Description: This is the first study to present the patterns and environmental controls of benthic biomass size spectra, carbon demand, and production along the entire bathymetric gradient from the shelf to the abyssal depths in the Arctic Ocean. The materials were collected at 17 stations (76 - 5561 m) in the eastern Fram Strait, in the Atlantic passage to the Arctic Ocean, in the vicinity of the productive Marginal Ice Zone, with concentrations of sediment-bound chloroplastic pigments (indicating food availability from phytodetritus sedimentation) higher than in other deep-sea localities at similar depths. Meiobenthic and macrobenthic individuals were measured using image analysis to assess their biovolume, biomass, annual production, and carbon demand. Benthic biomass in the area was clearly higher than that in the High Arctic locations and comparable to that in the lower-latitude North Atlantic. Biomass and annual production were significantly negatively correlated with water depth, with stronger bathymetric clines in macrofauna than in meiofauna and the increasing dominance of meiofauna with increasing depth. A bimodal shape in the size spectra was observed only at the shallow stations, while at depths below 2000 m, an additional trough was present in the macrofaunal part of the spectrum. The entire range of the spectra (i.e., the number of size classes) decreased with increasing depth, especially in the macrofaunal part of the spectrum. Similar slope values in the normalized spectra indicated that the distribution of the biomass across the present size classes was consistent from the shelf to the abyssal depths, irrespective of the decreasing amount of food availability. The fragmented macrofaunal size spectra documented at the two stations were probably due to physical disturbances at the sediment-water interface (e.g., intense bioturbation of holothurians and strong near-bottom currents). Benthic carbon demand declined from 50.7 gC m-2 y-1 at the shelf to 11.5 gC m-2 y-1 at the slope to 2.2 gC m-2 y-1 at the abyssal depths, and its partitioning among meiofauna and macrofauna changed with water depth, with meiofauna contributions increasing from 50 % at the shelf to over 90 % at the deepest station. The estimated total benthic carbon demand exceeded the vertical Corg fluxes, suggesting that the studied system can be particularly sensitive to future changes in productivity regimes and associated organic matter fluxes.

Description: Established in the Fram Strait in 1999, the LTER (Long-Term Ecological Research) observatory HAUSGARTEN enables us to study changes on the deep Arctic seafloor. Repeated deployments of a towed camera system (Ocean Floor Observation System) along the same tracks allowed us to build a time series longer than a decade (2004 - 2015). Here, we present the first time-series results from a northern and the southernmost station of the observatory (N3 and S3, ~2650m and 2350m depth respectively) obtained via the analysis of still imagery. We assess temporal variability in community structure, megafaunal densities and diversity, and use a range of biotic and abiotic factors to explain the patterns observed. There were significant temporal differences in megafaunal abundances, diversity and abiotic factors at both stations. A particularly high increase in megafaunal abundance was recorded at N3 from 12.08 (±0.39; 2004) individuals m-2 to 35.21 (±0.97; 2007) ind. m-2 alongside a ten-fold increase in (drop-)stones. At S3, megafaunal densities peaked in 2015 (22.74 ±0.61 ind. m-2) after an increasing trend since 2004 (12.44 ±0.32 ind. m-2). Holothurians showed particularly striking temporal differences: densities of the small sea cucumber Elpidia heckeri densities rose ten-fold from 0.31 ind. m-2 (±0.04; 2004) to 3.74 ind. m-2 (±0.14; 2015) at S3, coinciding with a sustained increase in phytodetritial matter (chloroplastic pigment equivalents) at the seafloor. Initially entirely absent from N3, densities of the larger holothurian Kolga hyalina peaked in 2007 (5.87 ±0.22 ind. m-2) and declined continuously since then. Overall diversity (γ) increased at both stations over the course of the study, however, with varying contributions of α and β diversities. Our results highlight the importance of time-series studies as megafaunal community composition is characterised by continuous changes. This indicates that epibenthic communities from the deep seafloor are reactive and dynamic, with no “null” community state. To continue to monitor them is therefore crucial in understanding natural and anthropogenic impacts in an area exposed to the effects of climate change.

Description: The LTER (Long-Term Ecological Research) observatory HAUSGARTEN, in the eastern Fram Strait, provides us the unique 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 transect, consisting of three different stations (N3, HG-IV, S3) between 78°30’N and 79°45’N (2500 m depth), obtained via the analysis of images acquired by a towed camera (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 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.

Description: Deep-sea benthic communities and their structural and functional characteristics are regulated by surface water processes. Our study focused on the impact of changes in water depth and food supplies on small-sized metazoan bottom-fauna (meiobenthos) along a bathymetric transect (1200–5500 m) in the western Fram Strait. The samples were collected every summer season from 2005 to 2009 within the scope of the HAUSGARTEN monitoring program. In comparison to other polar regions, the large inflow of organic matter to the sea floor translates into relatively high meiofaunal densities in this region. Densities along the bathymetric gradient range from approximately 2400 ind. 10 cm-2 at 1200 m to approximately 300 ind. 10 cm-2 at 4000 m. Differences in meiofaunal distribution among sediment layers (i.e., vertical profile) were stronger than among stations (i.e., bathymetric gradient). At all the stations meiofaunal densities and number of taxa were the highest in the surface sediment layer (0–1 cm), and these decreased with increasing sediment depth (down to 4–5 cm). However, the shape of the decreasing pattern differed significantly among stations. Meiofaunal densities and taxonomic richness decreased gradually with increasing sediment depth at the shallower stations with higher food availability. At deeper stations, where the availability of organic matter is generally lower, meiofaunal densities decreased sharply to minor proportions at sediment depths already at 2–3 cm. Nematodes were the most abundant organisms (60–98%) in all the sediment layers. The environmental factors best correlated to the vertical patterns of the meiofaunal community were sediment-bound chloroplastic pigments that indicate phytodetrital matter.

Description: Effects of increased near-bottom flow velocities on the sedimentary environment and its associated small biota were studied in a long-term in situ experiment at 2,500 m water depth at the Deep-Sea Observatory HAUSGARTEN in the eastern Fram Strait. In 2003, the Remotely Operated Vehicle (ROV) "Victor 6000" was used to install a stainless steel flume of about 8.5 m in length, consisting out of a 6 m long passageway with a cross-section of 50 x 50 cm and 3-4 m wide funnel-like doorways to increase bottom currents by a factor of approximately 6. Sediment sampling for biochemical sediment analyses, bacterial studies and meiofaunal investigations (with special focus on the nematode communities) was carried out four years after the installation of the flume using the ROV "Quest 4000". The data showed clearly reduced values for parameters indicating organic matter (food) availability in the sediments, and corresponding lower bacterial and meiofaunal densities inside the flume, compared to control sites outside the channel. Results suggest that increased near-bottom currents and food deficiency not only diminish sediment-inhabiting meiofaunal assemblages but also alter the meiobenthic composition. Compared to background sediments, the nematode community inside the flume evidently showed adaptations to the overall reduced food availability and a more heterogeneous environment due to generally increased and more turbulent flow velocities. The variable environmental conditions inside the flume have an effect not only on the number of genera present, but also on the identities of the genera and the functional composition of the nematode community.

Description: Epibenthic megafauna play an important role in the deep-sea environment and contribute significantly to benthic biomass, but their population dynamics are still understudied. We used a towed deep-sea camera system to assess the population densities of epibenthic megafauna in 2002, 2007, and 2012 at the shallowest station (HG I, ∼1300 m) of the deep-sea observatory HAUSGARTEN, in the eastern Fram Strait. Our results indicate that the overall density of megafauna was significantly lower in 2007 than in 2002, but was significantly higher in 2012, resulting in overall greater megafaunal density in 2012. Different species showed different patterns in population density, but the relative proportions of predator/scavengers and suspension-feeding individuals were both higher in 2012. Variations in megafaunal densities and proportions are likely due to variation in food input to the sea floor, which decreased slightly in the years preceding 2007 and was greatly elevated in the years preceding 2012. Both average evenness and diversity increased over the time period studied, which indicates that HG I may be food-limited and subject to bottom-up control. The community of HG I may be unique in its response to elevated food input, which resulted in higher evenness and diversity in 2012.

Description: In this paper we provide an overview of new knowledge on oxygen depletion (hypoxia) and related phenomena in aquatic systems resulting from the EU-FP7 project HYPOX (“In situ monitoring of oxygen depletion in hypoxic ecosystems of coastal and open seas, and landlocked water bodies”, www.hypox.net). In view of the anticipated oxygen loss in aquatic systems due to eutrophication and climate change, HYPOX was set up to improve capacities to monitor hypoxia as well as to understand its causes and consequences. Temporal dynamics and spatial patterns of hypoxia were analyzed in field studies in various aquatic environments, including the Baltic Sea, the Black Sea, Scottish and Scandinavian fjords, Ionian Sea lagoons and embayments, and Swiss lakes. Examples of episodic and rapid (hours) occurrences of hypoxia, as well as seasonal changes in bottom-water oxygenation in stratified systems, are discussed. Geologically driven hypoxia caused by gas seepage is demonstrated. Using novel technologies, temporal and spatial patterns of watercolumn oxygenation, from basin-scale seasonal patterns to meter-scale sub-micromolar oxygen distributions, were resolved. Existing multidecadal monitoring data were used to demonstrate the imprint of climate change and eutrophication on long-term oxygen distributions. Organic and inorganic proxies were used to extend investigations on past oxygen conditions to centennial and even longer timescales that cannot be resolved by monitoring. The effects of hypoxia on faunal communities and biogeochemical processes were also addressed in the project. An investigation of benthic fauna is presented as an example of hypoxia-devastated benthic communities that slowly recover upon a reduction in eutrophication in a system where naturally occurring hypoxia overlaps with anthropogenic hypoxia. Biogeochemical investigations reveal that oxygen intrusions have a strong effect on the microbially mediated redox cycling of elements. Observations and modeling studies of the sediments demonstrate the effect of seasonally changing oxygen conditions on benthic mineralization pathways and fluxes. Data quality and access are crucial in hypoxia research. Technical issues are therefore also addressed, including the availability of suitable sensor technology to resolve the gradual changes in bottom-water oxygen in marine systems that can be expected as a result of climate change. Using cabled observatories as examples, we show how the benefit of continuous oxygen monitoring can be maximized by adopting proper quality control. Finally, we discuss strategies for state-of-the-art data archiving and dissemination in compliance with global standards, and how ocean observations can contribute to global earth observation attempts.

Description: Spatial distribution patterns in macrobenthos were studied based on the material collected during the R.V. Polarstern expedition ARK-XXVII/2 in July 2012. Eleven stations along the latitudinal transect at the deep-sea observatory HAUSGARTEN in the Fram Strait were taken at depths of about 2500 m. Macrofauna was obtained using the USNEL box corer. A single core (0.25 m2) was taken at each station and four subcores (0.03 m2) were taken from each core and used for the quantitative analysis. The results suggest that the single highly variable macrobenthic community with the dominance of polychaetes Galathowenia fragilis and Myriochele heeri inhabits the studied area. The prevalence of a mosaic in the community structure with the grain size at least more than the size of a core was detected. However, several abundant species (e.g. the polychaetes Prionospio sp. and Galathowenia fragilis) tend to form patches at the scale less than a core (0.25 m2). Despite the lack of significant differences in species distribution patterns along the latitudinal transect, there is a slight difference in community structure between the northernmost and southernmost stations (~170 km apart), which can be explained by variations in environmental factors (e.g. higher food availability in the northern part of the transect).

Description: During a survey of the H°akon Mosby mud volcano (HMMV), located on the Bear Island fan in the southwest Barents Sea at �1250m water depth, different habitats inside the volcano caldera and outside it were hotographed using a towed camera platform, an Ocean Floor Observation System (OFOS). Three transects were performed across the caldera and one outside, in the background area, each transect was �2 km in length. We compared the density, taxa richness and diversity of nonsymbiotrophic megafauna in areas inside the volcano caldera with different bacterial mat and pogonophoran tubeworm cover. Significant variations in megafaunal composition, density and distribution were found between considered areas. Total megafaunal density was highest in areas of dense pogonophoran populations (mean 52.9 ind.m−2) followed by areas of plain light-coloured sediment that were devoid of bacterial mats and tube worms (mean 37.7 ind.m−2). The lowest densities were recorded in areas of dense bacterial mats (mean �1.4 ind.m−2). Five taxa contributed to most of the observed variation: the ophiuroid Ophiocten gracilis, lysianassid amphipods, the pycnogonid Nymphon macronix, the caprellid Metacaprella horrida and the fish Lycodes squamiventer. In agreement with previous studies, three zones within the HMMV caldera were distinguished, based on different habitats and megafaunal composition: “bacterial mats”, “pogonophoran fields” and “plain light-coloured sediments”. The zones were arranged almost concentrically around the central part of the caldera that was devoid of visible megafauna. The total number of taxa showed little variation inside (24 spp.) and outside the caldera (26 spp.). The density, diversity and composition of megafauna varied substantially between plain lightcoloured sediment areas inside the caldera and the HMMV background. Megafaunal density was lower in the background (mean 25.3 ind.m−2) compared to areas of plain light-coloured sediments inside the caldera. So the effect of the mud-volcano environment on benthic communities is expressed in increasing of biomass, changing of taxa composition and proportions of most taxonomic groups.

Description: This study compares the macrofaunal communities along two bathymetric transects (1000 – 2500 m water depth) in predominantly ice-covered western (offshore Greenland) and generally ice-free eastern (offshore Svalbard) regions of the Fram Strait. Material was collected using an USNEL 0.25 m2 box corer and all sediment samples were processed through a 500-μm sieve. A total of 1671 organisms from 169 species were found. Densities off Greenland were generally lower than those observed off Svalbard. On both sides of the Fram Strait, density, biomass and biodiversity generally decreased with increasing water depth. An exception was observed at the deepest station off Greenland (2500 m water depth), which was located within the Marginal Ice Zone. At this station, macrofaunal density was elevated (992 ± 281 ind. m−2) compared to the adjacent shallower sampling areas off Greenland (272 ± 208 ind. m−2 to 787 ± 172 ind. m−2) and the deeper stations (2000 and 2500 m water depth) off Svalbard (552 ± 155 ind. m−2 and 756 ± 182 ind. m−2). The most abundant species along both transects was the polychaete Galathowenia fragilis (off Greenland: 288 ind. m−2, off Svalbard: 740 ind. m−2). Sea ice coverage and water depth, as well as the associated food availability at the seafloor, seem to be crucial factors driving the macrofaunal community patterns. A strong pelago-benthic coupling is observed to be typical in Arctic deep-sea ecosystems, and is also confirmed by our study.