Meiofaunal abundance, biomass, taxon richness and indices of nematode diversity of samples from the Santa Maria di Leuca Bank (southern Adriatic margin)

Deep-water coral ecosystems are hot spots of biodiversity and provide habitats and refuges for several deep-sea species. However, their role in shaping the biodiversity of the surrounding open slopes is still poorly known. We investigated how meiofaunal biodiversity varies with and is related to the occurrence of deep-water living scleractinian corals and coral rubble in two deep-sea areas (the Rockall Bank, northeastern Atlantic) and the Santa Maria di Leuca (central Mediterranean). In both areas, replicated sampling on alive and dead coral areas and from the adjacent slope sediments without corals (at the same and increasing depths) allowed us to demonstrate that sediments surrounding the living corals and coral rubble were characterised by higher meiofaunal biodiversity (as number of higher taxa, and nematode species richness) than the slope sediments. Despite the soft sediments surrounding the living coral having a higher nutritional value than those not associated with corals, with the opposite seen for coral rubble, the presence of both alive and dead corals had a significant effect on nematode assemblages. Our data suggest that, due particularly to the effects on habitat heterogeneity/complexity, both living coral and coral rubble promoted higher biodiversity levels than in surrounding slope sediments. We conclude that the protection of deep-water corals can be crucial to preserve the biodiversity of surrounding open slopes, and that the protection of dead corals, a so-far almost neglected habitat in terms of biological conservation, can further contribute to the maintenance of a high deep-sea biodiversity along continental margins.

Stable Isotope composition of particulate organic matter and fauna in a deep Mediterranean cold-water coral bank

Cold-water corals (CWC) are frequently reported from deep sites with locally accelerated currents that enhance seabed food particle supply. Moreover, zooplankton likely account for ecologically important prey items, but their contribution to CWC diet remains unquantified. We investigated the benthic food web structure of the recently discovered Santa Maria di Leuca (SML) CWC province (300 to 1100 m depth) located in the oligotrophic northern Ionian Sea. We analyzed stable isotopes (delta13C and delta15N) of the main consumers (including ubiquitous CWC species) exhibiting different feeding strategies, zooplankton, suspended particulate organic matter (POM) and sedimented organic matter (SOM). Zooplankton and POM were collected 3 m above the coral colonies in order to assess their relative contributions to CWC diet. The delta15N of the scleractinians Desmophyllum dianthus, Madrepora oculata and Lophelia pertusa and the gorgonian Paramuricea cf. macrospinawere consistent with a diet mainly composed of zooplankton. The antipatharian Leiopathes glaberrima was more 15N- depletedthan other cnidarians, suggesting a lower contribution of zooplankton to its diet. Our delta13C data clearly indicate that the benthic food web of SML is exclusively fuelled by carbon of phytoplanktonic origin. Nevertheless, consumers feeding at the water sediment interface were more 13C-enriched than consumers feeding above the bottom (i.e. living corals and their epifauna). This pattern suggests that carbon is assimilated via 2 trophic pathways: relatively fresh phytoplanktonic production for 13C-depleted consumers and more decayed organic matter for 13C-enriched consumers. When the delta13C values of consumers were corrected for the influence of lipids (which are significantly 13C-depleted relative to other tissue components), our conclusions remained unchanged, except in the case of L. glaberrima which could assimilate a mixture of zooplankton and resuspended decayed organic matter.

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.

(Table 2) Chemical composition of skeletons of scleractinian non-zooxanthelate corals

The first data on chemical composition of nonreef-building non-zooxanthellate deep-sea corals presented in this publication allow us to identify following tendencies manifested in the biomineralization process. Comparison of concentration levels of some chemical elements in scleractinian corals and ambient ocean waters suggests that corals do not accumulate K in the process of biomineralization and weakly accumulate Mg, whereas Ca, Sr, Si, Al, Ti, Mn, Zn, Cu, Cd, Pb, and Fe are concentrated in skeletons of corals with enrichment coefficients of 10**3 to 10**7. Correlations between components contained in the skeletons of scleractinian corals suggest that the source of Al, Si, Fe, and Ti in them is the clayey constituent of bottom sediments and zooplankton, while trace elements are likely accumulated via bioassimilation from seawater. Such elements as Mn, Sr, Pb, and Cd can structurally substitute Ca in calcite and aragonite. Variations in concentrations of the elements in coral skeletons depending on their habitat depths are fairly significant. As could be expected Ca and Mg concentrations are prone to decrease with depth (R = -0.55 and -0.51, respectively), which can possibly be caused by partial dissolution of carbonate skeletons with increasing depth, whereas the Sr/Ca ratio does not depend on depth.

Corals, substrate and physical oceanography during the expedition Comau2012, Northern Chilean Patagonia

Cold-water corals provide an important habitat for a rich fauna along the continental margins and slopes. Although these azooxanthellate corals are considered particularly sensitive to ocean acidification, their responses to natural variations in pH and aragonite saturation are largely unknown due to the difficulty of studying their ecology in deep waters. Previous SCUBA investigations have shown an exceptionally shallow population of the cold-water coral Desmophyllum dianthus in near-surface waters of Comau Fjord, a stratified 480 m deep basin in northern Chilean Patagonia with suboxic deep waters. Here, we use a remotely operated vehicle to quantitatively investigate the distribution of D. dianthus and its physico-chemical drivers in so far uncharted naturally acidified waters. Remarkably, D. dianthus was ubiquitous throughout the fjord, but particularly abundant between 20 and 280 m depth in a pH range of 8.4 to 7.4. The persistence of individuals in aragonite-undersaturated waters suggests that present-day D. dianthus in Comau Fjord may show pre-acclimation or pre-adaptation to conditions of ocean acidification predicted to reach over 70% of the known deep-sea coral locations by the end of the century.

The HERMES cold-water coral database

The HERMES cold-water coral database is a combination of historical and published sclerectinia cold-water coral occurrences (mainly Lophelia pertusa) and new records of the HERMES project along the European margin. This database will be updated if new findings are reported. New or historical data can be sent to Ben De Mol (mailto:bendemol@ub.edu). Besides geocodes a second category indicates the coral species and if they are sampled alive or dead. If absolute dating is available of the corals this is provide together with the method. Only the framework building cold-water corals are selected: Lophelia pertusa, Madrepora oculata and common cold-water corals often associated with the framework builders like: Desmophyllum sp and Dendrophylia sp. in comments other observed corals are indicated. Another field indicates if the corals are part of a large build-up or solitary. A third category of parameters is referencing to the quality of the represented data. In this category are the following parameters indicated: source of reference, source type (such as Fishermen location, scientific paper, cruise reports). sample code and or name and sample type (e.g. rock dredge, grab, video line). These parameters must allow an assessment of the quality of the described parameters.

A deep-sea coral record of the North Atlantic

Our record of Younger Dryas intermediate-depth seawater D14C from North Atlantic deep-sea corals supports a link between abrupt climate change and intermediate ocean variability. Our data show that northern source intermediate water (~1700 m) was partially replaced by 14C-depleted southern source water at the onset of the event, consistent with a reduction in the rate of North Atlantic Deep Water formation. This transition requires the existence of large, mobile gradients of D14C in the ocean during the Younger Dryas. The D14C water column profile from Keigwin (2004) provides direct evidence for the presence of one such gradient at the beginning of the Younger Dryas (~12.9 ka), with a 100 per mil offset between shallow (<~2400 m) and deep water. Our early Younger Dryas data are consistent with this profile and also show a D14C inversion, with 35 per mil more enriched water at ~2400 m than at ~1700 m. This feature is probably the result of mixing between relatively well 14C ventilated northern source water and more poorly 14C ventilated southern source intermediate water, which is slightly shallower. Over the rest of the Younger Dryas our intermediate water/deepwater coral D14C data gradually increase, while the atmosphere D14C drops. For a very brief interval at ~12.0 ka and at the end of the Younger Dryas (11.5 ka), intermediate water D14C (~1200 m) approached atmospheric D14C. These enriched D14C results suggest an enhanced initial D14C content of the water and demonstrate the presence of large lateral D14C gradients in the intermediate/deep ocean in addition to the sharp vertical shift at ~2500 m. The transient D14C enrichment at ~12.0 ka occurred in the middle of the Younger Dryas and demonstrates that there is at least one time when the intermediate/deep ocean underwent dramatic change but with much smaller effects in other paleoclimatic records.

Neodymium isotope signature of Holocene corals

The neodymium isotopic composition of marine precipitates is increasingly recognized as a powerful tool for identifying changes in ocean circulation and mixing on million year to millennial timescales. Unlike nutrient proxies such as ?13C or Cd/Ca, Nd isotopes are not thought to be altered in any significant way by biological processes, and thus they can serve as a quasi-conservative water mass tracer. However, the application of Nd isotopes in understanding the role of thermohaline circulation in rapid climate change is currently hindered by the lack of direct constraints on the signature of the North Atlantic end-member through time. Here we present the first results of Nd isotopes measured in U-Th-dated deep-sea corals from the New England seamounts in the northwest Atlantic Ocean. Our data are consistent with the conclusion that the Nd isotopic composition of North Atlantic deep and intermediate water has remained nearly constant through the last glacial cycle. The results address long-standing concerns that there may have been significant changes in the Nd isotopic composition of the North Atlantic end-member during this interval and substantiate the applicability of this novel tracer on millennial timescales for paleoceanography research.

Rates of nitrogen cycling and chemo-autotrophy in the cold-water coral Lophelia pertusa

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.

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.

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.