Cold-water Coral Growth Research Articles

Resemblance of the global depth distribution of internal-tide generation and cold-water coral occurrences

Abstract. Internal tides are known to be an important source of mixing in the oceans, especially in the bottom boundary layer. The depth of internal-tide generation therefore seems important for benthic life and the formation of cold-water coral mounds, but internal-tide conversion is generally investigated in a depth-integrated sense. Using both idealized and realistic simulations on continental slopes, we found that the depth of internal-tide generation increases with increasing slope steepness and decreases with intensified shallow stratification. The depth of internal-tide generation also shows a typical latitudinal dependency related to Coriolis effects. Using a global database of cold-water corals, we found that, especially in Northern Hemisphere autumn and winter, the global depth pattern of internal-tide generation correlates (rautumn = 0.70, rwinter = 0.65, p < 0.01) with that of cold-water corals: shallowest near the poles and deepest around the Equator, with a decrease in depth around 25° S and N, and shallower north of the Equator than south. We further found that cold-water corals are situated significantly more often on topography that is steeper than the internal-tide beam (i.e. where supercritical reflection of internal tides occurs) than would be expected from a random distribution: in our study, in 66.9 % of all cases, cold-water corals occurred on a topography that is supercritical to the M2 tide whereas globally only 9.4 % of all topography is supercritical. Our findings underline internal-tide generation and the occurrence of supercritical reflection of internal tides as globally important for cold-water coral growth. The energetic dynamics associated with internal-tide generation and the supercritical reflection of internal tides likely increase the food supply towards the reefs in food-limited winter months. With climate change, stratification is expected to increase. Based on our results, this would decrease the depth of internal-tide generation, possibly creating new suitable habitat for cold-water corals shallower on continental slopes.

Cold-water coral energy reserves and calcification in contrasting fjord environments

The relationship between energy reserves of cold-water corals (CWCs) and their physiological performance remains largely unknown. In addition, it is poorly understood how the energy allocation to different metabolic processes might change with projected decreasing food supply to the deep sea in the future. This study explores the temporal and spatial variations of total energy reserves (proteins, carbohydrates and lipids) of the CWC Desmophyllum dianthus and their correlation with its calcification rate. We took advantage of distinct horizontal and vertical physico-chemical gradients in Comau Fjord (Chile) and examined the changes in energy reserves over one year in an in situ reciprocal transplantation experiment (20 m vs. 300 m and fjord head vs. mouth). Total energy reserves correlated positively with calcification rates. The fast-growing deep corals had higher and less variable energy reserves, while the slower-growing shallow corals showed pronounced seasonal changes in energy reserves. Novel deep corals (transplanted from shallow) were able to quickly increase both their calcification rates and energy reserves to similar levels as native deep corals. Our study shows the importance of energy reserves in sustaining CWC growth in spite of aragonite undersaturated conditions (deep corals) in the present, and potentially also future ocean.

Recovery and restoration potential of cold‐water corals: experience from a deep‐sea marine protected area

Cold‐water corals (CWCs) are important species that provide habitat for other taxa but are sensitive to mechanical damage from bottom trawling. CWC conservation has been implemented in the form of marine protected areas (MPAs), but recovery from impact may be particularly slow in the deep‐sea environment; consequently, the use of restoration techniques has been considered. To gain some insight into CWC recruitment and growth, in 2011 we deployed small seabed moorings in the Darwin Mounds MPA (~1,000 m water depth). This site hosts hundreds of CWC mounds, that had previously (until 2003) been impacted by deep‐water trawling. In 2019, we carried out in situ visual surveys of these moorings and the surrounding seabed environment, then recovered two of the moorings. The mooring buoys, glass floats with plastic covers, were extensively colonized by a diverse epifauna that included the CWCs Desmophyllum pertusum and D. dianthus. The presence of coral recruits indicated that environmental conditions, and larval supply, remained favorable for the settlement and growth of CWCs within the MPA. Based on our observations, we consider four possible restoration methods, together with a “do‐nothing” option, for the Darwin Mounds CWCs that have shown little, if any, natural recovery despite 16 years of protection. We conclude that seabed emplacement of high‐relief artificial substrata is likely to be the most efficient and cost‐efficient means of promoting enhanced recovery of the CWCs.

Decline in cold-water coral growth promotes molluscan diversity: A paleontological perspective from a cold-water coral mound in the western Mediterranean Sea

Framework-forming scleractinian cold-water corals (CWCs) act as ecosystem engineers, building and supporting biodiversity hotspots in the deep sea worldwide. While spatial patterns and drivers of species distributions have been evaluated on modern CWC reefs, little is known about how reef diversity is affected by habitat variability over geologic time – the scale at which CWC reefs initiate, thrive, and decline. Using three CWC reef sediment cores as species diversity archives, we investigated temporal trends of molluscan diversity over the last ~13 kyr from a CWC mound in the Alboran Sea (western Mediterranean Sea) to evaluate (a) how spatial patterns of CWC-associated diversity are recorded in reef sediments, (b) the potential of CWC reefs as biodiversity hotspots when coral growth is flourishing and when it is not, and (c) which palaeoceanographic conditions or habitat characteristics may be driving biodiversity. Our results reveal that at the ecosystem scale ecological differences between CWC habitats are more pronounced than ecological signatures of molluscan assemblages associated with intervals of CWC framework (flourishing growth) or non-framework (negligible CWC growth). However, within habitats, significant differences emerge between these assemblages with lower molluscan diversity associated with flourishing CWC growth. Significant negative correlations between molluscan diversity and palaeoceanographic conditions conducive for CWC growth (high food availability, strong hydrodynamics, optimal bottom-water temperatures and salinities, and high aggradation rates indicative of flourishing CWC growth also imply that CWC growth and relevant environmental conditions contribute to reduced molluscan diversity. Additionally, high coral volume, used here as a proxy for habitat structural complexity, is positively correlated with molluscan diversity just as high habitat complexity is in living CWC reefs. Altogether, these patterns detected over geologic time resemble those observed spatially across living CWC reefs today – where competition with resources, particularly food, prevents high reef biodiversity in the immediate vicinity of dense living CWC colonies. Overall, our study demonstrates that (1) ecological paradigms of living CWCs are preserved in their sedimentary record, (2) flourishing CWC growth and conditions promoting CWC growth drive habitat-scale diversity patterns, and (3) a geological approach can be applied to study long-term diversity dynamics in CWC ecosystems.

Cold-water coral framework architecture is selectively shaped by bottom current flow

The three-dimensional (3D) structure of habitat-forming corals has profound impacts on reef ecosystem processes. Elucidating coral structural responses to the environment is therefore crucial to understand changes in these ecosystems. However, little is known of how environmental factors shape coral structure in deep and dark waters, where cold-water coral (CWC) reefs thrive. Here, we attempt to infer the influence of current flow on CWC framework architecture, using 3D scanning to quantify colony shape traits (volume compactness and surface complexity) in the reef-building CWC Desmophyllum pertusum from adjacent fjord and offshore habitats with contrasting flow regimes. We find substantial architectural variability both between and within habitats. We show that corals are generally more compact in the fjord habitat, reflecting the prevailing higher current speeds, although differences in volume compactness between fjord and offshore corals are more subtle when comparing the fjord with the more exposed side of the offshore setting, probably due to locally intensified currents. Conversely, we observe no clear disparity in coral surface complexity between habitats (despite its positive correlation with volume compactness), suggesting it is not affected by current speed. Unlike volume compactness, surface complexity is similarly variable within a single colony as it is between colonies within the same habitat or between habitats and is therefore perhaps more dependent than volume compactness on microenvironmental conditions. These findings suggest a highly plastic, trait-specific and functionally relevant structural response of CWCs to current flow and underscore the importance of multiple concurrent sources of hydrodynamic forcing on CWC growth.

Local-scale feedbacks influencing cold-water coral growth and subsequent reef formation

Despite cold-water coral (CWC) reefs being considered biodiversity hotspots, very little is known about the main processes driving their morphological development. Indeed, there is a considerable knowledge gap in quantitative experimental studies that help understand the interaction between reef morphology, near-bed hydrodynamics, coral growth, and (food) particle transport processes. In the present study, we performed a 2-month long flume experiment in which living coral nubbins were placed on a reef patch to determine the effect of a unidirectional flow on the growth and physiological condition of Lophelia pertusa. Measurements revealed how the presence of coral framework increased current speed and turbulence above the frontal part of the reef patch, while conditions immediately behind it were characterised by an almost stagnant flow and reduced turbulence. Owing to the higher current speeds that likely promoted a higher food encounter rate and intake of ions involved in the calcification process, the coral nubbins located on the upstream part of the reef presented a significantly enhanced average growth and a lower expression of stress-related enzymes than the downstream ones. Yet, further experiments would be needed to fully quantify how the variations in water hydrodynamics modify particle encounter and ion intake rates by coral nubbins located in different parts of a reef, and how such discrepancies may ultimately affect coral growth. Nonetheless, the results acquired here denote that a reef influenced by a unidirectional water flow would grow into the current: a pattern of reef development that coincides with that of actual coral reefs located in similar water flow settings. Ultimately, the results of this study suggest that at the local scale coral reef morphology has a direct effect on coral growth thus, indicating that the spatial patterns of living CWC colonies in reef patches are the result of spatial self-organisation.

Living on the edge: environmental variability of a shallow late Holocene cold-water coral mound

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.

Glacio-eustatic variations and sapropel events as main controls on the Middle Pleistocene-Holocene evolution of the Cabliers Coral Mound Province (W Mediterranean)

Cold-water coral mounds are key hot-spots of deep ocean biodiversity and also important archives of past climatic conditions. Nonetheless, the paleo-oceanographic setting in which coral mounds developed in the Mediterranean Sea during the last 500 ka still needs to be properly understood. This study describes the coral deposits and corresponding ages of two on-mound gravity cores acquired from opposite sectors of the newly discovered Cabliers Coral Mound Province (CMP, Alboran Sea, W Mediterranean). U–Th data revealed Pleistocene-aged corals covering mound formation periods from >389 to 9.3 ka BP and from 13.7 to 0.3 ka BP in the southern and northern mounds respectively. The coral-rich deposits of the cores were mainly dominated by Desmophyllum pertusum, although in some sections concurrent with the Middle Pleistocene and the Holocene, other corals such as Dendrophyllia cornigera and Madrepora oculata also appeared as dominating species. Coral mound formation stages generally occurred during deglacials and temperate interstadial (3.5–4.1 δ18O‰) periods, whereas during interglacials (<3.5 δ18O‰) coral mound formation only occurred in the northern and shallower mound. We interpret this to indicate that the shoaling of the interface between Atlantic (AW) and Levantine Intermediate Waters (LIW) during interglacial periods prevented the corals in the southern CMP from acquiring sufficient food supply, thus causing periods of coral mound stagnation. Similarly, the interruption in LIW formation throughout sapropel events also coincides with coral mound stagnation phases. This suggests that sapropel-derived processes, which originated in the eastern Mediterranean, likely affected the entire Mediterranean basin and further supports the role of LIW as a conveyor belt facilitating cold-water coral growth in the Mediterranean Sea. Overall, we show that these coral mounds yield important insights into how local changes in oceanographic conditions can influence coral mound development.

Monsoonal forcing of cold-water coral growth off southeastern Brazil during the past 160 kyr

Abstract. Cold-water corals (CWCs) constitute important deep-water ecosystems that are under increasing environmental pressure due to ocean acidification and global warming. The sensitivity of these deep-water ecosystems to environmental change is demonstrated by abundant paleorecords drilled through CWC mounds that reveal characteristic alterations between rapid formation and dormant or erosive phases. Previous studies have identified several central parameters for driving or inhibiting CWC growth such as food supply, oxygenation, and the carbon saturation state of bottom water, yet there are still large uncertainties about the relative importance of the different environmental parameters. To advance this debate we have performed a multiproxy study on a sediment core retrieved from the 25 m high Bowie Mound, located at 866 m water depth on the continental slope off southeastern Brazil, a structure built up mainly by the CWC Solenosmilia variabilis. Our results indicate a multifactorial control on CWC growth at Bowie Mound during the past ∼ 160 kyr, which reveals distinct formation pulses during northern high-latitude glacial cold events (Heinrich stadials, HSs) largely associated with anomalously strong monsoonal rainfall over the continent. The ensuing enhanced runoff elevated the terrigenous nutrient and organic-matter supply to the continental margin and likely boosted marine productivity. The dispersal of food particles towards the CWC colonies during HSs was facilitated by the highly dynamic hydraulic conditions along the continental slope that prevailed throughout glacial periods. These conditions caused the emplacement of a pronounced nepheloid layer above Bowie Mound, thereby aiding the concentration and along-slope dispersal of organic matter. Our study thus emphasizes the impact of continental climate variability on a highly vulnerable deep-marine ecosystem.

Long term monitoring of cold-water coral growth shows response to episodic meteorological events in the NW Mediterranean

Lophelia pertusa and Madrepora oculata, two cosmopolitan coral species of the deep ocean, build reef-like structures that provide ecological niches and nurseries for many species. These cold-water corals (CWCs) are thought to grow slowly making associated communities vulnerable to disturbance. However, in situ growth dynamics remains poorly known. The aim of this study was thus to quantify the growth rates of L. pertusa and M. oculata and their variability in relation to environmental conditions that are expected to influence their physiological functions. Such data can be useful to ground-truth physiological rates and improve the prediction of their sensitivity to global change.As part of a long-term monitoring programme of the Lacaze-Duthiers canyon in the Northwestern Mediterranean Sea, fragments of the two species were repeatedly sampled and redeployed at 530 m depth, from November 2010 to May 2018. Budding rates and linear extensions of L. pertusa and M. oculata were hence quantified and their inter-annual variability documented. The data were analyzed in relation to in situ monitoring of water temperature, current speed and turbidity.This study presents the longest in situ survey of the growth patterns of L. pertusa and M. oculata. The observations revealed a strong inter-annual variability in growth rates, both for L. pertusa and M. oculata, and suggest an influence of hydrological conditions on the growth dynamics. Particularly, as regard to episodic water plumes events such as NW wind-induced dense water shelf cascades and E-SE storm-induced downwellings in autumn-winter. In this study, three types of responses were observed: (i) high budding rates but low colony linear extension for strong water plume events periods (ii) high budding rates and high linear extension associated to organic matter supply during medium intensity water plume events, and (iii) low budding rates, low linear extension and high mortality due to high sedimentation rates in the absence of water plume event. Lophelia pertusa appears to be more resilient to these hydrological changes. Disturbance of the regional circulation patterns, in addition to longer-term change in water mass properties such as warming, deoxygenation and acidification, are thus important parameters to consider for CWC reef stability. This is particularly important since increased storminess, and decreased cascading/downwelling intensities, are predicted to occur within the next century.

Cold-Water Corals in Gas Hydrate Drilling Cores from the South China Sea: Occurrences, Geochemical Characteristics and Their Relationship to Methane Seepages

Cold-water corals (CWCs) are frequently found at cold seep areas. However, the relationship between fluid seepage and CWC development is not clear. Here, for the first time, we report the occurrences, species identification, mineralogy, carbon and oxygen isotopes, as well as elemental compositions of fossil CWC skeletons from gas-hydrate-bearing sediment in drilling cores from the South China Sea (SCS). Three sites (GMGS-08, GMGS-09B, and GMGS-16) were investigated but CWCs were only found at one site (GMGS-09B). Interestingly, the CWCs were found in three horizons and they were all embedded with authigenic carbonates. Three genera of fossil CWCs (Crispatotrochus sp., Solenosmilia sp. and Enallopsammia sp.) were identified. The CWC fragments are predominantly aragonite. The CWCs exhibit δ13C values between −8.4‰ and −0.6‰ that are significantly higher than δ13C values of the associated seep carbonates (δ13C values with an average of −55.6‰, n = 19), which indicates a carbon source other than methane for the CWCs. It appears that authigenic carbonates provide a substratum for coral colonization. Bathymetric high points, appropriate water temperature and stronger bottom-water currents at site GMGS-09B might be crucial to keep conditions favorable for the growth of CWCs in the studied area. In addition, high trace-element concentrations of Cr, Ni, Pb, U, Ba, Th, and Sr suggest that the CWCs are influenced by strong fluid seepage that can reach the water-sediment interface, and associated microbial activity. Hence, it also becomes evident that CWCs in hydrocarbon-rich seepage areas not only provide a critical constraint on the impact of fluid emission on the bottom water chemistry, but also are likely to be very precise recorders of the end time of cold seep activity.

Environmental factors influencing benthic communities in the oxygen minimum zones on the Angolan and Namibian margins

Abstract. Thriving benthic communities were observed in the oxygen minimum zones along the southwestern African margin. On the Namibian margin, fossil cold-water coral mounds were overgrown by sponges and bryozoans, while the Angolan margin was characterized by cold-water coral mounds covered by a living coral reef. To explore why benthic communities differ in both areas, present-day environmental conditions were assessed, using conductivity–temperature–depth (CTD) transects and bottom landers to investigate spatial and temporal variations of environmental properties. Near-bottom measurements recorded low dissolved oxygen concentrations on the Namibian margin of 0–0.15 mL L−1 (≜0 %–9 % saturation) and on the Angolan margin of 0.5–1.5 mL L−1 (≜7 %–18 % saturation), which were associated with relatively high temperatures (11.8–13.2 ∘C and 6.4–12.6 ∘C, respectively). Semidiurnal barotropic tides were found to interact with the margin topography producing internal waves. These tidal movements deliver water with more suitable characteristics to the benthic communities from below and above the zone of low oxygen. Concurrently, the delivery of a high quantity and quality of organic matter was observed, being an important food source for the benthic fauna. On the Namibian margin, organic matter originated directly from the surface productive zone, whereas on the Angolan margin the geochemical signature of organic matter suggested an additional mechanism of food supply. A nepheloid layer observed above the cold-water corals may constitute a reservoir of organic matter, facilitating a constant supply of food particles by tidal mixing. Our data suggest that the benthic fauna on the Namibian margin, as well as the cold-water coral communities on the Angolan margin, may compensate for unfavorable conditions of low oxygen levels and high temperatures with enhanced availability of food, while anoxic conditions on the Namibian margin are at present a limiting factor for cold-water coral growth. This study provides an example of how benthic ecosystems cope with such extreme environmental conditions since it is expected that oxygen minimum zones will expand in the future due to anthropogenic activities.

Cold-water corals in a cold seep area on the northern continental slopes of the South China sea and their isotopic characteristics

Cold-water corals (CWCs) were found to occur in association with authigenic carbonates in a cold seep area on the northern continental slopes of the South China Sea (SCS). The taxa identified were: Balanophyllia (Balanophyllia) sp., Balanophyllia (Eupsammia) sp., Lochmaeotrochus sp., Enallopsammia sp., Crispatotrochus sp.1 and Crispatotrochus sp.2. The δ13C (−7.36‰ to −1.15‰, V-PDB) and 87Sr/86Sr ratios (0.709126–0.709184) indicated that CWC aragonite skeletons had been precipitated from seawater without the involvement of seeping fluids. The presence and growth of CWCs on the slopes of the submarine seamounts in the south-western (SW) Dongsha area could be directly linked with the hard substrates provided by exhumed hydrocarbon-imprinted authigenic carbonates and fed by the food particles enhanced by high-velocity internal tides and near-bottom currents. A multi-step process for CWC colonization was proposed that encompassed cold-seepage processes as a driver for hard-substrate generation of CWC, as well as the subsequent settlement and maintenance of CWC larvae under the persistent influence of bottom currents.

Experimental assessment of the effects of coldwater coral patches on water flow

Despite their importance for ecosystem functioning, little is known about flow patterns at the cold-water coral-patch scale (i.e. cm to m). In situ flow speed measurements using moorings or landers only provide general near-reef flow patterns, and experimental studies so far have focussed on single coral branches for feeding and behavioural studies. We addressed this knowledge gap by means of a flume study. The effect of different coral patches (4, 10 and 25 cm height), created from dead coral branches, on the ambient flow field and the refreshment rate of water within a patch was assessed using 3 realistic current speeds (0.05, 0.15 and 0.3 m s-1). High bottom roughness in combination with strong currents increased current velocity and turbulence in the wake of all patches, even with very low relief. The formation of 2 dynamically different environments was observed. The framework-water interface was characterised by high turbulence and enhanced vertical turbulent transport of momentum, while in the wake, reduced turbulence and vertical mixing activity was observed, characterised by near stagnant flows. Subsequently, water refreshment rates within a patch drastically decreased at current speeds 0.2 m s-1. Combining flume observations with available in situ data suggests that heterogeneity and patchiness of cold-water coral growth is likely induced by flow patterns at the coral-patch scale, influencing the fate of particulate and dissolved matter as well as oxygen exchange rates in and around the reef.

Macro- and microplastics affect cold-water corals growth, feeding and behaviour

Plastic contamination is now recognized as one of the most serious environmental issues for oceans. Both macro- and microplastic debris are accumulating in surface and deep waters. However, little is known about their impact on deep marine ecosystems and especially on the deep-sea reefs built by emblematic cold-water corals. The aim of this study was to investigate whether plastics affected the growth, feeding and behaviour of the main engineer species, Lophelia pertusa. Our experiments showed that both micro- and macroplastics significantly reduced skeletal growth rates. Macroplastics induced an increased polyp activity but decreased prey capture rates. They acted as physical barriers for food supply, likely affecting energy acquisition and allocation. Inversely, microplastics did not impact polyp behaviour or prey capture rates, but calcification was still reduced compared to control and in situ conditions. The exact causes are still unclear but they might involve possible physical damages or energy storage alteration. Considering the high local accumulation of macroplastics reported and the widespread distribution of microplastics in the world ocean, our results suggest that plastics may constitute a major threat for reef aggradation by inhibiting coral growth, and thus jeopardise the resilience of cold-water coral reefs and their associated biodiversity.

Imprint of Holocene Climate Variability on Cold‐Water Coral Reef Growth at the SW Rockall Trough Margin, NE Atlantic

AbstractU‐Th ages and temperatures derived from Li/Mg have been measured on coral fragments of Lophelia pertusa and Madrepora oculata collected from two sediment cores, which were taken from cold‐water coral (CWC) mounds at 700–790m water depth at the SW Rockall Trough margin. Our data, combined with previous published data, have allowed us to first estimate the occurrence of CWC at the SW Rockall Trough margin during the Holocene and, second, to better constrain the environmental conditions driving variability in CWC growth. CWC abundance is marked by a pronounced increase in the mid‐Holocene (∼6 ka) and is modulated by millennial‐scale variability throughout the late‐Holocene. The mid‐Holocene proliferation of CWC coincides with lowest IRD abundances and a major reorganization of the circulation at thermocline depth in the Rockall Trough, marked by the progressive replacement of the fresh‐cold Sub‐Arctic Intermediate Water (SAIW) by the saltier and nutrient‐rich Eastern North Atlantic Water (ENAW). This event must have established a modern‐like winter mixed layer and thermocline structure, generating suitable conditions for enhanced surface productivity, downslope transport of food particles, bottom current acceleration at mound depth and thus CWC growth. Several short time intervals of decreased CWC occurrences closely match prominent increases in North Atlantic drift ice and storminess in Northern Europe. We, therefore, propose that high detrital supply and/or changes in the vertical density gradient associated with millennial‐scale ice‐rafted detritus (IRD) events are the likely controlling factors for CWC growth and subsequent mound formation on the SW Rockall Trough margin.

Large-scale paleoceanographic variations in the western Mediterranean Sea during the last 34,000 years: From enhanced cold-water coral growth to declining mounds

Abstract Quantitative and qualitative analyses of cold-water coral (CWC) fragments from two sediment cores obtained from the Melilla Mounds Field (MMF) in the Alboran Sea, western Mediterranean Sea, reveal an alternation of periods dominated by distinct CWC species. The lower parts of the cores are dominated by the CWC species Lophelia pertusa, which is successively replaced in the upper parts by the species Madrepora oculata and Dendrophyllids. The transition in the macrofauna coincides with a characteristic change in the benthic foraminiferal assemblage. Benthic foraminiferal assemblage BFAglacial, in accordance with benthic (Cibicides lobatulus) and planktic (Globigerina bulloides) δ13C and δ18O values provide evidence for generally high surface productivity, cold and well-ventilated bottom waters lasting from the end of Marine Isotope Stage 3 (33.3 ka BP) reaching a maximum at the transition Allerod–Younger-Dryas. Together with δ13C of the organic carbon and Rock-Eval pyrolysis, benthic foraminiferal assemblage BFAinterglacial established since the Early Holocene indicates that the MMF experienced a decrease in bottom-water energy that caused an organic carbon-enrichment in the sediments and also depleted oxygen waters. Compared to the pre-Holocene interval dominated by the planktic foraminifera Neogloboquadrina incompta and the benthic foraminiferal assemblage BFAglacial, the organic carbon deposited during the Holocene in the MMF contains more refractory components in relation to sea-level rise and modern oceanographic configuration. Based on our data, we suggest that L. pertusa has a higher ecological requirement than M. oculata and Dendrophyllids especially with regard to oxygen and nutrient availability.

Sedimentary processes and cold-water coral mini-mounds at the Ferrol canyon head, NW Iberian margin

The western Iberian margin has a complex morphology controlled by both geological and oceanographic processes. Compared to the submarine canyons in the southern part of this margin (e.g. Nazaré and Setubal), the canyons in the northern part (e.g. Ferrol and A Coruña) have received little attention. This study maps the geomorphological features around the Ferrol canyon head and combines them with oceanographic observations to infer the sedimentary and oceanographic processes active in this area. Furthermore, the occurrence of a cold-water coral (CWC) mini-mound province near the canyon head and the Ortegal Spur pockmark field is investigated with regard to seepage processes, oceanographic conditions and anthropogenic impact.The Ferrol canyon head and outer Ortegal Spur are characterized by erosional (erosional and abraded surfaces, contourite channels and furrows), depositional (contourite drifts and sediment waves) and mixed (contourite terrace) features, indicating a dominant control of bottom currents on the sedimentary processes. Bottom currents are related to the interaction of the canyon head topography with both the Mediterranean Outflow Water (MOW) slope current and with M2 internal tides associated to the interface (pycnocline) between Eastern North Atlantic Central Water (ENACW) and MOW. Due to this interaction, the Ferrol canyon head is subject to active sediment resuspension and forms a potential source area for the intermediate nepheloid layers along the NW Iberian upper slope.The CWC mini-mounds (1.9–2.7 m high and 75–141 m in diameter) occur between 400 and 560 m depth. Seismic facies signatures indicative of fluid-flow and morphometric analysis implies that larger CWC mini-mounds adjacent to the pockmark field initiated by colonization of pre-existing pockmarks, while smaller more clustered mounds developed independently. The observed mini-mounds are relict features likely related to their position outside of the contemporary ENACW-MOW interface which favours CWC growth through (1) increased bottom currents, enhancing food supply and (2) the presence of a potential density envelope σθ = 27.35–27.65 kg/m3, proposed to control coral larvae transport along the NE Atlantic margin. Alternatively, the lack of contemporary coral growth might be related to habitat destruction by bottom trawling. However, a preliminary age constraint reveals CWC growth occurred during the early Holocene, coeval to relict CWC mini-mounds near canyon heads on the Armorican margin. Their existence suggests a regional shift in the NE Atlantic density profile where the ENACW-MOW interface occurred up to 200 m above its contemporary position.

Predictive habitat modeling in two Mediterranean canyons including hydrodynamic variables

Madrepora oculata and Lophelia pertusa are the two main ecosystem engineering, scleractinian cold-water corals (CWC) found in Mediterranean canyons. Factors controlling CWC distribution in the Mediterranean Sea are not yet fully understood in spite of such ecosystems being recognized as sensitive habitats by the General Fisheries Commission for the Mediterranean. As they are threatened by fishery activity, they are subject to management and protection measures. In order to contribute towards identifying the major drivers governing CWC distribution at local scale, which is a prerequisite for proper management, we focused our attention on two canyons: (1) the Cassidaigne canyon, located in the eastern part of the Gulf of Lion, in which CWC ecosystems have settled in an upwelling environment and form large colonies, and (2) the Bari Canyon System, in the southwestern Adriatic, a site of coral growth that has been hypothesized to respond to hydrographic processes, including the cascading of North Adriatic Dense Water.The objective of our study was to combine several ecological variables to describe the environmental conditions in favor of CWC settlement and growth: (1) CWC observations, extracted from geo-referenced underwater videos, (2) seafloor characteristics derived from high-resolution bathymetry, (3) data on local hydrodynamic conditions (from high resolution hydrodynamic models). Habitat suitability models were used to identify the main variables driving CWC distribution. Models based on presence-only data (Maxent and ENFA) and on presence-absence data (GLMs) were fitted and compared.Seafloor ruggedness was identified to be the major factor driving CWC distribution in both canyons with the three methods. Two hydrodynamic variables (mean temperature and current velocity) were the second most important predictors for explaining CWC settlement and growth. Suitable areas for CWC habitat occurrence were mapped for both canyons. Spatial distributions were generally predicted at the same locations, although the GLM gave less realistic results in the Bari canyon system probably due to the limited range cover of the entire environmental conditions by the absence points, suggesting that the Maxent and ENFA models were more efficient.These theoretical distributions will help in the assessment of potential habitat extent in the deep-sea and also in the scheme of the Marine Strategy Framework Directive (MSFD).

Upper Pleistocene cold-water corals from the Inner Sea of the Maldives: taphonomy and environment

Cold-water corals of the Late Pleistocene (21,400–22,500 BP) are recorded from the sea-bottom of two inter-atoll channels (Kardiva Channel at 457-m depth and Male Vaadhoo Channel at 443-m depth) of the eastern row of the Maldives archipelago. Coral assemblages are composed mainly by Lophelia pertusa and secondarily by Madrepora oculata and Enallopsammia rostrata. These cold-water coral patches are places where the benthic life, mainly sessile, is concentrated, which is clearly absent off-rubble patches. The main epibionts are tube-dwelling polychaetes (mainly Spirorbis and Serpula), bryozoans, siliceous sponges, barnacles, gorgonids, solitary corals, encrusting foraminifera, and microbial mats. The analysis of epibionts assemblages shows different biocoenoses between both studied sites as well as a dependency of the epibiont coverage with regard to the coral genus. Some living benthic organisms such as brachiopods, bivalves, gastropods, barnacles, and ophiuroids find refuge among coral branches. The common record of juvenile specimens of vagile organisms such as small ophiuroids, is probably related to the nursery function of the cold-water corals in spite they are fossils. Environmental requirements of Recent cold-water corals (Lophelia, Madrepora and Enallopsammia) differ of conditions at both sampling sites with sensibly lower oxygenation degree and density of waters than needed for cold-water corals. Therefore, it is proposed that the present-day oxygen and density conditions are the factors which inhibit modern cold-water coral growth in the inter-atoll channels.