Expanding our view of the cold-water coral niche and accounting of the ecosystem services of the reef habitat.

Coral reefs are iconic ecosystems that support diverse, productive communities in both shallow and deep waters. However, our incomplete knowledge of cold-water coral (CWC) niche space limits our understanding of their distribution and precludes a complete accounting of the ecosystem services they provide. Here, we present the results of recent surveys of the CWC mound province on the Blake Plateau off the U.S. east coast, an area of intense human activity including fisheries and naval operations, and potentially energy and mineral extraction. At one site, CWC mounds are arranged in lines that total over 150 km in length, making this one of the largest reef complexes discovered in the deep ocean. This site experiences rapid and extreme shifts in temperature between 4.3 and 10.7 °C, and currents approaching 1 m s-1. Carbon is transported to depth by mesopelagic micronekton and nutrient cycling on the reef results in some of the highest nitrate concentrations recorded in the region. Predictive models reveal expanded areas of highly suitable habitat that currently remain unexplored. Multidisciplinary exploration of this new site has expanded understanding of the cold-water coral niche, improved our accounting of the ecosystem services of the reef habitat, and emphasizes the importance of properly managing these systems.

Trophic ecology of Angolan cold-water coral reefs (SE Atlantic) based on stable isotope analyses.

Cold-water coral (CWC) reefs of the Angolan margin (SE Atlantic) are dominated by Desmophyllum pertusum and support a diverse community of associated fauna, despite hypoxic conditions. In this study, we use carbon and nitrogen stable isotope analyses (δ13C and δ15N) to decipher the trophic network of this relatively unknown CWC province. Although fresh phytodetritus is available to the reef, δ15N signatures indicate that CWCs (12.90 ± 1.00 ‰) sit two trophic levels above Suspended Particulate Organic Matter (SPOM) (4.23 ± 1.64 ‰) suggesting that CWCs are highly reliant on an intermediate food source, which may be zooplankton. Echinoderms and the polychaete Eunice norvegica occupy the same trophic guild, with high δ13C signatures (-14.00 ± 1.08 ‰) pointing to a predatory feeding behavior on CWCs and sponges, although detrital feeding on 13C enriched particles might also be important for this group. Sponges presented the highest δ15N values (20.20 ± 1.87 ‰), which could be due to the role of the sponge holobiont and bacterial food in driving intense nitrogen cycling processes in sponges' tissue, helping to cope with the hypoxic conditions of the reef. Our study provides first insights to understand trophic interactions of CWC reefs under low-oxygen conditions.

On the paradox of thriving cold-water coral reefs in the food-limited deep sea.

The deep sea is amongst the most food-limited habitats on Earth, as only a small fraction (<4%) of the surface primary production is exported below 200 m water depth. Here, cold-water coral (CWC) reefs form oases of life: their biodiversity compares with tropical coral reefs, their biomass and metabolic activity exceed other deep-sea ecosystems by far. We critically assess the paradox of thriving CWC reefs in the food-limited deep sea, by reviewing the literature and open-access data on CWC habitats. This review shows firstly that CWCs typically occur in areas where the food supply is not constantly low, but undergoes pronounced temporal variation. High currents, downwelling and/or vertically migrating zooplankton temporally boost the export of surface organic matter to the seabed, creating 'feast' conditions, interspersed with 'famine' periods during the non-productive season. Secondly, CWCs, particularly the most common reef-builder Desmophyllum pertusum (formerly known as Lophelia pertusa), are well adapted to these fluctuations in food availability. Laboratory and in situ measurements revealed their dietary flexibility, tissue reserves, and temporal variation in growth and energy allocation. Thirdly, the high structural and functional diversity of CWC reefs increases resource retention: acting as giant filters and sustaining complex food webs with diverse recycling pathways, the reefs optimise resource gains over losses. Anthropogenic pressures, including climate change and ocean acidification, threaten this fragile equilibrium through decreased resource supply, increased energy costs, and dissolution of the calcium-carbonate reef framework. Based on this review, we suggest additional criteria to judge the health of CWC reefs and their chance to persist in the future.

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.

Biodiversity, environmental drivers, and sustainability of the global deep-sea sponge microbiome.

In the deep ocean symbioses between microbes and invertebrates are emerging as key drivers of ecosystem health and services. We present a large-scale analysis of microbial diversity in deep-sea sponges (Porifera) from scales of sponge individuals to ocean basins, covering 52 locations, 1077 host individuals translating into 169 sponge species (including understudied glass sponges), and 469 reference samples, collected anew during 21 ship-based expeditions. We demonstrate the impacts of the sponge microbial abundance status, geographic distance, sponge phylogeny, and the physical-biogeochemical environment as drivers of microbiome composition, in descending order of relevance. Our study further discloses that fundamental concepts of sponge microbiology apply robustly to sponges from the deep-sea across distances of >10,000 km. Deep-sea sponge microbiomes are less complex, yet more heterogeneous, than their shallow-water counterparts. Our analysis underscores the uniqueness of each deep-sea sponge ground based on which we provide critical knowledge for conservation of these vulnerable ecosystems.

Bacterial precursors and unsaturated long-chain fatty acids are biomarkers of North-Atlantic deep-sea demosponges.

Sponges produce distinct fatty acids (FAs) that (potentially) can be used as chemotaxonomic and ecological biomarkers to study endosymbiont-host interactions and the functional ecology of sponges. Here, we present FA profiles of five common habitat-building deep-sea sponges (class Demospongiae, order Tetractinellida), which are classified as high microbial abundance (HMA) species. Geodia hentscheli, G. parva, G. atlantica, G. barretti, and Stelletta rhaphidiophora were collected from boreal and Arctic sponge grounds in the North-Atlantic Ocean. Bacterial FAs dominated in all five species and particularly isomeric mixtures of mid-chain branched FAs (MBFAs, 8- and 9-Me-C16:0 and 10- and 11-Me-C18:0) were found in high abundance (together ≥ 20% of total FAs) aside more common bacterial markers. In addition, the sponges produced long-chain linear, mid- and a(i)-branched unsaturated FAs (LCFAs) with a chain length of 24‒28 C atoms and had predominantly the typical Δ5,9 unsaturation, although the Δ9,19 and (yet undescribed) Δ11,21 unsaturations were also identified. G. parva and S. rhaphidiophora each produced distinct LCFAs, while G. atlantica, G. barretti, and G. hentscheli produced similar LCFAs, but in different ratios. The different bacterial precursors varied in carbon isotopic composition (δ13C), with MBFAs being more enriched compared to other bacterial (linear and a(i)-branched) FAs. We propose biosynthetic pathways for different LCFAs from their bacterial precursors, that are consistent with small isotopic differences found in LCFAs. Indeed, FA profiles of deep-sea sponges can serve as chemotaxonomic markers and support the concept that sponges acquire building blocks from their endosymbiotic bacteria.

Internal wave turbulence at a biologically rich Mid-Atlantic seamount.

The turbulence regime near the crest of a biologically rich seamount of the Mid-Atlantic Ridge southwest of the Azores was registered in high spatial and temporal resolution. Internal tides and their higher harmonics dominate the internal wave motions, producing considerable shear-induced turbulent mixing in layers of 10-50 m thickness. This interior mixing of about 100 times open-ocean interior values is observed both at a high-resolution temperature sensor mooring-site at the crest, 770 m water depth being nearly 400 m below the top of the seamount, and a CTD-yoyo site at the slope off the crest 400 m horizontally away, 880 m water depth. Only at the mooring site, additionally two times higher turbulence is observed near the bottom, associated with highly non-linear wave breaking. The highest abundance of epifauna, notably sponges, are observed just below the crest and 100 m down the eastern slope (700-800 m) in a cross-ridge video-camera transect. This sponge belt is located in a water layer of depressed oxygen levels (saturation 63±2%) with a local minimum centered around 700 m. Turbulent mixing supplies oxygen to this region from above and below and is expected to mix nutrients away from this biodegraded layer towards the depth of highest abundance of macrofauna.