Abstract
Similar to their tropical counterparts, cold-water corals (CWCs) are able to build large three-dimensional reef structures. These unique ecosystems are at risk due to ongoing climate change. In particular, ocean warming, ocean acidification and changes in the hydrological cycle may jeopardize the existence of CWCs. In order to predict how CWCs and their reefs or mounds will develop in the near future one important strategy is to study past fossil CWC mounds and especially shallow CWC ecosystems as they experience a greater environmental variability compared to other deep-water CWC ecosystems. We present results from a CWC mound off southern Norway. A sediment core drilled from this relatively shallow (~ 100 m) CWC mound exposes in full detail hydrographical changes during the late Holocene, which were crucial for mound build-up. We applied computed tomography, 230Th/U dating, and foraminiferal geochemical proxy reconstructions of bottom-water-temperature (Mg/Ca-based BWT), δ18O for seawater density, and the combination of both to infer salinity changes. Our results demonstrate that the CWC mound formed in the late Holocene between 4 kiloannum (ka) and 1.5 ka with an average aggradation rate of 104 cm/kiloyears (kyr), which is significantly lower than other Holocene Norwegian mounds. The reconstructed BWTMg/Ca and seawater density exhibit large variations throughout the entire period of mound formation, but are strikingly similar to modern in situ observations in the nearby Tisler Reef. We argue that BWT does not exert a primary control on CWC mound formation. Instead, strong salinity and seawater density variation throughout the entire mound sequence appears to be controlled by the interplay between the Atlantic Water (AW) inflow and the overlying, outflowing Baltic-Sea water. CWC growth and mound formation in the NE Skagerrak was supported by strong current flow, oxygen replenishment, the presence of a strong boundary layer and larval dispersal through the AW, but possibly inhibited by the influence of fresh Baltic Water during the late Holocene. Our study therefore highlights that modern shallow Norwegian CWC reefs may be particularly endangered due to changes in water-column stratification associated with increasing net precipitation caused by climate change.
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Introduction
Ecosystem engineering scleractinian cold-water corals (CWCs) create biodiversity and carbon cycling hotspots in the deep and cold regions in the world’s oceans (Roberts et al. 2009; Freiwald et al. 2021; White et al. 2012). These unique ecosystems are currently under threat due to ongoing climate change (Guinotte et al. 2006; Sweetman et al. 2017). In particular, climate change leads to an increase in atmospheric carbon dioxide (\(p_{{{\text{CO}}_{{2}} }}\)) causing a decrease in ocean pH, known as “ocean acidification” (e.g., Doney et al. 2009; Pachauri et al. 2014). This decrease in seawater pH leads to a shoaling of the aragonite saturation horizon (ASH), thereby limiting the ability of CWCs to build their skeletons (Guinotte et al. 2006). However, the oceans are not only a sink for CO2, but also store atmospheric heat being increasingly produced due to greenhouse effects (Glecker et al. 2016). Recent studies have highlighted that bathyal temperatures of the North Atlantic may increase by up to 3 °C by 2100 (Sweetman et al. 2017). Thus, the combined effect of decreasing seawater pH and increasing oceanic heat content may gradually jeopardize the existence of modern CWC reefs.
The main reef-forming CWC, Desmophyllum pertusum (formerly described as Lophelia pertusa, Addamo et al. 2016), is tolerant to a broad range of bottom water temperatures (BWT, 5–15 °C, Büscher et al. 2017; Dorey et al. 2020) suggesting that, to a certain degree, CWCs have the ability to cope with thermal stress. In order to obtain environmental and oceanographic thresholds controlling the occurrence of CWCs, ecological tolerance ranges have been studied by observation of modern reef systems (Dullo et al. 2008; Flögel et al. 2014; Juva et al. 2020, 2021) or by the correlation of CWC occurrences to global datasets (Freiwald 2002; Davies et al. 2008; Davies and Guinotte 2011). Additionally, cultivation experiments have been carried out in order to define parameters that inhibit or increase calcification or growth of CWCs (e.g., Form and Riebesell 2012; Büscher et al. 2017; Gammon et al. 2018; Maier et al. 2009). Based on these findings, a variety of parameters have been identified that control the distribution of CWCs and their growth: oxygen, temperature, seawater carbonate system parameters, nutrients, salinity and seawater density (Dodds et al. 2007; Davies et al. 2008; Dullo et al. 2008; Freiwald et al. 2009; White and Dorschel 2010; Flögel et al. 2014) as well as food supply, steered by surface productivity (White et al. 2005; Duineveld et al. 2007; White and Dorschel 2010). Other studies found CWC reef growth to be controlled by the interaction between increased bottom flow and local topography (Frederiksen et al. 1992; Mienis et al. 2007; Dorschel et al. 2007; Rüggeberg et al. 2011; Juva et al. 2020; Wienberg et al. 2020). Over the last 3 million years, CWCs have been able to build mound-like structures that are also composed of hemipelagic sediments and shells of associated fauna (e.g., De Mol et al. 2002; Kano et al. 2007; Raddatz et al. 2014; Rüggeberg et al. 2007). These “mounds” are predominantly formed by D. pertusum with contributions of Madrepora oculata, Solenosmilia variabilis, Batelia candina, and Enallopsammia profunda (Frank et al. 2011; Muñoz et al. 2012; Hebbeln et al. 2014; Raddatz et al. 2020). Here, we refer to the classification by Wienberg and Titschack (2017) highlighting that all three-dimensional seabed structures formed by CWCs are termed mounds.
On the Norwegian margin, CWC mound formation initiated after the Younger Dryas (YD) with the retreat of the Fennoscandian ice sheet at around 11 ka (Lindberg and Mienert 2005; Raddatz et al. 2016). During the Holocene their formation has been interrupted by regional climatic and oceanographic perturbations (López Correa et al. 2012; Douarin et al. 2014; Titschack et al. 2015; Raddatz et al. 2016). Modern CWC habitats off Norway extend from ~ 71° N to ~ 58° N (Fig. 1, Freiwald et al. 2021). These mounds are dominated by D. pertusum (Freiwald et al. 2004) and only to a minor degree by other scleractinian CWCs such as M. oculata. The size of these thriving mounds reaches up to 45 m in height and a lateral extent of several hundred meters (Lindberg and Mienert 2005), but often can be found in clusters with up to 35 km in length (Fosså et al. 2005). On the entire Norwegian margin CWC mounds occur in water depths mainly between 200 and 400 m (Flögel et al. 2014), but in a relatively large range of seawater salinities of 33 to 37 g/kg (Freiwald 2002) and BWTs between 4 and 8 °C (Dullo et al. 2008; Flögel et al. 2014). A very narrow seawater density (sigma theta, σΘ) envelope of 27.5 ± 0.15 can be found at living CWC mounds along the Norwegian margin (Dullo et al. 2008; Rüggeberg et al. 2011), that triggers the accumulation of organic matter and production of sediment suspended bottom water layers (nepheloid layers, Mazzini et al. 2012), thereby fostering the flourishing states of CWCs (Flögel et al. 2014; Rüggeberg et al. 2016).
a Distribution of the framework-building scleractinian cold-water coral Desmophyllum pertusum (red dots) along the Norwegian margin (after Freiwald et al. 2021). White square is the working area in the NE Skagerrak. Also shown are surface (dotted black) and subsurface or deep currents (dotted blue). NAC North Atlantic Current, AW Atlantic Water, BW Baltic Water, MSW Mixed Surface Skagerrak Water, SSW Skagerrak Surface Water, NCW Norwegian Coastal Water. Please note for simplification, only major current important for this study are shown. b Detailed bathymetric map of the NE Skagerrak (a) and the Søster mounds east of the Søster Islands with the core location POS391-576/1. Bathymetric maps were performed during RV ALKOR cruise AL232 (Pfannkuche 2004)
One way to reconstruct whether CWC mounds flourished in the past or not is to determine their aggradation rates (ARs, in centimetres per thousand years, cm/kyr, Frank et al. 2009; Titschack et al. 2015; Wienberg et al. 2018; Raddatz et al. 2020). During the Holocene, Norwegian mean ARs exceeded 1000 cm/kyr, close to the extension rates of individual D. pertusum specimens at 2.5 cm/yr (Gass and Roberts 2010). However, there is still a lack of knowledge on past temperature, density, and salinity variability in CWC mounds and such variability may have affected CWC mound formation, especially in shallow CWC habitats. Oceanographic and physicochemical properties of past water masses can be reconstructed using geochemical signatures in scleractinian CWCs (e.g., Montagna et al. 2014; Schleinkofer et al. 2019), but also in foraminifera captured in matrix sediment baffled by the coral framework (e.g., Rüggeberg et al. 2007; Margreth et al. 2009; Raddatz et al. 2011; Schönfeld et al. 2011; Stalder et al. 2014; Fentimen et al. 2020). In particular, benthic foraminiferal shells record changes in seawater temperatures in their magnesium to calcium ratios (Mg/Ca; e.g., Elderfield et al. 2010; Poggemann et al. 2018), seawater density variations in their stable oxygen isotope composition (δ18O; Lynch-Stieglitz et al. 1999b; Rüggeberg et al. 2016), and the combination of both proxies can be used to infer relative salinity changes (e.g., Lea et al. 2000; Nürnberg 2000). We applied these foraminifera-based proxy reconstructions to a sediment core retrieved from the shallow Søster CWC mound complex in the NE Skagerrak (Fig. 1), a CWC habitat which currently experiences large ranges in seawater temperature and salinity. The isotope and elemental proxy data are combined with computed tomography (CT) acquisitions and 230Th/U age determination in order to reconstruct past BWT, seawater density and salinity variability during late Holocene CWC mound formation off southern Norway.
Cold-water coral occurrences and the hydrographic setting of the NE Skagerrak
CWC occurrences in the eastern Skagerrak were first reported by Wahrberg and Eliason (1926). In particular, the Oslo fjord system with elevated sills and offshore islands provides dynamic environments, but also shelter from extreme and strong currents, enabling CWC mound formation (Freiwald et al. 2004) (Fig. 1). In contrast to the open Atlantic, the Skagerrak CWC mounds occur in relatively shallow water depth between 80 and 120 m (Figs. 1, 2). The most prominent CWC habitat in this area is the Tisler Reef extending over ~ 2 km (N–S) at 90–120 m water depth. The Tisler Reef connects the Kosterfjord via the Hvaler Deep with the Oslofjord (Fig. 1). The Søster Islands south of the Oslo fjord occur on submerged drumlins (hills build by a glacier, Freiwald et al. 2004) at which the Søster CWC mound complex (east/inshore of the islands) developed, characterized by two build-ups at 88 and 110 m water depth elevating from around 140 m (Figs. 1, 2). Both summits are covered by dead corals, while the amount of living CWCs and associated fauna increase continuously further downslope (Fig. 2). Overall, the environmental and oceanographic conditions at both, the Tisler and Søster Reefs, are similar to the open Skagerrak and characterized by a strong baroclinic stratification and a relatively large temperature range (Wisshak and Rüggeberg 2006; Guihen et al. 2012). Temperature changes of ~ 7 °C over the course of 4 yr and short-term temperature shocks of 4 °C within 24 h have been recorded at the Tisler Reef, possibly exerting a strong control on coral physiology (Guihen et al. 2012).
Images of Desmophyllum pertusum in the study area Søster mound, taken by the manned submersible JAGO. Mound top a dead D. pertusum framework at 88 m water depths, b few occurrences of living D. pertusum, but mostly dead coral framework plus white fan worm Filograna implexa. Upper slope c fishing line overgrown by tube worms, soft corals and bryozoans at 97 m water depth, d dead D. pertusum framework and coral rubble at 99 m water depths. Lower slope e flourishing D. pertusum colony extending tentacles at 107 m water depth, f patchy occurrences of live D. pertusum at 113 m water depth
The hydrography of the NE Skagerrak is mainly characterized by inflowing North Atlantic water from the North Sea overlain by brackish water from the Kattegat forming a strong haline stratification (Gustafsson 1999). The in- and outflow of North Atlantic water is intensified by a positive North Atlantic Oscillation (NAO), due to the stronger than average westerly winds, which in turn lead to higher precipitation and milder temperatures (Hurrell 1995; Hurrell et al. 2003; Winther and Johannessen 2006). Continental runoff enters the NE Skagerrak via many fjords, additionally lowering surface salinities (Winther and Johannsson 2006). Tidal currents are rather weak and residual flows are the main current strength driver along the channel axis from NE and SW dominantly controlled by either density or wind-driven sea-level height differences (Guihen et al. 2012; Lavaleye et al. 2009; Wagner et al. 2011; White et al. 2012; De Clippele et al. 2018). As a result of this the NE Skagerrak is characterized by three different water masses including the low salinity (20–32 g/kg) Skagerrak Surface Water (SSW) occupying the top ~ 30 m of the water column, the underlying Mixed Skagerrak Water (MSW) with salinities between 32 and 35 g/kg and below the saline Atlantic Water (AW > 35 g/kg, Andersson; 1996; Rohde 1996; Danielssen et al. 1997) (see Figs. 3, 4). In the transition zone of the MSW and the inflowing AW the Skagerrak CWC mounds are situated (Figs. 3, 4), but are separated from the surface by a low-salinity layer that forms an oceanographic barrier. Seasonal variations clearly affect the temperature and salinity of the upper water masses SSW and MSW, while the AW mass is more stable (Fig. 4).
Conductivity–temperature–depth (CTD) profile taken during research cruise POS391 adjacent to sediment core 576/1. Also plotted is the corresponding seawater density profile. Shading indicate the depth ranges of the different ambient water masses according to Danielssen et al. (1997). Skagerrak Surface Water (SSW), Mixed Skagerrak Water (MSW) and the Atlantic Water. For details see text
Temperature–salinity–density (TSD) plot based on annual, summer (July–September) and winter (January–March) World Ocean Atlas 2018 (0.25°) data (Boyer et al. 2018) highlighting the seasonal variability especially for the Skagerrak Surface Water (SSW) and the Mixed Skagerrak Water (MSW), while Atlantic Water (AW) stays relatively stable. Dashed box indicates the depth of CWC occurrence for the Oslofjord and Tisler Reef
Materials and methods
During RV POSEIDON cruise POS391 in 2009, the 212-cm long sediment core 576-1 was retrieved from the fossil Søster CWC mound complex in the NE Skagerrak east of the Søster Islands (Fig. 1, Raddatz et al. 2016) from 101 m water depth (59° 11′ N, 010° 98′ E). The coring location was chosen based on visual inspection by previous dives with the manned submersible JAGO (GEOMAR) as close as possible to the observed living CWC colonies (Fig. 2). In addition, conductivity–temperature–depth (CTD) profiles have been collected using a CTD Rosette equipped with a Seabird SBE 9plus device to measure physical properties (Fig. 3).
Computed tomography
X-ray CT acquisitions of whole round core sections were obtained with a multi-detector Siemens Somatom Sensation 64 (Siemens, Medical Solution AG, Erlangen, Germany) installed at the University Hospital Gasthuisberg, KU Leuven (Belgium). Core sections were scanned using an X-ray source with currents of 120 kV and 135 mA. Pixel resolution of the scanned slices is 0.6 mm and slice thickness 0.9 mm with reconstruction intervals of 0.6 mm (0.628906 mm pixel size). Reconstructed slices have been imported in the software Avizo (Thermo Fisher Scientific) for advanced segmentation and visualization. For both core sections, the same volume of interest has been extracted and individual slices have been filtered using a non-local means filter prior to segmentation to remove noise in the matrix sediments. Subsequently, corals were segmented from the matrix sediments using respective single thresholding followed by watershed segmentation. Prior to quantification and visualization, labeled objects smaller than 5 voxels have been removed. For each slice, the total volume % of coral fragments has been quantified (Supplementary Material, Table S2)
230Thorium/Uranium age determination
Supplementary to the four already dated coral fragments of Raddatz et al. (2016), 15 additional 230Th/U age determinations were carried out in order to improve the chronology of core 576-1 (Table S1). Samples were selected according to the depositional intervals of the coral abundances based on the CT images. Prior to the analyses, all coral samples were cleaned mechanically to remove any contaminants from the skeleton surface (e.g., epibionts, borings, ferro-manganese crusts and coatings) and were then chemically cleaned including an oxidative step (50/50 mixture of 30% H2O2 and 1 M NaOH) as well as brief leach in 1% HClO4 (50/50 mixture with 30% H2O2) according to the protocol of Cheng et al. (2000). The measurements were carried out at GEOMAR Helmholtz Centre for Ocean Research Kiel on a multi-collector inductively coupled plasma mass spectrometer (Thermo Fisher, Neptune plus) by adapting the multi-static MIC-ICP-MS approach after Fietzke et al. (2005). Age determinations (Fig. 5; Table 1S) are based on half-lives after Cheng et al. (2013) and values for reference material HU-1 given therein (for details refer to notes presented in Table S1). The 232Th concentrations of all analyzed corals were always < 6 ppb, which is indicative of minor residual contamination with Th either non-carbonate phases (detritus and coating) or seawater. The initial 234U/238U activity ratio of all analyzed corals (Table S1) plot, when transferred into δ234U notation (i.e., ‰ deviation from secular equilibrium), within uncertainty in a narrow band of ± 10‰ compared to the value of modern seawater (145.0 ± 1.5‰, Chutcharavan et al. 2018). This finding suggests a closed system behavior for the exchange of U between the skeletons and seawater or the embedding sediment matrix. In order to verify this assumption and to estimate the reliability of the Th/U system in this setting, corals with different external preservation state from the same core depth were analyzed. This analysis resulted within uncertainty in identical U isotope signatures and ages. Hence, all ages presented in this study (Table S1) are considered reliable, especially with respect to potential variations in seawater isotope signature due to the location at the interface between Atlantic and less saline Baltic water masses.
Downcore reconstruction on sediment core POS391-576/1. Computed tomography coral clast reconstructions, percentages of coral content, 230Th/U age determinations including ages from Raddatz et al. (2016, orange) and resulting ARs, foraminiferal Mg/Ca based bottom-water-temperature (BWT), salinity reconstruction and δ18O of seawater (dashed), as well seawater density reconstruction based on the method of Rüggeberg et al. (2016; solid line) and using the 4‰ fresher end member equation of Lynch-Stieglitz et al. (1999b, dashed line). The gray boxes indicate the intervals of identified coral clusters I–III. The green bars indicate the range of modern variability (e.g., Guihen et al. 2012). The plotted 2 SD uncertainties correspond to ± 1.1 °C for the Mg/Ca-based temperature reconstruction, ± 0.25 g/kg3 seawater density and ± 0.5 g/kg
Aggradation rates
Coral ages from distinct clusters can also be used to calculate coral mound ARs. These clusters are stratigraphically closely related and represent periods of active mound formation. Previous studies (Frank et al. 2009; Douarin et al. 2014; Titschack et al. 2015; Wienberg et al. 2018; Raddatz et al. 2020) have calculated ARs from chronologically occurring corals within clusters thereby representing minimum and maximum ARs. Here, ARs were calculated from the maximum and minimum core depths and the corresponding ages of those cluster based on the CT images, thus representing average ARs (Wienberg et al. 2018; Raddatz et al. 2020). A prominent issue in CWC mound sediments is the possible age offset between matrix sediment and coral ages (Eisele et al. 2014; Bahr et al. 2020). However, for Norwegian CWC mounds it has been shown, by the comparison of matrix (foraminifera) and coral ages, that these are coeval to active periods of mound growth with efficient baffling of sediment by the coral framework (López Correa et al. 2012). Therefore, we assume that (matrix) sediments represent similar time periods as the coral framework.
Geochemical proxy analyses on benthic foraminifera
Approximately 10–15 visually well-preserved foraminiferal specimens of Lobatula lobatula (Plate 1) were selected from the dry sediment for analyses of stable isotope and element to calcium ratios. In order to reduce ontogenetic effects on δ18OCalcite and Mg/Ca, we only used specimens > 315 μm (e.g., Elderfield et al. 2002). Prior to geochemical analyses, the tests were cracked carefully between glass plates and then split into two third for Mg/Ca measurements and one third for stable isotope analyses.
Elemental ratios of benthic foraminifera and bottom water temperature reconstructions
Prior to elemental analysis, the foraminiferal tests were cleaned following established protocols including oxidative and reductive cleaning steps (Barker et al. 2003; Martin and Lea 2002). The analyses were performed on an axial viewing Varian 720 inductively-coupled-plasma-optical-emission-spectrometer (ICP-OES) at GEOMAR. The sample solution was diluted with yttrium water (concentration 112.5 μmol/l) prior to measurement in order to detect possible matrix effects during the analyses. The element/Ca measurements were drift-corrected and standardized using an internal consistency standard (ECRM 752-1, 3.761 mmol/mol Mg/Ca, Greaves et al. 2008). The external reproducibility for Mg/Ca of the ECRM standard is ± 0.1 mmol/mol (2 SD). To monitor the cleaning success of the foraminifera, Al/Ca, and Fe/Ca ratios were measured. Accordingly, we observe in the present dataset relatively high Al/Ca and Fe/Ca ratios of ~ 0.3 mmol/mol and ~ 4.0 mmol/mol, respectively (Supplementary Material, Table S3). Relatively high Fe/Ca ratios were also detected in the near-by Little Belt (IODP Exp. 347, Site M0059), and were related to the formation of (Fe) sulfides (Kotthoff et al. 2017). However, an adjusted cleaning protocol by reversing the reduction and oxidation steps of the method by Martin and Lea (2002) to remove contamination by Fe-sulfides did not result in a significant difference in Fe/Ca ratios (Kotthoff et al. 2017). In this study the increased Fe/Ca ratios appear to be the result of reoccurring pyrite on the foraminiferal shell causing the observed discrepancies from common foraminiferal Fe/Ca ratios (Barker et al. 2003; Plate 1). As pyrite does not contain considerable amounts of Mg, the Mg/Ca paleo-thermometer is not biased (see also Nürnberg et al. 2015). Furthermore, all monitored ratios such as Fe/Ca and Al/Ca do not exhibit a significant relationship to the Mg/Ca ratios (R2 < 0.4) highlighting that the Mg/Ca ratios are not influenced (Fig. S1). The Mg/Ca-BWT calibration by Quillmann et al. (2012) can be described by a linear and an exponential relationship. An ongoing discussion exists if Mg/Ca-BWT calibrations in benthic foraminifera are linear or exponential (Lear et al. 2002; Marchitto et al. 2007; Elderfield et al. 2010; Hasenfratz et al. 2017), even though exponential regressions fully explain the expected relationships by thermodynamics (Rosenthal et al. 1997). This study used the species-specific exponential calibration by Quillmann et al. (2012) for the epibenthic foraminifer L. lobatula (exponential fit): Mg/Ca (mmol/mol) = 1.24 ± 0.04 e0.069 ± 0.005 * T. The reproducibility (2 SD) of our Mg/Ca measurements the results in uncertainties of the reconstructed BWT of ± 1.1 °C.
Stable oxygen isotope measurements
The δ18O measurements on L. lobatula (δ18OC) were conducted on a Thermo Scientific MAT 253 mass spectrometer equipped with a CARBO Kiel IV device at GEOMAR. Isotope ratios were calibrated against the NBS 19 (National Bureau of Standards) and the in-house “Standard-Bremen” (Solnhofen limestone). Values are reported relative to the Pee Dee Belemnite (PDB) standard (Supplementary Material, Fig. S1; Table S3). The external reproducibility of the foraminiferal samples based on the in-house standard is ± 0.06‰ for δ18O (2 SD). The stable carbon isotope (δ13C) results will be presented elsewhere.
Seawater density and salinity reconstructions
Regional variations of bottom water salinity were approximated from the reconstructed stable oxygen isotope ratios of the seawater (δ18OSW) (e.g., Nürnberg et al. 2021), that were calculated by combining the Mg/Ca-based temperatures and the corresponding δ18OC values from the same foraminiferal sample. In particular, the temperature effect of the measured foraminiferal δ18OC was removed by applying the temperature versus δ18OC equation for cosmopolitan epibenthic foraminifera of Marchitto et al. (2014) resolved towards δ18OSW. We refrain from calculating ice-volume free δ18OSW as the change of global ice volume over the last 4 ka appears to be negligible (Lambeck et al. 2014). The modern δ18OSW–salinity relationship (δ18OSW = 0.26 * S − 8.65; R2 = 0.87; see Supplementary Material Table S5) was used to calculate absolute salinity changes from reconstructed δ18OSW values and compared to salinity determinations based on combined Mg/Ca-temperatures and δ18OC-density reconstructions at depth of the Søster CWC mound (Tomczak 2000; based on algorithms of Fofonoff and Millard 1983).
Seawater density estimates were determined according to the established approach of Lynch-Stieglitz et al. (1999a, b) that has also been applied to CWC mounds (Rüggeberg et al. 2016; Raddatz and Rüggeberg 2021). This reconstruction is based on the fact that a change in both, seawater density and δ18OC is controlled by salinity and temperature (Lynch-Stieglitz et al. 1999a) and that this relation is constant throughout geological time. In this study, past seawater densities were estimated following the approach of Rüggeberg et al. (2016) on the basis of a regional calibration (σΘ = 25.64(± 0.26) + 1.43(± 0.03) * δ18OC + 0.21(± 0.02) * δ18OC2), which relies on recent δ18Osw, temperature, salinity, pressure and density data from Dullo et al. (2008), Harwood et al. (2008), Cefas CTD data of two surveys, “Cend 10/04” and “Cend 13/05” (BODC inventory) and this study (see Supplementary Material Tables S4, S5). This data set is compared to the application of Lynch-Stieglitz et al. (1999a) with 4‰ lower fresh end-member δ18OC–density relation of the global ocean (Fig. 5, σΘ = 25.7 + 1.0 * δ18OC + 0.12 * δ18OC2). The uncertainty of the combined δ18OC and Mg/Ca δ18OSW-reconstructions is ± 0.35‰ (see also Bahr et al. 2013; Nürnberg et al. 2015), whereas the uncertainties on the absolute salinity reconstruction are ± 0.5 g/kg following Fofonoff and Millard (1983) and the approach of Rüggeberg et al. (2016).
Results
CTD
The upper 25 m of the water column are characterized by a low-salinity layer with temperatures of ~ 15.3 °C and salinities as low as 27.2 g/kg (Fig. 3) which can be associated with the SSW. The seawater density also shows the low salinity layer down to a water depth of ~ 25 m, with values as low as 20 kg/m3. Below 25 m water depth salinity increases rapidly and reaches values of 31 g/kg. At similar depth the seawater temperature is characterized by an inversion, with temperatures increasing by up to 1 °C to maximum values of 16.3 °C. Below 40 m water depth salinity increases while temperature decreases gradually, indicating the presence of the MSW. Upper AW can be identified at ~ 100 m water depth with salinities of 35 g/kg accompanied with the presence of CWCs. The corresponding seawater density gradually increases with water depth reaching values of up 27.4 kg/m3. At the water depth of the sediment core 576-1 at 101 m, the temperature is 8.3 °C, the salinity 35 g/kg and the seawater density 27.4 kg/m3 (Fig. 3).
Coral content, age constraints and aggradation rates
The CT based coral content reconstruction in sediment core 576-1 exhibits a large range. On average the coral content is 7%, with maximal values of 30% and minimum values of 0% (Fig. 5). The ages obtained range from 1.64 ± 0.04 ka at 12 cm to 3.40 ± 0.1 ka at 212 cm (Raddatz et al. 2016), whereas the youngest sample of 1.44 ± 0.02 ka can be found at 24 cm and the oldest samples of 4.13 ± 0.04 ka at 211 cm. Overall the coral ages exhibit three age clusters. Cluster I ranges from 12 to 24 cm with ages between 1.44 ± 0.02 and 1.64 ± 0.04 ka. Cluster II reveals ages from 2.37 ± 0.03 to 2.89 ± 0.02 ka between 89 and 193 cm, whereas Cluster III exhibits a prominent sediment disturbance or age reversal of ~ 1 kyr (3.18 ± 0.03 to 4.13 ± 0.04 ka), between 207 and 212 cm (Fig. 5). Between 17 and 193 cm core depth, the mound sequence reveals a clear continuous aggradation with on average 107 cm (including the intervals with low coral content). ARs of the three identified clusters are I = 67 cm/ka, II = 223 cm/ka and III = 24 cm/ka, resulting in an average AR of 104 cm/ka, similar to the overall sedimentation rate of 107 cm/ka (Fig. 5).
Mg/Ca and δ18O ratios of benthic foraminifera
Mg/Ca ratios of the benthic foraminifera L. lobatula reveal large variations of 0.92 mmol/mol between 3.26 and 2.35 mmol/mol. This relatively large variability of Mg/Ca ratios is prominent throughout the entire core and varies around an average value of ~ 2.7 mmol/mol. The corresponding δ18OC ratios show a similar pattern with variations of 1.9‰ between 1.6 and − 0.3‰ resulting in an average of 0.4‰. Overall, the Mg/Ca and δ 18OC ratios have no clear relationship suggesting that δ18OC at ~ 100 m water depth at Søster mound is largely controlled by the δ18OSW (salinity, respectively) and not by BWT changes (Fig. 6; Fig. S2).
Linear relationship between δ18OC-based seawater density and δ18OC–Mg/Ca-based δ18OSW reconstructions. Also shown is the (non) relationship between Mg/Ca-based BWT and δ18OC-based seawater density (small plot, blue; R2 = 0.73) reconstructions. The 2 SD uncertainty on the δ18OSW is ± 0.35‰, ± 1.1 °C for the Mg/Ca-based BWT temperature and ± 0.25 g/kg3 for the seawater density reconstructions
Bottom-water-temperature, seawater δ18O, salinity and seawater density reconstructions
The Mg/Ca–BWT relation of Quillmann et al. (2012) has been calibrated for Mg/Ca ratios of L. lobatulus between 1.0 and 2.4 mmol/mol (0 and 10 °C, respectively). Most of the Late Holocene Mg/Ca ratios of this study (Fig. S1) are at or outside the warmer end of the calibration line of Quillmann et al. (2012) leading to unrealistic high bottom water temperatures when using the linear calibration curve. However, their exponential fit results in reconstructed BWT with values between 7.1 and 12.5 °C and an average of 10.1 °C (Fig. 5), which is in overall agreement with instrumental data of the near-by Tisler Reef (Guihen et al. 2012).
The BWT record does not show a clear trend throughout the entire sediment core but a tendency towards higher values at the core top (Fig. 5). The δ18OSW reconstruction reveals variations of 2.18‰ from − 1.55 to 0.63‰, with an average of − 0.6‰. Similar to the BWT reconstruction the δ18OSW record does not exhibit clear trends but is rather characterized by the relatively large variability (Fig. 5). The seawater density reconstruction based on the foraminiferal δ18OC also exhibits a relatively large variability of 2.3 kg/m3 between 25.1 and 27.4 (Fig. 5). Reconstructed salinity values differ according to the method: those reconstructed on the basis of the present-day regional S–δ18OSW relation (δ18OSW = 0.2547 * S − 8.6493; R2 = 0.87; see Supplementary Data 2) vary between 27.87 and 36.45 g/kg (Δ = 8.57 g/kg) and have a large uncertainty of ± 3.2 g/kg, while the ones reconstructed on the basis of Mg/Ca, seawater density and water depth (pressure) following Fofonoff and Millard (1983) show a smaller variability of 3.28 g/kg ranging between 32.25 and 35.63 g/kg with an accuracy of ± 0.5 g/kg (Fig. 5, Supplementary Material). Interestingly, similar variations have been also measured today in the nearby Tisler-Reef (e.g., BWT = 6.44–12.39 °C, salinity = 33.57–35.10 g/kg; seawater density = 25.46–27.41 kg/m3; Guihen and White 2012), highlighting the reliability of the foraminiferal-based reconstructions.
Discussion
Søster mound development during the Late Holocene
Accumulation rates identify periods of active coral mound formation (Titschack et al. 2015; Wienberg et al. 2018; Raddatz et al. 2020), whereas appraisals of the percent composition of corals gives insights into the importance of corals as mound builders (Foubert and Henriet 2009; Van der Land et al. 2011; Douarin et al. 2013; Titschack et al. 2015; Raddatz et al. 2020). In comparison to other CWC mounds the average ARs of Søster mound (Fig. 5) are higher (on average 104 cm/kyr) than those determined for the Rost Reef (off Norway, ~ 52 cm/kyr, Titschack et al. 2015), the Rockall Bank (NE Atlantic, 37 cm/kyr, Frank et al. 2009), and offshore Brazil (30 cm/kyr, S. variabailis, Raddatz et al. 2020), but rather similar to those observed in the Porcupine Seabight (83 cm/kyr, Frank et al. 2009) and Urania Bank/Mediterranean Sea (111 cm/kyr, Titschack et al. 2016). However, the AR found here are significantly lower than those determined in glacial CWC mounds off Mauretania and Holocene CWC mounds off northern Norway (Traenadjupet, Stjernsund) with ARs of 444 cm/kyr (Wienberg et al. 2018; Titschack et al. 2015). High ARs can be attributed to favorable environmental conditions and may represent the growth rates of individual scleractinian CWCs (Titschack et al. 2015; Raddatz et al. 2020). In particular, off Norway and Mauretania CWC mound ARs are on a similar order of magnitude as growth rate estimates of CWCs and therefore, appear to represent the upper limit of CWC mound aggradation, assuming that CWC mound growth never outpaces CWC growth (Titschack et al. 2015). A lower AR of the Søster CWC mound relative to other Holocene Norwegian CWC mounds implies that environmental conditions were not optimal for rapid mound aggregation. Differences in ARs of Clusters I, II and II sequence may thus be attributed to changes in environmental conditions. Cluster II is characterized by the highest ARs, but shows variations in coral quantities of similar magnitude as for the entire mound sequence (Fig. 5). This suggest that even within one period of enhanced formation environmental conditions were not optimal.
In general, CWC habitats are characterized by a strong current regime (Davies et al. 2009a, b; Rüggeberg et al. 2005; Mienis et al. 2007) that maintain large amounts of particles in suspension. Coral mound constructors benefit from an increased sediment supply (Wheeler et al. 2008), whereas reduced current intensities accompanied with high sedimentation may bury the corals (Dorschel et al. 2005; Rüggeberg et al. 2007; Mienis et al. 2007). Therefore, periods of enhanced sediment accumulation with only limited amounts of corals may be related to protracted periods of reduced current strength that favored the settlement of fine-grained material and/or periods of enhanced sedimentation (runoff/erosion).
Changes in sedimentation pattern may also partly be associated with changes in water depth and hence with the postglacial isostatic uplift of the Fennoscandian shield. The shoreline tilt gradient indicates that most of this uplift occurred prior to 4 ka bp and uplift occurred linearly in the NE Skagerrak after 4 ka bp with a rate of ~ 5 mm/yr (Lindberg et al. 2007; Rosentau et al. 2012). Assuming this uplift rate to be constant for the last 3.5 kyr the Søster mound would have initiated in ~ 17.5 m deeper water depth and thus would have been bathed directly in the strong current flow of the North Atlantic water (Figs. 3, 4) suggesting a strong water mass control on CWC mound formation.
Environmental control of Søster mound
Temperature control
The distribution of CWCs and their reefs/mounds has been shown by predictive habitat modelling to be controlled by seawater temperature and salinity (Davies et al. 2008; Davies and Guinotte 2011). Modern CWC habitats off Norway in the aphotic Zone are accompanied by a strong hydrodynamic regime and experience relatively large variations in seawater temperature (Godø et al. 2012; Van Engeland et al. 2019; Juva et al 2020; Rüggeberg et al. 2011). For example, at the Hola Reef seasonal temperature variations of 4.5 °C (5.5–9.0 °C) were recorded and even 4 °C at Tisler Reef within 24 h (Guihen et al. 2012). How these temperature variations impact coral physiology and reef growth is still poorly constrained. Short-term experiments only found increased mortality rates at temperatures > 14 °C (Brooke et al. 2013; Lunden et al. 2014), whereas long-term experiments did not observe a change in respiration rate at similar temperatures (6–12 °C) as observed in a study over a period of three months (Naumann et al. 2014). A recent on-board experiment with D. pertusum did not find an increase in mortality with increasing temperatures from 5 to 15 °C (Dorey et al. 2020). Past BWT variability at Søster mound of < 6 °C are similar in all clusters (Fig. 5). As such a variability is within the known range of D. pertusum occurrences, we suggest that temperature is not a primary controlling parameter for CWC mound formation. Furthermore, our BWT reconstruction supports the newly performed temperature experiments by Dorey et al. (2020) implying that CWCs off Norway are acclimatized and/or have the ability to acclimatize to large temperature variations.
Seawater density and salinity changes
Throughout the entire mound sequence, salinity inferred from δ18OSW exhibit large variations, suggesting that salinity variability exerts a strong control on the water mass stratification over the CWC habitats. Seawater salinity has been shown to be one of the main parameters for the prediction of suitable CWC habitats (Davies and Guinotte 2011), where different species of scleractinian CWCs prefer a limited range of seawater salinity from 34 to 37 g/kg. However, in situ observations at different flourishing D. pertusum reefs exhibit relatively large ranges from 31.7 to 38.8 g/kg (Freiwald et al. 2004) implying that salinity is not a primary control on the growth of CWC reefs (Dullo et al. 2008). Estimating the overall relative seawater salinity variability at Søster mound by using the calculated 2.5% range in δ18OSW results in ~ 5 g/kg (Fig. 6, Schmidt et al. 1999), which is smaller than the tolerated range of modern CWCs and CWC reefs (Dullo et al. 2008; Freiwald et al. 2004).
Nevertheless, our dataset demonstrates that salinity exerts a strong control on the density of seawater in the region (Fig. 6). In the NE Skagerrak bottom-waters are dominated either by a warm saline North Atlantic source or a cold and fresh Baltic source, even though short-term, rapid events with warm and less saline waters have been documented at the close-by Tisler Reef (Guihen et al. 2012). Between 4 ka and 1.5 ka these different water mass sources likely resulted in the observed variations in seawater density of up to 2.2 kg/m3 at Søster mound (Figs. 5, 6). Today the inflow of dense AW is accompanied by an increase in current strength and a supply of fresh oxygen beneficial for the corals (Guihen et al. 2018). Furthermore, the North Sea and the Northeast Atlantic are the only sources of coral larvae migrating into the NE Skagerrak (Dahl 2013; Fox et al. 2016). These regions are atmospherically and oceanographically strongly connected via the NAO (Winther and Johannessen 2006; Fox et al. 2019). Therefore, periodically (NAO-driven) inflow of larvae from the North Atlantic CWC provinces may have supported continuous CWC growth throughout the Holocene (Mikkelsen et al. 1982; Raddatz et al. 2016). In contrast, periods of lower seawater density are accompanied by a decrease current strength resulting in a reduced replenishment of oxygen being strongly unfavorable for CWCs. During periods of reduced current flow only little water mass movement across the coral reef results into an increased descent of zooplankton mass and organic matter (Diesing et al. 2021; Guihen et al. 2018). Scleractinian CWC have been shown to consume zooplankton (e.g., Artemia salina, Naumann et al. 2011), which may suggest that the increased zooplankton load during times of reduced current strength may counteract (at least partly) unfavorable conditions.
However, the supply of organic matter and food may instead be associated with periods with a higher seawater density and the build-up of a water mass boundary. Generally, at flourishing CWC habitats seawater density is characterized by a steep gradient within a short bathymetric range (Dullo et al. 2008; Hebbeln et al. 2014). Large CWC occurrences associated with a strong density gradient have also been reported from the Mediterranean Sea (Freiwald et al. 2009) and the Gulf of Mexico (Davies et al. 2010; Hebbeln et al. 2014), although the absolute sigma-theta values differ (higher and lower) from those initially reported by Dullo et al. (2008). Nevertheless, paleoceanographic reconstructions using oxygen isotopes of benthic foraminifera from CWC mounds on the Irish margin reveal that mound formation always occurred within a very narrow seawater density envelope of σΘ between 27.3 and 27.7 kg/m3 (Rüggeberg et al. 2016; Raddatz and Rüggeberg 2021). Such narrow envelopes in seawater density highlight that flourishing CWC reefs and fast mound build-up appear preferentially near water mass boundaries characterized by a strong density gradient, where organic matter is carried along bottom and intermediate nepheloid layers and where internal waves propagate and reflect (e.g., White et al. 2007; Dullo et al. 2008; De Mol et al. 2011; van der Kaaden et al. 2020; White and Dorschel 2010). The ARs of NE Atlantic CWC mounds are smaller (15 cm/kyr, Titschack et al. 2009) than those observed at Søster mound. However, at the shallow Søster mound environmental conditions and especially sediment supply are not comparable to those of the CWC mounds at > 800–1000 m water depth in the North Atlantic. Nevertheless, we suggest that the interplay between inflowing Atlantic and the outflowing low saline Baltic water generated short-periods of relatively stable water mass boundary conditions between the MSW and the AW at ~ 100 m water depth (Figs. 3, 4), which in turn favored the accumulation of organic matter and thus CWC growth and CWC mound formation.
Conclusion
Our results reveal that the Søster mounds have been influenced by relatively large temperature, salinity and seawater density variations between 4 ka and 1.5 ka. In this respect, bottom water temperature is not the prime controlling factor for mound formation. Instead, pronounced seawater density variations induced by high amplitude salinity variations likely caused by the interplay between the Atlantic inflow and cold-fresh Baltic seawater have been the primary control on CWCs in the region. The Atlantic Inflow is characterized by a high current flow leading to an enhanced supply of oxygen and also carries larvae from other Atlantic CWC provinces into the NE Skagerrak and have formed a water mass stratification beneficial for the food supply for CWCs. Whereas a stronger influence of fresh Baltic Water may have hampered CWC growth.
Data availability
The datasets as well as additional figures for this study can be found in the Supplementary Material associated to this article and will be made available at www.pangaea.de.
Change history
11 May 2022
A Correction to this paper has been published: https://doi.org/10.1007/s00338-022-02270-7
References
Addamo AM, Vertino V, Stolarski J, García-Jiménez R, Taviani M, Machordom A (2016) Merging scleractinian genera: the overwhelming genetic similarity between solitary Desmophyllum and colonial Lophelia. BMC Evol Biol 16:108
Andersson L (1996) Trends in nutrient and oxygen concentrations in the Skagerrak-Kattegat. J Sea Res 35:63–71
Bahr A, Nürnberg D, Karas C, Grützner J (2013) Millennial-scale versus long-term dynamics in the surface and subsurface of the western North Atlantic Subtropical Gyre during marine isotope stage 5. Glob Planet Change 111:77–87
Bahr A, Doubrawa M, Titschack J, Austermann G, Koutsodendris A, Nürnberg D, Albuquerque AL, Friedrich O, Raddatz J (2020) Monsoonal forcing of cold-water coral growth off southeastern Brazil during the past 160 kyr. Biogeosciences 17:5883–5908
Barker S, Elderfield H, Greaves M (2003) A study of cleaning procedure used for foraminiferal Mg/Ca paleothermometry. Geochem Geophys Geosyst 4:8407
Boyer TP, Garcia HE, Locarnini RA, Zweng MM, Mishonov AV, Reagan JR, Weathers KA, Baranova OK, Seidov D, Smolyar IV (2018) World ocean atlas 2018, dataset. NOAA National Centers for Environmental Information. https://accession.nodc.noaa.gov/NCEI-WOA18
Brooke S, Ross SW, Bane JM, Seim HE, Young CM (2013) Temperature tolerance of the deep-sea coral Lophelia pertusa from the southeastern United States. Deep Sea Res II 92:240–248
Büscher JV, Form AU, Riebesell U (2017) Interactive effects of ocean acidification and warming on growth, fitness and survival of the cold-water coral Lophelia pertusa under different food availabilities. Front Mar Sci 4:101
Cheng H, Adkins J, Edwards RL, Boyle EA (2000) U-Th dating of deep-sea corals. Geochim Cosmochim Acta 64:2401–2416
Cheng H, Edwards RL, Shen C-C, Polyak VJ, Asmerom A, Woodhead J, Hellstrom J, Wang Y, Kong X, Spötl C, Wang X, Alexander EC Jr (2013) Improvements in 230Th dating, 230Th and 234U half-life values, and U-Th isotopic measurements by multi-collector inductively coupled plasma mass spectrometry. Earth Planet Sci Lett 371–372:82–91
Chutcharavan PM, Dutton A, Ellwood MJ (2018) Seawater 234U/238U recorded by modern and fossil corals. Geochem Cosmochim Acta 224:1–17
Dahl MP (2013) Conservation genetics of Lophelia pertusa. Dissertation Thesis, University of Gothenburg
Danielssen DS, Edler L, Fonselius S, Hernroth L, Ostrowski M, Svendsen E, Talsepp L (1997) Oceanographic variability in the Skagerrak and Northern Kattegat, May–June, 1990. ICES J Mar Sci 54:753–773
Davies AJ, Guinotte JM (2011) Global habitat suitability for framework-forming cold-water corals. PLoS ONE 6:e18483
Davies AJ, Wisshak M, Orr JC, Roberts JM (2008) Predicting suitable habitat for the cold-water coral Lophelia pertusa (Scleractinia). Deep Sea Res I 55:1048–1062
Davies AJ, Duineveld GCA, Lavaleye MSS, Bergman MJN, van Haren H, Roberts JM (2009a) Downwelling and deep-water bottom currents as food supply mechanisms to the cold-water coral Lophelia pertusa (Scleractinia) at the Mingulay Reef Complex. Limnol Oceanogr 54:620–629
Davies AJ, Duineveld GCA, Lavaleye MSS, Bergman MJN, van Haren H, Roberts JM (2009b) Downwelling and deep-water bottom currents as food supply mechanisms to the cold-water coral Lophelia pertusa (Scleractinia) at the Mingulay Reef Complex. Limnol Oceanogr 54(2):620–629
Davies AJ, Duineveld GCA, van Weering TCE, Mienis F, Quattrini AM, Seim HE, Bane JM, Ross SW (2010) Short-term environmental variability in cold-water coral habitat at Viosca Knoll, Gulf of Mexico. Deep Sea Res I 57:199–212
De Clippele LH, Huvenne VAI, Orejas C, Lundälv T, Fox A, Hennige SJ, Roberts JM (2018) The effect of local hydrodynamics on the spatial extent and morphology of cold-water coral habitats at Tisler Reef, Norway. Coral Reefs 37:253–266
De Mol B, Van Rensbergen P, Pillen S, Van Herreweghe K, Van Rooij D, McDonnell A, Huvenne V, Ivanov M, Swennen R, Henriet JP (2002) Large deep-water coral banks in the Porcupine Basin, southwest of Ireland. Mar Geol 188:193–231
De Mol L, Van Rooij D, Pirlet H, Greinert J, Frank N, Quemmerais F, Henriet J-P (2011) Cold-water coral habitats in the Penmarc’h and Guilvinec Canyons (Bay of Biscay): deep-water versus shallow-water settings. Mar Geol 282:40–52
Diesing M, Thorsnes T, Bjarnadóttir LR (2021) Organic carbon densities and accumulation rates in surface sediments of the North Sea and Skagerrak. Biogeosciences 18:2139–2160
Dodds LA, Roberts JM, Taylor AC, Marubini F (2007) Metabolic tolerance of the cold-water coral Lophelia pertusa (Scleractinia) to temperature and dissolved oxygen change. J Exp Mar Biol Ecol 349:205–214
Doney S, Fabry V, Feely R, Kleypas J (2009) Ocean acidification: the other CO2 problem. Annu Rev Mar Sci 1:169–192
Dorey N, Gjelsvik Ø, Kutti T, Büscher JV (2020) Broad thermal tolerance in the cold-water coral Lophelia pertusa from Arctic and Boreal Reefs. Front Physiol 10:1636
Dorschel B, Hebbeln D, Rüggeberg A, Dullo W-C, Freiwald A (2005) Growth and erosion of a cold-water coral covered carbonate mound in the Northeast Atlantic during the Late Pleistocene and Holocene. Earth Planet Sci Lett 233:33–44
Dorschel B, Hebbeln D, Foubert A, White M, Wheeler AJ (2007) Hydrodynamics and cold-water coral facies distribution related to recent sedimentary processes at Galway Mound west of Ireland. Mar Geol 244:184–195
Douarin M, Elliot M, Noble S, Sinclair D, Henry L-A, Long D, Moreton S, Roberts M (2013) Growth of north-east Atlantic cold-water coral reefs and mounds during the Holocene: a high resolution U-series and 14C chronology. Earth Planet Sci Lett 375:176–187
Douarin M, Sinclair D, Elliot M, Henry LA, Long D, Mitchison F, Roberts JM (2014) Changes in fossil assemblage in sediment cores from Mingulay Reef Complex (NE Atlantic): implications for coral reef build-up. Deep Sea Res II 99:286–296
Duineveld GCA, Lavaleye MSS, Bergman MJN, Stigter HCD, Mienis F (2007) Trophic structure of a cold-water coral mound community (Rockall Bank, NE Atlantic) in relation to the near-bottom particle supply and current regime. Bull Mar Sci 81:449–467
Dullo W-C, Flögel S, Rüggeberg A (2008) Cold-water coral growth in relation to the hydrography of the Celtic and Nordic European continental margin. Mar Ecol Prog Ser 371:165–176
Eisele M, Frank N, Wienberg C, Titschack J, Mienis F, Beuck L, Tisnerat-Laborde N, Hebbeln D (2014) Sedimentation patterns on a cold-water coral mound off Mauritania. Deep Sea Res II 99:307–315
Elderfield H, Vautravers M, Cooper M (2002) The relationship between shell size and Mg/Ca, Sr/Ca, δ18O, and δ13C of species of planktonic foraminifera. Geochem Geophys Geosyst 3:8
Elderfield H, Greaves M, Barker S, Hall IR, Tripati A, Ferretti P, Crowhurst S, Booth L, Daunt C (2010) A record of bottom water temperature and seawater δ18O for the Southern Ocean over the past 440 kyr based on Mg/Ca of benthic foraminiferal Uvigerina spp. Quat Sci Rev 29:1–2
Fentimen R, Feenstra E, Rüggeberg A, Vennemann T, Hajdas I, Adatte T, Van Rooij D, Foubert A (2020) Cold-water coral mound archive provides unique insights into intermediate water mass dynamics in the Alboran Sea during the Last Deglaciation. Front Mar Sci 7:354
Fietzke J, Liebetrau V, Eisenhauer A, Dullo W-C (2005) Determination of uranium isotope ratios by multi-static MIC-ICP-MS: method and implementation for precise U- and Th series isotope measurements. J Anal At Spectrom 20:395–401
Flögel S, Dullo W-C, Pfannkuche O, Kiriakoulakis K, Rüggeberg A (2014) Geochemical and physical constraints for the occurrence of living cold-water corals. Deep Sea Res II 99:19–26
Fofonoff NP, Millard RC Jr (1983) Algorithms for computation of fundamental properties of seawater. UNESCO Tech Pap Mar Sci 44:53
Form AU, Riebesell U (2012) Acclimation to ocean acidification during long-term CO2 exposure in the cold-water coral Lophelia pertusa. Glob Change Biol 18:843–853
Fosså JH, Lindberg B, Christensen O, Lundälv T, Svellingen I, Mortensen PB, Alvsvåg J (2005) Map** of Lophelia reefs in Norway: experiences and survey methods. In: Freiwald A, Roberts JM (eds) Cold-water corals and ecosystems. Springer, Berlin, pp 359–391
Foubert A, Henriet J-P (2009) Nature and significance of the recent carbonate mound record. Springer, Berlin
Fox AD, Henry LA, Corne DW, Roberts JM (2016) Sensitivity of marine protected area network connectivity to atmospheric variability. R Soc Open Sci 3(11):160494
Frank N, Ricard E, Lutringer-Paquet A, van der Land C, Colin C, Blamart D, Foubert A, Van Rooij D, Henriet J-P, de Haas H, van Weering T (2009) The Holocene occurrence of cold water corals in the NE Atlantic: implications for coral carbonate mound evolution. Mar Geol 266:129–142
Frank N, Freiwald A, López Correa M, Wienberg C, Eisele M, Hebbeln D, Van Rooij D, Henriet JP, Colin C, van Weering T, de Haas H, Buhl-Mortensen P, Roberts JM, De Mol B, Douville E, Blamart D, Hatte C (2011) Northeastern Atlantic cold-water coral reefs and climate. Geology 39:743–746
Frederiksen R, Jensen A, Westerberg H (1992) The distribution of the scleractinian coral Lophelia pertusa around the Faroe Islands and the relation to internal tidal mixing. Sarsia 77:157–171
Freiwald A (2002) Reef-forming cold-water corals. In: Wefer G, Billett D, Hebbeln D, Jørgensen BB, Schlüter M, Weering TV (eds) Ocean margin systems. Springer, Berlin, pp 365–385
Freiwald A, Fosså JH, Grehan A, Koslow T, Roberts JM (2004) Cold-water coral reefs. UNEP-WCMC, Cambridge, p 84
Freiwald A, Beuck L, Rüggeberg A, Taviani M, Hebbeln D, RV Meteor Cruise Participants (2009) The white coral community in the Central Mediterranean Sea revealed by ROV surveys. Oceanography 22:58–74
Freiwald, A., Rogers, A., Hall-Spencer, J., Guinotte, J.M,, Davies, A.J., Yesson, C, Martin, C.S., Weatherdon, L.V. (2021). Global distribution of cold-water corals (version 5.1). Fifth update to the dataset. In: Freiwald A et al (2004) by UNEP-WCMC, in collaboration with Andre Freiwald and John Guinotte. UN Environment Programme World Conservation Monitoring Centre, Cambridge
Gammon MJ, Tracey DM, Marriott PM, Cummings VJ, Davy SK (2018) The physiological response of the deep-sea coral Solenosmilia variabilis to ocean acidification. PeerJ 6:e5236
Gass SE, Roberts JM (2010) Growth and branching patterns of Lophelia pertusa (Scleractinia) from the North Sea. J Mar Biol Assoc UK 90:1–5
Glecker PJ, Durack PJ, Stouffer RJ, Johnson GC, Forest CE (2016) Industrial-era global ocean heat uptake doubles in recent decades. Nat Clim Change 6:394–398
Godø OR, Tenningen E, Ostrowski M, Kubilius R, Kutti T, Korneliussen R et al (2012) The Hermes Lander Project—the technology, the data and evaluation of concept and results. Havforskningsinstituttet, Bergen
Greaves M, Caillon N, Rebaubier H, Bartoli G, Bohaty S, Cacho I, Clarke L, Cooper M, Daunt C, Delaney M, DeMenoca Pl, Dutton A, Eggins S, Elderfield H, Garbe-Schoenberg D, Goddard E, Green D, Groeneveld J, Hastings D, Hathorne E, Kimoto K, Klinkhammer G, Labeyrie L, Lea DW, Marchitto T, Martínez-Botí MA, Mortyn PG, Ni Y, Nuernberg D, Paradis G, Pena L, Quinn T, Rosenthal Y, Russell A, Sagawa T, Sosdian S, Stott L, Tachikawa K, Tappa E, Thunell R, Wilson PA (2008) Interlaboratory comparison study of calibration standards for foraminiferal Mg/Ca thermometry. Geochem Geophys Geosyst 9:Q08010
Guihen D, White M (2012) Physical oceanography from mooring TR_Micro3 at Tisler Cold-Water Coral Reef, Norway. . https://doi.org/10.1594/PANGAEA.774544National University of Ireland, Galway
Guihen D, White M, Lundälv T (2012) Temperature shocks and ecological implications at a cold-water coral reef. Mar Biodivers Rec 5:e68
Guihen D, White M, Lundälv T (2018) Zooplankton drive diurnal changes in oxygen concentration at Tisler Cold-Water Coral Reef. Coral Reefs 37:1013–1025
Guinotte JM, Orr J, Cairns S, Freiwald A, Morgan L, George R (2006) Will human-induced changes in seawater chemistry alter the distribution of deep-sea scleractinian corals? Front Ecol Environ 4:141–146
Gustafsson B (1999) High frequency variability of the surface layers in the Skagerrak during SKAGEX. Cont Shelf Res 19(8):1021–1047
Harwood AJP, Dennis PF, Marca AD, Pilling GM, Millner RS (2008) The oxygen isotope composition of water masses within the North Sea. Estuar Coast Shelf Sci 78:353–359
Hasenfratz AP, Martínez-García A, Jaccard SL, Vance D, Wälle M, Greaves M, Haug GH (2017) Determination of the Mg/Mn ratio in foraminiferal coatings: an approach to correct Mg/Ca temperatures for Mn-rich contaminant phases. Earth Planet Sci Lett 457:335–347
Hebbeln D, Wienberg C, Wintersteller P, Freiwald A, Becker M, Beuck L, Dullo C, Eberli GP, Glogowski S, Matos L, Forster N, Reyes-Bonilla H, Taviani M (2014) Environmental forcing of the Campeche Cold-Water Coral Province, southern Gulf of Mexico. Biogeosciences 11:1799–1815
Hurrell JW (1995) Decadal trends in the North Atlantic Oscillation. Science (New York NY) 269:676–679
Hurrell JW, Kushnir Y, Ottersen G, Visbeck M (2003). An Overview of the North Atlantic Oscillation. In: Hurrell JW, Kushnir Y, Ottersen G, Visbeck M (eds) The North Atlantic Oscillation: climatic significance and environmental impact, vol 134. American Geophysical Union, pp 1–36
Juva K, Flögel S, Karstensen J, Linke P, Dullo W-C (2020) Tidal dynamics control on cold-water coral growth: a high-resolution multivariable study on eastern Atlantic cold-water coral site. Front Mar Sci 7:132
Juva K, Kutti T, Chierici M, Dullo W-C, Flögel S (2021) Cold-water coral reefs in the Langenuen Fjord, southwestern Norway—a window into future environmental change. Oceans 2:583–610
Kano A, Ferdelman TG, Williams T, Henriet J-P, Ishikawa T, Kawagoe N, Takashima C, Kakizaki Y, Abe K, Sakai S, Browning E, Li X, IODPE Scientists (2007) Age constraints on the origin and growth history of a deep-water coral mound in the northeast Atlantic drilled during Integrated Ocean Drilling Program Expedition 307. Geology 35:1051–1054
Kotthoff U, Groeneveld J, Ash JL, Fanget A-S, Krupinski NQ, Peyron O, Stepanova A, Warnock J, Van Helmond NAGM, Passey BH, Clausen OR, Bennike O, Andrén E, Granoszewski W, Andrén T, Filipsson HL, Seidenkrantz M-S, Slomp CP, Bauersachs T (2017) Reconstructing Holocene temperature and salinity variations in the western Baltic Sea region: a multi-proxy comparison from the Little Belt (IODP Expedition 347, Site M0059). Biogeosciences 14:5607–5632
Lambeck K, Rouby H, Purcell A, Sun Y, Sambridge M (2014) Sea level and global ice volumes from the Last Glacial Maximum to the Holocene. Proc Natl Acad Sci USA 111(43):15296–15303
Lavaleye M, Duineveld G, Lundälv T, White M, Guihen D, Kiriakoulakis K, Wolff G (2009) Cold-water corals on the Tisler Reef—preliminary observations on the dynamic reef environment. Oceanography 22:76–84
Lea DW, Pak DK, Spero HJ (2000) Climate impact of Late Quaternary Equatorial Pacific sea surface temperature variations. Science 289:1719–1724
Lear CH, Rosenthal Y, Slowey N (2002) Benthic foraminiferal Mg/Ca-paleothermometry: a revised core-top calibration. Geochim Cosmochim Acta 66:3375–3387
Lindberg B, Mienert J (2005) Sedimentological and geochemical environment of the Fugløy Reef off northern Norway. In: Freiwald A, Roberts JM (eds) Cold-water corals and ecosystems. Springer, Berlin, pp 633–650
Lindberg B, Berndt C, Mienert J (2007) The Fugløy Reef at 70° N; acoustic signature, geologic, geomorphologic and oceanographic setting. Int J Earth Sci 96:201–213
López Correa M, Montagna P, Joseph N, Rüggeberg A, Fietzke J, Flögel S, Dorschel B, Goldstein SL, Wheeler A, Freiwald A (2012) Preboreal onset of cold-water coral growth beyond the Arctic Circle revealed by coupled radiocarbon and U-series dating and neodymium isotopes. Quat Sci Rev 34:24–43
Lunden JJ, McNicholl CG, Sears CR, Morrison CL, Cordes EE (2014) Acute survivorship of the deep-sea coral Lophelia pertusa from the Gulf of Mexico under acidification, warming, and deoxygenation. Front Mar Sci 1:78
Lynch-Stieglitz J, Curry WB, Slowey N (1999a) Weaker gulf stream in the Florida Straits during the Last Glacial Maximum. Nature 402:644–648
Lynch-Stieglitz J, Curry WB, Slowey N (1999b) A geostrophic transport estimate for the Florida Current from the oxygen isotope composition of benthic foraminifera. Paleoceanography 14:360–373
Maier C, Hegeman J, Weinbauer MG, Gattuso P (2009) Calcification of the cold-water coral Lophelia pertusa under ambient and reduced pH. Biogeosciences 6:1671–1680
Marchitto TM, Bryan SP, Curry WB, McCorkle DC (2007) Mg/Ca temperature calibration for the benthic foraminifer Cibicidoides pachyderma. Paleoceanography 22:PA1203
Marchitto TM, Curry WB, Lynch-Stieglitz J, Bryan SP, Cobb KM, Lund DC (2014) Improved oxygen isotope temperature calibrations for cosmopolitan benthic foraminifera. Geochim Cosmochim Acta 130:1–11
Margreth S, Rüggeberg A, Spezzaferri S (2009) Benthic foraminifera as bioindicator for cold-water coral reef ecosystems along the Irish margin. Deep Sea Res I 56:2216–2234
Martin P, Lea D (2002) A simple evaluation of cleaning procedures on fossil benthic foraminiferal Mg/Ca. Geochem Geophys Geosyst 3(10):1–8
Mazzini A, Akhmetzhanov A, Monteys X, Ivanov M (2012) The Porcupine Bank Canyon coral mounds: oceanographic and topographic steering of deep-water carbonate mound development and associated phosphatic deposition. Geomar Lett 32:205–225
Mienis F, de Stigter HC, White M, Duineveld G, de Haas H, van Weering TCE (2007) Hydrodynamic controls on cold-water coral growth and carbonate-mound development at the SW and SE Rockall Trough Margin, NE Atlantic Ocean. Deep Sea Res I 54:1655–1674
Mikkelsen N, Erlenkeuser H, Killingley JS, Berger WH (1982) Norwegian corals: radiocarbon and stable isotopes in Lopehlia pertusa. Boreas 11:163–171
Montagna P, McCulloch M, Douville E, López Correa M, Trotter J, Rodolfo-Metalpa R, Dissard D, Ferrier-Pagés C, Frank N, Freiwald A, Goldstein S, Mazzoli C, Reynaud S, Rüggeberg A, Russo S, Taviani M (2014) Li/Mg systematics in scleractinian corals: calibration of the thermometer. Geochim Cosmochim Acta 132:288–310
Muñoz A, Cristobo J, Rios P, Druet M, Polonio V, Uchupi E, Acosta J, Group A (2012) Sediment drifts and cold-water coral reefs in the Patagonian upper and middle continental slope. Mar Pet Geol 36:70–82
Naumann MS, Orejas C, Wild C, Ferrier-Pagès C (2011) First evidence for zooplankton feeding sustaining key physiological processes in a scleractinian cold-water coral. J Exp Biol 214:3570–3576
Naumann MS, Orejas C, Ferrier-Pagès C (2014) Species-specific physiological response by the cold-water corals Lophelia pertusa and Madrepora oculata to variations within their natural temperature range. Deep Sea Res II 99:36–41
Nürnberg D (2000) Taking the temperature of past ocean surfaces. Science 289:1698–1699
Nürnberg D, Böschen T, Doering K, Mollier-Vogel E, Raddatz J, Schneider R (2015) Sea surface and subsurface circulation dynamics off equatorial Peru during the last ~ 17 kyr. Paleoceanography 30:984–999
Nürnberg D, Riff T, Bahr A, Karas C, Meier K, Lippold J (2021) Western boundary current in relation to Atlantic Subtropical Gyre dynamics during abrupt glacial climate fluctuations. Glob Planet Change 201:103497
Pachauri RK, Allen MR, Barros VR, Broome J, Cramer W et al (2014) In: Pachauri R Meyer L (eds) Climate Change 2014: Synthesis Report. Contribution of Working Groups I, II and III to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change. IPCC, Geneva. ISBN 978-92-9169-143-2
Pfannkuche O, Cruise Participants (2004) FS ALKOR Cruise No. 232—Cruise Report. Leibniz-Institut für Meereswissenschaften an der Universität Kiel Germany, p 35.
Poggemann D-W, Nürnberg D, Hathorne EC, Frank M, Rath W, Reißig S, Bahr A (2018) Deglacial heat uptake by the Southern Ocean and rapid northward redistribution via Antarctic intermediate water. Paleoceanogr Paleoclimatol 33:1292–1305
Quillmann U, Marchitto TM, Jennings AE, Andrews JT, Friestad BF (2012) Cooling and freshening at 8.2 ka on the NW Iceland Shelf recorded in paired δ18O and Mg/Ca measurements of the benthic foraminifer Cibicides lobatulus. Quat Res 78:528–539
Raddatz J, Rüggeberg A (2021) Constraining past environmental changes of cold-water coral mounds with geochemical proxies in corals and foraminifera. Depos Rec 7(2):200–222
Raddatz J, Rüggeberg A, Margreth S, Dullo W-C, The IODP, Expedition 307 Scientific Party (2011) Paleoenvironmental reconstruction of Challenger Mound initiation in the Porcupine Seabight, NE Atlantic. Mar Geol 282:79–90
Raddatz J, Rüggeberg A, Flögel S, Hathorne EC, Liebetrau V, Eisenhauer A, Dullo W-C (2014) The influence of seawater pH on U/Ca ratios in the scleractinian cold-water coral Lophelia pertusa. Biogeosciences 11:1863–1871
Raddatz J, Liebetrau V, Trotter J, Rüggeberg A, Flögel S, Dullo W-C, Eisenhauer A, Voigt S, McCulloch M (2016) Environmental constraints on Holocene cold-water coral reef growth off Norway: insights from a multiproxy approach. Paleoceanography 31(10):1350–1367
Raddatz J, Titschack J, Frank N, Freiwald A, Conforti A, Osborne A, Skornitzke S, Stiller W, Rüggeberg A, Voigt S, Albuquerque ALS, Vertino A, Schröder-Ritzrau A, Bahr A (2020) Solenosmilia variabilis-bearing cold-water coral mounds off Brazil. Coral Reefs 39:69–83
Roberts JM, Wheeler A, Freiwald A, Cairns S (2009) Cold-water corals: the biology and geology of deep-sea coral habitats. Cambridge University Press, Cambridge
Rohde J (1996) On the dynamics of the large-scale circulation of the Skagerrak. J Sea Res 35:9–21
Rosentau A, Harff J, Meyer M, Oja T (2012) Postglacial rebound and relative sea level changes in the Baltic Sea since the Litorina transgression. Baltica 25:113–120
Rosenthal Y, Boyle EA, Slowey N (1997) Temperature control on the incorporation of magnesium, strontium, fluorine, and cadmium into benthic foraminiferal shells from Little Bahama Bank: prospects for thermocline paleoceanography. Geochim Cosmochim Acta 61:3633–3643
Rüggeberg A, Dorschel B, Dullo W-C, Hebbeln D (2005) Sedimentary patterns in the vicinity of a carbonate mound in the Hovland Mound Province, Northern Porcupine Seabight. In: Freiwald A, Roberts JM (eds) Cold-water corals and ecosystems. Springer, Berlin, pp 87–112
Rüggeberg A, Dullo C, Dorschel B, Hebbeln D (2007) Environmental changes and growth history of Propeller Mound, Porcupine Seabight: evidence from benthic foraminiferal assemblages. Int J Earth Sci 96:57–72
Rüggeberg A, Flögel S, Dullo W-C, Hissmann K, Freiwald A (2011) Water mass characteristics and sill dynamics in a subpolar cold-water coral reef setting at Stjernsund, northern Norway. Mar Geol 282:5–12. https://doi.org/10.1016/j.margeo.2010.05.009
Rüggeberg A, Flögel S, Dullo W-C, Raddatz J, Liebetrau V (2016) Paleo-seawater density reconstruction and its implication for cold-water coral carbonate mounds in the northeast Atlantic through time. Paleoceanography 31:365–379
Schleinkofer N, Raddatz J, Freiwald A, Evans D, Beuck L, Rüggeberg A, Liebetrau V (2019) Environmental and biological controls on Na/Ca ratios in scleractinian cold-water corals. Biogeosciences 16:3565–3582
Schmidt G, Bigg G, Rohling E (1999) Global seawater oxygen-18 database. http://data.giss.nasa.gov/o18data/
Schönfeld J, Dullo W-C, Pfannkuche O, Freiwald A, Rüggeberg A, Schmidt S, Weston J (2011) Recent benthic foraminiferal assemblages from cold-water coral mounds in the Porcupine Seabight. Facies 57:187–213
Stalder C, Spezzaferri S, Rüggeberg A, Pirkenseer C, Gennari G (2014) Late Weichselian deglaciation and early Holocene development of a cold-water coral reef along the Lopphavet Shelf (Northern Norway) recorded by benthic foraminifera and ostracoda. Deep Sea Res II 99:249–169
Sweetman AK, Thurber AR, Smith CR, Levin LA, Mora C, Wei C-L, Gooday AJ, Jones DOB, Rex M, Yasuhara M, Ingels J, Ruhl HA, Frieder CA, Danovaro R, Würzberg L, Baco A, Grupe BM, Pasulka A, Meyer KS, Dunlop KM, Henry L-A, Roberts JM (2017) Major impacts of climate change on deep-sea benthic ecosystems. Elem Sci Anthr 5:4
Titschack J, Thierens M, Dorschel B, Schulbert C, Freiwald A, Kano A, Takashima C, Kawagoe N, Li X, Expedition IODP, 307 Scientific Party (2009) Carbonate budget of a cold-water coral mound (Challenger Mound, IODP Expedition 307). Mar Geol 259:36–46. https://doi.org/10.1016/j.margeo.2008.12.007
Titschack J, Baum D, De Pol-Holz R, López Correa M, Forster N, Flögel S, Hebbeln D, Freiwald A (2015) Aggradation and carbonate accumulation of Holocene Norwegian cold-water coral reefs. Sedimentology 62:1873–1898
Titschack J, Fink HG, Baum D, Wienberg C, Hebbeln D, Freiwald A (2016) Mediterranean cold-water corals—an important regional carbonate factory? Depos Rec 2(1):74–96
Tomczak M (2000) Seawater density calculator. http://www.mt-oceanography.info/Utilities/density.html
van der Kaaden A-S, van Oevelen D, Rietkerk M, Soetaert K, van de Koppel J (2020) Spatial self-organization as a new perspective on cold-water coral mound development. Front Mar Sci 7:631
van der Land C, Mienis F, de Haas H, de Stigter HC, Swennen R, Reijmer JJG, van Weering TCE (2011) Paleo-redox fronts and their formation in carbonate mound sediments from the Rockall Trough. Mar Geol 284:86–95
Van Engeland T, Godø OR, Johnsen E, Duineveld GCA, van Oevelen D (2019) Cabled ocean observatory data reveal food supply mechanisms to a cold-T water coral reef. Prog Oceanogr 172:51–64
Wagner H, Purser A, Thomsen L, Jesus CC, Lundälv T (2011) Particulate organic matter fluxes and hydrodynamics at the Tisler Cold-Water Coral Reef. J Mar Syst 85:19–29
Wahrberg R, Eliason A (1926) Ny local för levande Lophelia prolifera (PALLAS) vid svensk kust. Fauna Och Flora 1926:256–260
Wheeler A, Kozachenko M, Masson DG, Huvenne VAI (2008) Influence of benthic sediment transport on cold-water coral bank morphology and growth: the example of the Darwin Mounds, north-east Atlantic. Sedimentology 55(6):1875–1887
White M, Dorschel B (2010) The importance of the permanent thermocline to the cold water coral carbonate mound distribution in the NE Atlantic. Earth Planet Sci Lett 296:395–402
White M, Mohn C, de Stigter H, Mottram G (2005) Deep-water coral development as a function of hydrodynamics and surface productivity around the submarine banks of the Rockall Trough, NE Atlantic. In: Freiwald A, Roberts JM (eds) Cold-water corals and ecosystems. Springer, Berlin, pp 503–514
White M, Roberts JM, Van Weering TCE (2007) Do bottom-intensified diurnal tidal currents shape the alignment of carbonate mounds in the NE Atlantic? Geomar Lett 27:391–397
White M, Wolff GA, Lundälv T, Guihen D, Kiriakoulakis K, Lavaleye M, Duineveld G (2012) Cold-water coral ecosystem (Tisler Reef, Norwegian Shelf) may be a hotspot for carbon cycling. Mar Ecol Prog Ser 465:11–23
Wienberg C, Titschack J (2017) Framework-forming scleractinian cold-water corals through space and time: a Late Quaternary North Atlantic perspective. In: Rossi S (ed) Marine animal forests. Springer, Cham, pp 1–34
Wienberg C, Titschack J, Freiwald A, Frank N, Lundälv T, Taviani M, Beuck L, Schröder-Ritzrau A, Krengel T, Hebbeln D (2018) The giant Mauritanian cold-water coral mound province: oxygen control on coral mound formation. Quat Sci Rev 185:135–152. https://doi.org/10.1016/j.quascirev.2018.02.012
Wienberg C, Titschack J, Frank N, De Pol-Holz R, Fietzke J, Eisele M, Kremer A, Hebbeln D (2020) Deglacial upslope shift of NE Atlantic intermediate waters controlled slope erosion and cold-water coral mound formation (Porcupine Seabight, Irish margin). Quat Sci Rev 237:106310
Winther NG, Johannessen JA (2006) North Sea circulation: Atlantic inflow and its destination. J Geophys Res Oceans 111:C12018
Wisshak M, Rüggeberg A (2006) Colonisation and bioerosion of experimental substrates by benthic foraminiferans from euphotic to aphotic depths (Kosterfjord, SW Sweden). Facies 52:1–17
Acknowledgements
This work is dedicated to Dr. Volker Liebetrau who unexpectedly passed away during the review process of this manuscript. He was a great inspirer, a passionate geochemist and a good friend. The authors thank captains, crews and cruise participants of research vessels POSEIDON, ALKOR and POLARSTERN of cruises POS 391, POS 325, ALK 275 and ARK XXII/1a. Cefas CTD data of “Cend 10/04” and “Cend 13/05” data are being provided under Open Government Licence v3.0 with no limitations. Nadine Gehre is thanks for her support in the lab. Walter Coudyzer (Department of Radiology, UZ Leuven) is acknowledged for his support during CT-scanning.
Funding
Open Access funding enabled and organized by Projekt DEAL. Research Cruise POS391 was realized through DFG Project RI 598/4-1. JR acknowledges funding from DFG Project RA 2156 1/1. SF would like to thank SFB 754, Subproject A7. AF and AR acknowledge support from Swiss National Science Foundation Project Number SNF 200021_149247.
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JR, VL, and AR designed the research. JR and VL participated in Cruise POS 391 and AR in Cruise ALK 275. JR analyzed the CTD data. AF performed the computer tomography scans. Coral sampling and dating was carried out by JR, VL and TJG. JM picked under supervision of JR the foraminifera. Geochemical analyses were conducted by DN and JR. KH analyzed the underwater video and image footage. JR wrote the manuscript and all authors contributed to the discussion and writing of the manuscript.
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Raddatz, J., Liebetrau, V., Rüggeberg, A. et al. Living on the edge: environmental variability of a shallow late Holocene cold-water coral mound. Coral Reefs 41, 1255–1271 (2022). https://doi.org/10.1007/s00338-022-02249-4
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DOI: https://doi.org/10.1007/s00338-022-02249-4