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.

Cape Lookout cold-water coral area, coral ages and environmental parameters for the last glacial cycle

Climatic and oceanographic changes, as occurring at a glacial-interglacial scale, may alter the environmental conditions needed for the development of prolific cold-water coral reefs and mounds. Studies constraining the temporal distribution of cold-water corals in the NE Atlantic suggested the cyclic changes of the Atlantic Meridional Overturning Circulation as the main driver for the development and dispersal of cold-water coral ecosystems. However, conclusions were hindered by lack of data from the NW Atlantic. Aiming to overcome this lack of data, the temporal occurrence of cold-water corals in the Cape Lookout area along the southeastern US margin was explored by U-series dating. Furthermore, the local influence of the regional water masses, namely the Gulf Stream, on cold-water coral proliferation and occurrence since the Last Glacial Maximum was examined. Results suggest that the occurrence of cold-water corals in the Cape Lookout area is restricted to interglacial periods, with corals being present during the last ~7 kyr and also during the Eemian (~125 ka). The reconstructed local environmental conditions suggest an offshore displacement of the Gulf Stream and increased influence from the Mid-Atlantic Bight shelf waters during the last glacial period. During the deglacial sea level rise, the Gulf Stream moved coastward providing present-day-like conditions to the surface waters. Nevertheless, present-day conditions at the ocean sea floor were not established before 7.5 cal ka BP once the ultimate demise of the Laurentide ice-sheet caused the final sea level rise and the displacement of the Gulf Stream to its present location. Occasional presence of the Gulf Stream over the site during the Mid- to Late Holocene coincides with enhanced bottom current strength and a slightly higher bottom water temperature, which are environmental conditions that are favorable for cold-water coral growth.

Mediterranean cold-water coral measurements

This study presents aggradation rates supplemented for the first time by carbonate accumulation rates from Mediterranean cold-water coral sites considering three different regional and geomorphological settings: (i) a cold-water coral ridge (eastern Melilla coral province, Alboran Sea), (ii) a cold-water coral rubble talus deposit at the base of a submarine cliff (Urania Bank, Strait of Sicily) and (iii) a cold-water coral deposit rooted on a predefined topographic high overgrown by cold-water corals (Santa Maria di Leuca coral province, Ionian Sea). The mean aggradation rates of the respective cold-water coral deposits vary between 10 and 530 cm kyr?1 and the mean carbonate accumulation rates range between 8 and 396 g cm?2 kyr?1 with a maximum of 503 g cm?2 kyr?1 reached in the eastern Melilla coral province. Compared to other deep-water depositional environments the Mediterranean cold-water coral sites reveal significantly higher carbonate accumulation rates that were even in the range of the highest productive shallow-water Mediterranean carbonate factories (e.g. Cladocora caespitosa coral reefs). Focusing exclusively on cold-water coral occurrences, the carbonate accumulation rates of the Mediterranean cold-water coral sites are in the lower range of those obtained for the prolific Norwegian coral occurrences, but exhibit much higher rates than the cold-water coral mounds off Ireland. This study clearly indicates that cold-water corals have the potential to act as important carbonate factories and regional carbonate sinks within the Mediterranean Sea. Moreover, the data highlight the potential of cold-water corals to store carbonate with rates in the range of tropical shallow-water reefs. In order to evaluate the contribution of the cold-water coral carbonate factory to the regional or global carbonate/carbon cycle, an improved understanding of the temporal and spatial variability in aggradation and carbonate accumulation rates and areal estimates of the respective regions is needed.

Stable isotope composition of carbonate components of Rhaetian shallow-water limestones of the Wombat Plateau

Stable oxygen- and carbon-isotope ratios of Rhaetian (upper Triassic) limestone samples from the Wombat Plateau, northwest Australia, were measured to explore possible diagenetic pathways that the material underwent after deposition in a shallow-water environment, before plateau submergence in the Early Cretaceous. Host sediment isotopic values cluster near typical marine carbonate values (d18O ranging from -2.57 per mil to +1.78 per mil and d13C, from +2.45 per mil to +4.01 per mil). Isotopic values of equant clear calcite lining or filling rock pores also plot in the field of marine cements (d18O = +1.59 per mil to -2.24 per mil and d13C = +4.25 per mil to +2.57 per mil), while isotopic values for neomorphic calcites replacing skeletal (megalodontid shell) carbonate material show a wider scatter of oxygen and carbon values, d18O ranging from +2.73 per milo to -6.2 per mil and d13C, from +5.04 per mil to +1.22 per mil. Selective dissolution of metastable carbonate phases (aragonite?) and neomorphic replacement of skeletal material probably occurred in a meteoric phreatic environment, although replacement products (inclusion-rich microspar, clear neomorphic spar, etc.) retained the original marine isotopic signature because transformation probably occurred in a closed system dominated by the composition of the dissolving phases (high rock/water ratio). The precipitation of late-stage equant (low-Mg?) calcite cement in the pores occurred in the presence of normal marine waters, probably in a deep-water environment, after plateau drowning. Covariance of d18O and d13C toward negative values indeed suggests influence of meteorically modified fluids. However, none of the samples shows negative carbon values, excluding the persistence of organic-rich soils on subaerial karstic surfaces (Caribbean-style diagenesis). Petrographical and geochemical data are consistent with the sedimentological evidence of plateau drowning in post-Rhaetian times and with a submarine origin of the >70-m.y.-long Jurassic hiatus.

Sedimentology and stable oxygen isotope ratios of foraminifera from cores of the Porcupine Seabight, Northeast Atlantic

Large carbonate mound structures have been discovered in the northern Porcupine Seabight (Northeast Atlantic) at depths between 600 and 1000 m. These mounds are associated with the growth of deep-sea corals Lophelia pertusa and Madrepra oculata. In this study, three sediment cores have been analysed. They are from locations close to Propeller Mound, a 150 m high ridge-like feature covered with a cold-water coral ecosystem at its upper flanks. The investigations are concentrated on grain-size analyses, carbon measurements and on the visual description of the cores and computer tomographic images, to evaluate sediment content and structure. The cores portray the depositional history of the past ~31 kyr BP, mainly controlled by sea-level fluctuations and the climate regime with the advance and retreat of the Irish Ice Sheet onto the Irish Mainland Shelf. A first advance of glaciers is indicated by a turbiditic release slightly older than 31 kyr BP, coherent with Heinrich event 3 deposition. During Late Marine Isotope Stage 3 (MIS 3) and MIS 2 shelf erosion prevailed with abundant gravity flows and turbidity currents. A change from glaciomarine to hemipelagic contourite sedimentation during the onset of the Holocene indicates the establishment of the strong, present-day hydrodynamic regime at intermediate depths. The general decrease in accumulation of sediments with decreasing distance towards Propeller Mound suggests that currents (turbidity currents, gravity flows, bottom currents) had a generally stronger impact on the sediment accumulation at the mound base for the past ~31 kyr BP, respectively.

Benthic foraminifera, stable isotope ratios and carbon concentrations of sediment cores from Propeller Mound and Porcupine Srabight

On- and off-mound sediment cores from Propeller Mound (Hovland Mound province, Porcupine Seabight) were analysed to understand better the evolution of a carbonate mound. The evaluation of benthic foraminiferal assemblages from the off-mound position helps to determine the changes of the environmental controls on Propeller Mound in glacial and interglacial times. Two different assemblages describe the Holocene and Marine Isotope Stage (MIS) 2 and late MIS 3 (~31 kyr BP). The different assemblages are related to changes in oceanographic conditions, surface productivity and the waxing and waning of the British Irish Ice Sheet (BIIS) during the last glacial stages. The interglacial assemblage is related to a higher supply of organic material and stronger current intensities in water depth of recent coral growth. During the last glaciation the benthic faunas showed high abundances of cassidulinid species, implying cold bottom waters and a reduced availability of organic matter. High sedimentation rates and the domination of Elphidium excavatum point to shelf erosion related to sea-level lowering (~50 m) and the progradation of the BIIS onto the shelf. A different assemblage described for the on-mound core is dominated by Discanomalina coronata, Gavelinopsis translucens, Planulina ariminensis, Cibicides lobatulus and to a lower degree by Hyrrokkin sarcophaga. These species are only found or show significantly higher relative abundances in on-mound samples and their maximum contribution in the lower part of the record indicates a higher coral growth density on Propeller Mound in an earlier period. They are less abundant during the Holocene, however. This dataset portrays the boundary conditions of the habitable range for the cold-water coral Lophelia pertusa, which dominates the deep-water reefal ecosystem on the upper flanks of Propeller Mound. The growth of this ecosystem occurs during interglacial and interstadial periods, whereas a retreat of corals is documented in the absence of glacial sediments on-mound. Glacial conditions with cold intermediate waters, a weak current regime and high sedimentation rates provide an unfavourable environmental setting for Lophelia corals to grow. A Late Pleistocene decrease is observed in the mound growth for Propeller Mound, which might face its complete burial in the future, as it already happened to the buried mounds of the Magellan Mound province further north.

Stable oxygen isotope ratios and grain size analyses of sediment cores from the Propeller Mound, Northeast Atlantic

The first detailed stratigraphic record from a deep-water carbonate mound in the Northeast Atlantic based on absolute datings (U/Th and AMS 14C) and stable oxygen isotope records reveals that its top sediment sequences are condensed by numerous hiatuses. According to stable isotope data, mainly sediments with an intermediate signal are preserved on the mound, while almost all fully glacial and interglacial sediments have either not been deposited or have been eroded later. The resulting hiatuses reduce the Late Pleistocene sediment accumulation at Propeller Mound to amounts smaller than the background sedimentation. The hiatuses most likely result due to the sweeping of the mound in turn with the re-establishment of vigour interglacial circulation patterns after sluggish current regimes during glacials. Thus, within the discussion if internal, fluid-driven or external environmentally driven processes control the evolution of such carbonate mounds, our findings for Propeller Mound clearly point to environmental forcing as the dominant mechanism shaping deep-water carbonate mounds in the NE Atlantic during the Late Pleistocene and Holocene.

Deep thermohaline Circulation in the low-latitude Atlantic during the Last Glacial

Present-day low-latitude eastern and western Atlantic basins are geochemically distinct below the sill depth of the Mid-Atlantic Ridge. While Antarctic Bottom Water (AABW) circulates freely in the western Atlantic, flow into the eastern Atlantic is restricted below 4 km which results in filling the abyssal depths of this basin with water of geochemical similarity to nutrient depleted North Atlantic Deep Water. Using carbon isotopes and Cd/Ca ratios in benthic foraminifera we reconstruct the geochemistry of these basins during the last glacial maximum. Results indicate that deep eastern and western Atlantic basins became geochemically identical during the last glacial. This was achieved by shoaling of the upper surface of AABW above the sill depth of the Mid-Atlantic Ridge, which allowed bottom waters in both basins to be filled with the same water mass. Although AABW became the dominant water mass in the deep eastern Atlantic basin during the glacial, Holocene-glacial delta13C-PO4 shifts in this basin are in Redfield proportions, unlike the disproportionate Holocene-glacial delta13C-PO4 shifts observed in the Southern Ocean. By examining the composition of deep and intermediate waters throughout the Atlantic, we show that this effect was induced by a change in gradient of the delta13C-PO4 deepwater mixing line during glacial times. Evidence from high-latitude planktonic data suggests that the change in gradient of the deepwater mixing line was brought about through a significant reduction in the thermodynamic effect on Southern Ocean surface waters. By using coupled delta13C-PO4 data to constrain the composition of end member water masses in the glacial Atlantic, we conclude that deep waters in the low-latitude glacial Atlantic were composed of a mixture of northern and southern source waters in a ratio of 1:3.

Plutonium and americium concentrations in sea water and in sinking particles

Vertical fluxes of 239+240Pu and 241Am and temporal changes in their inventories in the northwestern Mediterranean Sea have been examined through high-resolution water column sampling coupled with direct measurements of the vertical flux of particle-bound transuranics using time-series sediment traps. Water column profiles of both radionuclides showed well-defined sub-surface maxima (2391240Pu between 100-400 m; 241Am at 100-200 m and 800 m), the depths of which are a result of the different biogeochemical scavenging behavior of the two radionuclides. Comparison of deep water column (0-2,000 m) transuranic inventories with those derived from earlier measurements demonstrate that the total 2391240Pu inventory had not substantially changed between 1976-1990 whereas 241Am had decreased by approximately 24%. Enhanced scavenging of 241Am and a resultant, more rapid removal from the water column relative to 239+240Pu was also supported by the observation of elevated Am/Pu activity ratios in sinking particles collected in sediment traps at depth. Direct measurements of the downward flux of particulate 239+240Pu and 241Am compared with transuranic removal rates derived from observed total water column inventory differences over time, show that particles sinking out of deep waters (1,000-2,000 m) could account for 26-72% of the computed total annual 239+240Pu loss and virtually all of the 241Am removal from the water column. Upper water column (0-200 m) residence times based on direct flux measurements ranged from 20-30 yr for 239+240Pu and 5-10 yr for 241Am. The observation that 241Am/239+240Pu activity ratios in unfiltered Mediterranean seawater are six times lower than those in the north Pacific suggests the existence of a specific mechanism for enhanced scavenging and removal of 241Am from the generally oligotrophic waters of the open Mediterranean. It is proposed that atmospheric inputs of aluminosilicate particles transported by Saharan dust events which frequently occur in the Mediterranean region could enhance the geochemical scavenging and resultant removal of 241Am to the sediments.

Quantitative component analysis of the coarse fraction of surface sediments from the Persian Gulf

In the Persian Gulf and the Gulf of Oman marl forms the primary sediment cover, particularly on the Iranian side. A detailed quantitative description of the sediment components > 63 µ has been attempted in order to establish the regional distribution of the most important constituents as well as the criteria governing marl sedimentation in general. During the course of the analysis, the sand fraction from about 160 bottom-surface samples was split into 5 phi° fractions and 500 to 800 grains were counted in each individual fraction. The grains were cataloged in up to 40 grain type catagories. The gravel fraction was counted separately and the values calculated as weight percent. Basic for understanding the mode of formation of the marl sediment is the "rule" of independent availability of component groups. It states that the sedimentation of different component groups takes place independently, and that variation in the quantity of one component is independent of the presence or absence of other components. This means, for example, that different grain size spectrums are not necessarily developed through transport sorting. In the Persian Gulf they are more likely the result of differences in the amount of clay-rich fine sediment brought in to the restricted mouth areas of the Iranian rivers. These local increases in clayey sediment dilute the autochthonous, for the most part carbonate, coarse fraction. This also explains the frequent facies changes from carbonate to clayey marl. The main constituent groups of the coarse fraction are faecal pellets and lumps, the non carbonate mineral components, the Pleistocene relict sediment, the benthonic biogene components and the plankton. Faecal pellets and lumps are formed through grain size transformation of fine sediment. Higher percentages of these components can be correlated to large amounts of fine sediment and organic C. No discernable change takes place in carbonate minerals as a result of digestion and faecal pellet formation. The non-carbonate sand components originate from several unrelated sources and can be distinguished by their different grain size spectrum; as well as by other characteristics. The Iranian rivers supply the greatest amounts (well sorted fine sand). Their quantitative variations can be used to trace fine sediment transport directions. Similar mineral maxima in the sediment of the Gulf of Oman mark the path of the Persian Gulf outflow water. Far out from the coast, the basin bottoms in places contain abundant relict minerals (poorly sorted medium sand) and localized areas of reworked salt dome material (medium sand to gravel). Wind transport produces only a minimal "background value" of mineral components (very fine sand). Biogenic and non-biogenic relict sediments can be placed in separate component groups with the help of several petrographic criteria. Part of the relict sediment (well sorted fine sand) is allochthonous and was derived from the terrigenous sediment of river mouths. The main part (coarse, poorly sorted sediment), however, was derived from the late Pleistocene and forms a quasi-autochthonous cover over wide areas which receive little recent sedimentation. Bioturbation results in a mixing of the relict sediment with the overlying younger sediment. Resulting vertical sediment displacement of more than 2.5 m has been observed. This vertical mixing of relict sediment is also partially responsible for the present day grain size anomalies (coarse sediment in deep water) found in the Persian Gulf. The mainly aragonitic components forming the relict sediment show a finely subdivided facies pattern reflecting the paleogeography of carbonate tidal flats dating from the post Pleistocene transgression. Standstill periods are reflected at 110 -125m (shelf break), 64-61 m and 53-41 m (e.g. coare grained quartz and oolite concentrations), and at 25-30m. Comparing these depths to similar occurrences on other shelf regions (e. g. Timor Sea) leads to the conclusion that at this time minimal tectonic activity was taking place in the Persian Gulf. The Pleistocene climate, as evidenced by the absence of Iranian river sediment, was probably drier than the present day Persian Gulf climate. Foremost among the benthonic biogene components are the foraminifera and mollusks. When a ratio is set up between the two, it can be seen that each group is very sensitive to bottom type, i.e., the production of benthonic mollusca increases when a stable (hard) bottom is present whereas the foraminifera favour a soft bottom. In this way, regardless of the grain size, areas with high and low rates of recent sedimentation can be sharply defined. The almost complete absence of mollusks in water deeper than 200 to 300 m gives a rough sedimentologic water depth indicator. The sum of the benthonic foraminifera and mollusca was used as a relative constant reference value for the investigation of many other sediment components. The ratio between arenaceous foraminifera and those with carbonate shells shows a direct relationship to the amount of coarse grained material in the sediment as the frequence of arenaceous foraminifera depends heavily on the availability of sand grains. The nearness of "open" coasts (Iranian river mouths) is directly reflected in the high percentage of plant remains, and indirectly by the increased numbers of ostracods and vertebrates. Plant fragments do not reach their ultimate point of deposition in a free swimming state, but are transported along with the remainder of the terrigenous fine sediment. The echinoderms (mainly echinoids in the West Basin and ophiuroids in the Central Basin) attain their maximum development at the greatest depth reached by the action of the largest waves. This depth varies, depending on the exposure of the slope to the waves, between 12 to 14 and 30 to 35 m. Corals and bryozoans have proved to be good indicators of stable unchanging bottom conditions. Although bryozoans and alcyonarian spiculae are independent of water depth, scleractinians thrive only above 25 to 30 m. The beginning of recent reef growth (restricted by low winter temperatures) was seen only in one single area - on a shoal under 16 m of water. The coarse plankton fraction was studied primarily through the use of a plankton-benthos ratio. The increase in planktonic foraminifera with increasing water depth is here heavily masked by the "Adjacent sea effect" of the Persian Gulf: for the most part the foraminifera have drifted in from the Gulf of Oman. In contrast, the planktonic mollusks are able to colonize the entire Persian Gulf water body. Their amount in the plankton-benthos ratio always increases with water depth and thereby gives a reliable picture of local water depth variations. This holds true to a depth of around 400 m (corresponding to 80-90 % plankton). This water depth effect can be removed by graphical analysis, allowing the percentage of planktonic mollusks per total sample to be used as a reference base for relative sedimentation rate (sedimentation index). These values vary between 1 and > 1000 and thereby agree well with all the other lines of evidence. The "pteropod ooze" facies is then markedly dependent on the sedimentation rate and can theoretically develop at any depth greater than 65 m (proven at 80 m). It should certainly no longer be thought of as "deep sea" sediment. Based on the component distribution diagrams, grain size and carbonate content, the sediments of the Persian Gulf and the Gulf of Oman can be grouped into 5 provisional facies divisions (Chapt.19). Particularly noteworthy among these are first, the fine grained clayey marl facies occupying the 9 narrow outflow areas of rivers, and second, the coarse grained, high-carbonate marl facies rich in relict sediment which covers wide sediment-poor areas of the basin bottoms. Sediment transport is for the most part restricted to grain sizes < 150 µ and in shallow water is largely coast-parallel due to wave action at times supplemented by tidal currents. Below the wave base gravity transport prevails. The only current capable of moving sediment is the Persian Gulf outflow water in the Gulf of Oman.