, long-term exposures to elevated CO2 and temperature. Here we report the first coastal transplant experiment designed to investigate the effects of naturally acidified seawater on the rates of net calcification and dissolution of the branched calcitic bryozoan Myriapora truncata (Pallas, 1766). Colonies were transplanted to normal (pH 8.1), high (mean pH 7.66, minimum value 7.33) and extremely high CO2 conditions (mean pH 7.43, minimum value 6.83) at gas vents off Ischia Island (Tyrrhenian Sea, Italy). The net calcification rates of live colonies and the dissolution rates of dead colonies were estimated by weighing after 45 days (May-June 2008) and after 128 days (July-October) to examine the hypothesis that high CO2 levels affect bryozoan growth and survival differently during moderate and warm water conditions. In the first observation period, seawater temperatures ranged from 19 to 24 °C; dead M. truncata colonies dissolved at high CO2 levels (pH 7.66), whereas live specimens maintained the same net calcification rate as those growing at normal pH. In extremely high CO2 conditions (mean pH 7.43), the live bryozoans calcified significantly less than those at normal pH. Therefore, established colonies of M. truncata seem well able to withstand the levels of ocean acidification predicted in the next 200 years, possibly because the soft tissues protect the skeleton from an external decrease in pH. However, during the second period of observation a prolonged period of high seawater temperatures (25-28 °C) halted calcification both in controls and at high CO2, and all transplants died when high temperatures were combined with extremely high CO2 levels. Clearly, attempts to predict the future response of organisms to ocean acidification need to consider the effects of concurrent changes such as the Mediterranean trend for increased summer temperatures in surface waters. Although M. truncata was resilient to short-term exposure to high levels of ocean acidification at normal temperatures, our field transplants showed that its ability to calcify at higher temperatures was compromised, adding it to the growing list of species now potentially threatened by global warming.

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: 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: Ocean acidification represents a pervasive environmental change that is predicted to affect a wide range of species, yet our understanding of the emergent ecosystem impacts is very limited. Many studies report detrimental effects of acidification on single species in lab studies, especially those with calcareous shells or skeletons. Observational studies using naturally acidified ecosystems have shown profound shifts away from such calcareous species, and there has been an assumption that direct impacts of acidification on sensitive species drive most ecosystem responses. We tested an alternative hypothesis that species interactions attenuate or amplify the direct effects of acidification on individual species. Here, we show that altered competitive dynamics between calcareous species and fleshy seaweeds drive significant ecosystem shifts in acidified conditions. Although calcareous species recruited and grew at similar rates in ambient and low pH conditions during early successional stages, they were rapidly overgrown by fleshy seaweeds later in succession in low pH conditions. The altered competitive dynamics between calcareous species and fleshy seaweeds is probably the combined result of decreased growth rates of calcareous species, increased growth rates of fleshy seaweeds, and/or altered grazing rates. Phase shifts towards ecosystems dominated by fleshy seaweed are common in many marine ecosystems, and our results suggest that changes in the competitive balance between these groups represent a key leverage point through which the physiological responses of individual species to acidification could indirectly lead to profound ecosystem changes in an acidified ocean.