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Over the last few decades the exploration of deep-water
environments using new technologies has revealed insights
into parts of our planet that challenge conventional
wisdom. Coral reefs, once thought to be restricted to warm
shallow waters in tropical and subtropical regions, have
been found found in dark, cold, nutrient-rich waters off the
coasts...
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Citations
... For example, these species are often referred to as non-constructional (previously ahermatypic), not habitat forming, found only in deep or cold-water habitats, and solitary living 17 . However, while many species within these groups align with these descriptions, some have the capacity to form constructional (previously hermatypic) colonial reef systems 14,[17][18][19] and maintain similar calcification rates to photoendosymbiotic species 20 . The absence of photoendosymbiotic dinoflagellates within these coral species removes light constraints on growth, enabling these species to exploit broad geographical and bathymetric ranges. ...
... As such, coral species living without reliance on photoendosymbioses are found across polar, temperate, sub-tropical and tropical regions, exhibiting immense depth variation, from the intertidal zone to abyssal depths greater than 6000 m 17,21 . In deep and/or cold-water habitats, coral reefs can play host to highly diverse biological assemblages similar to those of tropical coral reefs 9,18 but these ecosystems are largely understudied due to the complexities and cost associated with deep-water research, a perceived lack of environmental significance and little information on potential environmental threats 19 . However, these ecosystems cover vast areas, are environmentally important including facilitating speciation within the deep sea 18 and have significant socio-economic roles, such as hosting breeding grounds for fished species 22 . ...
... Approximately 20 of the deep sea Scleractinia form structural reef systems 22 , including species such as; Desmophyllum pertusum, Madrepora oculata, Oculina varicosa, Solenosmilia variabilis, Enallopsammia profunda, Goniocorella dumosa and Bathelia candida, whilst the majority of species records in the deep sea are for solitary species. Desmophyllum pertusum (Linneaus 1758) is the most commonly occurring reef-building coral at depth 19 , significantly contributing to reef structures that have been found to span several kilometres 76 . Desmophyllum pertusum has a broad geographical range, extending throughout the North Atlantic Ocean 77 and the Pacific Ocean 78,79 and can be found anywhere from 50 to 3000 m depth 77 . ...
Globally tropical Scleractinian corals have been a focal point for discussions on the impact of a changing climate on marine ecosystems and biodiversity. Research into tropical Scleractinian corals, particularly the role and breakdown of photoendosymbiosis in response to warming, has been prolific in recent decades. However, research into their subtropical, temperate, cold- and deep-water counterparts, whose number is dominated by corals without photoendosymbiosis, has not been as prolific. Approximately 50% of Scleractinian corals (> 700 species) do not maintain photoendosymbiosis and as such, do not rely upon the products of photosynthesis for homeostasis. Some species also have variable partnerships with photendosymbionts depending on life history and ecological niche. Here we undertake a systematic map of literature on Scleractinian corals without, or with variable, photoendosymbiosis. In doing so we identify 482 publications spanning 5 decades. In mapping research effort, we find publications have been sporadic over time, predominately focusing on a limited number of species, with greater research effort directed towards deep-water species. We find only 141 species have been studied, with approximately 30% of the total identified research effort directed toward a single species, Desmophyllum pertusum, highlighting significant knowledge gaps into Scleractinian diversity. We find similar limitations to studied locations, with 78 identified from the global data, of which only few represent most research outputs. We also identified inconsistencies with terminology used to describe Scleractinia without photoendosymbiosis, likely contributing to difficulties in accounting for their role and contribution to marine ecosystems. We propose that the terminology requires re-evaluation to allow further systematic assessment of literature, and to ensure it’s consistent with changes implemented for photoendosymbiotic corals. Finally, we find that knowledge gaps identified over 20 years ago are still present for most aphotoendosymbiotic Scleractinian species, and we show data deficiencies remain regarding their function, biodiversity and the impacts of anthropogenic stressors.
... and OBIS (2022). From the NOAA database we selected all records of the main CWC reef-building species Desmophyllum pertusum (previously Lophelia pertusa), Enallopsammia profunda, E. pusilla, E. rostrata, Goniocorella dumosa, Madrepora carolina, M. oculata, and Solenosmilia variabilis (Freiwald et al., 2004;Maier et al., 2023), below 100 m depth, recorded from 1900 or later, with a horizontal location accuracy of 1000 m or less, and between the critical latitudes for the M2 tide of 70°N and S (15 629 records). ...
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.
... Their colony morphology varies from branched, bush-like, and feather-like to whip types (Wagner et al. 2012). The dense canopies they form can change local physical conditions and generate a three-dimensional habitat, which provide shelter to associated species and, ultimately, increase biodiversity (Freiwald et al. 2004;Buhl-Mortensen et al. 2010;De Clippele et al. 2019). One of the main components of the visible black coral forests-associated biodiversity is accounted by epifauna (i.e., crustaceans, polychaetes and molluscs, Lavelle 2012; Wagner et al. 2012), which can find habitat (Herler 2007), food (Angel 1990;Bo et al. 2012), and protection against predators (Lavelle 2012). ...
Black corals are important components of mesophotic and deep-water marine habitats. Their presence at great depths (e.g., 50 to 200 m) makes accessibility difficult, limiting our understanding of the associated biodiversity. Amphipods dominate vagile epifauna in marine habitats around the world, fulfilling important ecosystem functions. However, there are no studies on amphipods exclusively associated with black corals, including relationships between their ecological patterns (e.g., abundances) and the size of coral colonies. We investigated the epifaunal composition and abundance associated with black coral colonies of Antipathella wollastoni in the subtropical eastern Atlantic Ocean. In total, 1,736 epifaunal individuals were identified, of which 1,706 (98.27%) were amphipods, belonging to 6 taxa. We identified and described a new amphipod genus and species within the Stenothoidae family, Wollastenothoe minuta gen. nov., sp. nov., which outnumbered the amphipod assemblage (86.15%) and provided a complete taxonomic key of Stenothoidae family including this new finding. For the first time, the association between an amphipod species and a black coral was described, including a strong correlation between coral colony size and amphipod abundances. This study demonstrates that epifauna associated with mesophotic black corals remains largely undescribed.
... 18 The next depth zone found at between 70-140 m include similar assemblages to the upper-mid assemblages of anemones, so corals, hydroids, tube worms and barnacles, as well as the hard bodies deep water cold coral, L. pertusa. At greater depths assemblages can become sparser, but this isn't necessarily the case, L. pertusa for example has been found at depths greater than 1000 m. 46 Birds including gules, garnets and kittiwakes 47 colonise the topsides (structures above sea level) as structures such as drilling derricks provide many places to build nests and raise chicks and deposit guano. Fig. 5 illustrates the evolution of the species assemblages over time from emplacement to the point at which the OGI MMS are ready for decommissioning. ...
Oil and gas industry manmade structures (OGIMMS) in the marine environment can support thriving and biodiverse ecosystems. ‘Clear seabed’ policies require that all OGIMMS are removed once commercial activity has ceased, thereby removing a large proportion of the ecosystem and leaving the remaining community degraded beyond repair. The environmental impact of this method of decommissioning is huge and wide ranging, but no studies have to date looked at the possible impacts that removing these ecosystems has to climate change. This first of its kind study modelled biomass associated with OGIMMS and the potential Blue Carbon (BC) that these ecosystems may represent. The study found that in the UK North Sea (UKNS) there is currently 1.75 MtC of BC adhered to OGIMMS and globally there is 64 MtC. The study investigated the consequences of removing this BC and found that if it is allowed to degrade in landfill, up to 96 MtCO2e of greenhouse gas (GHG) emissions will be released from UK BC and globally up to 2,730 MtCO2e of GHG emissions will be released. Furthermore, forward modelling techniques were uniquely employed to look at future potential growth of these ecosystem and found that in the UKNS if the ecosystem was allowed to remain in place by decommissioning in situ, at 100 years since installation, UKNS OGIMMS BC could grow to 27 MtC and global BC could grow to 264 MtC. The study demonstrates the vast potential of BC associated with OGIMMS in the marine environment to sequester carbon over the long term and that current clear seabed practises damage these important ecosystems beyond repair, destroying current BC stocks and the vast potential BC stocks that could develop over time, as well as releasing large volumes of GHG emissions from the degradation of the biomass.
... Scleractinian corals D. pertusum, M. oculata, E. rostrata, G. dumosa, O. varicosa, and S. variabilis are all reef-framework forming species with cosmopolitan distributions (Davies and Guinotte, 2011). Desmophyllum pertusum often occurs as the dominant framework-forming Scleractinian or as isolated thickets mainly in the North Atlantic (Freiwald et al., 2004;Tong et al., 2016), while M. oculata has mainly been found as a secondary framework-forming species within D. pertusum or G. dumosa reefs (Freiwald et al., 2004;Roberts et al., 2006). Goniocorella dumosa has been observed as the dominant reef-builder in New Zealand waters (Tracey et al., 2011). ...
... Scleractinian corals D. pertusum, M. oculata, E. rostrata, G. dumosa, O. varicosa, and S. variabilis are all reef-framework forming species with cosmopolitan distributions (Davies and Guinotte, 2011). Desmophyllum pertusum often occurs as the dominant framework-forming Scleractinian or as isolated thickets mainly in the North Atlantic (Freiwald et al., 2004;Tong et al., 2016), while M. oculata has mainly been found as a secondary framework-forming species within D. pertusum or G. dumosa reefs (Freiwald et al., 2004;Roberts et al., 2006). Goniocorella dumosa has been observed as the dominant reef-builder in New Zealand waters (Tracey et al., 2011). ...
... Goniocorella dumosa has been observed as the dominant reef-builder in New Zealand waters (Tracey et al., 2011). Enallopsammia rostrata is also observed associated with D. pertusum, M. oculata, and S. variabilis (Freiwald et al., 2004). Oculina varicosa can inhabit both shallow and deep waters (Reed, 2002). ...
Species distribution models (SDMs) are useful tools for describing and predicting the distribution of marine species in data-limited environments. Outputs from SDMs have been used to identify areas for spatial management, analyzing trawl closures, quantitatively measuring the risk of bottom trawling, and evaluating protected areas for improving conservation and management. Cold-water corals are globally distributed habitat-forming organisms that are vulnerable to anthropogenic impacts and climate change, but data deficiency remains an ongoing issue for the effective spatial management of these important ecosystem engineers. In this study, we constructed 11 environmental seabed variables at 500 m resolution based on the latest multi-depth global datasets and high-resolution bathymetry. An ensemble species distribution modeling method was used to predict the global habitat suitability for 10 widespread cold-water coral species, namely, 6 Scleractinian framework-forming species and 4 large gorgonian species. Temperature, depth, salinity, terrain ruggedness index, carbonate saturation state, and chlorophyll were the most important factors in determining the global distributions of these species. The Scleractinian Madrepora oculata showed the widest niche breadth, while most other species demonstrated somewhat limited niche breadth. The shallowest study species, Oculina varicosa, had the most distinctive niche of the group. The model outputs from this study represent the highest-resolution global predictions for these species to date and are valuable in aiding the management, conservation, and continued research into cold-water coral species.
... Model outputs have indicated that continental slopes are one of the main suitable coral sites worldwide 8 . The bathyal (~200-3000 m depth) region 12 has large protruding topographical features, interfering with bottom current flow and creating colonizable hard substrates at multiple depths; this enhances seabed erosion and resuspends particulates, which provides an elevated supply of suspended food particles for corals 13 . Along the Brazilian margin, five main water masses with distinct intensity and transport directions influence the continental slope 14 , which may contribute to regional species diversity pools. ...
Deep-sea coral distribution and composition are unknown in much of the global ocean, but repurposing ocean industry surveys can fill that gap. In Santos Basin, southeast Brazil, areas (241-963 m depth) were surveyed during seven Petrobras cruises, mapping bottom topography with multibeam sonar, then collecting video with remotely operated vehicles. Here, we defined deep-sea coral species distribution and richness, using these surveys, correlating them to physical oceanographic properties. Solenosmilia variabilis was the most prevalent colonial species in coral mounds. Overall, 67% of species were Octocorallia. Coral assemblage structure, abundance, and richness varied among sites both within and among depths, with higher density and richness in the northernmost Santos basin. Depth was the strongest predictor for scleractinian coral distribution, with depth ranges varying by species. Assemblage differences corresponded to changes in water mass. Desmophyllum pertusum was more abundant in South Atlantic Central Water and S. variabilis in Antarctic Intermediate Water influenced areas.
... In the intertidal area, brackish mangroves as well as sand-and mudflats (tidal flats) emerge, whereas permanently submerged ecosystems such as seagrass beds and coral reefs arise close to the open ocean. At higher latitudes, tidal marshes (i.e., salt marshes) displace mangroves, and cold-water corals can thrive in place of warm-water corals but in water depths of usually >40 m (Freiwald et al., 2004). ...
Data were obtained from the literature to identify past changes in and the present status of the coastal carbon cycle. They indicate that marine coastal ecosystems driving the coastal carbon cycle cover, on average, 5.8% of the Earth’s surface and contributed 55.2% to carbon transport from the climate-active carbon cycle to the geological carbon cycle. The data suggest that humans not only increase the CO 2 concentration in the atmosphere but also mitigate (and before 1860 even balanced) their CO 2 emissions by increasing CO 2 storage within marine coastal ecosystems. Soil degradation in response to the expansion and intensification of agriculture is assumed to be a key process driving the enhanced CO 2 storage in marine coastal ecosystems because it increases the supply of lithogenic matter that is known to favour the burial of organic matter in sediments. After 1860, rising CO 2 concentrations in the atmosphere indicate that enhanced CO 2 emissions caused by land-use changes and the burning of fossil fuel disturbed what was a quasi-steady state before. Ecosystem restoration and the potential expansion of forest cover could mitigate the increase of atmospheric CO 2 concentrations, but this carbon sink to the atmosphere is much too weak to represent an alternative to the reduction of CO 2 emission in order to keep global warming below 1.5–2.°C. Although the contribution of benthic marine coastal ecosystems to the global CO 2 uptake potential of ecosystem restoration is only around 6%, this could be significant given national carbon budgets. However, the impact on climate is still difficult to quantify because the associated effects on CH 4 and N 2 O emissions have not been established. Addressing these uncertainties is one of the challenges faced by future research, as are related issues concerning estimates of carbon fluxes between the climate-active and the geological carbon cycle and the development of suitable methods to quantify changes in the CO 2 uptake of pelagic ecosystems in the ocean.
... Deep-sea corals (Phylum Cnidaria) are of particular importance within deep-sea faunal groups (Roberts and Hirshfield 2004). Also referred to as cold-water corals, deep-sea corals are found globally at temperatures between 4 and 12°C and most commonly between 200 and 2000 m depth (Freiwald et al. 2004;Roberts et al. 2006). Several species in this group can create large, complex, and fragile habitats (Huehnerbach et al. 2007). ...
Seabed-contact activities operating in the deep sea can generate sediment plumes that pose varying levels of threat to benthic fauna. Corals are important components of deep-sea ecosystems and can be particularly sensitive to elevated suspended sediment concentrations. In this study, we exposed colony fragments of the New Zealand deep-sea scleractinian Goniocorella dumosa to four-day pulses of four target sediment concentrations: 0 mg l−1 (representing control conditions) and 45, 102 and 435 mg l−1 (targeting concentrations expected from mining and trawling disturbance). All coral fragments survived. Oxygen consumption rates were not affected by treatments and time. No visible detrimental effects on coral health were noted after the first pulse of sediment exposure. However, both a loss of coenosarc and instances of polyp mortality were observed on fragments exposed to suspended sediments during the following sediment pulses. This observed decline in coral health over time indicates that G. dumosa could cope with sediment disturbance from human activities that disturb the seafloor for periods of up to four days, but that repeated, or prolonged sediment exposure could cause a deterioration in the health condition of this species. This hypothesis should be further investigated in following studies.
... Cold-water coral (CWC) reefs are biodiversity-rich ecosystems found in a vast range of water depths and habitats, down to over 1000 m of depth on seamounts and continental slopes (Freiwald et al. 2004), and up to 39 m in Norwegian fjords (Fosså et al. 2002). These ecosystems rely on the three-dimensional (3D) skeletal frameworks formed by a few species of azooxanthellate scleractinians such as the widely distributed Desmophyllum pertusum (previously known as Lophelia pertusa), the primary framework builder in the North Atlantic Ocean (Freiwald et al. 2004). ...
... Cold-water coral (CWC) reefs are biodiversity-rich ecosystems found in a vast range of water depths and habitats, down to over 1000 m of depth on seamounts and continental slopes (Freiwald et al. 2004), and up to 39 m in Norwegian fjords (Fosså et al. 2002). These ecosystems rely on the three-dimensional (3D) skeletal frameworks formed by a few species of azooxanthellate scleractinians such as the widely distributed Desmophyllum pertusum (previously known as Lophelia pertusa), the primary framework builder in the North Atlantic Ocean (Freiwald et al. 2004). The majority of the CWC reef framework consists of dead coral skeleton (devoid of living tissue) (Vad et al. 2017), which supports key ecological functions modulated by its physical branching structure, including habitat provision for most of the numerous reef-associated species (Mortensen et al. 1995) and baffling and retention of suspended sediment and particulate organic matter (Wheeler et al. 2005;Maier et al. 2021;Wang et al. 2021). ...
... The marked architectural diversity of D. pertusum at local scales has important implications for CWC reef ecosystems, suggesting partitioning of the coral's ecological functions not solely among different reef zones (living coral, dead framework and coral rubble; Freiwald et al. 2004), but also between different portions of the dead framework. In terms of habitat provision, this architectural variability may induce locally heterogeneous distribution patterns of reef-dwelling organisms that thrive in different framework configurations, contributing to complex local structuring of CWC reef communities (Henry et al. 2013). ...
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.
... Humans often exploit these ecosystem services without a good understanding of the impacts on the processes that sustain them (Levin et al., 2020;Mejjad and Rovere, 2021). For example, fisheries bottom trawls remove coral colonies, which can be hundreds to thousands of years old (Watling and Norse, 1998;Roberts et al., 2005) and provide nursery and breeding grounds for the same species that are targeted by the fisheries (Freiwald et al., 2004;Clark et al., 2016). In another example, the Deepwater Horizon (DWH) oil spill caused significant damage to coral colonies that were hundreds of years old (Prouty et al., 2016), and complete recovery of the ecosystem and its services may require time scales on the same order-of-magnitude (Girard & Fisher, 2018). ...
The deep ocean comprises complex ecosystems made up of numerous community and habitat types that provide multiple services that benefit humans. As the industrialization of the deep sea proceeds, a standardized and robust set of methods and metrics need to be developed to monitor the baseline conditions and any anthropogenic and climate change-related impacts on biodiversity, ecosystem function, and ecosystem services. Here, we review what we have learned from studies involving offshore-energy industries, including state-of-the-art technologies and strategies for obtaining reliable metrics of deep-sea biodiversity and ecosystem function. An approach that includes the detection and monitoring of ecosystem services, with open access to baseline data from multiple sectors, can help to improve our global capacity for the management of the deep ocean.
