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 acidification on the biogeochemistry of unvegetated and seagrass marine sediments.

Many studies addressed ocean acidification (OA) effects on marine life, whereas its effects on sedimentary organic matter (OM) have received less attention. We investigated differences in OM features in sediments from unvegetated and seagrass (Posidonia oceanica) beds in a shallow hydrothermal area (Aeolian Archipelago, Mediterranean Sea), under natural (8.1-8.0) and acidified (7.8-7.9) conditions. We show that a pH difference of -0.3 units have minor effects on OM features in unvegetated sediments, but relevant consequences within acidified seagrass meadows, where OM quantity and nutritional quality are lower than those under natural pH conditions. Effects of acidified conditions on OM biogeochemistry vary between unvegetated and seagrass sediments, with lower C degradation rates and longer C turnover time in the former than in the latter. We conclude that OA, although with effects not consistent between unvegetated and vegetated sediments, can affect OM quantity, composition, and degradation, thus having possible far-reaching consequences for benthic trophic webs.

Long-term patterns and drivers of microbial organic matter utilization in the northernmost basin of the Mediterranean Sea.

Marine heterotrophic prokaryotes degrade, transform, and utilize half of the organic matter (OM) produced by photosynthesis, either in dissolved or particulate form. Microbial metabolic rates are affected by a plethora of different factors, spanning from environmental variables to OM composition. To tease apart the environmental drivers underlying the observed organic matter utilization rates, we analysed a 21 year-long time series from the Gulf of Trieste (NE Adriatic Sea). Heterotrophic carbon production (HCP) time series analysis highlighted a long-term structure made up by three periods of coherent observations (1999-2007; 2008-2011; 2012-2019), shared also by OM concentration time series. Temporal patterns of HCP drivers, extracted with a random forest approach, demonstrated that a period of high salinity anomalies (2002-2008) was the main driver of this structure. The reduced river runoff and the consequent depletion of river-borne inorganic nutrients induced a long-term Chl a decline (2006-2009), followed by a steady increase until 2014. HCP driving features over the three periods substantially changed in their seasonal patterns, suggesting that the years following the draught period represented a transition between two long-term regimes. Overall, temperature and particulate organic carbon concentration were the main factors driving HCP rates. The emergence of these variables highlighted the strong control exerted by the temperature-substrate co-limitation on microbial growth. Further exploration revealed that HCP rates did not follow the Arrhenius' linear response to temperature between 2008 and 2011, demonstrating that microbial growth was substrate-limited following the draught event. By teasing apart the environmental drivers of microbial growth on a long-term perspective, we demonstrated that a substantial change happened in the biogeochemistry of one of the most productive areas of the Mediterranean Sea. As planktonic microbes are the foundation of marine ecosystems, understanding their past dynamics may help to explain present and future changes.

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.

Benthic nutrient cycling at the sediment-water interface in a lagoon fish farming system (northern Adriatic Sea, Italy).

Metabolism and carbon, oxygen, and nutrient fluxes (DIC, DOC, DO2, NO2-, NO3-, NH4+, PO43- and SiO44-) were studied during three surveys at two sites (VN1 and VN3) located at a fish farm at the Marano and Grado Lagoon (northern Adriatic Sea), using an in situ benthic chamber. Field experiments were conducted in July and October 2015 and March 2016 at a depth of approximately 2 m along the main channels of the fish farm. Water samples were collected by a scuba diver every 2 h in order to investigate daily fluxes of solutes across the sediment-water interface (SWI). Regarding the solid phase, Corg/Ntot and Corg/Porg molar ratios suggested an autochthonous marine origin of the organic matter and a minor preservation of P in the sediments, respectively; high values of sulphur (Stot) were also encountered (0.8-2%). The conditions at VN3 were mostly anoxic with high NH4+ levels (30-1027 µM) and the absence of NO3-. Substantial daily patterns of all solutes occurred especially in autumn and winter. On the contrary, fluxes at VN1 were less pronounced. Usually, inverse correlations appeared between dissolved O2 and DIC trends, but in our system this was observed only at VN3 in autumn and accomplished by a parallel increase in NH4+, PO43- and SiO44- during intense nutrient regeneration. These results are significantly different than those reported for open lagoon environments, where nutrient regeneration at the SWI and in surface sediments is the primary source of nutrients available for assimilation processes, especially during the warmer period of the year when the natural nutrient input by fresh water inflows is limited. Due to the importance of this site for aquaculture, biodiversity and ecosystem services, useful suggestions have been provided from this study in order to improve the quality of this unique aquatic system.

CO2 leakage alters biogeochemical and ecological functions of submarine sands.

Subseabed CO2 storage is considered a future climate change mitigation technology. We investigated the ecological consequences of CO2 leakage for a marine benthic ecosystem. For the first time with a multidisciplinary integrated study, we tested hypotheses derived from a meta-analysis of previous experimental and in situ high-CO2 impact studies. For this, we compared ecological functions of naturally CO2-vented seafloor off the Mediterranean island Panarea (Tyrrhenian Sea, Italy) to those of nonvented sands, with a focus on biogeochemical processes and microbial and faunal community composition. High CO2 fluxes (up to 4 to 7 mol CO2 m-2 hour-1) dissolved all sedimentary carbonate, and comigration of silicate and iron led to local increases of microphytobenthos productivity (+450%) and standing stocks (+300%). Despite the higher food availability, faunal biomass (-80%) and trophic diversity were substantially lower compared to those at the reference site. Bacterial communities were also structurally and functionally affected, most notably in the composition of heterotrophs and microbial sulfate reduction rates (-90%). The observed ecological effects of CO2 leakage on submarine sands were reproduced with medium-term transplant experiments. This study assesses indicators of environmental impact by CO2 leakage and finds that community compositions and important ecological functions are permanently altered under high CO2.

Microphytobenthic community composition and primary production at gas and thermal vents in the Aeolian Islands (Tyrrhenian Sea, Italy).

Sediment samplings were performed to investigate the microphytobenthic community and photosynthetic activity adaptations to gas emissions and higher temperature in the Aeolian Islands during a three-year period (2012-2014). Higher microphytobenthic densities were recorded at the vent stations and values were even more pronounced in relation with high temperature. The gross primary production estimates strongly depended on microphytobenthic abundance values reaching up to 45.79 ± 6.14 mgC m(-2) h(-1). High abundances were coupled with low community richness and diversity. Motile diatom living forms were predominant at all stations and the greatest differences among vent and reference stations were detected on the account of the tychopelagic forms. Morphological deformities and heavily silicified diatom frustules were also observed. A significant influence of the gas emission and high temperature on the phototrophic community was highlighted suggesting the Aeolian Islands as a good natural laboratory for studies on high CO2 and global warming effects.

Biomonitoring of a polluted coastal area (Bay of Muggia, Northern Adriatic Sea): A five-year study using transplanted mussels.

The subcellular effects of pollution were evaluated using two lysosomal biomarkers in mussels, Mytilus galloprovincialis, deployed periodically over a period of 5 years in a harbour area in the Bay of Muggia (Gulf of Trieste, North Adriatic Sea) that is strongly influenced by anthropogenic activities. Mussels were collected from a clean marine farm and analysed (sample T0). A sub-sample was transplanted to the harbour site (sample M) and analysed after about 12 weeks. An additional sub-sample was relocated within the farm as a control and was also tested at the end of the 12-week period (sample T1). The transplantation procedures were repeated twice yearly for 5 consecutive years, starting in 2009. Two well-established lysosomal biomarkers, i.e. lysosomal membrane stability and lipofuscin accumulation, were evaluated in hepatopancreas cells. The body condition index and mortality rate were also assessed. Moreover, various pollutants were determined in both mussel flesh, for a better comprehension of the biological response, and sediments, for a general characterization of the study area. As a whole, the applied biomarkers were found to be appropriate for determining the responses of mussels to environmental pollutant loads over time. Variations in lysosomal membrane stability and lipofuscin content were mostly related to total PAHs and metals respectively. Our results confirm the usefulness of active biomonitoring in evaluating pollution trends in marine coastal areas and in particular the value of lysosomal biomarkers as a rapid screening tool for highlighting pollutant effects at least at organism level.

Recent evolution of the physical-chemical characteristics of a Site of National Interest-the Mar Piccolo of Taranto (Ionian Sea)-and changes over the last 20 years.

The Mar Piccolo of Taranto, classified as a 'Site of National Interest' (SIN), is a semi-enclosed basin divided into two inlets with lagoon features and sea influences, seriously affected by anthropic activities. In the framework of the RITMARE project, a study has been carried out to evaluate the functionality of this ecosystem. As part of this work, measurements of the water abiotic parameters were performed in order to assess the physical-chemical features of this area after the activation, in the last decade, of treatment plants for various urban and industrial dumping. Seawater intrusions and continental inputs, as well as several submarine freshwater springs, clearly affect physical-chemical characteristics of the water column in the two inlets. This finding suggests that small-scale patterns in water circulation have the potential to influence the chemical properties of the seawater. The comparison with a 20-year dataset reveals a drastic decrease in nutrient concentrations after the year 2000, validating the functionality of the treatment plants. The reduction of nutrient inputs into the basin (up to -90 % in the first inlet characterized by lower hydraulic residence time) has changed the biogeochemical characteristics of the Mar Piccolo from being relatively eutrophic to moderately oligotrophic.

Oxygen, carbon, and nutrient exchanges at the sediment-water interface in the Mar Piccolo of Taranto (Ionian Sea, southern Italy).

In the shallow environment, the nutrient and carbon exchanges at the sediment-water interface contribute significantly to determine the trophic status of the whole water column. The intensity of the allochthonous input in a coastal environment subjected to strong anthropogenic pressures determines an increase in the benthic oxygen demand leading to depressed oxygen levels in the bottom waters. Anoxic conditions resulting from organic enrichment can enhance the exchange of nutrients between sediments and the overlying water. In the present study, carbon and nutrient fluxes at the sediment-water interface were measured at two experimental sites, one highly and one moderately contaminated, as reference point. In situ benthic flux measurements of dissolved species (O2, DIC, DOC, N-NO3 (-), N-NO2 (-), N-NH4 (+), P-PO4 (3-), Si-Si(OH)4, H2S) were conducted using benthic chambers. Furthermore, undisturbed sediment cores were collected for analyses of total and organic C, total N, and biopolymeric carbon (carbohydrates, proteins, and lipids) as well as of dissolved species in porewaters and supernatant in order to calculate the diffusive fluxes. The sediments were characterized by suboxic to anoxic conditions with redox values more negative in the highly contaminated site, which was also characterized by higher biopolymeric carbon content (most of all lipids), lower C/N ratios and generally higher diffusive fluxes, which could result in a higher release of contaminants. A great difference was observed between diffusive and in situ benthic fluxes suggesting the enhancing of fluxes by bioturbation and the occurrence of biogeochemically important processes at the sediment-water interface. The multi-contamination of both inorganic and organic pollutants, in the sediments of the Mar Piccolo of Taranto (declared SIN in 1998), potentially transferable to the water column and to the aquatic trophic chain, is of serious concern for its ecological relevance, also considering the widespread fishing and mussel farming activities in the area.

The application of a plug-flow reactor to measure the biodegradable dissolved organic carbon (BDOC) in seawater.

Most of the ambient dissolved organic carbon (DOC) is refractory to microbial degradation; bacteria can consume a minor but variable part of the DOC pool over periods of hours and days. It is important to increase our knowledge of the dynamics of the biodegradable fraction of DOC (BDOC) to understand the global carbon budget. Several methods for determining BDOC have been developed; however, the problem of most of them is the time (days/weeks) required for the colonization and/or determination. In this paper, we describe the application to seawater of a plug-flow bioreactor to measure BDOC within 3-4 h. The bioreactor was built following Søndergaard and Worm [Søndergaard, M., Worm, J., 2001. Measurement of biodegradable dissolved organic carbon (BDOC) in lake water with a bioreactor. Water Res. 35, 2505-2513.] protocols for the measurement of BDOC in lake water. We analyzed BDOC on samples collected in the Gulf of Trieste during autumn-winter and summer 2003-2004. BDOC concentrations varied from 8 to 24 microM and represented from 10.3% to 25.5% of the total DOC. To evaluate the effectiveness of this method, we compared bioreactor BDOC measurement with data obtained from batch cultures. The results indicate that BDOC in coastal seawater can be measured rapidly and reliably with this bioreactor.