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Cold-water corals are important habitat formers in deep-water ecosystems and at high latitudes. Ocean acidification and the resulting change in aragonite saturation are expected to affect these habitats and impact coral growth. Counter to expectations, the deep water coral Lophelia pertusa has been found to be able to sustain growth even in undersaturated conditions. However, it is important to know whether such undersaturation modifies the skeleton and thus its ecosystem functioning. Here we used Synchrotron X-Ray Tomography and Raman spectroscopy to examine changes in skeleton morphology and fibre orientation. We combined the morphological assessment with boron isotope analysis to determine if changes in growth are related to changes in control of calcification pH. We compared the isotopic composition and structure formed in their natural environment to material grown in culture at lower pH conditions. Skeletal morphology is highly variable but shows no distinctive differences between natural and low pH conditions. Raman investigations found no difference in macromorphological skeletal arrangement of early mineralization zones and secondary thickening between the treatments. The δ11B analyses show that L. pertusa up-regulates the internal calcifying fluid pH (pHcf) during calcification compared to ambient seawater pH and maintains a similar elevated pHcf at increased pCO2 conditions. We suggest that as long as the energy is available to sustain the up-regulation, i.e. individuals are well fed, there is no detrimental effect to the skeletal morphology.

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.

In many marine biogeographic realms, bioeroding sponges dominate the internal bioerosion of calcareous substrates such as mollusc beds and coral reef framework. They biochemically dissolve part of the carbonate and liberate so-called sponge chips, a process that is expected to be facilitated and accelerated in a more acidic environment inherent to the present global change. The bioerosion capacity of the demosponge Cliona celata Grant, 1826 in subfossil oyster shells was assessed via alkalinity anomaly technique based on 4 days of experimental exposure to three different levels of carbon dioxide partial pressure (pCO(2)) at ambient temperature in the cold-temperate waters of Helgoland Island, North Sea. The rate of chemical bioerosion at present-day pCO(2) was quantified with 0.08-0.1 kg m(-2) year(-1). Chemical bioerosion was positively correlated with increasing pCO(2), with rates more than doubling at carbon dioxide levels predicted for the end of the twenty-first century, clearly confirming that C. celata bioerosion can be expected to be enhanced with progressing ocean acidification (OA). Together with previously published experimental evidence, the present results suggest that OA accelerates sponge bioerosion (1) across latitudes and biogeographic areas, (2) independent of sponge growth form, and (3) for species with or without photosymbionts alike. A general increase in sponge bioerosion with advancing OA can be expected to have a significant impact on global carbonate (re)cycling and may result in widespread negative effects, e.g. on the stability of wild and farmed shellfish populations, as well as calcareous framework builders in tropical and cold-water coral reef ecosystems.

Coral reefs are under threat, exerted by a number of interacting effects inherent to the present climate change, including ocean acidification and global warming. Bioerosion drives reef degradation by recycling carbonate skeletal material and is an important but understudied factor in this context. Twelve different combinations of pCO2 and temperature were applied to elu- cidate the consequences of ocean acidification and global warming on the physiological response and bioerosion rates of the zooxanthellate sponge Cliona orientalis —one of the most abundant and effective bioeroders on the Great Barrier Reef, Australia. Our results confirm a significant amplifi- cation of the sponges' bioerosion capacity with increasing pCO2, which is expressed by more car- bonate being chemically dissolved by etching. The health of the sponges and their photosymbionts was not affected by changes in pCO2, in contrast to temperature, which had significant negative impacts at higher levels. However, we could not conclusively explain the relationship between temperature and bioerosion rates, which were slightly reduced at both colder as well as warmer temperatures than ambient. The present findings on the effects of ocean acidification on chemical bioerosion, however, will have significant implications for predicting future reef carbonate budgets, as sponges often contribute the lion's share of internal bioerosion on coral reefs.

Coralline algae (Corallinales, Rhodophyta) that form rhodoliths are important ecosystem engineers and carbonate producers in many polar coastal habitats. This study deals with rhodolith communities from Floskjeret (78°18′N), Krossfjorden (79°08′N), and Mosselbukta (79°53′N), off Spitsbergen Island, Svalbard Archipelago, Norway. Strong seasonal variations in temperature, salinity, light regime, sea-ice coverage, and turbidity characterize these localities. The coralline algal flora consists of Lithothamnion glaciale and Phymatolithon tenue. Well-developed rhodoliths were recorded between 27 and 47 m water depth, while coralline algal encrustations on lithoclastic cobbles were detected down to 77 m water depth. At all sites, ambient waters were saturated with respect to both aragonite and calcite, and the rhodolith beds were located predominately at dysphotic water depths. The rhodolith-associated macrobenthic fauna included grazing organisms such as chitons and echinoids. With decreasing water depth, the rhodolith pavements were regularly overgrown by non-calcareous Polysiphonia-like red algae. The corallines are thriving and are highly specialized in their adaptations to the physical environment as well as in their interaction with the associated benthic fauna, which is similar to other polar rhodolith communities. The marine environment of Spitsbergen is already affected by a climate-driven ecological regime shift and will lead to an increased borealization in the near future, with presently unpredictable consequences for coralline red algal communities.

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).

Polar coralline red algae (Corallinales, Rhodophyta) that form rhodoliths have received little attention concerning their potential as ecosystem engineers and carbonate factories; although, recent findings revealed that they are much more widespread in polar waters than previously thought. The present study deals with the northernmost rhodolith communities currently known, discovered in 2006 at 80°31'N in Nordkappbukta (North Cape Bay) at Nordaustlandet, Svalbard. These perennial coralline algae must be adapted to extreme seasonality in terms of light regime (c. 4 months winter darkness), sea ice coverage, nutrient supply, turbidity of the water column, temperature and salinity. The rhodolith communities and their environment were investigated using multibeam swath bathymetry, CTD measurements, recordings of the photosynthetic active radiation (PAR) and determination of the water chemistry, seabed imaging and targeted sampling by means of the manned submersible JAGO as well as benthic collections with a dredge. The coralline flora was composed mainly of Lithothamnion glaciale, with a lesser amount of Phymatolithon tenue. Based on their distribution and development at different depth levels, a facies model was developed. Rhodoliths occurred between 30 and 51 m, while coralline algae attached to cobbles were present as deep as 78 m. Measurements of the PAR indicated their adaptation to extreme low light levels. Ambient waters were always saturated with reference to calcite and aragonite for the whole area. The rhodolith-associated macrobenthic fauna samples yielded 59 species, only one of which was typically Arctic, and the concomitant appearance of corallines and grazers kept the corallines free from epiphytes and coequally provided feeding grounds for the grazers. Overall, L. glaciale and P. tenue appeared to be well adapted to the extreme environment of the Arctic.

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 (1 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 (6 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.

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.

The rugged submarine topography of the Azores supports a diverse heterozoan association resulting in intense biotically-controlled carbonate-production and accumulation. In order to characterise this cold-water (C) factory a 2-year experiment was carried out in the southern Faial Channel to study the biodiversity of hardground communities and for budgeting carbonate production and degradation along a bathymetrical transect from the intertidal to bathyal 500 m depth. Seasonal temperatures peak in September (above a thermocline) and bottom in March (stratification diminishes) with a decrease in amplitude and absolute values with depth, and tidal-driven short-term fluctuations. Measured seawater stable isotope ratios and levels of dissolved nutrients decrease with depth, as do the calcium carbonate saturation states. The photosynthetic active radiation shows a base of the euphotic zone in ~70 m and a dysphotic limit in ~150 m depth. Bioerosion, being primarily a function of light availability for phototrophic endoliths and grazers feeding upon them, is ~10 times stronger on the illuminated upside versus the shaded underside of substrates in the photic zone, with maximum rates in the intertidal (−631 g/m2/yr). Rates rapidly decline towards deeper waters where bioerosion and carbonate accretion are slow and epibenthic/endolithic communities take years to mature. Accretion rates are highest in the lower euphotic zone (955 g/m2/yr), where the substrate is less prone to hydrodynamic force. Highest rates are found – inversely to bioerosion – on down-facing substrates, suggesting that bioerosion may be a key factor governing the preferential settlement and growth of calcareous epilithobionts on down-facing substrates. In context of a latitudinal gradient, the Azores carbonate cycling rates plot between known values from the cold-temperate Swedish Kosterfjord and the tropical Bahamas, with a total range of two orders in magnitude. Carbonate budget calculations for the bathymetrical transect yield a mean 266.9 kg of epilithic carbonate production, −54.6 kg of bioerosion, and 212.3 kg of annual net carbonate production per metre of coastline in the Azores C factory.

The ability of coral reefs to engineer complex three-dimensional habitats is central to their success and the rich biodiversity they support. In tropical reefs, encrusting coralline algae bind together substrates and dead coral framework to make continuous reef structures, but beyond the photic zone, the cold-water coral Lophelia pertusa also forms large biogenic reefs, facilitated by skeletal fusion. Skeletal fusion in tropical corals can occur in closely related or juvenile individuals as a result of non-aggressive skeletal overgrowth or allogeneic tissue fusion, but contact reactions in many species result in mortality if there is no ‘self-recognition’ on a broad species level. This study reveals areas of ‘flawless’ skeletal fusion in Lophelia pertusa, potentially facilitated by allogeneic tissue fusion, are identified as having small aragonitic crystals or low levels of crystal organisation, and strong molecular bonding. Regardless of the mechanism, the recognition of ‘self’ between adjacent L. Pertusa colonies leads to no observable mortality, facilitates ecosystem engineering and reduces aggression-related energetic expenditure in an environment where energy conservation is crucial. The potential for self-recognition at a species level, and subsequent skeletal fusion in framework-forming cold-water corals is an important first step in understanding their significance as ecological engineers in deep-seas worldwide.

It is important to understand how marine calcifying organisms may acclimatize to ocean acidification to assess their survival over the coming century. We cultured the cold water coralline algae, Lithothamnion glaciale, under elevated pCO2 (408, 566, 770, and 1024 μatm) for 10 months. The results show that the cell (inter and intra) wall thickness is maintained, but there is a reduction in growth rate (linear extension) at all elevated pCO2. Furthermore a decrease in Mg content at the two highest CO2 treatments was observed. Comparison between our data and that at 3 months from the same long-term experiment shows that the acclimation differs over time since at 3 months, the samples cultured under high pCO2 showed a reduction in the cell (inter and intra) wall thickness but a maintained growth rate. This suggests a reallocation of the energy budget between 3 and 10 months and highlights the high degree plasticity that is present. This might provide a selective advantage in future high CO2 world.

SummaryCoral reefs face multiple anthropogenic threats, from pollution and overfishing to the dual effects of greenhouse gas emissions: rising sea temperature and ocean acidification [1]. While the abundance of coral has declined in recent decades [2, 3], the implications for humanity are difficult to quantify because they depend on ecosystem function rather than the corals themselves. Most reef functions and ecosystem services are founded on the ability of reefs to maintain their three-dimensional structure through net carbonate accumulation [4]. Coral growth only constitutes part of a reef’s carbonate budget; bioerosion processes are influential in determining the balance between net structural growth and disintegration [5, 6]. Here, we combine ecological models with carbonate budgets and drive the dynamics of Caribbean reefs with the latest generation of climate models. Budget reconstructions using documented ecological perturbations drive shallow (6–10 m) Caribbean forereefs toward an increasingly fragile carbonate balance. We then projected carbonate budgets toward 2080 and contrasted the benefits of local conservation and global action on climate change. Local management of fisheries (specifically, no-take marine reserves) and the watershed can delay reef loss by at least a decade under “business-as-usual” rises in greenhouse gas emissions. However, local action must be combined with a low-carbon economy to prevent degradation of reef structures and associated ecosystem services.

In the recent discussion how biotic systems may react to ocean acidification caused by the rapid rise in carbon dioxide partial pressure (pCO(2)) 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-CO(2) 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 pCO(2) confirms a significant enforcement of the sponges' bioerosion capacity with increasing pCO(2) 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.