Description: In the recent discussion how biotic systems may react to ocean acidification caused by the rapid rise in carbon dioxide partial pressure (pCO2) in the marine realm, substantial research is devoted to calcifiers such as stony corals. The antagonistic process-biologically induced carbonate dissolution via bioerosion- has largely been neglected. Unlike skeletal growth, we expect bioerosion by chemical means to be facilitated in a high-CO2 world. This study focuses on one of the most detrimental bioeroders, the sponge Cliona orientalis, which attacks and kills live corals on Australia's Great Barrier Reef. Experimental exposure to lowered and elevated levels of pCO2 confirms a significant enforcement of the sponges' bioerosion capacity with increasing pCO2 under more acidic conditions. Considering the substantial contribution of sponges to carbonate bioerosion, this finding implies that tropical reef ecosystems are facing the combined effects of weakened coral calcification and accelerated bioerosion, resulting in critical pressure on the dynamic balance between biogenic carbonate build-up and degradation.

Description: Physiological sensitivity of cold-water corals to ocean change is far less understood than of tropical corals and very little is known about the impacts of ocean acidification and warming on degradative processes of dead coral framework. In a 13-month laboratory experiment, we examined the interactive effects of gradually increasing temperature and pCO2 levels on survival, growth, and respiration of two prominent color morphotypes (colormorphs) of the framework-forming cold-water coral Lophelia pertusa, as well as bioerosion and dissolution of dead framework. Calcification rates tended to increase with warming, showing temperature optima at ~ 14°C (white colormorph) and 10–12°C (orange colormorph) and decreased with increasing pCO2. Net dissolution occurred at aragonite undersaturation (ΩAr 〈 1) at ~ 1000 μatm pCO2. Under combined warming and acidification, the negative effects of acidification on growth were initially mitigated, but at ~ 1600 μatm dissolution prevailed. Respiration rates increased with warming, more strongly in orange corals, while acidification slightly suppressed respiration. Calcification and respiration rates as well as polyp mortality were consistently higher in orange corals. Mortality increased considerably at 14–15°C in both colormorphs. Bioerosion/dissolution of dead framework was not affected by warming alone but was significantly enhanced by acidification. While live corals may cope with intermediate levels of elevated pCO2 and temperature, long-term impacts beyond levels projected for the end of this century will likely lead to skeletal dissolution and increased mortality. Our findings further suggest that acidification causes accelerated degradation of dead framework even at aragonite saturated conditions, which will eventually compromise the structural integrity of cold-water coral reefs.

Description: Ocean acidity has increased by 30% since preindustrial times due to the uptake of anthropogenic CO2 and is projected to rise by another 120% before 2100 if CO2 emissions continue at current rates. Ocean acidification is expected to have wide-ranging impacts on marine life, including reduced growth and net erosion of coral reefs. Our present understanding of the impacts of ocean acidification on marine life, however, relies heavily on results from short-term CO2 perturbation studies. Here we present results from the first long-term CO2 perturbation study on the dominant reef-building cold-water coral Lophelia pertusa and relate them to results from a short-term study to compare the effect of exposure time on the coral's responses. Short-term (one week) high CO2 exposure resulted in a decline of calcification by 26-29% for a pH decrease of 0.1 units and net dissolution of calcium carbonate. In contrast, L. pertusa was capable to acclimate to acidified conditions in long-term (six months) incubations, leading to even slightly enhanced rates of calcification. Net growth is sustained even in waters sub-saturated with respect to aragonite. Acclimation to seawater acidification did not cause a measurable increase in metabolic rates. This is the first evidence of successful acclimation in a coral species to ocean acidification, emphasizing the general need for long-term incubations in ocean acidification research. To conclude on the sensitivity of cold-water coral reefs to future ocean acidification further ecophysiological studies are necessary which should also encompass the role of food availability and rising temperatures.

Description: The uptake of anthropogenic emission of carbon dioxide is resulting in a lowering of the carbonate saturation state and a drop in ocean pH. Understanding how marine calcifying organisms such as coralline algae may acclimatize to ocean acidification is important to understand their survival over the coming century. We present the first long-term perturbation experiment on the cold-water coralline algae, which are important marine calcifiers in the benthic ecosystems particularly at the higher latitudes. Lithothamnion glaciale, after three months incubation, continued to calcify even in undersaturated conditions with a significant trend towards lower growth rates with increasing pCO2. However, the major changes in the ultra-structure occur by 589 µatm (i.e. in saturated waters). Finite element models of the algae grown at these heightened levels show an increase in the total strain energy of nearly an order of magnitude and an uneven distribution of the stress inside the skeleton when subjected to similar loads as algae grown at ambient levels. This weakening of the structure is likely to reduce the ability of the alga to resist boring by predators and wave energy with severe consequences to the benthic community structure in the immediate future (50 years).

Description: Cold-water corals are important bioengineers that provide structural habitat for a diverse species community. About 70 % of the presently known scleractinian cold-water corals are expected to be exposed to corrosive waters by the end of this century due to ocean acidification. At the same time, the corals will experience a steady warming of their environment. Studies on the sensitivity of cold-water corals to climate change mainly concentrated on single stressors in short-term incubation approaches, thus not accounting for possible long-term acclimatisation and the interactive effects of multiple stressors. Besides, preceding studies did not test for possible compensatory effects of a change in food availability. In this study a multifactorial long-term experiment (6 months) was conducted with end-of-the-century scenarios of elevated pCO2 and temperature levels in order to examine the acclimatisation potential of the cosmopolitan cold-water coral Lophelia pertusa to future climate change related threats. For the first time multiple ocean change impacts including the role of the nutritional status were tested on L. pertusa with regard to growth, 'fitness', and survival. Our results show that while L. pertusa is capable of calcifying under elevated CO2 and temperature, its condition (fitness) is more strongly influenced by food availability rather than changes in seawater chemistry. Whereas growth rates increased at elevated temperature (+ 4°C), they decreased under elevated CO2 concentrations (800 µatm). No difference in net growth was detected when corals were exposed to the combination of increased CO2 and temperature compared to ambient conditions. A 10-fold higher food supply stimulated growth under elevated temperature, which was not observed in the combined treatment. This indicates that increased food supply does not compensate for adverse effects of ocean acidification and underlines the importance of considering the nutritional status in studies investigating organism responses under environmental changes.

Description: Cold-water corals (CWC) are widely distributed around the world forming extensive reefs at par with tropical coral reefs. They are hotspots of biodiversity and organic matter processing in the world's deep oceans. Living in the dark they lack photosynthetic symbionts and are therefore considered to depend entirely on the limited flux of organic resources from the surface ocean. While symbiotic relations in tropical corals are known to be key to their survival in oligotrophic conditions, the full metabolic capacity of CWC has yet to be revealed. Here we report isotope tracer evidence for efficient nitrogen recycling, including nitrogen assimilation, regeneration, nitrification and denitrification. Moreover, we also discovered chemoautotrophy and nitrogen fixation in CWC and transfer of fixed nitrogen and inorganic carbon into bulk coral tissue and tissue compounds (fatty acids and amino acids). This unrecognized yet versatile metabolic machinery of CWC conserves precious limiting resources and provides access to new nitrogen and organic carbon resources that may be essential for CWC to survive in the resource-depleted dark ocean.

Description: Rising atmospheric CO2 concentrations could cause a calcium carbonate subsaturation of Arctic surface waters in the next 20 yr, making these waters corrosive for calcareous organisms. It is presently unknown what effects this will have on Arctic calcifying organisms and the ecosystems of which they are integral components. So far, acidification effects on crustose coralline red algae (CCA) have only been studied in tropical and Mediterranean species. In this work, we investigated calcification rates of the CCA Lithothamnion glaciale collected in northwest Svalbard in laboratory experiments under future atmospheric CO2 concentrations. The algae were exposed to simulated Arctic summer and winter light conditions in 2 separate experiments at optimum growth temperatures. We found a significant negative effect of increased CO2 levels on the net calcification rates of L. glaciale in both experiments. Annual mean net dissolution of L. glaciale was estimated to start at an aragonite saturation state between 1.1 and 0.9 which is projected to occur in parts of the Arctic surface ocean between 2030 and 2050 if emissions follow 'business as usual' scenarios (SRES A2; IPCC 2007). The massive skeleton of CCA, which consist of more than 80% calcium carbonate, is considered crucial to withstanding natural stresses such as water movement, overgrowth or grazing. The observed strong negative response of this Arctic CCA to increased CO2 levels suggests severe threats of the projected ocean acidification for an important habitat provider in the Arctic coastal ocean.

Description: In a 13-months laboratory experiment conducted in 2014/2015, the interactive effects of gradually increasing temperature and pCO2 levels on survival, growth and respiration of two prominent colour morphotypes (white and orange) of the framework-forming cold-water coral Lophelia pertusa (also known as Desmophyllum pertusum), as well as bioerosion and dissolution of dead coral framework were assessed. In six-week intervals, three treatments (T1: acidification, T2: warming, T3: combined acidification and warming) were gradually increased in their respective manipulated parameters by 1°C and/or 200 µatm pCO2 after an initial two intervals under ambient (near in-situ) conditions. Each treatment consisted of 7 replicates that were manipulated over the course of the experiment and 3 control replicates that remained at ambient conditions throughout the entire duration of the experiment. Each replicate tank consisted of one live coral fragment of the white morphotype, one fragment of the orange morphotype and one dead framework fragment (naturally bioeroded framework material). Dead framework was examined with regard to attached bioeroders and calcifying organisms, the latter being removed prior to the experiment. All coral samples were collected from an inshore Norwegian cold-water coral habitat in the outer Trondheim-Fjord near Nord-Leksa (63°36.4'N, 09°22.7'E) between 150 to 230 m water depth using the manned submersible JAGO (GEOMAR, 2017; doi:10.17815/jlsrf-3-157) during RV POSEIDON (GEOMAR, 2015; doi:10.17815/jlsrf-1-62) cruise POS455 in June/July 2013. In situ conditions at the time of sampling near the corals were 7.7°C in temperature, 35.2 in salinity and ~6 mL/L oxygen concentration. Prior to the experiment, corals were kept in a closed recirculating system of 1,700 L in a climate-controlled laboratory facility at GEOMAR in Kiel at near in situ conditions of temperature and salinity (7.8 145 ± 0.2 °C and 35.8 ± 0.6) for half a year. Calcification/dissolution rates of live corals and bioerosion/dissolution rates of dead coral framework were determined using the buoyant weighing technique (Davies, 1989; doi:10.1007/BF00428135) with a high precision analytical balance (Sartorius CPA225D, readability = 0.1 mg) placed above every individual aquarium for each measurement. Respiration rates were determined via oxygen consumption measurements using an optode-based oxygen analyser (Oxy-10 mini, PreSens GmbH). Mortality was examined during every six-week interval by visual inspection of all live fragments. Dead polyp counts were calculated as percentage of total polyps counts of every individual fragment. Carbonate system parameters were calculated from the two measured parameters total alkalinity (TA) and dissolved inorganic carbon (DIC). TA and DIC samples were taken at the end of every 6-week interval and analyzed via potentiometric open-cell titration (862 Compact Titrosampler, Metrohm) in case of TA and by infrared detection of CO2 using an Automated Infra-Red Inorganic Carbon Analyzer (AIRICA with LI-COR 7000, Marianda) in case of DIC. TA and DIC were corrected against Certified Reference Material from A.G. Dickson (Scripps Institution of Oceanography) and density-corrected. The purpose of this study was to examine thresholds and optima of live corals under gradual increases of ocean acidification and warming and to quantify dissolution and bioerosion rates of dead coral framework to ultimately assess the balance between live coral calcification and degradation of dead coral framework under future ocean conditions.

Description: All parameters assessed at the end of the experiment (dry weight of the corals/dead coral framework fragments, ash-free dry mass (AFDM), total polyp count, bacterial background respiration in experimental tanks (no corals incubations).

Description: Measured parameters (net calcification/dissolution, net dissolution/bioerosion, respiration, mortality, temperature, salinity, total alkalinity (TA), dissolved inorganic carbon (DIC)) throughout the 6-week experiment intervals under gradual alterations of the manipulation parameters (temperature, pCO2).