Microplastics and cellulosic microparticles in North Atlantic deep waters and in the cold-water coral Lophelia pertusa.

This study explores microplastic and cellulosic microparticle occurrences in the NE Atlantic, focusing on the Porcupine Bank Canyon and Porcupine Seabight. Water samples from depths ranging between 605 and 2126 m and Lophelia pertusa coral samples from 950 m depth were analysed. Microparticles were detected in deep-water habitats, with concentrations varying from 2.33 to 9.67 particles L-1 in the Porcupine Bank Canyon, notably lower at greater depths. This challenges the assumption of deeper habitats solely acting as microplastic sinks. We also found evidence of microparticle adsorption and ingestion by L. pertusa. The presence of microparticles in cold-water corals underscores their vulnerability to pollutants. Furthermore, the dominance of rayon microparticles in both water and coral samples raises questions about marine pollution sources, potentially linked to terrestrial origins. This research emphasises the critical need for comprehensive exploration and conservation efforts in deep-sea environments, especially to protect vital ecosystems like L. pertusa reefs.

Do cold water corals provide an essential habitat for Helicolenus dactylopterus (Delaroche, 1809) in the Northwest Africa?

Cold water corals (CWC) provide habitats for many organisms including demersal fish. Bottom trawl observations have indicated a co-occurrence of the fish Helicolenus dactylopterus with CWC reefs, but a detailed understanding of this relation is lacking. To better understand the nature of this relation we have analyzed 85 video-lines from ROV dives conducted at 25-1700 m depth off Morocco, Mauritania, and Senegal in 2020 and 2021. We annotated abundance, size, and behavior of the 552 specimens observed (32% juveniles and 68% adults), of these 82% occurred in CWC habitats at 400-600 m depth. Both juveniles and adults were observed standing on the seafloor. Our observations are discussed considering available knowledge on feeding ecology and life cycle of H. dactylopterus. Our findings show that CWC provides an essential habitat for this species at least during parts of its lifecycle, however, more behavioral studies are needed for an in-depth understanding of this association.

Incorporation of abandoned and lost fishing gear into the structure of Dendrophyllia ramea in the Atlantic coast of Portugal.

Plastic pollution poses global and societal concerns, especially from discarded fishing gear, threatening seabed environments like coral reefs. This study examines the incorporation of lost and/or abandoned fishing gear - specifically synthetic lines, and filaments - into the structure of orange tree coral, Dendrophyllia ramea along the coast of Portugal, in the North-East Atlantic Ocean. The specimens were inadvertently captured by local fishers (Sines and Cascais), with 6 % showing filaments inside their structure, raising questions about their potential impact on coral health. We discuss the implications of understanding the interactions between plastics, fishing gear, and corals, which is important for developing conservation strategies. We address the need for improved of measures aimed at reducing the impact of fishing gear on corals, emphasizing the importance of endorsing biodegradable fishing materials and supporting lost gear retrieval initiatives. Furthermore, we emphasize the urgent need to communicate these issues to both fishers and stakeholders.

Implications for the conservation of deep-water corals in the face of multiple stressors: A case study from the New Zealand region.

The waters around New Zealand are a global hotspot of biodiversity for deep-water corals; approximately one sixth of the known deep-water coral species of the world have been recorded in the region. Deep-water corals are vulnerable to climate-related stressors and from the damaging effects of commercial fisheries. Current protection measures do not account for the vulnerability of deep-water corals to future climatic conditions, which are predicted to alter the distribution of suitable habitat for them. Using recently developed habitat suitability models for 12 taxa of deep-water corals fitted to current and future seafloor environmental conditions (under different future climatic conditions: SSP2 - 4.5 and SSP3 - 7.0) we explore possible levels of spatial protection using the decision-support tool Zonation. Specifically, we assess the impact of bottom trawling on predictions of current distributions of deep-water corals, and then assess the effectiveness of possible protection for deep-water corals, while accounting for habitat refugia under future climatic conditions. The cumulative impact of bottom trawling was predicted to impact all taxa, but particularly the reef-forming corals. Core areas of suitable habitat were predicted to decrease under future climatic conditions for many taxa. We found that designing protection using current day predictions alone, having accounted for the impacts of historic fishing impacts, was unlikely to provide adequate conservation for deep water-corals under future climate change. Accounting for future distributions in spatial planning identified areas which may provide climate refugia whilst still providing efficient protection for current distributions. These gains in conservation value may be particularly important given the predicted reduction in suitable habitat for deep-water corals due to bottom fishing and climate change. Finally, the possible impact that protection measures may have on deep-water fisheries was assessed using a measure of current fishing value (kg km-2 fish) and future fishing value (predicted under future climate change scenarios).

Cold-water coral diversity along the continental shelf margin of northwestern South China Sea.

Scleractinian cold-water corals (CWCs) are one of the most important habitat engineers of the deep sea. Although the South China Sea (SCS) abuts the biodiversity center of scleractinian CWCs in the western Pacific, only a few sporadic records are available. We discovered new CWC sites by means of trawl sampling and video observation along the continental shelf of the northwestern SCS. All trawled scleractinian CWC specimens were identified to species level according to skeleton morphology and structure. The living CWCs and associated fauna recorded in the video were -identified to a higher level of classification. Scleractinian corals were identified to genus level, while non-scleractinian CWCs were identified to family level and given general names such as gorgonian corals, bamboo corals and black corals. Associated benthic dwellers were divided into major categories. A total of 28 scleractinian CWC species were identified to 7 families, 15 genera, and 1 additional subgenus. Among them, 13 species were colonial, including important habitat-forming species in the genera Eguchipsammia, Dendrophyllia and Cladopsammia. Non-scleractinian CWCs were identified to 7 families, including 4 families gorgonian corals, 1 family bamboo corals, and 2 families black corals. Gorgonian corals were the most abundant non-scleractinian CWCs in this region. Meanwhile, starfish, sea anemones, fish, gastropods, echinoderms, and other associated benthic fauna were recorded in the CWC habitats, with starfish belonging to the order Brisingida being most common. New scleractinian CWC assemblages were discovered along the continental seabed mounds in the northwestern SCS. This study highlights the remarkable diversity of cold-water scleractinian corals in the whole SCS, and shows the potential widespread distribution and conservation prospect of CWC habitats in this region.

Predicting the distribution and abundance of abandoned, lost or discarded fishing gear (ALDFG) in the deep sea of the Azores (North Atlantic).

Abandoned, lost, or discarded fishing gear (ALDFG), represents a significant percentage of the global plastic pollution, currently considered one of the major sources from sea-based activities. However, there is still limited understanding of the quantities of ALDFG present on the seafloor and their impacts. In this study, data on the presence of ALDFG was obtained from a large archive of seafloor video footage (351 dives) collected by different imaging platforms in the Azores region over 15 years (2006-2020). Most ALDFG items observed in the images relate to the local bottom longline fishery operating in the region, and include longlines but also anchors, weights, cables and buoys. A generalized additive mixed model (GAMM) was used to predict the distribution and abundance of ALDFG over the seafloor within the limits of the Azores Exclusive Economic Zone (EEZ) using a suite of environmental and anthropogenic variables. We estimated an average of 113 ± 310 items km-2 (597 ± 756 per km-2 above 1000 m depth), which could imply that over 20 million ALDFG items are present on the deep seafloor of the Azores EEZ. The resulting model identified potential hotspots of ALDFG along the seabed, some of them located over sensitive benthic habitats, such as specific seamounts. In addition, the interactions between ALDFG and benthic organisms were also analysed. Numerous entanglements were observed with several species of large anthozoans and sponges. The use of predictive distribution modelling for ALDFG should be regarded as a useful tool to support ecosystem-based management, which can provide indirect information about fishing pressure and allow the identification of potential high-risk areas. Additional knowledge about the sources, amounts, fates and impacts of ALDFG will be key to address the global issue of plastic pollution and the effects of fishing on marine ecosystems.

A health survey of the reef forming scleractinian cold-water corals Lophelia pertusa and Madrepora oculata in a remote submarine canyon on the European continental margin, NE Atlantic.

Monitoring of cold-water corals (CWCs) for pathogens and diseases is limited due to the environment, protected nature of the corals and their habitat and as well as the challenging and sampling effort required. It is recognised that environmental factors such as temperature and pH can expedite the ability of pathogens to cause diseases in cold-water corals therefore the characterisation of pathogen diversity, prevalence and associated pathologies is essential. The present study combined histology and polymerase chain reaction (PCR) diagnostic techniques to screen for two significant pathogen groups (bacteria of the genus Vibrio and the protozoan Haplosporidia) in the dominant NE Atlantic deep-water framework corals Lophelia pertusa (13 colonies) and Madrepora oculata (2 colonies) at three sampling locations (canyon head, south branch and the flank) in the Porcupine Bank Canyon (PBC), NE Atlantic. One M. oculata colony and four L. pertusa colonies were collected from both the canyon flank and the south branch whilst five L. pertusa colonies were collected from the canyon head. No pathogens were detected in the M. oculata samples. Neither histology nor PCR detected Vibrio spp. in L. pertusa, although Illumina technology used in this study to profile the CWCs microbiome, detected V. shilonii (0.03%) in a single L. pertusa individual, from the canyon head, that had also been screened in this study. A macroborer was observed at a prevalence of 0.07% at the canyon head only. Rickettsiales-like organisms (RLOs) were visualised with an overall prevalence of 40% and with a low intensity of 1 to 4 (RLO) colonies per individual polyp by histology. L. pertusa from the PBC canyon head had an RLO prevalence of 13.3% with the highest detection of 26.7% recorded in the south branch corals. Similarly, unidentified cells observed in L. pertusa from the south branch (20%) were more common than those observed in L. pertusa from the canyon head (6.7%). No RLOs or unidentified cells were observed in corals from the flank. Mean particulate organic matter concentration is highest in the south branch (2,612 µg l-1) followed by the canyon head (1,065 µg l-1) and lowest at the canyon flank (494 µg l-1). Although the route of pathogen entry and the impact of RLO infection on L. pertusa is unclear, particulate availability and the feeding strategies employed by the scleractinian corals may be influencing their exposure to pathogens. The absence of a pathogen in M. oculata may be attributed to the smaller number of colonies screened or the narrower diet in M. oculata compared to the unrestricted diet exhibited in L. pertusa, if ingestion is a route of entry for pathogen groups. The findings of this study also shed some light on how environmental conditions experienced by deep sea organisms and their life strategies may be limiting pathogen diversity and prevalence.

Seasonal and diel variations in the vertical distribution, composition, abundance and biomass of zooplankton in a deep Chilean Patagonian Fjord.

Comau Fjord is a stratified Chilean Patagonian Fjord characterized by a shallow brackish surface layer and a >400 m layer of aragonite-depleted subsurface waters. Despite the energetic burden of low aragonite saturation levels to calcification, Comau Fjord harbours dense populations of cold-water corals (CWC). While this paradox has been attributed to a rich supply of zooplankton, supporting abundance and biomass data are so far lacking. In this study, we investigated the seasonal and diel changes of the zooplankton community over the entire water column. We used a Nansen net (100 µm mesh) to take stratified vertical hauls between the surface and the bottom (0-50-100-200-300-400-450 m). Samples were scanned with a ZooScan, and abundance, biovolume and biomass were determined for 41 taxa identified on the web-based platform EcoTaxa 2.0. Zooplankton biomass was the highest in summer (209 g dry mass m-2) and the lowest in winter (61 g dry mass m-2). Abundance, however, peaked in spring, suggesting a close correspondence between reproduction and phytoplankton spring blooms (Chl a max. 50.86 mg m-3, 3 m depth). Overall, copepods were the most important group of the total zooplankton community, both in abundance (64-81%) and biovolume (20-70%) followed by mysids and chaetognaths (in terms of biovolume and biomass), and nauplii and Appendicularia (in terms of abundance). Throughout the year, diel changes in the vertical distribution of biomass were found with a daytime maximum in the 100-200 m depth layer and a nighttime maximum in surface waters (0-50 m), associated with the diel vertical migration of the calanoid copepod family Metridinidae. Diel differences in integrated zooplankton abundance, biovolume and biomass were probably due to a high zooplankton patchiness driven by biological processes (e.g., diel vertical migration or predation avoidance), and oceanographic processes (estuarine circulation, tidal mixing or water column stratification). Those factors are considered to be the main drivers of the zooplankton vertical distribution in Comau Fjord.

Microbiomes and Obligate Symbiosis of Deep-Sea Animals.

Microbial communities associated with deep-sea animals are critical to the establishment of novel biological communities in unusual environments. Over the past few decades, rapid exploration of the deep sea has enabled the discovery of novel microbial communities, some of which form symbiotic relationships with animal hosts. Symbiosis in the deep sea changes host physiology, behavior, ecology, and evolution over time and space. Symbiont diversity within a host is often aligned with diverse metabolic pathways that broaden the environmental niche for the animal host. In this review, we focus on microbiomes and obligate symbionts found in different deep-sea habitats and how they facilitate survival of the organisms that live in these environments. In addition, we discuss factors that govern microbiome diversity, host specificity, and biogeography in the deep sea. Finally, we highlight the current limitations of microbiome research and draw a road map for future directions to advance our knowledge of microbiomes in the deep sea.

Dense cold-water coral garden of Paragorgia johnsoni suggests the importance of the Mid-Atlantic Ridge for deep-sea biodiversity.

Mid-ocean ridges generate a myriad of physical oceanographic processes that favor the supply of food and nutrients to suspension- and filter-feeding organisms, such as cold-water corals and deep-sea sponges. However, the pioneering work conducted along the Mid-Atlantic Ridge failed to report the presence of large and dense living coral reefs, coral gardens, or sponge aggregations. Here, we describe the densest, near-natural, and novel octocoral garden composed of large red and white colonies of Paragorgia johnsoni Gray, 1862 discovered at 545-595 m depth on the slopes of the Mid-Atlantic Ridge, in the Azores region. This newly discovered octocoral garden is a good candidate for protection since it fits many of the FAO criteria that define what constitutes a Vulnerable Marine Ecosystem. The observations described here corroborate the existence of a close relationship between the octocoral structure and the ambient currents on ridge-like topographies, providing new insights into the functioning of mid-ocean ridges' ecosystems. The ubiquitous presence of biogenic and geological topographies associated with mid-ocean ridges, which could act as climate refugia, suggests their global importance for deep-sea biodiversity. A better understanding of the processes involved is, therefore, required. Our observations may inspire future deep-sea research initiatives to narrow existing knowledge gaps of biophysical connections with benthic fauna at small spatial scales along mid-ocean ridges.

The modeled distribution of corals and sponges surrounding the Salas y Gómez and Nazca ridges with implications for high seas conservation.

The Salas y Gómez and Nazca ridges are two adjacent seamount chains off the west coast of South America that collectively contain more than 110 seamounts. The ridges support an exceptionally rich diversity of benthic and pelagic communities, with the highest level of endemism found in any marine environment. Despite some historical fishing in the region, the seamounts are relatively pristine and represent an excellent conservation opportunity to protect a global biodiversity hotspot before it is degraded. One obstacle to effective spatial management of the ridges is the scarcity of direct observations in deeper waters throughout the region and an accompanying understanding of the distribution of key taxa. Species distribution models are increasingly used tools to quantify the distributions of species in data-poor environments. Here, we focused on modeling the distribution of demosponges, glass sponges, and stony corals, three foundation taxa that support large assemblages of associated fauna through the creation of complex habitat structures. Models were constructed at a 1 km2 resolution using presence and pseudoabsence data, dissolved oxygen, nitrate, phosphate, silicate, aragonite saturation state, and several measures of seafloor topography. Highly suitable habitat for each taxa was predicted to occur throughout the Salas y Gómez and Nazca ridges, with the most suitable habitat occurring in small patches on large terrain features such as seamounts, guyots, ridges, and escarpments. Determining the spatial distribution of these three taxa is a critical first step towards supporting the improved spatial management of the region. While the total area of highly suitable habitat was small, our results showed that nearly all of the seamounts in this region provide suitable habitats for deep-water corals and sponges and should therefore be protected from exploitation using the best available conservation measures.

Living benthic foraminifera from cold-water coral ecosystems in the eastern Alboran Sea, Western Mediterranean.

Benthic foraminifera (protists with biomineralized tests) coupled with geochemical proxies are used for the first time to characterize present oceanographic conditions occurring in cold-water coral ecosystems (CWC) in the eastern Alboran Sea (Brittlestar Ridge and Cablier Mound), western Mediterranean Sea. Quantitative data on living (stained) benthic foraminifera from 5 box cores retrieved during the MD194 cruise on the RV Marion Dufresne reveal that these organisms are more diverse in presence of corals, where more numerous ecological niches occur than they are in pelagic adjacent sediments. These data confirm that CWC can be considered as "diversity hotspots" also for benthic foraminifera. Geochemical characterization shows that these sediments contain relatively fresh (labile) organic matter but also a reworked refractory component. In particular, the total organic carbon and the δ 13Corg values suggest that some of the organic matter may be a mixture of marine and reworked particulate organic matter, compared to typical values from temperate phytoplankton. The δ 15N of the organic fraction suggests that important atmospheric N2-fixation and degradation processes occur in the region. Finally, our results show that a more effective advection of freshly exported particulate organic matter from the surface waters occur at the mound top rather than at the mound base or off-mound allowing some coral colonies to survive on the top of mounds in this region. The mud layer covering the coral rubble debris may suggest that the Brittlestar Ridge is today exposed to siltation preventing the growth of corals at the mound base or off-mound.

Embryo and larval biology of the deep-sea octocoral Dentomuricea aff. meteor under different temperature regimes.

Deep-sea octocorals are common habitat-formers in deep-sea ecosystems, however, our knowledge on their early life history stages is extremely limited. The present study focuses on the early life history of the species Dentomuricea aff. meteor, a common deep-sea octocoral in the Azores. The objective was to describe the embryo and larval biology of the target species under two temperature regimes, corresponding to the minimum and maximum temperatures in its natural environment during the spawning season. At temperature of 13 ±0.5 °C, embryos of the species reached the planula stage after 96h and displayed a median survival of 11 days. Planulae displayed swimming only after stimulation, swimming speed was 0.24 ±0.16 mm s-1 and increased slightly but significantly with time. Under a higher temperature (15 °C ±0.5 °C) embryos reached the planula stage 24 h earlier (after 72 h), displayed a median survival of 16 days and had significantly higher swimming speed (0.3 ±0.27 mm s-1). Although the differences in survival were not statistically significant, our results highlight how small changes in temperature can affect embryo and larval characteristics with potential cascading effects in larval dispersal and success. In both temperatures, settlement rates were low and metamorphosis occurred even without settlement. Such information is rarely available for deep-sea corals, although essential to achieve a better understanding of dispersal, connectivity and biogeographical patterns of benthic species.

Seafloor litter sorting in different domains of Cap de Creus continental shelf and submarine canyon (NW Mediterranean Sea).

We analyzed litter occurrence in 68 underwater video transects performed on the middle/outer continental shelf and submarine canyon off Cap de Creus (NW Mediterranean), an area recently declared Site of Community Importance (SCI). Low densities of urban litter were registered on the shelf (7.2 items ha-1), increasing in abundance towards the deepest part of the submarine canyon, with 188 items ha-1 below 1000 m depth. We hypothesize that the strong bottom currents that recurrently affect this area efficiently move litter objects from the shelf towards the deep. Of all litter items, approximately 50% had a fishing-related origin, mostly longlines entangled on rocks in the canyon head and discarded trawl nets in deeper areas. Over 10% of cold-water colonies observed had longlines entangled, indicating the harmful effects of such practices over benthic habitats. These results should be considered when designing mitigation measures to reduce litter pollution in Cap de Creus SCI.

Rapid deep ocean deoxygenation and acidification threaten life on Northeast Pacific seamounts.

Anthropogenic climate change is causing our oceans to lose oxygen and become more acidic at an unprecedented rate, threatening marine ecosystems and their associated animals. In deep-sea environments, where conditions have typically changed over geological timescales, the associated animals, adapted to these stable conditions, are expected to be highly vulnerable to any change or direct human impact. Our study coalesces one of the longest deep-sea observational oceanographic time series, reaching back to the 1960s, with a modern visual survey that characterizes almost two vertical kilometers of benthic seamount ecosystems. Based on our new and rigorous analysis of the Line P oceanographic monitoring data, the upper 3,000 m of the Northeast Pacific (NEP) has lost 15% of its oxygen in the last 60 years. Over that time, the oxygen minimum zone (OMZ), ranging between approximately 480 and 1,700 m, has expanded at a rate of 3.0 ± 0.7 m/year (due to deepening at the bottom). Additionally, carbonate saturation horizons above the OMZ have been shoaling at a rate of 1-2 m/year since the 1980s. Based on our visual surveys of four NEP seamounts, these deep-sea features support ecologically important taxa typified by long life spans, slow growth rates, and limited mobility, including habitat-forming cold water corals and sponges, echinoderms, and fish. By examining the changing conditions within the narrow realized bathymetric niches for a subset of vulnerable populations, we resolve chemical trends that are rapid in comparison to the life span of the taxa and detrimental to their survival. If these trends continue as they have over the last three to six decades, they threaten to diminish regional seamount ecosystem diversity and cause local extinctions. This study highlights the importance of mitigating direct human impacts as species continue to suffer environmental changes beyond our immediate control.

Constraining the formation of authigenic carbonates in a seepage-affected cold-water coral mound by lipid biomarkers.

Cold-water coral (CWC) mounds are build-ups comprised of coral-dominated intervals alternating with a mixed carbonate-siliciclastic matrix. At some locations, CWC mounds are influenced by methane seepage, but the impact of methane on CWC mounds is poorly understood. To constrain the potential impact of methane on CWC mound growth, lipid biomarker investigations were combined with mineralogical and petrographic analyses to investigate the anaerobic oxidation of methane (AOM) and authigenic carbonate formation in sediment from a seep-affected CWC mound in the Gulf of Cadiz. The occurrence of AOM was confirmed by characteristic lipids found within a semi-lithified zone (SLZ) consisting of authigenic aragonite, high-magnesium calcite and calcium-excess dolomite. The formation of high-Mg calcite is attributed to AOM, acting as a lithifying agent. Aragonite is only a minor phase. Ca-excess dolomite in the SLZ and upper parts may be formed by organoclastic sulphate reduction, favouring precipitation by increased alkalinity. The AOM biomarkers in the SLZ include isoprenoid-based archaeal membrane lipids, such as abundant glycerol dibiphytanyl glycerol tetraethers (GDGTs) dominated by GDGT-2. The δ13 C values of GDGT-2, measured as ether-cleaved monocyclic biphytanes, are as low as -100‰ versus V-PDB. Further, bacterial dialkyl glycerol diethers with two anteiso-C15 alkyl chains and δ13 C values of -81‰ are interpreted as biomarkers of sulphate-reducing bacteria. The lipid biomarker signatures and mineralogical patterns suggest that anaerobic methane-oxidizing archaea of the ANME-1 group thrived in the subsurface at times of slow and diffusive methane seepage. Petrographic analyses revealed that the SLZ was exhumed at some point (e.g. signs of bioerosion of the semi-lithified sediment), providing a hard substrate for CWC larval settlement. In addition, this work reveals that AOM-induced semi-lithification likely played a role in mound stabilization. Lipid biomarker analysis proves to be a powerful tool to disentangle early diagenetic processes induced by microbial metabolisms.

Climate-induced changes in the suitable habitat of cold-water corals and commercially important deep-sea fishes in the North Atlantic.

The deep sea plays a critical role in global climate regulation through uptake and storage of heat and carbon dioxide. However, this regulating service causes warming, acidification and deoxygenation of deep waters, leading to decreased food availability at the seafloor. These changes and their projections are likely to affect productivity, biodiversity and distributions of deep-sea fauna, thereby compromising key ecosystem services. Understanding how climate change can lead to shifts in deep-sea species distributions is critically important in developing management measures. We used environmental niche modelling along with the best available species occurrence data and environmental parameters to model habitat suitability for key cold-water coral and commercially important deep-sea fish species under present-day (1951-2000) environmental conditions and to project changes under severe, high emissions future (2081-2100) climate projections (RCP8.5 scenario) for the North Atlantic Ocean. Our models projected a decrease of 28%-100% in suitable habitat for cold-water corals and a shift in suitable habitat for deep-sea fishes of 2.0°-9.9° towards higher latitudes. The largest reductions in suitable habitat were projected for the scleractinian coral Lophelia pertusa and the octocoral Paragorgia arborea, with declines of at least 79% and 99% respectively. We projected the expansion of suitable habitat by 2100 only for the fishes Helicolenus dactylopterus and Sebastes mentella (20%-30%), mostly through northern latitudinal range expansion. Our results projected limited climate refugia locations in the North Atlantic by 2100 for scleractinian corals (30%-42% of present-day suitable habitat), even smaller refugia locations for the octocorals Acanella arbuscula and Acanthogorgia armata (6%-14%), and almost no refugia for P. arborea. Our results emphasize the need to understand how anticipated climate change will affect the distribution of deep-sea species including commercially important fishes and foundation species, and highlight the importance of identifying and preserving climate refugia for a range of area-based planning and management tools.

Middle Holocene expansion of Pacific Deep Water into the Southern Ocean.

The Southern Ocean is a key region for the overturning and mixing of water masses within the global ocean circulation system. Because Southern Ocean dynamics are influenced by the Southern Hemisphere westerly winds (SWW), changes in the westerly wind forcing could significantly affect the circulation and mixing of water masses in this important location. While changes in SWW forcing during the Holocene (i.e., the last ∼11,700 y) have been documented, evidence of the oceanic response to these changes is equivocal. Here we use the neodymium (Nd) isotopic composition of absolute-dated cold-water coral skeletons to show that there have been distinct changes in the chemistry of the Southern Ocean water column during the Holocene. Our results reveal a pronounced Middle Holocene excursion (peaking ∼7,000-6,000 y before present), at the depth level presently occupied by Upper Circumpolar Deep Water (UCDW), toward Nd isotope values more typical of Pacific waters. We suggest that poleward-reduced SWW forcing during the Middle Holocene led to both reduced Southern Ocean deep mixing and enhanced influx of Pacific Deep Water into UCDW, inducing a water mass structure that was significantly different from today. Poleward SWW intensification during the Late Holocene could then have reinforced deep mixing along and across density surfaces, thus enhancing the release of accumulated CO2 to the atmosphere.

In situ growth and bioerosion rates of Lophelia pertusa in a Norwegian fjord and open shelf cold-water coral habitat.

Coral reef resilience depends on the balance between carbonate precipitation, leading to reef growth, and carbonate degradation, for example, through bioerosion. Changes in environmental conditions are likely to affect the two processes differently, thereby shifting the balance between reef growth and degradation. In cold-water corals estimates of accretion-erosion processes in their natural habitat are scarce and solely live coral growth rates were studied with regard to future environmental changes in the laboratory so far, limiting our ability to assess the potential of cold-water coral reef ecosystems to cope with environmental changes. In the present study, growth rates of the two predominant colour morphotypes of live Lophelia pertusa as well as bioerosion rates of dead coral framework were assessed in different environmental settings in Norwegian cold-water coral reefs in a 1-year in situ experiment. Net growth (in weight gain and linear extension) of live L. pertusa was in the lower range of previous estimates and did not significantly differ between inshore (fjord) and offshore (open shelf) habitats. However, slightly higher net growth rates were obtained inshore. Bioerosion rates were significantly higher on-reef in the fjord compared to off-reef deployments in- and offshore. Besides, on-reef coral fragments yielded a broader range of individual growth and bioerosion rates, indicating higher turnover in live reef structures than off-reef with regard to accretion-bioerosion processes. Moreover, if the higher variation in growth rates represents a greater variance in (genetic) adaptations to natural environmental variability in the fjord, inshore reefs could possibly benefit under future ocean change compared to offshore reefs. Although not significantly different due to high variances between replicates, growth rates of orange branches were consistently higher at all sites, while mortality was statistically significantly lower, potentially indicating higher stress-resistance than the less pigmented white phenotype. Comparing the here measured rates of net accretion of live corals (regardless of colour morphotype) with net erosion of dead coral framework gives a first estimate of the dimensions of both processes in natural cold-water coral habitats, indicating that calcium carbonate loss through bioerosion amounts to one fifth to one sixth of the production rates by coral calcification (disregarding accretion processes of other organisms and proportion of live and dead coral framework in a reef). With regard to likely accelerating bioerosion and diminishing growth rates of corals under ocean acidification, the balance of reef accretion and degradation may be shifted towards higher biogenic dissolution in the future.