Seawater carbonate chemistry and calcification rate of cold-water coral Lophelia pertusa during experiments, 2011

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.

Effects of high temperature and CO2 on intracellular DMSP in the cold-water coral Lophelia pertusa

Significant warming and acidification of the oceans is projected to occur by the end of the century. CO2 vents, areas of upwelling and downwelling, and potential leaks from carbon capture and storage facilities may also cause localised environmental changes, enhancing or depressing the effect of global climate change. Cold-water coral ecosystems are threatened by future changes in carbonate chemistry, yet our knowledge of the response of these corals to high temperature and high CO2 conditions is limited. Dimethylsulphoniopropionate (DMSP), and its breakdown product dimethylsulphide (DMS), are putative antioxidants that may be accumulated by invertebrates via their food or symbionts, although recent research suggests that some invertebrates may also be able to synthesise DMSP. This study provides the first information on the impact of high temperature (12 °C) and high CO2 (817 ppm) on intracellular DMSP in the cold-water coral Lophelia pertusa from the Mingulay Reef Complex, Scotland (56°49' N, 07°23' W), where in situ environmental conditions are meditated by tidally induced downwellings. An increase in intracellular DMSP under high CO2 conditions was observed, whilst water column particulate DMS + DMSP was reduced. In both high temperature treatments, intracellular DMSP was similar to the control treatment, whilst dissolved DMSP + DMS was not significantly different between any of the treatments. These results suggest that L. pertusa accumulates DMSP from the surrounding water column; uptake may be up-regulated under high CO2 conditions, but mediated by high temperature. These results provide new insight into the biotic control of deep-sea biogeochemistry and may impact our understanding of the global sulphur cycle, and the survival of cold-water corals under projected global change.

Differential response of two Mediterranean cold-water coral species to ocean acidification

Cold-water coral (CWC) reefs constitute one of the most complex deep-sea habitats harboring a vast diversity of associated species. Like other tropical or temperate framework builders, these systems are facing an uncertain future due to several threats, such as global warming and ocean acidification. In the case of Mediterranean CWC communities, the effect may be exacerbated due to the greater capacity of these waters to absorb atmospheric CO2 compared to the global ocean. Calcification in these organisms is an energy-demanding process, and it is expected that energy requirements will be greater as seawater pH and the availability of carbonate ions decrease. Therefore, studies assessing the effect of a pH decrease in skeletal growth, and metabolic balance are critical to fully understand the potential responses of these organisms under a changing scenario. In this context, the present work aims to investigate the medium- to long-term effect of a low pH scenario on calcification and the biochemical composition of two CWCs from the Mediterranean, Dendrophyllia cornigera and Desmophyllum dianthus. After 314 d of exposure to acidified conditions, a significant decrease of 70 % was observed in Desmophyllum dianthus skeletal growth rate, while Dendrophyllia cornigera showed no differences between treatments. Instead, only subtle differences between treatments were observed in the organic matter amount, lipid content, skeletal microdensity, or porosity in both species, although due to the high variability of the results, these differences were not statistically significant. Our results also confirmed a heterogeneous effect of low pH on the skeletal growth rate of the organisms depending on their initial weight, suggesting that those specimens with high calcification rates may be the most susceptible to the negative effects of acidification.

Radiocarbon dating on cold-water corals and planktonic foraminifera, grain size analysis, stable oxygen isotopes, and XRF data of sediment cores from the Alboran Sea

Cold-water corals are common along the Moroccan continental margin off Melilla in the Alboran Sea (western Mediterranean Sea), where they colonise and largely cover mound and ridge structures. Radiocarbon ages of the reef-forming coral species Lophelia pertusa and Madrepora oculata sampled from those structures, reveal that they were prolific in this area during the last glacial-interglacial transition with pronounced growth periods covering the Bølling-Allerød interstadial (13.5-12.8 ka BP) and the Early Holocene (11.3-9.8 ka BP). Their proliferation during these periods is expressed in vertical accumulation rates for an individual coral ridge of 266-419 cm ka**-1 that consists of coral fragments embedded in a hemipelagic sediment matrix. Following a period of coral absence, as noted in the records, cold-water corals re-colonised the area during the Mid-Holocene (5.4 ka BP) and underwater photographs indicate that corals currently thrive there. It appears that periods of sustained cold-water coral growth in the Melilla Coral Province were closely linked to phases of high marine productivity. The increased productivity was related to the deglacial formation of the most recent organic rich layer in the western Mediterranean Sea and to the development of modern circulation patterns in the Alboran Sea.

Seawater carbonate chemistry, nutrients and calcification during experiments with cold-water scleractinian corals (Lophelia pertusa, Madrepora oculata and Desmophyllum dianthus), 2011

Global environmental changes, including ocean acidification, have been identified as a major threat to scleractinian corals. General predictions are that ocean acidification will be detrimental to reef growth and that 40 to more than 80 per cent of present-day reefs will decline during the next 50 years. Cold-water corals (CWCs) are thought to be strongly affected by changes in ocean acidification owing to their distribution in deep and/or cold waters, which naturally exhibit a CaCO3 saturation state lower than in shallow/warm waters. Calcification was measured in three species of Mediterranean cold-water scleractinian corals (Lophelia pertusa, Madrepora oculata and Desmophyllum dianthus) on-board research vessels and soon after collection. Incubations were performed in ambient sea water. The species M. oculata was additionally incubated in sea water reduced or enriched in CO2. At ambient conditions, calcification rates ranged between -0.01 and 0.23% d-1. Calcification rates of M. oculata under variable partial pressure of CO2 (pCO2) were the same for ambient and elevated pCO2 (404 and 867 µatm) with 0.06 ± 0.06% d-1, while calcification was 0.12 ± 0.06% d-1 when pCO2 was reduced to its pre-industrial level (285 µatm). This suggests that present-day CWC calcification in the Mediterranean Sea has already drastically declined (by 50%) as a consequence of anthropogenic-induced ocean acidification.

Physiological response of the cold-water coral Desmophyllum dianthus to thermal stress and ocean acidification

Rising temperatures and ocean acidification driven by anthropogenic carbon emissions threaten both tropical and temperate corals. However, the synergistic effect of these stressors on coral physiology is still poorly understood, in particular for cold-water corals. This study assessed changes in key physiological parameters (calcification, respiration and ammonium excretion) of the widespread cold-water coral Desmophyllum dianthus maintained for 8 months at two temperatures (ambient 12 °C and elevated 15 °C) and two pCO2 conditions (ambient 390 ppm and elevated 750 ppm). At ambient temperatures no change in instantaneous calcification, respiration or ammonium excretion rates was observed at either pCO2 levels. Conversely, elevated temperature (15 °C) significantly reduced calcification rates, and combined elevated temperature and pCO2 significantly reduced respiration rates. Changes in the ratio of respired oxygen to excreted nitrogen (O:N), which provides information on the main sources of energy being metabolized, indicated a shift from mixed use of protein and carbohydrate/lipid as metabolic substrates under control conditions, to less efficient protein-dominated catabolism under both stressors. Overall, this study shows that the physiology of D. dianthus is more sensitive to thermal than pCO2 stress, and that the predicted combination of rising temperatures and ocean acidification in the coming decades may severely impact this cold-water coral species.

The influence of seawater pH on U/ Ca ratios in the scleractinian cold-water coral Lophelia pertusa

The increasing pCO2 in seawater is a serious threat for marine calcifiers and alters the biogeochemistry of the ocean. Therefore, the reconstruction of past-seawater properties and their impact on marine ecosystems is an important way to investigate the underlying mechanisms and to better constrain the effects of possible changes in the future ocean. Cold-water coral (CWC) ecosystems are biodiversity hotspots. Living close to aragonite undersaturation, these corals serve as living laboratories as well as archives to reconstruct the boundary conditions of their calcification under the carbonate system of the ocean. We investigated the reef-building CWC Lophelia pertusa as a recorder of intermediate ocean seawater pH. This species-specific field calibration is based on a unique sample set of live in situ collected L. pertusa and corresponding seawater samples. These data demonstrate that uranium speciation and skeletal incorporation for azooxanthellate scleractinian CWCs is pH dependent and can be reconstructed with an uncertainty of ±0.15. Our Lophelia U / Ca-pH calibration appears to be controlled by the high pH values and thus highlighting the need for future coral and seawater sampling to refine this relationship. However, this study recommends L. pertusa as a new archive for the reconstruction of intermediate water mass pH and hence may help to constrain tipping points for ecosystem dynamics and evolutionary characteristics in a changing ocean.

Calcification is not the Achilles'heel of cold-water corals in an acidifying ocean

Ocean acidification is thought to be a major threat to coral reefs: laboratory evidence and CO2 seep research has shown adverse effects on many coral species, although a few are resilient. There are concerns that cold-water corals are even more vulnerable as they live in areas where aragonite saturation (Omega ara) is lower than in the tropics and is falling rapidly due to CO2 emissions. Here, we provide laboratory evidence that net (gross calcification minus dissolution) and gross calcification rates of three common cold-water corals, Caryophyllia smithii, Dendrophyllia cornigera, and Desmophyllum dianthus, are not affected by pCO2 levels expected for 2100 (pCO2 1058 µatm, Omega ara 1.29), and nor are the rates of skeletal dissolution in D. dianthus. We transplanted D. dianthus to 350 m depth (pHT 8.02; pCO2 448 µatm, Omega ara 2.58) and to a 3 m depth CO2 seep in oligotrophic waters (pHT 7.35; pCO2 2879 µatm, Omega ara 0.76) and found that the transplants calcified at the same rates regardless of the pCO2 confirming their resilience to acidification, but at significantly lower rates than corals that were fed in aquaria. Our combination of field and laboratory evidence suggests that ocean acidification will not disrupt cold-water coral calcification although falling aragonite levels may affect other organismal physiological and/or reef community processes.

Seawater carbonate chemistry and calcification of two mediterranean cold-water coral species in a laboratory experiment

Deep-water ecosystems are characterized by relatively low carbonate concentration values and, due to ocean acidification (OA), these habitats might be among the first to be exposed to undersaturated conditions in the forthcoming years. However, until now, very few studies have been conducted to test how cold-water coral (CWC) species react to such changes in the seawater chemistry. The present work aims to investigate the mid-term effect of decreased pH on calcification of the two branching CWC species most widely distributed in the Mediterranean, Lophelia pertusa and Madrepora oculata. No significant effects were observed in the skeletal growth rate, microdensity and porosity of both species after 6 months of exposure. However, while the calcification rate of M. oculata was similar for all colony fragments, a heterogeneous skeletal growth pattern was observed in L. pertusa, the younger nubbins showing higher growth rates than the older ones. A higher energy demand is expected in these young, fast-growing fragments and, therefore, a reduction in calcification might be noticed earlier during long-term exposure to acidified conditions.

Planktonic foraminiferal faunas and stable isotope ratios of sediments from the Benguela Upwelling System

Two sediment cores retrieved from the continental slope in the Benguela Upwelling System, GeoB 1706 (19°33.7'S 11°10.5'E) and GeoB 1711 (23°18.9'S, 12°22.6'E), reveal striking variations in planktonic foraminiferal abundances during the last 160,000 years. These fluctuations are investigated to assess changes in the intensity and position of the upwelling centres off Namibia. Four species make up over 95% of the variation within the core, and enable the record to be divided into episodes characterized by particular planktonic foraminiferal assemblages. The fossil assemblages have meaningful ecological significance when compared to those of the modern day and the relationship to their environment. The cold-water planktonic foraminifer, Neogloboquadrina pachyderma sinistral [N. pachyderma (s)], dominates the modern-day, coastal upwelling centres, and Neogloboquadrina pachyderma dextral and Globigerina bulloides characterize the fringes of the upwelling cells. Globorotalia inflata is representative of the offshore boundary between newly upwelled waters and the transitional, reduced nutrient levels of the subtropical waters. In the fossil record, episodes of high N. pachyderma (s) abundances are interpreted as evidence of increased upwelling intensity, and the associated increase in nutrients. The N. pachyderma (s) record suggests temporal shifts in the intensity of upwelling, and corresponding trophic domains, that do not follow the typical glacial-interglacial pattern. Periods of high N. pachyderma (s) abundance describe rapid, discrete events dominating isotope stages 3 and 2. The timing of these events correlates to the temporal shifts of the Angola-Benguela Front (Jansen et al., 1997) situated to the north of the Walvis Ridge. Absence of high abundances of N. pachyderma (s) from the continental slope of the southern Cape Basin indicates that Southern Ocean surface water advection has not exerted a major influence on the Benguela Current System. The coincidence of increased upwelling intensity with the movement of the Angola-Benguela Front can be interpreted mainly by changes in strength and zonality of the trade wind system.

Short-term metabolic and growth responses of the cold-water coral lophelia pertusa to ocean acidification

Cold-water corals are amongst the most three-dimensionally complex deep-sea habitats known and are associated with high local biodiversity. Despite their importance as ecosystem engineers, little is known about how these organisms will respond to projected ocean acidification. Since preindustrial times, average ocean pH has already decreased from 8.2 to ~ 8.1. Predicted CO2 emissions will decrease this by up to another 0.3 pH units by the end of the century. This decrease in pH may have a wide range of impacts upon marine life, and in particular upon calcifiers such as cold-water corals. Lophelia pertusa is the most widespread cold-water coral (CWC) species, frequently found in the North Atlantic. Data here relate to a short term data set (21 days) on metabolism and net calcification rates of freshly collected L. pertusa from Mingulay Reef Complex, Scotland. These data from freshly collected L. pertusa from the Mingulay Reef Complex will help define the impact of ocean acidification upon the growth, physiology and structural integrity of this key reef framework forming species.

Sediment and ice characteristics at a buried ice-wedge system at Barrow

Barrow, the northernmost point in Alaska, is one of the most intensively studied areas in the Arctic. However, paleoenvironmental evidence is limited for northern Alaska for the Lateglacial-Holocene transition. For a regional paleoenvironmental reconstruction, we investigated a permafrost ice-wedge tunnel near Barrow, Alaska. The studied site was first excavated in the early 1960s and intercepts a buried ice-wedge system at 3-6 m depth below the surface. A multi-methodological approach was applied to this buried ice-wedge system and the enclosing sediments, which in their combination, give new insight into the Late Quaternary environmental and climate history. Results of geochronological, sedimentological, cryolithological, paleoecological, isotope geochemical and microbiological studies reflect different stages of mid to late Wisconsin (MW to LW), Allerod (AD), Younger Dryas (YD), Preboreal (PB), and Late Holocene paleoenvironmental evolution. The LW age of the site is indicated by AMS dates in the surrounding sediments of 21.7 kyr BP at the lateral contact of the ice-wedge system as well as 39.5 kyr BP below the ice-wedge system. It is only recently that in this region, stable isotope techniques have been employed, i.e. to characterize different types of ground ice. The stable isotope record (oxygen: d18O; hydrogen: dD) of two intersecting ice wedges suggests different phases of the northern Alaskan climate history from AD to PB, with radiocarbon dates from 12.4 to 9.9 kyr BP (ranging from 14.8 to 10.6 kyr cal BP). Stable isotope geochemistry of ice wedges reveals winter temperature variations of the Lateglacial-Holocene transition including a prominent YD cold period, clearly separated from the warmer AD and PB phases. YD is only weakly developed in summer temperature indicators (such as pollen) for the northern Alaska area, and by consequence, the YD cold stadial was here especially related to the winter season. This highlights that the combination of winter and summer indicators comprehensively describes the seasonality of climate-relevant processes in discrete time intervals. The stable isotope record for the Barrow buried ice-wedge system documents for the first time winter climate change at the Lateglacial-Holocene transition continuously and at relatively high (likely centennial) resolution.