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Until recently, benthic habitats dominated by deep-sea corals (DSC) appeared to be less extensive on the slope of the Gulf of Mexico (GOM) than in the northeast Atlantic Ocean or off the southeastern US. There are relatively few bioherms (i.e., coral-built mounds) in the northern GOM, and most DSCs are attached to existing hard substrata (e.g., authigenically formed carbonate). The primary structure-forming, DSC in the GOM is Lophelia pertusa , but structure is also provided by other living and dead scleractinians, antipatharians (black corals), octocorals (gorgonians, soft corals), hydrocorals and sponges, as well as abundant rocky substrata. The best development of DSCs in the GOM was previously documented within Viosca Knoll oil and gas lease blocks 826 and 862/906 (north-central GOM) and on the Campeche Bank (southern GOM in Mexican waters). This paper documents extensive deep reef ecosystems composed of DSC and rocky hard-bottom recently surveyed on the West Florida Slope (WFS, eastern GOM) during six research cruises (2008–2012). Using multibeam sonar, CTD casts, and video from underwater vehicles, we describe the physical and oceanographic characteristics of these deep reefs and provide size or area estimates of deep coral and hardground habitats. The multibeam sonar analyses revealed hundreds of mounds and ridges, some of which were subsequently surveyed using underwater vehicles. Mounds and ridges in <525 m depths were usually capped with living coral colonies, dominated by L . pertusa . An extensive rocky scarp, running roughly north-south for at least 229 km, supported lower abundances of scleractinian corals than the mounds and ridges, despite an abundance of settlement substrata. Areal comparisons suggested that the WFS may exceed other parts of the GOM slope in extent of living deep coral coverage and other deep-reef habitat (dead coral and rock). The complex WFS region warrants additional studies to better understand the influences of oceanography and geology on the occurrence of DSC and associated organisms. Protection measures are being considered to ensure the long-term integrity of this diverse ecosystem. [ABSTRACT FROM AUTHOR]

Whale falls are considered important habitats contributing to biodiversity, evolutionary novelty and connectivity of deep-sea environments. The organic input of a whale carcass increases the community standing stock and changes the species composition in comparison with the surrounding benthic infauna. However, endofauna living inside the bone matrix seems to contribute to the overall whale-fall biodiversity. Benthic infauna is distributed in bathymetric ranges, but the contribution of endofauna from whale bones in different depth regions is still not accessed. We investigated endofauna composition of whale vertebrae of a humpback whale (Megaptera novaeangliae) in two distinct depths, bathyal (1500 m, stations SP1500 and ES1500) and abyssal (3300 m, stations SP3300, RJ3300 and ES3300), deployed in the SW Atlantic Ocean. Even though each bone represented the same energetic input, a higher density of endofauna was found at 1500 m while diversity was higher at 3300 m. The whale-fall endofaunal assemblages were distinct according to the depth zones. SP3300 site was distinct from bones placed at 1500 m depth, while the assemblages at RJ3300 and ES3300 sites consisted of a mixture between both depths. This pattern seems to be related to the water currents of the region, once SP3300 was under influence of the Antarctic Bottom Water (AABW), while SP1500 and ES1500 were dominated by the North Atlantic Deep Water (NADW). • This paper investigates whalebone communities of bathyal and abyssal sites of the SW Atlantic Ocean. • Whalebone endofaunal community was different between bathyal and abyssal sites. • Endofauna of whale bones is more abundant in bathyal sites, but more diverse in abyssal ones. • Bone communities are highly influenced by deep-water current circulation. [ABSTRACT FROM AUTHOR]

Three hundred and twenty-nine species of micronektonic fishes were identified in 1040 midwater trawl collections taken between 1963 and 1974 from the North and South Atlantic, the Mediterranean, the Caribbean and the Gulf of Mexico. The target of most tows was the deep scattering layer, and consequently the dominant species in the material were those that were concentrated in the layer. The results only generally confirm the 11 Atlantic mesopelagic ecoregions previously recognized. The geometric mean of the proportion of joint occurrences (GMPJO) of species with tows within each ecoregion was used to characterize the faunas of the ecoregions. The ecoregion affinities of fishes were compared to those of decapod shrimp in the same collections. The fish and shrimp faunas of ecoregions could be distinguished by GMPJO values, but the ranges of species favoring each ecoregion varied widely in extent and did not conform well to ecoregion boundaries or features of circulation. This suggests that co-occurring species respond differently to the physical properties and resulting biological factors defining mesopelagic ecoregions. [ABSTRACT FROM AUTHOR]

High habitat diversity was observed on the Seco de los Olivos Seamount (SW Mediterranean Sea), a Site of Community Importance belonging to the Spanish marine Natura 2000 Network. Thirteen epibenthic habitats were identified by analysing 55 Remotely Operated Vehicle (ROV) transects from 76 m to 700 m depth and derived data from multibeam bathymetry and high resolution seismic profiles. Habitat identification was based on a combination of assemblages of habitat-forming species and the environmental characteristics supporting their distribution. Depth and slope were identified as the main significant factors structuring epibenthic assemblages. The high diversity and patchiness of habitats found on the Seco de los Olivos Seamount can be explained by the high environmental variability resulting from its wide geomorphologic diversity, where flat summits, steep flanks, rocky outcrops and sedimentary moats are combined. The distribution of benthic habitats at this seamount is likely a combination of suitable ecological conditions, local recruitment, feeding strategies and attachment mechanisms. Knowledge on the occurrence of habitats in areas of natural importance is crucial to species and habitats conservation and to develop proper monitoring and management programs aimed at fulfilling European regulation requirements. [ABSTRACT FROM AUTHOR]

The Rio Grande Rise (RGR) is a large and geomorphologically complex structure of the deep SW Atlantic Ocean. In 2013, the 600−1200 m deep plateau of the most prominent topographic component of the RGR (named Alpha) was explored during two dives of the manned submersible Shinkai 6500 (30°22′15′′S − 36°02′02′′W and 31°05′58′′S − 34°02′40′′W). Video profiles recorded during these dives were analyzed for description of benthopelagic megafauna (fish and crustaceans) assemblages, and quantitative assessment of structuring factors (depth, topography and habitat types). Fishes represented over 92% (462) of all benthopelagic megafauna, divided into 11 orders and 17 families. Over half of fish records were Macrouridae, Synaphobranchidae and Chaunacidae. Megafauna abundance varied at different spatial scales, being higher in shallower habitats (~600 m) dominated by branched suspension feeders (mostly sponges and cnidarians). Beta-diversity and community structure were related to habitat diversity. Because the RGR is vast and may comprise numerous distinctive habitats associated with depth, topography and water mass dynamics, fauna diversity may be high and patchy. [ABSTRACT FROM AUTHOR]

The deep-sea is increasingly viewed as a lucrative environment for the growth of resource extraction industries. To date, our ability to study deep-sea species lags behind that of those inhabiting the photic zone limiting scientific data available for management. In particular, knowledge of horizontal movements is restricted to two locations; capture and recapture, with no temporal information on absolute animal locations between endpoints. To elucidate the horizontal movements of a large deep-sea fish, a novel tagging approach was adopted using the smallest available prototype satellite tag – the mark-report pop-up archival tag (mrPAT). Five Greenland sharks ( Somniosus microcephalus ) were equipped with multiple mrPATs as well as a standard archival satellite tag (miniPAT) that were programmed to release in sequence at 8–10 day intervals. The performance of the mrPATs was quantified. The tagging approach provided multiple locations per individual and revealed a previously unknown directed migration of Greenland sharks from the Canadian high Arctic to Northwest Greenland. All tags reported locations, however, the accuracy and time from expected release were variable among tags (average time to an accurate location from expected release = 30.8 h, range: 4.9–227.6 h). Average mrPAT drift rate estimated from best quality messages (LQ1,2,3) was 0.37 ± 0.09 m/s indicating tags were on average 41.1 ± 63.4 km (range: 6.5–303.1 km) from the location of the animal when they transmitted. mrPATs provided daily temperature values that were highly correlated among tags and with the miniPAT (70.8% of tag pairs were significant). In contrast, daily tilt sensor data were variable among tags on the same animal (12.5% of tag pairs were significant). Tracking large-scale movements of deep-sea fish has historically been limited by the remote environment they inhabit. The current study provides a new approach to document reliable coarse scale horizontal movements to understand migrations, stock structure and habitat use of large species. Opportunities to apply mrPATs to understand the movements of medium size fish, marine mammals and to validate retrospective movement modeling approaches based on archival data are presented. [ABSTRACT FROM AUTHOR]

The absorption coefficient of chromophoric dissolved organic matter (CDOM) is a major variable used in developing robust bio‐optical models and understanding biogeochemical processes. Over the last decade, the optical properties of CDOM in the open sea have been intensely studied. However, their variations in clear water are poorly documented, particularly in the Red Sea, owing to the absence of in situ measurements. We performed several cruises in the Red Sea to investigate the spatial distribution of the absorption coefficient of CDOM. The spectral absorption coefficients were determined from 400 nm to 740 nm using a WETLabs ac-s hyper-spectral spectrophotometer. In general, we found a latitudinal gradient in the CDOM absorption coefficient at 443 nm (a CDOM (443)) from south to north that is likely influenced by the exchange of water through the strait of Bab-el-Mandeb and the thermohaline circulation of the Red Sea. However, high a CDOM (443) values were observed in the northern Red Sea due to the existence of a sub-mesoscale feature that may induce an increase in phytoplankton production and lead to CDOM production. The a CDOM (443) covaried with the chlorophyll a concentration ([Chl a],) despite a high scatter. Furthermore, the a CDOM (443) for a given [Chl a] concentration was higher than those predicted by global ocean bio-optical models. This study advances our understanding of CDOM concentration in the Red Sea and may help improve the accuracy of the algorithms used to obtain CDOM absorption from ocean color. [ABSTRACT FROM AUTHOR]

Specific mechanisms, driving trophic interactions between seamount associated fishes and the pelagic community may be highly variable in different seamount systems. This study investigated the trophic structure and the main prey of benthopelagic fishes from the summit and slope regions of Ampère and Senghor, two shallow seamounts in the subtropical and tropical NE Atlantic, and the adjacent deep-sea plains. For the identification of food sources and nutritional links to the pelagic realm a combination of stomach content and stable isotope ratio (δ 13 C and δ 15 N) analyses was used. δ 13 C ranged from −22.2‰ to −15.4‰ and δ 15 N covered a total range of 8.0–15.9‰. Feeding types of fish species comprised mainly zooplanktivores and mixed feeders, but also benthivores, piscivores, and predator-scavengers. Based on epipelagic particulate organic matter, they occupied trophic positions between the 2nd and 4th trophic level. Differences in stomach contents and stable isotope signatures indicate a resource partitioning among the benthopelagic fish fauna through distinct habitat choice, vertical feeding positions and prey selection. Topographic trapping of vertically migrating zooplankton on the summit seemed to be of minor importance for food supply of the resident near-bottom fishes, rather horizontal current-driven advection of the planktonic prey was assumed as major factor. Vertically migrating micronekton and mesopelagic fishes show up as key players within the food webs at Ampère and Senghor Seamounts and the adjacent deep-sea plains. [ABSTRACT FROM AUTHOR]

Understanding ecosystem functioning and stability of continental slope environments requires assessing the effects of habitat heterogeneity (canyon and non-canyon habitats) on benthic macrofaunal communities and ecosystem functioning. We evaluated the importance of habitat heterogeneity and phytodetrital enrichment in canyon, channel, and inter-canyon habitats of the Gulf of Maine, Northwest Atlantic. Using an ROV, we collected sediment cores at five seafloor sites in the Gulf of Maine (663–969 m depth). Over ∼24 h, we incubated ∼6 cores per site, three of which were enriched with the diatom Chaetoceros muelleri. For all core treatments, we measured macrofaunal composition, functional diversity indices (functional richness, evenness, and divergence), and benthic nutrient fluxes (ammonium, nitrate, phosphate, and silicate), as well as oxygen consumption. We also dedicated two non-incubated cores to quantifying sedimentary and chemical-physical characteristics. We found that benthic communities differed among the three habitats, but these differences and our phytodetrital additions did not translate to significant differences in rates of benthic fluxes among habitats., expect for nitrate and phosphate fluxes. Total organic carbon and chlorophyll -a explained most of the variation in oxygen consumption, which was similar in enriched and control cores. We conclude that large-scale habitat heterogeneity contributes to variation in benthic community composition, but with no consistent effect on nutrient fluxes. These results also suggest that phytodetrital deposition influences oxygen consumption but plays a lesser role in short-term nutrient cycling in deep-sea heterogeneous habitats, perhaps masked by other environmental variables. Moreover, sediments with high functional richness and relatively high functional divergence may sometimes lack the specific trait types that rapidly enhance nutrient regeneration and thus contribute to ecosystem functioning of deep-sea sediments. • Habitat heterogeneity affected infaunal community assemblages but had less effect on benthic nutrient fluxes. • Deposit feeders, surficial modifiers, and freely mobile organisms within the sediment matrix dominated the macrofauna. • Seafloor sediments in our study region acted more as a sink than a source of nutrients. • Biological and environmental variables did not explain overall variation in benthic nutrient fluxes. • Phytodetrital addition resulted in different flux responses only for nitrate (Inter-canyon) and phosphate (Channel). [ABSTRACT FROM AUTHOR]

Many biological and physical variables influence microbes at the sediment-water interface, whose response to those variables affects the overall fate and residence time of organic matter (OM) in the marine environment. In addition to the many ecological roles microbes play in deep-sea sediments, further studies are needed to address microbial community diversity and its influence on seafloor organic matter remineralization rates. We explored microbial diversity, distribution, and associated organic matter remineralization among contrasting habitats and following 24-h experimental phytodetrital enrichment of sediments from Canyon, Inter-Canyon, and Channel habitats of the Gulf of Maine, Northwest Atlantic. Based on multivariate analyses of community composition we found that habitat heterogeneity influenced microbial community composition but not microbial diversity. Similarly, the phytodetrital addition did not translate to significant differences in microbial diversity but altered nitrate flux in Inter-canyon sediments and phosphate flux in Channel sediments. Bacteria were dominant over Archaea and, of the environmental variables we measured, quantity and quality of organic matter best explained overall microbial community variation. Proteobacteria, Bacteroidae, Acidobacteriota, Planctomycetota, and NB1-J Deltaproteobacteria dominated all habitats, but variation in the microbial community at our study sites explained only 7% of nutrient fluxes. Our exploratory study did not suggest a strong contribution of microbial community composition to OM remineralization variation both among contrasting habitats and in response to phytodetrital addition, and thus to this aspect of deep-sea ecosystem functioning, at least over short-term incubations. • Contrasting habitats affected microbial community composition, but not diversity, in the Gulf of Maine. • Phytodetrital addition did not translate to significant differences in benthic microbial diversity in short-term incubations. • Quantity and quality of organic matter best explained overall microbial community variation. • Microbial community composition variation explained only 7% of benthic nutrient fluxes. • Benthic nutrient fluxes did not vary significantly among contrasting habitats. [ABSTRACT FROM AUTHOR]

Habitat-forming cold-water corals (CWCs) represent a key component of deep-sea benthic communities and a priority target for conservation. Although research efforts have been mounting to try and identify the drivers of CWC distributions, progress has been limited by the scarcity of ecological data. The present work employs habitat suitability models (HSMs) to estimate spatial distributions, environmental drivers and co-existence patterns of 14 habitat-forming CWCs in the Azores, an area considered a hotspot of coral diversity in the Atlantic. The modelled CWCs showed a strong bathymetric zonation, which seems to be determined by the vertical stratification of water masses in the region. In particular, the modelled CWCs can be clustered in four groups named after the isopycnal (vertical) layers in which Atlantic water masses are organized: species restricted to upper water masses, species extending down from upper water masses, species restricted to intermediate water masses and species extending up from deep water masses. Horizontal patterns further indicate that the Azores Current and different production regimes north and south of the archipelago likely influence the distribution of CWCs in sub-surface waters. Such results have important implications for the regional management of deep-sea benthic communities and, in particular, for the design of representative networks of protected areas. The combined habitat of all modelled species covered only 11%. Given that they all possess the characteristics of benthic foundation organisms and represent indicator taxa of vulnerable marine ecosystems all the modelled species should be viewed as important targets for conservation. The lace coral Errina dabneyi deserves particular attention since this species appears to be endemic to the Azores and has a very limited estimated distribution. • Different pools of CWC taxa exist in the Azores. • The vertical stratification of water masses is a driver of CWC distributions. • Different production regimes in the archipelago influence the distribution of CWCs. • The estimated habitat of key CWCs species covered only 11% of the study area. • Spatial protection is advised for all the considered CWC taxa. [ABSTRACT FROM AUTHOR]

The yellow coral Dendrophyllia cornigera (Lamarck, 1816) is a NE Atlantic-Mediterranean scleractinian. It is considered a typical hard bottom species, generally reported on outcropping rocks from mesophotic to upper bathyal depths. Several evidences suggest that this species is able to tolerate a broad range of temperatures, which allows it to colonize numerous environments in a wide depth range. In the present study, we first provided a detailed ecological characterization of the D. cornigera dense aggregation thriving on the Mantice Shoal (NW Ligurian Sea, Mediterranean Sea). Information on substrate type and inclination, average extension and density, colonies size-class frequency distribution and associated fauna are reported. Then, we presented an extensive review of the available information on the ecology of this species, including 142 new ROV records from the Italian coast (40–1820 m). Results indicated that D. cornigera occurs on a wide range of substrates, including soft bottoms and hardgrounds (outcropping rocks, coralligenous rock and dead cold-water coral frameworks), with significant differences in colony density and size among different substrates. Dendrophyllia cornigera creates three main facies, each characterized by a specific combination of substrate, inclination, depth, and associated fauna. Scattered living colonies, as well as large thanatocoenoses, display a wide geographical and bathymetric distribution. Differently, the facies represented by dense meadows on horizontal soft-bottoms results rare, being reported only from the Mantice Shoal and the Amendolara Bank (Ionian Sea). The radiocarbon age of the thanatocoenoses varies between 400 (Corsica Channel) and 13000 (Vercelli Seamount) years before present. This study highlights the wide adaptability of D. cornigera in terms of environmental settings, changing the current view on the ecology of this species and providing essential insights for the implementation of international deep-sea habitat classification schemes and conservation measures. • The ecology of structuring species is poorly known, minimizing conservation efforts. • Dendrophyllia cornigera is considered a hard bottom, temperature-tolerant coral. • This study evidences the wide ecological adaptability of this deep habitat-former. • Deep thanatocoenoses hold paleo-information on past Mediterranean biocoenoses. • Results support the implementation of deep-sea habitat classification systems. [ABSTRACT FROM AUTHOR]

Abyssal polymetallic nodule fields represent a unique deep-sea habitat. The hard substratum provided by nodules is thought to increase habitat complexity, promoting the occurrence of highly diverse abyssal assemblages. However, hard substratum in these habitats is also available as outcropping rock fragments, but their contribution to habitat heterogeneity has been largely overlooked. Here, we assessed if the type and size of hard substratum can modulate benthic biodiversity at local scales within the Clarion Clipperton Zone (N Pacific abyss). We compared megafauna (animals >10 mm) assemblages in nodule bearing habitats with those in areas containing rocks in addition to nodules. We found a lower faunal density but a clearly higher diversity and more heterogenous composition in the assemblages of rock bearing areas compared to those with only nodules. In addition, hard substratum patch size appeared to positively influence the size of some taxa, like bamboo corals. These results suggest that rocks, in addition to nodules, constitute keystone structures enhancing habitat heterogeneity at local scales within nodule fields. Rock areas appear common in some abyssal plains and may make regionally-important contributions to abyssal biodiversity, suggesting that they may be relevant to include in conservation efforts. • We compared megafauna from Nodule and Rocks areas in an abyssal nodule field. • Rock bearing areas exhibit a higher diversity than those with only nodules. • Hard substratum patch size positively influences the size of some taxa. • Rocks and nodules are keystone structures enhancing abyssal habitat heterogeneity. • Rock areas in abyssal environments should be considered in conservation efforts. [ABSTRACT FROM AUTHOR]

Since the discovery of deep-sea hydrothermal vents in 1977, chemosynthesis-based ecosystems have been found in a wide range of habitats, such as hydrocarbon seeps, coastal sediments and terrestrial caves. Several invertebrates found in dark ecosystems harbor chemosynthetic bacteria (epibionts) on the surfaces of their specialised tissues; in particular, a vent galatheid crab Shinkaia crosnieri consumes the epibionts thriving on their setae as the primary nutritional source. In this study, we found that the water stream is the key to the nutritional symbiosis between S. crosnieri and chemosynthetic epibionts. The chemosynthetic functions of epibionts were highly activated by the water flow, and observations in the laboratory supported rheotaxis by S. crosnieri . In addition, endogenous water flow generated by respiratory water ventilation of S. crosnieri stimulated the chemosynthetic activity of epibionts under static conditions. These results point out that the epibionts are cultivated by the active behaviors of S. crosnieri , potentially representing the first example of a cultivation mutualism in chemosynthetic ectosymbiosis. [ABSTRACT FROM AUTHOR]

The association of organisms with particular habitats and habitat-forming organisms, can strongly influence species distributions, interactions and wider ecosystem services. At the sub-Antarctic Prince Edward Islands, the caridean shrimp Nauticaris marionis is a principal part of the benthic ecosystem, occurring between ca. 50 m and 600 m. Its role as a trophic link between the primary productivity and higher predators is established, but little is understood of its in situ habitat usage and associations or of how these structure patterns of abundance. We investigated these aspects directly using a benthic camera sled, sampling 27 stations between 50 m and 500 m. Substratum type was characterised, and estimates of percentage cover of the 13 main groups of habitat-forming epibenthic taxa were made, alongside absolute counts of N. marionis within ‘digital quadrats’ drawn from 300 m transects. The distribution of N. marionis was influenced by depth, substratum type and overall biogenic cover, being limited to habitats between 50 and 160 m depth on mud or gravel substrata only, and having > 50% biogenic cover. The presence/absence of N. marionis related to significantly different epibenthic assemblages (termed biogenic habitats), but this effect was contingent on depth. Likewise, densities of N. marionis were significantly affected by biogenic habitat type, identifying an association with two biogenic habitat groups, one dominated by red-algae, the other by structurally complex bryozoan species. These associations likely relate to the structural complexity of the two habitat groups, rather than the specific taxa involved. The apparent absence of N. marionis at depths > 160 m contrasts with earlier records and poses questions about the trophic importance of the shrimp in deeper habitats. [ABSTRACT FROM AUTHOR]

Species distribution modelling can be applied to identify potentially suitable habitat for species with largely unknown distributions, such as many deep-water corals. Important variables influencing species occurrence in the deep sea, e.g. substrate composition, are often not included in these modelling approaches because high-resolution data are unavailable. We investigated the relationship between substrate composition and the occurrence of the two deep-water octocoral species Primnoa resedaeformis and Paragorgia arborea , which require hard substrate for attachment. On a scale of 10s of metres, we analysed images of the seafloor taken at two locations inside the Northeast Channel Coral Conservation Area in the Northwest Atlantic. We interpolated substrate composition over the sampling areas and determined the contribution of substrate classes, depth and slope to describe habitat suitability using maximum entropy modelling (Maxent). Substrate composition was similar at both sites - dominated by pebbles in a matrix of sand (>80%) with low percentages of suitable substrate for coral occurrence. Coral abundance was low at site 1 (0.9 colonies of P. resedaeformis per 100 m 2 ) and high at site 2 (63 colonies of P. resedaeformis per 100 m 2 ) indicating that substrate alone is not sufficient to explain varying patterns in coral occurrence. Spatial interpolations of substrate classes revealed the difficulty to accurately resolve sparsely distributed boulders (3–5% of substrate). Boulders were by far the most important variable in the habitat suitability model (HSM) for P. resedaeformis at site 1, indicating the fundamental influence of a substrate class that is the least abundant. At site 2, HSMs identified cobbles and sand/pebble as the most important variables for habitat suitability. However, substrate classes were correlated making it difficult to determine the influence of individual variables. To provide accurate information on habitat suitability for the two coral species, substrate composition needs to be quantified so that small fractions (<20% contribution of certain substrate class) of suitable substrate are resolved. While the collection and analysis of high-resolution data is costly and spatially limited, the required resolution is unlikely to be achieved in coarse-scale interpolations of substrate data. [ABSTRACT FROM AUTHOR]

We describe the seasonal changes in the horizontal and vertical distribution and abundance of euphausiid species associated with seven physical and 61 biological variables in the Gulf of California (24–31°N). Euphausiid community structure was explored in the epipelagic habitat (<200 m) in January, July, and October 2007 and in epipelagic to bathypelagic habitats (<1400 m depth) in May 2015. Twelve euphausiid species comprising two distinct regional assemblages were identified. Nyctiphanes simplex and Nematoscelis difficilis were the most abundant species (>90%) in all cruises carried out in the 26–31°N region and Euphausia distinguenda in the 24–26°N region (mostly in October >90%). We confirmed that Euphausia gibboides and Nematobrachion flexipes inhabit the mesopelagic habitat, adapted to <1 ml O 2 l –1 environmental condition. Although Euphausia lamelligera and N. simplex populations were concentrated in well-oxygenated water (>3 ml O 2 l –1 ) near the surface (<50 m), they were also detected in low densities in the hypoxic mesopelagic habitat (250–800 m), but only at oxygen levels less than 0.09 ml O 2 l –1 . Stylocheiron affine and Stylocheiron carinatum were numerous well below the seasonal thermocline (100–350 m). Nematoscelis difficilis and N. simplex extended into the hypoxic mesopelagic habitat, but at low densities. Multivariate analyses showed six seasonal and regional krill assemblages, which are characterized by different species (Similarity percentage analysis). Canonical Correspondence Analysis show that, of 68 variables, three abiotic factors (temperature, dissolved oxygen, and OMZ depth) and eight biotic factors (accessory pigments; zeaxanthin and 19'-hexanoyloxyfucoxanthin, and the abundance of six copepod species; Candacia pectinata , Canthocalanus pauper , Centropages furcatus , Rhincalanus nasutus , Scolecithrix danae , and Temora discaudata ), were the most influential variables associated with the vertical distribution and abundance of euphausiids. Krill‒copepod (predator–prey) relationships may be important determinants of daily and seasonal vertical distribution patterns. We conclude that the euphausiid vertical habitat reaches down to 900 m depth (with historical records of six species collected between 1000 and 2280 m) but with lower diversity and abundance in the mesopelagic and bathypelagic habitats than in the epipelagic habitat. [ABSTRACT FROM AUTHOR]

The sediment-covered deep-sea floor was initially thought to be environmentally homogeneous. Recent work has shown otherwise, and deep-sea ecologists have been searching for ecologically important environmental heterogeneities on different spatial and temporal scales, with particular interest in canyons. Here we report results for harpacticoid copepods from a site at 3262 m depth in the axis of Monterey Canyon and one on an escarpment 46 km away at 3090 m depth. Multivariate community analyses revealed significant differences between sites in community structure. Absolute abundance, the ratio of subadult copepodites to adults, species density, the proportion of the harpacticoid individuals that emerged, and the proportion that lived in tubes were significantly lower at the canyon site than at the escarpment site. The proportion of the harpacticoid individuals that belonged to the surface-dweller life-style group was significantly higher than at the escarpment site. These marked differences imply that ecologically important environmental heterogeneities exist. We speculate that differences between the sites in food conditions and sediment grain-size distributions are among them. [ABSTRACT FROM AUTHOR]

Ecological theory predicts that heterogeneous habitats allow more species to co-exist in a given area, but to date, knowledge on relationships between habitat heterogeneity and biodiversity of coral forests in the outer shelf and upper slope along continental margins is rather limited. We investigated biodiversity of coral forests from 8 sites spread over two different geomorphological settings (namely, pinnacles vs . canyons) in the outer shelf along Sardinian continental margin. Using a combination of multivariate statistical analyses, we show here that differences in the composition of coral assemblages among contrasting geomorphological settings were not statistically significant, whereas significant differences emerged among sites within similar geomorphologies (i.e. among pinnacles and among canyons). Our results reveal that environmental and bathymetric factors such as sediment coverage, slope of the substrate, terrain ruggedness, bathymetric positioning index and aspect were important drivers of the observed patterns of coral biodiversity, in both settings. Spatial variability of coral forests' biodiversity is affected by environmental factors that act at the scale of each geomorphological setting (i.e. within each pinnacle and canyon) rather than by the contrasting geomorphological settings themselves. This result allows us to suggest that simple categorization of benthic communities according topographically defined habitat is unlikely to be sufficient for addressing conservation purposes. [ABSTRACT FROM AUTHOR]

Food web configuration is shaped by many factors, including primary production patterns and oceanographic features, such as upwelling events. In this study, we investigate variability in the trophic position, food web interlinks and energy pathways of four planktivorous demersal fish in the Southern Bay of Biscay- NE Atlantic. The study area is exposed to upwelling events of varying intensity and shows a significant spatial gradient along the coast. The two sampling years were characterized by markedly different conditions, with weak summer upwelling in 2012 and an intense upwelling season in 2013. We used a complementary approach based on stomach content analysis (SCA) and stable isotope analysis (SIA) to test the effects of upwelling intensity and persistence on the food-web. In particular we investigated whether different intensities shift the main flow of energy between the pelagic and benthic energy pathways. We found conspicuous interannual changes in the isotopic POM, whose δ 15 N was 2.5‰ lower during the productive 2013 season. Interannual changes in the isotopic signature of the fish species were also detected, although their values did not mirror variability at the basal level. The SCA results did not match the isotopic changes, which likely reflected dietary adjustments of the species during summer. The upwelling intensity gradient along the coast did not affect the nitrogen isotopic ratio of any group, however there was a significant effect of such gradient on the carbon isotopic ratio of the fish and euphausiid species. This effect was likely related to the higher primary production associated with intense upwelling conditions. [ABSTRACT FROM AUTHOR]

The ecological study of corals in their habitat is essential to determine the effects of global change and to develop strategies for reef conservation. Based on mark and recovery experiments, we investigated skeletal growth patterns of two reef-building cold-water coral species, Lophelia pertusa and Madrepora oculata , in the Lacaze-Duthiers canyon in the northwestern Mediterranean Sea. Coral fragments were collected, stained and deployed for short-term (2.5 months) and long-term (15 months) growth experiments at two sites located 4.5 km and 6.8 km from the canyon head. The analysis of distinct growth parameters (budding, new polyp growth and linear extension of the coral fragments) revealed that growth patterns are consistent among branches of different sizes, but discrepancies arose from the different types of staining used. Calcein appeared more suitable than alizarin red, which strongly limited growth by delaying coral recovery, for short-term experiments at least. Both species grew rapidly when redeployed in their habitat. Effects of long-term experiments could not be observed because corals were exposed to harmful environmental stress, particularly the lethal effect of sedimentation on the fragments. Despite limited in situ deployment, the growth analysis from the short-term experiment highlighted species-specific responses according to the location along a longitudinal gradient in the canyon and were likely related to the local environmental conditions. Consistent with the observed species distributions, M. oculata showed optimal growth at the site closer to the canyon head compared with L. pertusa , which had optimal growth at the deeper site. In situ experiments are difficult to conduct in deep-sea ecosystems, but with the use of remotely operated vehicles (ROVs), such a simple approach may be of interest to managers of cold-water coral ecosystems. [ABSTRACT FROM AUTHOR]

The Northeast Pacific continental margin is characterized by strong seasonal upwelling, which drives high primary productivity, and supports high diversity and biomass of benthic megafauna. The recent occurrence of a marine heat wave ("The Blob", sensu Kintisch, 2015) in 2013–2016 resulted in changes to phytoplankton community composition and loss of coastal kelp abundance and diversity, reducing gross primary productivity in the region. However, cumulative effects of marine heat waves and ongoing basin-scale deoxygenation in deep-sea ecosystems remain poorly understood. Here, we use a 7-year time series of physicochemical and video imagery data from Ocean Networks Canada's NEPTUNE observatory to investigate temporal dynamics of the deep-sea pink urchin Strongylocentrotus fragilis in relation to multi-year environmental variability. Using generalized additive models, we show that local S. fragilis density at Barkley Upper Slope (420 m) fluctuated over time and was partially explained by changes in dissolved oxygen concentration and suspended particulate matter in the benthic boundary layer (ADCP backscatter), with high urchin density corresponding to high oxygen and low backscatter. Seafloor dissolved oxygen ranged from 0.80 to 1.89 mL/L and varied seasonally, exhibiting a clear negative correlation with sea surface primary productivity (MODIS satellite Chl-a data), corresponding with the onset of yearly upwelling conditions. However, during the anomalously warm years affected by 'The Blob', dampened upwelling maintained higher dissolved oxygen conditions near the seafloor. S. fragilis density declined during 'Blob' conditions, likely in response to reduced kelp and phytodetritus subsidies from coastal waters. We propose a foraging-respiration trade-off hypothesis, whereby S. fragilis forages in deeper water during weak upwelling and migrates to shallower habitats during low oxygen conditions. S. fragilis is an important bioturbator and detritivore; changes in the density and distribution of this species may directly affect sediment turnover rates and nutrient cycling on the continental margin, with consequences for surface and coastal productivity. https://github.com/rjcommand/. • 7 years of video imagery and oceanographic data and with 14 years of trawl surveys. • Pink urchin densities mainly controlled by oxygen and availability of phytodetritus. • S. fragilis densities were reduced during the blob when detritus subsidies declined. • Shoaling of S. fragilis on the shelf may affect other seafloor nutrient cycling. [ABSTRACT FROM AUTHOR]

Deep-sea coral habitats, comprising mostly Lophelia pertusa (Linnaeus 1758), are well developed on the upper and middle continental slope off the southeastern United States (SEUS). These habitats support a diverse and abundant invertebrate fauna, yet ecology and biology of most of these species are poorly known. Ten cruises conducted off the SEUS (Summer–Fall; Cape Lookout, NC–Cape Canaveral, FL) from 2000 to 2005, and in 2009 provided an opportunity to investigate abundance and distribution of Eumunida picta Smith1883, a large-sized species of squat lobster commonly associated with these deep-water coral habitats. Video analysis from 70 manned-submersible dives documented occurrence, density, location on the coral colony, and behavioral observations for 5774 individuals of E. picta. Individuals collected (n = 178) from coral and adjacent habitats (e.g., rubble, soft sediments) were measured and their sex determined. Males and females were comparable in size (to 53.5 mm carapace length) and exhibited a sex ratio of approximately 1:1. Eumunida picta were most frequently observed as solitary individuals on high-profile coral matrix and were noted only infrequently on coral rubble, or rarely on soft substratum. Presence of coral habitat (i.e., live/dead L. pertusa), geographic region within the sampling area, and depth significantly influenced abundances of E. picta. Additionally, coral habitat (dead versus live coral), vertical position on the coral (upper, middle, or lower zone), as well as horizontal position in relation to the coral matrix (outer surface versus embedded in coral matrix) were significant factors influencing E. picta distributions within the coral habitat. More individuals were found on dead versus live coral, and most frequently occurred on the outer surfaces of coral branches located on the upper portion or near the tops of coral colonies. Eumunida picta were most often observed with claws extended into the water column. This unique hunting stance provides this squat lobster the opportunity to capture prey from the water column. An active predator, this species utilizes both pelagic (i.e., fishes, pyrosomes) and benthic (e.g., scavenging and grazing) food resources, and may function as an important trophic link between the water column and the benthos. Although considered a facultative reef associate in the strictest sense of the term, E. picta has a complex and intimate relationship with L. pertusa. Based on observations from dive videos, E. picta is a dominant and ecologically important member of the invertebrate assemblage associated with deep-sea coral habitats off the SEUS. As such, this species figures prominently in the structure and function of this ecosystem. • Most comprehensive dataset examining demography and behavior of Eumunida picta. • Eumunida picta has a complex and intimate relationship with Lophelia pertusa. • Eumunida picta abundance correlates primarily with abundance of L. pertusa habitat. • The claws-extended behavior is a unique hunting stance for capturing pelagic prey. • Eumunida picta is an important trophic link between the benthos and water column. [ABSTRACT FROM AUTHOR]

In the deep-sea, cold-water corals (CWCs) and other structure-forming fauna locally increase habitat complexity and are host to many fish species. While many studies have focused on hard-bottom CWCs, very little is known about fish associations with soft-bottom CWC habitats. To understand the small-scale linkages between fish, benthic and non-bio-structural habitats in a soft-sediment deep-sea environment, we analysed remotely operated vehicle (ROV) survey videos and recorded the occurrence of fish, invertebrate habitat-forming species and substratum along transects, in the Laurentian Channel Marine Protected Area (MPA) in the Canadian Northwest Atlantic. Almost 13,300 individual fish were recorded, of which all but 65 specimens were identified to family or lower taxonomic rank. The five numerically dominant fish taxa were Redfish (Sebastes spp.), Witch Flounder (Glyptocephalus cynoglossus), Marlin-Spike Grenadier (Nezumia bairdii), Longfin Hake (Phycis chesteri) and one Teleostei morphotype. Teleostei sp1 represents three fish genera (Sandlance (Ammodytes spp.) and two Barracudinas (Arctozenus sp. and Paralepis spp.)) morphologically difficult to differentiate on video footage. Multivariate analysis revealed four habitats based on epibenthic invertebrate densities and diversity. These were dominated by various combinations of sea anemones (Actiniaria), nephtheid soft coral, solitary cup corals (Scleractinia) and three taxa of sea pen (Pennatula spp., Kophobelemnon sp. and Anthoptilum spp.). Univariate analyses performed on dominant fish densities revealed the local influence of hard-bottom substrates and the common influence of soft-sediment micro-features for three fish taxa. All fish densities were correlated with invertebrate density, negatively for Redfish and positively for all other taxa. Our models also predicted the association of four fish taxa with one to two benthic habitats within their preferred depth range. Our results suggest that small-scale habitat heterogeneity in a low relief soft-sediment environment, provided by both physical and biological structures, has a measurable species-specific influence on fish communities. This influence was weaker than typical fish-habitat relationships found in hard-bottom systems, suggesting fish-invertebrate relationships are not obligate. Additionally, we provided evidence that Redfish continue occupying sea pen habitats months after spawning. Our study supports the necessity to continue monitoring the MPA using in-situ video systems to understand the links between fish and habitats in the Laurentian Channel. • Evidence of taxon-specific influence of specific habitat biotic and abiotic features on fish density • Evidence of taxon-specific influence of specific CWC taxa on fish density • Results suggest a facultative relationship between fish and invertebrates in soft-bottom ecosystems. • Evidence of occupation and multiple use of specific sea pen habitats by Redfish for months after spawning. • ROV is an efficient tool for detecting in-situ small-scale associations of fish in their habitat. [ABSTRACT FROM AUTHOR]

To understand the spatial distribution of phytoplankton communities in various habitats in the Amundsen Sea, western Antarctica, a field survey was conducted at 15 stations during the austral summer, from December 2013 to January 2014. Water samples were analyzed by microscopy. We found high phytoplankton abundance and biomass in the Amundsen Sea polynya (ASP). Their strong positive correlation with water temperature suggests that phytoplankton biomass accumulated in the surface layer of the stratified polynya. In the ASP, the predominant phytoplankton species was Phaeocystis antarctica , while diatoms formed a major group in the sea ice zone, especially Fragilariopsis spp., Chaetoceros spp., and Proboscia spp. Although this large diatom abundance sharply decreased just off the marginal sea ice zone, weakly silicified diatoms, due to their high buoyancy, were distributed at almost all stations on the continental shelf. Dictyocha speculum appeared to favor the area between the marginal sea ice zone and the ASP in contrast to cryptophytes and picophytoplankton, whose abundance was higher in the area between the continental shelf and the open ocean of Amundsen Sea. Several environmental factors were found to affect the spatial variation of phytoplankton species, but the community structure appeared to be controlled mainly by the seawater density related to sea-ice melting and water circulation in the Amundsen Sea. [ABSTRACT FROM AUTHOR]

Seamounts are commercially important fishing grounds. Yet, little is known about their physical characteristics as fish habitat, important for informing conservation and ecosystem-based management. This study examines how multiscale seabed spatial heterogeneity influences commercially important fish families at three Southwest Indian Ridge seamounts (Coral Seamount, Melville Bank and Atlantis Bank). We quantified seascape heterogeneity from bathymetry and geomorphological habitat maps and identified 15 focal fish families from video data. Fish-habitat associations were examined using spatial pattern metrics that measured terrain morphology, seascape composition (variety and relative abundance of patch types) and seascape configuration (spatial arrangement of patches). Broader seascape context was characterised by geographic location and water depth. Multivariate regression trees and random forests modelled fish-habitat associations and identified the most influential explanatory variables. Assemblage characteristics and individual families were strongly influenced by geographic location and depth, and at finer scales (500 m buffers) seascape composition and configuration helped explain fish-habitat associations. Spatially continuous summit habitat and complex shaped ridge features supported high abundance and diversity of commercial fish families. Metrics of seascape composition and configuration (i.e., habitat size, shape and structural connectivity) had higher predictive power than the terrain metrics commonly used in developing proxies for deep-water fish species and biodiversity. These outcomes indicate that seascape metrics, commonly applied on land and in shallow marine environments, are also relevant environmental predictors of fish distributions in deep-sea environments. We highlight strong context dependency and depth-specific associations that hinder attempts to draw wider generalisations on fish-seascape linkages for seamounts. • Fish assemblages on three Southwest Indian Ridge seamounts are mainly driven by location and depth. • Landscape ecology metrics identified environmental predictors of fish families and assemblages. • Metric combinations shape ecological patterns, with similarities between co-occurring families. • Main drivers were the proportion of summit and ridge habitat, and their spatial continuity and shape. [ABSTRACT FROM AUTHOR]

Deep-sea environments are harsh habitats; however, many animal taxa flourish in deep-sea ecosystems such as vents and seeps. The success of those animals in such habitats has been attributed to their symbiotic relationships with diverse microorganisms. Previous studies extensively examined the roles of chemoautotrophic bacteria in symbiotic systems, revealing that they are essential to the ecological adaptation of animal holobionts. Some recent studies have focused on the influence of viral partners on the bacterial symbionts in deep-sea animals, which shows that bacterial viruses (bacteriophages/phages) might also play important roles in the deep ocean symbiosis. In the present review, we deduce phage-bacteria interactions from existing literature, newer comparative analyses, and considerations of microbial adaptation to extreme environments. In the analyses, we specifically investigated viral signals in hologenomes of animals in hydrothermal vents and cold seeps and discussed the potential roles of phages in symbiosis maintenance. Metagenome investigation reveals diverse phage families in the holobionts of mussels, clams, sponges, tubeworms, and snails. Phages in symbiotic systems of vents and seeps may undergo a lysogenic life cycle. The metabolic genes in phage genomes can complement or compensate bacterial symbiont metabolism. In seep mussels and tubeworms, the presence of phage-born immunity-related genes may aid bacterial symbionts in immune evasion by reducing animal phagocytosis. Furthermore, lysis by phages may mediate nutritional supply to deep-sea animals by lysing bacterial cells. For the sponges with relatively high microbial diversity, the diversity may be sustained through phage infection of dominant bacterial symbionts, thereby contributing to animal fitness. We propose a mutualistic tripartite symbiosis model to illustrate the relationships of phages, bacteria, and animals in deep-sea vent/seep ecosystems. • Phages in animal holobionts in deep-sea hydrothermal vents and cold seeps have received increasing attention. • Phages are widely distributed in a wide range of vent/seep animal species and are genotypically diverse. • Phages appear to have a lysogenic lifestyle in vent/seep animal holobionts. • Phages play potential roles in metabolic regulation, immune evasion, nutrition supply, and microbial diversity maintenance. [ABSTRACT FROM AUTHOR]

Here we describe the methods and results for biological characterization of the benthos on a previously unexplored area of central California, USA seafloor. We conducted 40 remotely operated vehicle dives from 371 to 1173 m water depth. Seafloor habitats and megafauna (fish and invertebrates) were documented from 46.8 km of seafloor video footage. Our expanded development and analysis of biotopes from quantitative data allowed us to describe detailed biological communities, along with the physical characteristics and habitat associations within the study area. This method provides a framework for potential monitoring, detection of future environmental change (natural and anthropogenic) and comprehensive comparison to other deep water regions. From 185 h of observational video at 25 sites, nearly 120,000 annotations of organisms, habitat characters, biological detritus and anthropogenic debris were recorded and analyzed. We identified a total of 228 taxa, with 173 of them present on linear quantitative transects. Species richness for transects ranged from 0.04 to 0.28 m-2 (8–55 taxa), with densities ranging from 0.07 to 5.20 ind. m−2. Both were highest on hard substrate with greatest surface area. Densities decreased with depth. Within soft substratum zones was a large field of pockmarks, which are seafloor depressions averaging 175 m in diameter and 5 m in depth. Pockmarks have sometimes been associated with seafloor gas seepage, but here we found no biological evidence of chemosynthetic organisms. No significant differences were found in either density nor species richness at pockmark sites vs. non-pockmark sites. Mud draped greenish-black coarse sand occurred only in low oxygen areas, while hummocky, rugose mud supported somewhat different species than flat mud plains. Seventy percent of the transects occurred inside the oxygen minimum zone. We conclude that high rugosity, slope, and the presence of hard substratum were better predictors of species richness and density than oxygen concentration in this specific study. Abundant biological detritus, in the form of dead and dying pelagic pyrosomes and salps, created a large, presumably ephemeral flux of carbon to the seafloor during the study period. • Multi-factored biotopes was an effective method to detail biological and environmental differences within the survey area. • Rugose, hard substrata on slopes supported the highest number of taxa and densities. • Fauna from outside vs. within pockmark fields were not significantly different in density, species richness, or uniqueness. • Mud draped greenish-black coarse sand was a unique habitat which occurred only in a low oxygen area on Santa Lucia Bank. • Future studies may be effected by high ephemeral carbon flux to the seafloor and changes in dissolved oxygen levels. [ABSTRACT FROM AUTHOR]

In the deep sea, biological data are often sparse; hence models capturing relationships between observed fauna and environmental variables (acquired via acoustic mapping techniques) are often used to produce full coverage species assemblage maps. Many statistical modelling techniques are being developed, but there remains a need to determine the most appropriate mapping techniques. Predictive habitat modelling approaches (redundancy analysis, maximum entropy and random forest) were applied to a heterogeneous section of seabed on Rockall Bank, NE Atlantic, for which landscape indices describing the spatial arrangement of habitat patches were calculated. The predictive maps were based on remotely operated vehicle (ROV) imagery transects high-resolution autonomous underwater vehicle (AUV) sidescan backscatter maps. Area under the curve (AUC) and accuracy indicated similar performances for the three models tested, but performance varied by species assemblage, with the transitional species assemblage showing the weakest predictive performances. Spatial predictions of habitat suitability differed between statistical approaches, but niche similarity metrics showed redundancy analysis and random forest predictions to be most similar. As one statistical technique could not be found to outperform the others when all assemblages were considered, ensemble mapping techniques, where the outputs of many models are combined, were applied. They showed higher accuracy than any single model. Different statistical approaches for predictive habitat modelling possess varied strengths and weaknesses and by examining the outputs of a range of modelling techniques and their differences, more robust predictions, with better described variation and areas of uncertainties, can be achieved. As improvements to prediction outputs can be achieved without additional costly data collection, ensemble mapping approaches have clear value for spatial management. [ABSTRACT FROM AUTHOR]

We studied seasonal and regional change in vertical distribution and DVM patterns of four euphausiid species ( Euphausia pacifica, Thysanoessa inspinata, Thysanoessa longipes, and Tessarabrachion oculatum ) from two years of surveys using MOCNESS above 1500 m depth across a transect in 3 regions of the northwestern (NW) Pacific, off east of Japan; Oyashio, Kuroshio, and Oyashio–Kuroshio Mixed Water Regions (MWR). The four euphausiid species exhibited a regional change in vertical distribution, i.e., slightly deeper in the MWR and much deeper in the Kuroshio region than in the Oyashio region. They found in higher and wider temperature ranges in the MWR than in the Oyashio region, which demonstrated that the four species were able to adapt to different temperatures in different regions. In the MWR and Oyashio regions, E. pacifica is a surface migrant (differences between day and night mean median depths, D-N, were ca. 300 m) and T. oculatum is a moderate subsurface migrant that performs short DVM in the upper mesopelagic zone (D-N ca. 100 m). The other two morphologically similar Thysanoessa species ( T. inspinata and T. longipes ) segregated vertically between E. pacifica and T. oculatum at night in the Oyashio region, suggesting vertical habitat partitioning with the former two species but not with themselves. However, a seasonal pattern was observed in the vertical distribution and DVM of T. longipes in the Oyashio region. It behaves as a surface migrant in May, whereas most of individuals were found in the mesopelagic layer in September. In contrast, T. inspinata did not exhibit a clear DVM throughout the year (i.e., a moderate subsurface migrant). This seasonal difference might be a strategy to minimize competition between related species. Among the four species, only E. pacifica was found in higher temperatures at night than during the daytime, and the highest temperatures at the median depth varied among species (from 7.5 °C to 13.7 °C) although the lowest temperature did not vary greatly (from 1.0 °C to 1.8 °C), which indicates high temperatures act as a limiting factor as opposed to low temperatures. Furthermore, the integrated chlorophyll a values exhibited significant negative correlation with median depths of only E. pacifica at night. These results indicate a strategy which makes E. pacifica the dominant species in the area, that is, it has a trade-off of long migrations and a warmer environment that accelerates metabolism, in return for obtaining a food-rich environment. [ABSTRACT FROM AUTHOR]

The Long-Term Ecological Research (LTER) observatory HAUSGARTEN, in the eastern Fram Strait, provides us the valuable ability to study the composition of benthic megafaunal communities through the analysis of seafloor photographs. This, in combination with extensive sampling campaigns, which have yielded a unique data set on faunal, bacterial, biogeochemical and geological properties, as well as on hydrography and sedimentation patterns, allows us to address the question of why variations in megafaunal community structure and species distribution exist within regional (60–110 km) and local (<4 km) scales. Here, we present first results from the latitudinal HAUSGARTEN gradient, consisting of three different stations (N3, HG-IV, S3) between 78°30′N and 79°45′N (2351–2788 m depth), obtained via the analysis of images acquired by a towed camera (OFOS-Ocean Floor Observation System) in 2011. We assess variability in megafaunal densities, species composition and diversity as well as biotic and biogenic habitat features, which may cause the patterns observed. While there were significant regional-scale differences in megafaunal composition and densities between the stations (N3=26.74±0.63; HG-IV=11.21±0.25; S3=18.34±0.39 individuals m −2 ), significant local differences were only found at HG-IV. Regional-scale variations may be due to the significant differences in ice coverage at each station as well as the different quantities of protein available, whereas local-scale differences at HG-IV may be a result of variation in bottom topography or factors not yet identified. [ABSTRACT FROM AUTHOR]

Growing economic interest in seabed resources in the deep-sea highlights the need for information about the spatial distribution and vulnerability to disturbance of benthic habitats and fauna. Categorisation of seabed habitats for management is often based on topographic features such as canyons and seamounts that can be distinguished using regional bathymetry ('mega-habitats'). This is practical but because such habitats are contiguous with others, there is potential for overlap in the communities associated with them. Because concepts of habitat and community vulnerability are based on the traits of individual taxa, the nature and extent of differences between communities have implications for strategies to manage the environmental effects of resource use. Using towed video camera transects, we surveyed mega-epifaunal communities of three topographically-defined habitats (canyon, seamount or knoll, and continental slope) and two physico-chemically defined meso-scale habitats (cold seep and hydrothermal vent) in two regions off New Zealand to assess whether each supports a distinct type of community. Cold seep and hydrothermal vent communities were strongly distinct from those in other habitats. Across the other habitats, however, distinctions between communities were often weak and were not consistent between regions. Dissimilarities among communities across all habitats were stronger and the density of filter-feeding taxa was higher in the Bay of Plenty than on the Hikurangi Margin, whereas densities of predatory and scavenging taxa were higher on the Hikurangi Margin. Substratum diversity at small spatial scales (<1 km) and trawl history were significantly correlated with community composition in both regions. We conclude that, (1) a lack of consistent distinction between communities raises questions about the general utility of topographically-defined mega-habitats in environmental management, (2) fine-scale survey of individual features is necessary to identify the locations, characteristics, and extents of ecologically important or vulnerable seabed communities, and (3) evaluation of habitat vulnerability to future events should be in the context of previous and current disturbances. [ABSTRACT FROM AUTHOR]

The deep-sea mussel Bathymodiolus thermophilus is a dominant species in the East Pacific Rise (EPR) hydrothermal vent fields. On the EPR volcanically unstable area, this late colonizer reaches high biomass within 4–5 years on new habitats created by lava flows. The environmental conditions and growth rates characterizing the reestablishment of B. thermophilus populations are however largely unknown, leaving unconstrained the role of this foundation species in the ecosystem dynamics. A typical example from the vent field at 9°50’N that was affected by the last massive eruption was the Bio-9 hydrothermal vent site. Here, six years later, a large mussel population had reestablished. The von Bertalanffy growth model estimates the oldest B. thermophilus specimens to be 1.3 year-old in March 2012, consistent with the observation of scarce juveniles among tubeworms in 2010. Younger cohorts were also observed in 2012 but the low number of individuals, relatively to older cohorts, suggests limited survival or growth of new recruits at this site, that could reflect unsuitable habitat conditions. To further explore this asumption, we investigated the relationships between mussel growth dynamics and habitat properties. The approach combined sclerochronology analyses of daily shell growth with continuous habitat monitoring for two mussel assemblages; one from the Bio-9 new settlement and a second from the V-vent site unreached by the lava flow. At both vent sites, semi-diurnal fluctuations of abiotic conditions were recorded using sensors deployed in the mussel bed over 5 to 10 days. These data depict steep transitions from well oxygenated to oxygen-depleted conditions and from alkaline to acidic pH, combined with intermittent sulfide exposure. These semi-diurnal fluctuations exhibited marked changes in amplitude over time, exposing mussels to distinct regimes of abiotic constraints. The V-vent samples allowed growth patterns to be examined at the scale of individual life and compared to long-term records of habitat temperature and oceanographic mooring data in the years following the eruption. Both shell growth and habitat temperature at V-vent varied over the spring-neap tidal cycle and over longer periods of c.a. 60 days. The correlation of growth rate with temperature and, for some individuals, with current velocities supports the idea that tidal forcing impacts growth. Its influence on habitat conditions includes the spring-neap cycle, which is not reflected in current velocities but influences the venting rate. Additionally, it is expected that mesoscale eddies periodically passing across the ridge imprint shell growth through the influence of bottom current on the decimeter-thick mixing interface where mussels thrive. We conclude that diurnal-semidiurnal tidal fluctuations exert major abiotic constraints on B. thermophilus mussels and that low-frequency fluctuations act as significant determinants on growth. Finally, we postulate that the modulation of tidal fluctuations by large-scale hydrodynamic forcing ultimately constrains the capacity of this mussel species to form high biomass aggregations. This study indeed shows that the absence of these strong hydrodynamic drivers would limit the alternance of oxic and sulfidic conditions and significantly affect the growth rate of this species over time. [ABSTRACT FROM AUTHOR]

In February 2019, seamounts of the Discovery Rise, SE Atlantic (41–45°S, 3°W - 3°E), were explored in support of the application of the Ecosystem Approach to Fisheries Management in the Southeast Atlantic Fisheries Organization (SEAFO) Convention Area. Video records of the seafloor were produced by the Video-Assisted Multisampler System (VAMS) along 15 valid transects conducted in plateau and flank areas of Shannon, Tablemount, Discovery and Heardman seamounts at depths ranging from 394 to 1839 m. Nine benthic seascapes were classified and described based on substrate hardness, texture, slope, physical and biological modifiers as observed in the video images. Predominant water masses were estimated from temperature, salinity and dissolved oxygen vertical profiles obtained by CTD casts in the vicinity of each transect. A total of 366 fishes were seen (total observation time = 25.0 h) and classified in 32 morphotypes, included in 9 orders and 12 families. Most fish morphotypes (14) were included in the Macrouridae family. Family Moridae, on the other hand, included 65.3% of all fish records (239), with two particularly abundant morphotypes: Laemonema sp. (116) and Guttigadus sp. (92). Thirteen cephalopods were observed and classified in five morphotypes; the oegopsid squid Moroteuthopsis ingens was the most abundant of them (6). Despite the taxonomic uncertainties associated with video identifications, the explored region was found to contain a mixture of tropical – subtropical and subantarctic faunas of the Atlantic. Similarities of fauna composition and non-directional beta diversity estimates revealed some degree of seamount identity, but 57.5–61.9% of morphotypes were shared among seamounts (Jaccard = 0.425, Sørensen = 0.381). Fishes and cephalopods were more frequently observed on the shallower plateau areas under the influence of warmer and more oxygenated Antarctic Intermediate Waters. Depth and related factors did not influence richness, but dissimilarities in fauna composition between video transects increased with increasing depth intervals. Spatial habitat heterogeneity may have accounted for the increased beta diversity within seamounts. The most widely explored Tablemount seamount contained the most diverse observed sites, but also a considerable spatial variability likely associated with seascape heterogeneity. In two sites (transects 11 and 14), there were exceptionally high aggregations of the morid cods Laemonema sp., at the shallowest 'gravely' site explored (397 m), and Guttigadus sp., at a site of intermediate depth (1020 m) covered mostly by soft sediments. Neither of these species (or congenerics) have been previously classified as 'seamount-aggregating" species. On the other hand, the Patagonian toothfish (Dissostichus eleginoides) and other commercial species known to aggregate in these seamounts were not found in the explored areas. • The Discovery Rise seamounts comprise a variety of contrasting seascapes. • Observed benthopelagic fauna was dominated by macrourids and morid cods. • The region included tropical-subtropical and subantarctic species of the Atlantic. • Fishes and cephalopods were more frequently observed on the shallower plateau areas. • Fauna composition varied (beta-diversity) within and between the explored seamounts. [ABSTRACT FROM AUTHOR]

The Lacaze-Duthiers Canyon is located in the western Mediterranean Sea and is long known for hosting cold-water coral colonies in the canyon head region at depths ranging from 250 to 550 m. In 2019 during the CALADU cruise, three kinds of 3D-reconstructions were applied to better understand the distribution of coral colonies, their habitat and their skeleton morphologies. The canyon's flanks were mapped using a hull-mounted echosounder and an ROV multibeam echosounder. Digital terrain models were built with resolutions of 5 and 1 m and examined in three dimensions. ROV bathymetric data collected on the canyon's flanks made it possible to highlight a series of sub-parallel structures identified as lithified sedimentary strata along which coral colonies grow. Coral assemblages were explored at four locations and photographic images were assembled using structure from motion techniques to build photogrammetric models. Coral assemblages reconstructed in 3D enabled geo-localizing and recreating coral colonies on 16 models over a total area of 4370 m2. Two colonial species, Madrepora oculata and Desmophyllum pertusum were plotted and reported on bathymetric models to interpret their location at the scale of the canyon. The coordinates and depth of the colonies were used to calculate the vertical distribution (limited to our small bathymetric exploration, between 339 and 214 m depth) and density of populations (up to 4.3 colonies per m2). The spatial coverage of the 16 assemblages measured between 100 and 600 m2 each. The sizes of the colonies were measured to analyze the population structures of both species (mean sizes of 28 cm for D. pertusum and 18 cm for M. oculata , maximum sizes 1 m and 0.5 m, respectively, bushes 2.5 m long). In addition, lost fishing gears were quantified, longlines measured and their densities calculated (0.16 m/m2, up to 0.30 m/m2). An area with exuberant orange colonies of D. pertusum was discovered for the first time in the Lacaze-Duthiers Canyon. Five deep-sea scleractinian species were collected and micro-tomographic scans computed to view their intrinsic skeleton organization. Micro-CT scans of M. oculata , D. pertusum , Desmophyllum dianthus , Caryophyllia smithii , and Dendrophyllia cornigera enabled longitudinal and transversal cuts, highlighting morphological criteria for species identification and the multidirectional examination of specimens. We observed a thin canal connecting calices along the axis of D. pertusum colonies, and separate calices along the axis of M. oculata colonies. • Canyon walls were mapped with a MBES installed with a 45° angle. • A surface of 4370 m2 was reconstructed on Cold-Water Corals ecosystem. • The highest density of coral colonies among the Mediterranean canyons. • Coral colonies faced prevailing currents coming from canyon head incisions. • A thin canal connects buds of successive generation in Desmophyllum pertusum. [ABSTRACT FROM AUTHOR]

The shrimp Rimicaris exoculata is the most conspicuous component of vent communities developing around hydrothermal fluid emissions below 2000 m on the northern Mid-Atlantic Ridge (nMAR). Its high genetic connectivity suggests a remarkable ability to produce dispersing larval stages. However, so far brooding females have been rarely observed and reproduction remained enigmatic. Spatially complex population structures related to the heterogeneity of local habitat conditions are described for many vent species, this information being fundamental to gain a better understanding of their life history. Here our aim was to assess such complexity along with reproductive development in R. exoculata populations within two vent fields, TAG and Snake Pit (3620m and 3470m depth respectively). We compared samples collected in January–February 2014 among visually distinct assemblages with different degrees of exposure to vent fluids. Dense aggregations located near active venting included mostly females and immature individuals, while inactive peripheries harbored low density assemblages of large males. Small juveniles gathering around low temperature diffusions belonged to another species, Rimicaris chacei. One third of the sexually mature females were ovigerous at the two vent fields during our sampling period, with lower fecundities and egg sizes in the TAG population. Overall, the observed shrimp distribution patterns were consistent across both vent fields, although a high degree of heterogeneity in population structure was observed locally within dense aggregations, probably reflecting micro-scale variations in environmental conditions. Our results thus highlight spatially complex population structures where R. exoculata females brood eggs within dense aggregations exposed to vent fluids, while peripheral inactive areas may be important mating grounds for adults. In addition, we suggest temporal variability in reproductive activity, increasing in the winter season, which questions potential seasonality in a deep-sea species. • Shrimps show contrasting population structures between habitats within vent fields. • Females and immature individuals dominate dense R. exoculata aggregations. • Scattered shrimps in the inactive vent peripheries are mostly R. exoculata males. • One third of the sexually mature females were brooding in winter 2014. • R. exoculata broods near vent fluids, whereas mating may occur in the periphery. [ABSTRACT FROM AUTHOR]

The Rio Grande Rise (RGR) is a large and geomorphologically complex feature located in the Southwest Atlantic, with great commercial and scientific interest due to its potential for mining rare earth elements that are critical for low-carbon technologies. Brazilian interest in this area led to the submission of a petition to the UN Commission on the Limits of the Continental Shelf in 2018 to include RGR on the limits of its continental shelf beyond 200 nautical miles. However, mining activities are potentially harmful to deep-sea ecosystems and will likely cause some extent of biodiversity loss. Thus, baseline and continuous environmental studies in the RGR are important to address potential conflicts between mineral extraction and the conservation of deep-sea biodiversity. The RGR is characterized by a series of summit plateaus of ∼600 m deep divided NE-SW by a rift valley, up to 2000 m deep. In 2018, the plateaus and rift of a small area in RGR (30°35′S – 31°03′S, 35°36′W – 36°16′W) were explored through 13 dives of the robotic underwater vehicle (RUV) HyBIS. Videos were analyzed for the description of structuring factors (topography and habitat types) and to record benthic megafauna occurrences. Video transects revealed highly heterogeneous and rapidly changing habitats. Eleven habitats, five in the rift and six in the plateaus are proposed based on geomorphology, slope, and substrate textures. We recorded 17,008 megabenthic organisms classified in 83 morphotypes and six different phyla, from which Porifera (42.7%) and Cnidaria (41.5%) were the most representative. Samples were characterized by a high dominance and the dissimilarities result chiefly from differences in abundance scores. PERMANOVA tests indicated that Habitat and Region variables were the most important to explain structure within the community data, followed by depth and slope. The rift floor exhibited a low abundance of megabenthic epifauna, except in a sinkhole in the northern part of the rift. The lower and upper rift wall were characterized by different communities delimited by the transition between the Antarctic Intermediate Water and the Upper Circumpolar Deep Water. The habitats formed by Fe–Mn deposits were dominated by distinct communities, which were rarely observed elsewhere. Additionally, we found variations in community structure at regional scales (20–30 km), with distinct communities on each side of the rift and at the southwest of the study area. Our results contribute toward understanding the diversity, biogeography, and environmental drivers of the RGR. Fauna distribution is patchy, linked to habitats with potential mining resources, and dominated by Vulnerable Marine Ecosystems (VMEs) indicator taxa. Extensive community analysis should occur at a given site prior to consideration for the exploitation of natural resources. • New insights into sessile megafaunal abundance and diversity from the RGR. • Patchy megafaunal abundance found across the habitats of RGR. • Community structure is driven by habitat, geographical locations, depth, and slope. • Fe–Mn crust areas had distinct communities compared with other hard substrates. [ABSTRACT FROM AUTHOR]

Habitat-forming deep-sea scleractinian and alcyonacean corals from around the southwest Pacific were analysed for their calcium carbonate mineralogy. Scleractinian coral species Solenosmilia variabilis, Enallopsammia rostrata, Goniocorella dumosa, Madrepora oculata and Oculina virgosa were all found to be 100% aragonitic, while some members of the alcyonacean taxa Keratoisis spp. , Lepidisis spp., and Paragorgia spp. were determined to be high magnesium (Mg) calcite (with 8–11 mol% MgCO 3 ) and Primnoa sp. is bimineralic with both aragonite and Mg calcite. The majority of these habitat-forming deep-sea corals are found at intermediate depths (800–1200 m) in the Antarctic Intermediate Waters (AAIW) with low salinities (~34.5), temperatures of 4–8 °C and high oxygen concentrations (>180 μmol/kg) and currently sitting above the aragonite saturation horizon (ASH). However, habitat-forming corals have been recorded from greater depths, in cooler waters (2–4 °C) that are undersaturated with respect to aragonite (Ω aragonite <1), but with oxygen levels still >160 μmol/kg. To address the sampling depth bias the coral records were normalised by the number of benthic stations (sampling effort) in the same depth range. This shows that the highest number of corals per sampling effort is between 1000 and 1400 m with corals present in over 5% of the stations at these depths. The normalised records and Boot Strap analyses suggests that scleractinian corals, especially S. variabilis should be present in >1% of stations down to 1800 m water depth, with E. rostrata, M. oculata and G. dumosa slightly shallower. While alcyonacean corals are found in >1% down to 2600 m, with Keratoisis spp. the deepest down to 2600 m, while Lepidisis spp. and Paragorgia spp. found down to 1800 m. This suggests that most species can probably tolerate some undersaturation of aragonite (Ω aragonite =0.8–0.9), with several species/genera ( S. variabilis ; Keratoisis spp.) even more tolerant of lower carbonate concentrations ([ CO 3 2 − ]), down to Ω aragonite of 0.7. With this tolerance for some carbonate undersaturation it is unclear how deep sea habitat-forming corals might respond to future ocean acidification. It is likely that some species/genera will cope better than others. However, future changes in oxygen concentrations and food availability, are also going to have a strong influence on the depth and spatial distribution of deep-sea corals in the southwest Pacific. [ABSTRACT FROM AUTHOR]

We simulated transport of Pacific sardine eggs captured offshore of California in spring of 2001–2012 using a regional ocean circulation model. Eggs were assumed to have developed into larvae within a few days and were modeled using five behavioral patterns: passive transport, diel vertical migration, diel vertical migration combined with swimming against the current, diel migration combined with migration toward shore, and diel migration combined with migration toward the best habitat. Simulated larvae with no swimming behavior were advected far offshore to poor habitat where they were unlikely to survive. Diel vertical migration resulted in less offshore transport because larvae were less affected by surface currents during the day. However, in half the years simulated nearly all juveniles were also located in poor habitat by late summer in this scenario. Swimming against the current combined with diel vertical migration resulted in similar transport patterns to the diel-vertical-migration scenario because currents dominated the transport of eggs and small larvae during the spring and early summer. Migration toward shore resulted in a large fraction of juveniles being located in appropriate habitat during late summer in all years. Migration toward the best habitat was the best strategy modeled. This strategy resulted in a slightly greater proportion of larvae being located in appropriate habitat at the end of summer than the swimming-toward-shore scenario, despite the fact that most larvae were located farther offshore. These results suggest that larval sardine might use directed horizontal swimming behavior to remain in suitable habitat conditions. A large fraction of larvae were transported south into Mexican waters by late summer in all five scenarios. Surveying juvenile sardines in fall near the border of the U.S. and Mexico may be an efficient means of estimating recruitment because the advection pattern of eggs and larvae to the south is opposite the adult migration pattern to the north. This pattern may cause juveniles to be spatially segregated from adults at the time they are being recruited. [ABSTRACT FROM AUTHOR]

Sharks are a major component of the top predator guild in oceanic ecosystems, but the trophic relationships of many populations remain poorly understood. We examined chemical tracers of diet and habitat (δ 15 N and δ 13 C, respectively) and total mercury (Hg) concentrations in muscle tissue of seven pelagic sharks: blue shark ( Prionace glauca ), short-fin mako shark ( Isurus oxyrinchus ), oceanic whitetip shark ( Carcharhinus longimanus ), scalloped hammerhead shark ( Sphyrna lewini ), pelagic thresher shark ( Alopias pelagicus ), crocodile shark ( Pseudocarcharias kamoharai ) and silky shark ( Carcharhinus falciformis ), from the data poor south-western tropical Indian Ocean. Minimal interspecific variation in mean δ 15 N values and a large degree of isotopic niche overlap – driven by high intraspecific variation in δ 15 N values – was observed among pelagic sharks. Similarly, δ 13 C values of sharks overlapped considerably for all species with the exception of P. glauca , which had more 13 C-depleted values indicating possibly longer residence times in purely pelagic waters. Geographic variation in δ 13 C, δ 15 N and Hg were observed for P. glauca and I. oxyrinchus. Mean Hg levels were similar among species with the exception of P. kamoharai which had significantly higher Hg concentrations likely related to mesopelagic feeding. Hg concentrations increased with body size in I. oxyrinchus , P. glauca and C. longimanus . Values of δ 15 N and δ 13 C varied with size only in P. glauca , suggesting ontogenetic shifts in diets or habitats. Together, isotopic data indicate that – with few exceptions – variance within species in trophic interactions or foraging habitats is greater than differentiation among pelagic sharks in the south-western Indian Ocean. Therefore, it is possible that this group exhibits some level of trophic redundancy, but further studies of diets and fine-scale habitat use are needed to fully test this hypothesis. [ABSTRACT FROM AUTHOR]

Cold-water corals and sponges aggregations are important features of the deep sea, recently classified as vulnerable marine ecosystems (VMEs). VMEs increase habitat complexity, believed to act as feeding, reproductive, nursery and refuge areas for a high number of invertebrates and fish species. In the Azores archipelago (NE Atlantic), VMEs are prevalent but their ecological role has not received much attention. The objective of this study was to investigate the importance of VMEs in influencing the distribution of demersal fish in the Azores. With data collected during experimental longline surveys , we modeled the catch of six demersal fish species of commercial value ( Helicolenus dactylopterus, Pagellus bogaraveo , Mora moro , Conger conger , Phycis phycis , Pontinus kuhlii ) in relation to the presence of VMEs and other environmental factors using General Additive Models (GAMs). Our study demonstrated that total fish catch was higher inside VMEs but the relationship between fish and VMEs varied among fish species. Species specific models showed that catch was strongly influenced by environmental factors, mainly depth, whilst the presence of VMEs was only important for two rockfish species; juvenile and adult P. kuhlii and juvenile H. dactylopterus . Although the association between deep-sea demersal fish and VMEs may be an exception to the rule, we suggest that VMEs act as an important habitat for two commercially important species in the Azores. [ABSTRACT FROM AUTHOR]

High densities of ice krill Euphausia crystallorophias were observed along six acoustic transects within the Amundsen Sea Coastal Polynya, Antarctica. Two-frequency acoustic backscatter data was examined in the austral summers of January 2011 and February 2012. A dB identification window ( S v120−38 ) identified ice krill dominating the acoustic backscatter. The density of ice krill, calculated with the stochastic distorted-wave born approximation model, ranged between 4.5 and 30 g wet mass m −2 for each transect (a mean of 16 g wet mass m −2 for all transects), these high values are an order of magnitude higher than recorded previously in the Ross Sea Polynya. High densities were detected along the ice shelf and near the boundary between pack ice and coastal polynya, and we postulate that these could be important habitats for ice krill. The high densities observed along the transects make ice krill a potentially important, but poorly known contributor to these high-latitude shelf food webs. [ABSTRACT FROM AUTHOR]

The eastern tropical north Pacific Ocean (ETNP) contains one of the world’s most severe oxygen minimum zones (OMZs), where oxygen concentrations are less than 2 µmol kg −1 . OMZs cause habitat compression, whereby species intolerant of low oxygen are restricted to near-surface oxygenated waters. Copepods belonging to the family Eucalanidae are dominant zooplankters in this region and inhabit a variety of vertical habitats within the OMZ. The purpose of this study was to compare the metabolic responses of three species of eucalanoid copepods, Eucalanus inermis, Rhincalanus rostrifrons , and Subeucalanus subtenuis , to changes in temperature and environmental oxygen concentrations. Oxygen consumption and urea, ammonium, and phosphate excretion rates were measured via end-point experiments at three temperatures (10, 17, and 23 °C) and two oxygen concentrations (100% and 15% air saturation). S. subtenuis , which occurred primarily in the upper 50 m of the water column at our study site, inhabiting well-oxygenated to upper oxycline conditions, had the highest metabolic rates per unit weight, while E. inermis , which was found throughout the water column to about 600 m depth in low oxygen waters, typically had the lowest metabolic rates. Rates for R. rostrifrons (found primarily between 200 and 300 m depth) were intermediate between the other two species and more variable. Metabolic ratios suggested that R. rostrifrons relied more heavily on lipids to fuel metabolism than the other two species. S. subtenuis was the only species that demonstrated a decrease in oxygen consumption rates (at intermediate 17 °C temperature treatment) when environmental oxygen concentrations were lowered. The percentage of total measured nitrogen excreted as urea (% urea-N), as well as overall urea excretion rates, responded in a complex manner to changes in temperature and oxygen concentration. R. rostrifrons and E. inermis excreted a significantly higher % of urea-N in low oxygen treatments at 10 °C. At 17 °C, the opposite trend was observed as E. inermis and S. subtenuis excreted a higher % of urea-N in the high oxygen treatment. This unique relationship has not been documented previously for crustacean zooplankton, and warrants additional research into regulation of metabolic pathways to better understand nitrogen cycling in marine systems. This study also compared metabolic data for E. inermis individuals captured near the surface versus those that were resident in the deeper OMZ. Deeper-dwelling individuals had significantly higher nitrogen excretion rates and O:N ratios, suggesting an increased reliance on lipids for energy while residing in the food-poor waters of the OMZ. [ABSTRACT FROM AUTHOR]

Xenophyophores are a group of exclusively deep-sea agglutinating rhizarian protozoans, at least some of which are foraminifera. They are an important constituent of the deep-sea megafauna that are sometimes found in sufficient abundance to act as a significant source of habitat structure for meiofaunal and macrofaunal organisms. This study utilised maximum entropy modelling (Maxent) and a high-resolution environmental database to explore the environmental factors controlling the presence of Xenophyophorea and two frequently sampled xenophyophore species that are taxonomically stable: Syringammina fragilissima and Stannophyllum zonarium . These factors were also used to predict the global distribution of each taxon. Areas of high habitat suitability for xenophyophores were highlighted throughout the world׳s oceans, including in a large number of areas yet to be suitably sampled, but the Northeast and Southeast Atlantic Ocean, Gulf of Mexico and Caribbean Sea, the Red Sea and deep-water regions of the Malay Archipelago represented particular hotspots. The two species investigated showed more specific habitat requirements when compared to the model encompassing all xenophyophore records, perhaps in part due to the smaller number and relatively more clustered nature of the presence records available for modelling at present. The environmental variables depth, oxygen parameters, nitrate concentration, carbon-chemistry parameters and temperature were of greatest importance in determining xenophyophore distributions, but, somewhat surprisingly, hydrodynamic parameters were consistently shown to have low importance, possibly due to the paucity of well-resolved global hydrodynamic datasets. The results of this study (and others of a similar type) have the potential to guide further sample collection, environmental policy, and spatial planning of marine protected areas and industrial activities that impact the seafloor, particularly those that overlap with aggregations of these conspicuously large single-celled eukaryotes. [ABSTRACT FROM AUTHOR]

Release of hydrocarbons from sediments is important in increasing habitat heterogeneity on deep ocean margins. Heterogeneity arises from variation in abiotic and biotic conditions, including changes in substratum, geochemistry, fluid flow, biological communities and ecological interactions. The seepage of heavy hydrocarbons to the seafloor is less well studied than most other cold seep systems and may lead to the formation of asphalt mounds. These have been described from several regions, particularly the Gulf of Mexico. Here, we describe the structure, potential formation and biology of a large asphalt mound province in Block 31SE Angola. A total of 2254 distinct mound features was identified by side-scan sonar, covering a total area of 3.7 km 2 of seafloor. The asphalt mounds took a number of forms from small (<0.5 m diameter; 13% observations) mounds to large extensive (<50 m diameter) structures. Some of the observed mounds were associated with authigenic carbonate and active seepage (living chemosynthetic fauna present in addition to the asphalt). The asphalt mounds are seabed accumulations of heavy hydrocarbons formed from subsurface migration and fractionation of reservoir hydrocarbons primarily through a network of faults. In Angola these processes are controlled by subsurface movement of salt structures. The asphalt mounds were typically densely covered with epifauna (74.5% of mounds imaged had visible epifauna) although individual mounds varied considerably in epifaunal coverage. Of the 49 non-chemosynthetic megafaunal taxa observed, 19 taxa were only found on hard substrata (including asphalt mounds), 2 fish species inhabited the asphalt mounds preferentially and 27 taxa were apparently normal soft-sediment fauna. Antipatharians (3.6±2.3% s.e.) and poriferans (2.6±1.9% s.e.) accounted for the highest mean percentage of the observed cover, with actinarians (0.9±0.4% s.e.) and alcyonaceans (0.4±0.2% s.e.) covering smaller proportions of the area. Asphalt mounds represent a common and important habitat on several margin systems globally and should be recognised in future environmental assessment and management of these areas. [ABSTRACT FROM AUTHOR]

Research on marine mammal habitat-associations often uses satellite remote sensing of sea surface temperature, chlorophyll concentration, and sea surface height to map mesoscale features, which may indicate areas of enhanced productivity and prey availability. However, for species that feed at depths >400 m, the increased productivity associated with mesoscale features observed near the surface may have little or no immediate effect on habitat-associations at depth. As a result, previous studies have found a weak correlation between mesoscale features and the movements of marine mammals. The advantage of biologging is that hydrographic variables are recorded in situ and at foraging depths using animal-borne instruments with sensors for temperature, conductivity (salinity), and dissolved oxygen. The goal of this study was to characterize the habitat-associations of female southern elephant seals (SES) from Península Valdés, Argentina during the post-breeding foraging trip. Although female SES exhibited significant habitat-associations with sea surface height anomaly and chlorophyll concentrations, the presence or absence of eddies was not predictive of foraging behavior, and the majority of foraging dives (74%) and prey encounters (77%) occurred in the absence of eddies. The strongest habitat-association was with deep (>500 m) and cold (3.73 ± 1.29 °C) subantarctic water, primarily during foraging dives from dusk to dawn. Female SES made most of their foraging dives (68%, mean maximum depth of 539 ± 226 m) and had the most prey encounters (67%) in Antarctic Intermediate Water (AAIW), which is formed near the Subantarctic Front on the northern flank of the Antarctic Circumpolar Current. Our results suggest that AAIW is the principal foraging habitat of female SES from Península Valdés, which may not be directly associated with near-surface mesoscale features. Future research on the habitat-associations for SES and other deep-diving marine mammals should focus on indices of foraging success and the hydrographic features of water masses at foraging depths, not mesoscale features observed near the surface. • Female southern elephant seals foraged mainly in Antarctic Intermediate Water. • The strongest habitat-association was with deep and cold subantarctic water. • The presence or absence of eddies was not predictive of foraging behavior or success. • Mesoscale features can affect depth of subantarctic water masses where elephant seals forage. [ABSTRACT FROM AUTHOR]

During the period 1992–2014, we quantified 79,233 deep-sea corals (DSC; azooxanthellate corals occurring in cold or deep waters, generally >50 m) at depths 23–867 m from 1281 visual transects in areas of rocky substrata off northern, central, and southern California. DSC were present on 68% of transects at all depths and substrata types. Thirty-nine taxa and morphotypes from at least 19 families and 24 genera, including 20 species, were identified. Survey depths differed among study regions, overlapping between north and south at >575 m, and between central and south at <325 m. Within study regions, we examined taxa-specific depth and temperature ranges and quantified the spatial distribution of DSC densities. Across study regions, we interpreted species distributions in the context of the literature and biogeographic provinces off California, examined size distributions, and characterized assemblages relative to latitude and depth. We discovered depth range extensions for seven species, and provide the first taxa-specific in situ temperature data for the U.S. West Coast. Overall densities (number of colonies/100 m2) of DSC within study regions were 105.8 (north), 5.0 (central), and 7.3 (south), and highest densities occurred on mixed and hard substrata. We did not observe a change in density with depth. Density 'hot' spots (p < 0.05) were identified at multiple locations in the central and south regions for DSC (other than sea pens) and for sea pens, respectively. Despite survey depth differences, three species with large depth ranges were observed in all three study regions, and most species were observed in the biogeographical province(s) reported for them in the literature. Most DSC were ≤20 cm high and ≤10 cm wide; colonies >50 cm high and >40 cm wide were rare and found primarily in the south. Six DSC assemblages (p < 0.05) were identified. Mid-depth assemblages in the central and south differed considerably, while deep assemblages in the north and south were similar. A shallow assemblage comprised taxa shared by central and south regions. As the relative habitat value of DSC to fishes becomes more apparent, our results will provide invaluable information for the identification of essential fish habitat and priority areas in need of protection. [ABSTRACT FROM AUTHOR]

Fin whales and sei whales are two migratory baleen whale species sighted every year across the waters of the Azores. Improved understanding of the ecological niche and habitat requirements of these baleen whales is needed to identify persistent or predictable oceanographic events that may set the time of their migration, as well as local or ephemeral oceanographic features that may aggregate their prey in a particular area. In dynamic environments such as the open ocean, mesoscale and submesoscale features can become decisive to determine the distributions of highly mobile species such as baleen whales. In this study, we analyse the habitat preferences of fin whales and sei whales around São Miguel Island (Azores) using environmental variables at different temporal and spatial scales. For both species, model results showed a clear influence of variables linked with primary production and therefore, prey availability; as well as a noticeable preference for oceanographically dynamic areas which directly affect distribution and aggregation of prey. Those environmental choices may indicate different levels of foraging habitat use for both species. Differences were found between the species, highlighting preferences for colder waters in fin whales and areas with stronger sea surface temperature gradients in sei whales. Model results obtained for fin whales were similar with those previously published for blue whales, suggesting that both species make similar use of the waters around São Miguel, often foraging during the migration across these waters. Results for sei whale, however, emphasize dynamic variables, indicating that travelling may prevail over feeding behaviour during their migration by the Azores. • Fin whales and sei whales are sighted in sympatry every year in Azores. • Fin whale models indicate preference for dynamic areas and colder waters. • Sei whale habitat suitability increases with strong temperature gradients. • Highly mobile species habitat is analysed at different spatio-temporal scales. • Long-term opportunistic whale watching data provides valuable cetacean information. [ABSTRACT FROM AUTHOR]

Micronekton of the mesopelagic zone (200 m–1000 m depth) play a central role in most aspects of global marine ecology linking epipelagic and deep scattering layers as planktivores, carnivores and principal prey to both shallow diving and deep living apex predators. Yet, despite this critical role and conspicuous presence, little is known regarding most aspects of mesopelagic community ecology, including habitat associations at sub-basin spatial scales. Here we address several aspects of these data deficiencies in the eastern Bering Sea mesopelagic – a highly productive subarctic marine ecosystem home to multiple protected predator species and large commercial fisheries. Through 41 midwater trawls conducted at 250 m, 500 m and 1000 m depths during May of 1999 and 2000, and a series of generalized linear models and cluster analyses, we describe the influence of remotely sensed oceanographic habitat variables on the diel distribution, relative abundance and community composition of 30 species of mesopelagic fish and squid in 3 contrasting study areas. We then project the total energy available to predators at depth and across the eastern Bering Sea Basin based on the identified habitat constraints. Fishes from the families Bathylagidae and Myctophidae, and squids from the Gonatidae dominated the 72 species catch biomass (n = 225,000, 2,100 kg). The six most abundant species were equally divided between each of the three dominant families and are also the most highly represented mesopelagic species in eastern Bering Sea pelagic predator diets. Their association with surface habitat variables describe a complex and multifaceted relationship between physical/biological predictors and the total mesopelagic energy field during the spring bloom period in this ecosystem. Leuroglossus schmidti , Bathylagus pacificus , Stenobrachius leucopsaurus, S. nannochir, Gonatopsis borealis and Eogonatus tinro contrast in the suite of physical and biological variables that they respond to, and also in the character of their response. Although L. schmidti is of moderate caloric value, it expressed the greatest overall energy signature at all depths due to its overwhelming biomass, and response to the presence of specific, variable habitat conditions across extensive portions of the study area. Interannual variability in habitat variables forecasted a pronounced redistribution of dominant species and thereby total mesopelagic energy content – providing a potential mechanism for shifts in central-place predator habitat use, energy expenditure, and diet. The known relevance of mesopelagic species as prey and the energetic potential for apex predators foraging in the Basin illustrates a cautionary narrative regarding unregulated harvest of biota from this poorly understood and critically important oceanic zone. • Community composition of mesopelagic fish and cephalopod species dielly at depth in the eastern Bering Sea basin. • Relationships between physical and biological habitat variables and mesopelagic species distributions. • Dynamic distribution of mesopelagic species and subsequent energetic potential to predators at depth. [ABSTRACT FROM AUTHOR]

Pennatulacean octocorals (sea pens), one of the primary conservation targets of the Laurentian Channel Marine Protected Area (MPA) in eastern Canada, occur at high densities in some deep-sea sedimentary habitats. Considered important habitat-forming organisms for many megafaunal organisms, the effect of sea pens on nearby sedimentary macrofaunal communities remains unexplored. During two cruises in September 2017 and July 2018, we sampled 9 stations within the MPA (336-445 m depth), including sea pen fields and bare sedimentary habitats, targeting individual sea pens in a subset of the cores to assess small-scale effects. We evaluated macrofaunal density, taxonomic diversity, vertical distribution, community composition, and biological trait expression, and investigated variation between sea pen fields and other (bare) sedimentary habitats, as well as between cores with and without sea pen specimens. Using multivariate analyses, we identified abiotic and biotic drivers of macrofaunal community composition and biological trait expression. Enhancement of macrofaunal density and taxonomic diversity and higher percentages of organisms in the upper sediment layers characterized sea pen fields in autumn, with more variable results in summer. Community composition and biological trait expression consistently differed in sea pen fields compared to bare sedimentary habitats, with Pennatula sea pen density as one of the primary drivers of variation especially for community composition, along with other environmental drivers (depth, grain size, and organic matter quantity and quality). We also detected small-scale enhancement of macrofaunal diversity in cores containing sea pens at stations characterized by predominantly bare sediments. Our results indicate that macrofaunal communities within sea pen fields differ from those in bare sediments and we propose sea pens play a role in influencing those patterns by increasing food availability, stability, and small-scale heterogeneity in sedimentary habitats, acknowledging that other environmental factors might also play a role in determining the observed patterns. Characteristics of macrofaunal communities within sea pen fields also suggested potentially higher sensitivity to disturbance, which amplifies the need for protection of sea pen fields in deep-sea sedimentary environments. [ABSTRACT FROM AUTHOR]