Beschreibung: 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.

Beschreibung: Deep-sea regions provide vast ecosystem services such as biological habitat and nutrient cycling. Even though being threatened by climate change and facing possible biodiversity loss, these deep-sea ecosystems are poorly understood. So are macrobenthic communities and their functions within these ecosystems. Biodiversity and ecosystem function relationships as well as their link to environmental drivers can be assessed with the biological trait analysis. We used this approach for the first time for macrofauna assemblages across the deep Fram Strait between Greenland and Svalbard (1000–5500 m water depth) to evaluate their community-specific function from the upper continental slope down to the deepest known Arctic depression, the Molloy Deep. We aimed to investigate whether there are changes in benthic functioning along the bathymetric gradient and if so, which environmental stressors may drive these changes. In total, 16 stations were sampled with a giant box corer (0.25 m2) in 2016 and 2018. Sediments were sieved through a 0.5 mm mesh size sieve and fauna was identified to lowest possible taxonomic entity. Functions of species were characterized by using six traits split in 24 modalities gathered in a fuzzy coded species × traits array. Environmental parameters shaping the benthic habitat and reflecting food availability were gathered from remote sensing, mooring deployments, and sediment sampling. A distance-based redundancy analysis indicated near-bottom water temperature, seabed inclination, water depth as well as phytodetritial matter at the sea surface and seafloor (indicating food availability) to be the best variables explaining the trait and station distribution. Stations clustered into three groups based on their trait composition. Shallower stations characterized by high chlorophyll a concentration with large organisms, living within the sediment as well as predating specimens clustered in one group. A second group was characterized by stations with low chlorophyll a concentration and medium-sized, suspension feeding, epifaunal living macrofauna. A third group comprised stations with water depths ≥ 3000 m and was dominated by medium sized, surface deposit feeding and infaunal living specimens. Overall, the functional structure of macrofauna communities in the Fram Strait followed a food availability-driven gradient. Based on the relationship between sea ice, surface water primary production and food availability at the seafloor, these results point to macrobenthos being sensible to predicted anthropogenically generated environmental variations in polar regions. Alterations in benthic ecosystem functions might be expected when environmental conditions change.

Beschreibung: 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.

Beschreibung: Bathymetric patterns in standing stocks and diversity are a major topic of investigation in deep-sea biology. From the literature, responses of metazoanmeiofauna and nematodes to bathymetric gradients arewell studied, with a general decrease in biomass and abundance with increasing water depth, while bathymetric diversity gradients often, although it is not a rule, show a unimodal pattern. Spatial distribution patterns of nematode communities along bathymetric gradients are coupled with surface-water processes and interacting physical and biological factors within the benthic system. We studied the nematode communities at the Long-TermEcological Research (LTER) observatory HAUSGARTEN, located in the FramStrait at theMarginal Ice Zone, with respect to their standing stocks as well as structural and functional diversity.We evaluated whether nematode density, biomass and diversity indices, such asH0, Hinf, EG(50), Θ−1, are linkedwith environmental conditions along a bathymetric transect spanning from 1200mto 5500mwater depth. Nematode abundance, biomass and diversity, as well as food availability from phytodetritus sedimentation (indicated by chloroplastic pigments in the sediments), were higher at the stations located at upper bathyal depths (1200–2000 m) and tended to decrease with increasing water depth. A faunal shift was found below 3500 m water depth, where genus composition and trophic structure changed significantly and structural diversity indices markedly decreased. A strong dominance of very fewgenera and its high turnover particularly at the abyssal stations (4000–5500 m) suggests that environmental conditions were rather unfavorable for most genera. Despite the high concentrations of sediment-bound chloroplastic pigments and elevated standing stocks found at the deepest station (5500 m), nematode genus diversity remained the lowest compared to all other stations. This study provides a further insight into the knowledge of deep-sea nematodes, their diversity patterns and a deeper understanding of the environmental factors shaping nematodes communities at bathyal and abyssal depths.

Beschreibung: Benthic fauna constantly modifies their physical, chemical and biological environment. The permanent biological reworking of surface sediments mediates biogeochemical processes at the seafloor and is, therefore, of global importance. There are numerous studies measuring the rate and extent of bioturbation worldwide, however, information on mixing rates in the deep ocean and especially in the Polar Regions are extremely scarce; to our knowledge there is, by now, only a single study providing bioturbation rates from the deep Arctic Ocean. The present study presents mixing rates and mixed layer depths for the deep seafloor at the LTER (Long-Term Ecological Research) observatory HAUSGARTEN in Fram Strait, Arctic Ocean. Two stations at similar water depths (2400 m and 2500 m water depth, respectively) but approx. 55 km apart from each other were chosen to carry out long-term (2 and 4 years, respectively) in situ bioturbation experiments using luminophores as a tracer. Biodiffusion-like mixing rates Db at the experimental sites were rather similar (0.26 cm2 a-1 at HG-IV; 0.28 cm2 a-1 at S3); slightly (non-significantly) higher Db values at the southern HAUSGARTEN site S3 could be explained by more favorable environmental conditions and related differences in the faunal composition. Indications for a non-local transport of sediment particles from the surface to deeper parts of the sediment, resulting in higher values for the Non-Local Index (NLI), could only be found for the central HAUSGARTEN site HG-IV. Elevated densities of burrowing megafauna at HG-IV, compared to S3, might be responsible for the subsurface maxima in luminophore distribution and comparably higher NLI values at the central HAUSGARTEN site (5.37 at HG-IV; 3.26 at S3). Mixed layer depths L at the two sites were almost identical; considerable mixing of surface sediments occurred down to max. 6-7 cm sediment depth.

Beschreibung: In this paper the concept of resilience is discussed on the base of 13 case studies from the German branch of the International Long-Term Ecological Research Program. In the introduction the resilience approach is presented as one possibility to describe ecosystem dynamics. The relations with the concepts of adaptability and ecological integrity are discussed and the research questions are formulated. The focal research objectives are related to the conditions of resilient behaviour of ecosystems, the role of spatio-temporal scales, the differences between short- or long-term dynamics, the basic methodological requirements to exactly define resilience, the role of the reference state and indicators and the suitability of resilience as a management concept. The main part of the paper consists of 13 small case study descriptions, which demonstrate phase transitions and resilient dynamics of several terrestrial and aquatic ecosystems at different time scales. In the discussion, some problems arising from the interpretation of the time series are highlighted and discussed. The topics of discussion are the conceptual challenges of the resilience approach, methodological problems, the role of indicator selection, the complex interactions between different disturbances, the significance of time scales and a comparison of the case studies. The article ends with a conclusion which focuses on the demand to link resilience with adaptability, in order to support the long-term dynamics of ecosystem development.

Beschreibung: 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.

Beschreibung: 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.