Weak seasonality in benthic food web structure within an Arctic inflow shelf region

The Arctic Ocean is characterized by pronounced seasonality in the quantity and quality of organic matter exported from the surface ocean. While it is well established that changes in food availability can alter the abundance, biomass and function of benthic organisms, the impact on food web structure is not well studied. We used bulk carbon and nitrogen stable isotope analysis to assess the quantity and quality of sediment organic matter and structure of the benthic food web in four seasons within the Northern Barents Sea (76°N − 82 °C). Despite a highly seasonal vertical flux, we found that the organic carbon and chlorophyll-a content of surface sediments was seasonally stable, suggesting a lack of seasonality in food availability at the seafloor. However, organic biomarkers indicate that the quality of sediment organic matter increased to a maximum in August and December, up to 6 months after the spring bloom. The seasonal stability of food quantity was mirrored in food-web structure (e.g., total isotopic range, number of trophic levels) which did not change significantly across sampling periods. We expected that suspension and deposit feeders would respond more readily to seasonal changes in food quality compared to predators. However, we observed no significant seasonal changes in the trophic levels or isotopic niche areas of benthic functional groups. The centroids of isotopic niches of all benthic functional groups shifted seasonally by <2 ‰ along the δ13C-axis, suggesting minimal shifts in carbon resource use. Because the northern Barents Sea experiences significant changes in seasonal sea ice cover, we expected that stable-isotope ratios of benthic organisms would show an increased consumption of sympagic-derived organic matter through less negative δ13C values in early spring and summer. However, only two taxa (the soft coral Gersemia spp. and bivalves in the family Yoldiidae) showed 13C-enrichment in spring or summer consistent with the assimilation of sympagic-derived organic matter, despite previous evidence suggesting widespread use of this carbon source. Overall, our results show that there is an apparent de-coupling in time between pelagic processes and benthic food-webs in which the accumulation and assimilation of high-quality organic matter occurs for benthos during the fall and early winter months when there is little to no fresh organic matter generated at the surface. This temporal mismatch highlights the importance of considering the timescales over which components of the marine ecosystem respond to short-term environmental changes and the methods employed to assess seasonality.Weak seasonality in benthic food web structure within an Arctic inflow shelf regionpublishedVersio

Isotopic turnover of polar cod (Boreogadus saida) muscle determined through a controlled feeding experiment

Polar cod (Boreogadus saida) is an important trophic link within Arctic marine food webs and is likely to experience diet shifts in response to climate change. One important tool for assessing organism diet is bulk stable isotope analysis. However, key parameters necessary for interpreting the temporal context of stable isotope values are lacking, especially for Arctic species. This study provides the first experimental determination of isotopic turnover (as half-life) and trophic discrimination factors (TDFs) of both δ13C and δ15N in adult polar cod muscle. Using a diet enriched in both 13C and 15N, we measured isotopic turnover times of 61 and 49 days for δ13C and δ15N, respectively, with metabolism accounting for >94% of the total turnover. These half-life estimates are valid for adult polar cod (>3 years) experiencing little somatic growth. We measured TDFs in our control of 2.6‰ and 3.9‰ for δ13C and δ15N, respectively, and we conclude that applying the commonly used TDF of ~1‰ for δ13C for adult polar cod may lead to misrepresentation of dietary carbon source, while the use of 3.8‰ for δ15N is appropriate. Based on these results, we recommend that studies investigating seasonal shifts in the diet of adult polar cod sample at temporal intervals of at least 60 days to account for isotopic turnover in polar cod muscle. Although isotopic equilibrium was reached by the fish in this study, it was at substantially lower isotope values than the diet. Additionally, the use of highly enriched algae in the experimental feed caused very high variability in diet isotope values which precluded accurate calculation of TDFs from the enriched fish. As a result of the challenges faced in this study, we discourage the use of highly enriched diets for similar experiments and provide recommendations to guide the design of future isotopic turnover experiments.publishedVersio

High Arctic Mytilus spp.: occurrence, distribution and history of dispersal

Many marine species are known to change their distribution in response to changing climatic conditions. One such example is the blue mussel Mytilus spp., spreading northward coincident with an increase in ocean temperatures. On Svalbard, the frst living specimens of Mytilus spp. were discovered in 2004. Here we present an analysis of the current distribution of Mytilus spp. on Svalbard, with a focus on the west coast of Spitsbergen where strong Atlantifcation has been documented over the last few decades. We conducted diver-based surveys to develop a distributional map and to compare the current distribution with that of the Holocene. Furthermore, we investigate the recent history of recruitment of mussels on Svalbard to help identify invasion pathways. Our results show that blue mussels have been present on the archipelago at least since 2000 and are widespread along the west coast today. We also present evidence of local reproduction in one of the sites explored

Effects of Detached Seaweeds on Structure and Function of Arctic Intertidal Soft-Bottom Communities

Expected lower sea-ice cover and increased storm frequency have led to projections of an increase in seaweed detritus in Arctic marine systems in the near future. To assess whether detached seaweed affects structural and functional traits of species assemblages in soft-bottom habitats, comparable experiments were run in two intertidal sites (Longyearbyen and Thiisbukta) on Svalbard. At each site, we fixed nets containing the locally dominating seaweeds Saccharina latissima and Desmarestia aculeata (Thiisbukta) or Fucus sp. (Longyearbyen) to intertidal mud flats. Empty nets were fixed as procedural controls at both sites. After 2.5 months, one sediment core was taken from each manipulated plot and the number of individuals, dry mass, and average length of each encountered animal taxon were recorded. The same measurements were taken from cores collected from unmanipulated areas at each site, both at the start and end of the experiment. Abundance data were used to calculate estimates of diversity (Shannon-index, evenness, and taxon richness), while initial and final average length measurements were used to estimate taxon-specific growth. Log response ratios of initial and final abundance in unmanipulated areas were used to estimate magnitude and direction of the effect of change in community traits over time, serving as a reference to log response ratios estimating manipulated seaweed effects. In Longyearbyen, the presence of detached seaweeds reduced abundance and dry mass by, on average, 46 and 70%, respectively, compared to unmanipulated benthic communities. In Thiisbukta, the presence of seaweeds enhanced evenness, on average by 16%, but reduced abundance and growth of benthic fauna by, on average, 31 and 86%, respectively. Seaweed effects were generally smaller in Thiisbukta than in Longyearbyen. At both sites, time effects were generally opposite in direction to those caused by the seaweed treatments, yet of similar or larger magnitude. Through reversing temporal dynamics of several of the tested community traits, detached seaweeds strongly modified the structure and functioning of soft-bottom species assemblages at both intertidal sites. We suggest that the detected effects possibly result from seaweed-induced changes in environmental conditions and/or physical disturbance as underlying processes

Limited predatory effects on infaunal macrobenthos community patterns in intertidal soft-bottom of Arctic coasts

Predation shapes marine benthic communities and affects prey species population dynamics in tropic and temperate coastal systems. However, information on its magnitude in systematically understudied Arctic coastal habitats is scarce. To test predation effects on the diversity and structure of Arctic benthic communities, we conducted caging experiments in which consumers were excluded from plots at two intertidal sedimentary sites in Svalbard (Longyearbyen and Thiisbukta) for 2.5 months. Unmanipulated areas served as controls and partial (open) cages were used to estimate potential cage effects. At the end of the experiment, we took one sediment core from each plot and quantified total biomass and the number of each encountered taxon. At both sites, the experimental exclusion of predators slightly changed the species composition of communities and had negligible effects on biomass, total abundance, species richness, evenness, and Shannon Index. In addition, we found evidence for cage effects, and spatial variability in the intensity of the predation effects was identified. Our study suggests that predators have limited effects on the structure of the studied intertidal macrobenthic Arctic communities, which is different from coastal soft-bottom ecosystems at lower latitudes

Lack of strong seasonality in macrobenthic communities from the northern Barents Sea shelf and Nansen Basin

The Barents Sea has been coined ‘the Arctic hotspot’ of climate change due to the rapidity with which environmental changes are taking place. This transitional domain from Atlantic to Arctic waters is home to highly productive benthic communities. This system strongly fluctuates on a seasonal basis in its sympagic-pelagicbenthic coupling interactions, with potential effects on benthic standing stocks and production. Recent discoveries have questioned the marked seasonality for several high Arctic seafloor communities in coastal waters of Svalbard. Still, the seasonal variability of benthic process in the extensive Barents Sea open shelf remains poorly understood. Therefore, we studied the seasonality of macrofauna communities along a transect in the northwestern Barents Sea comprising two hydrographic domains (Arctic vs. Atlantic Water, across the Polar Front) and three geomorphological settings (shelf, continental slope and abyssal plain). Overall, we did not find strong signs of seasonal variation in taxonomic community structure and functional diversity. However, we found some weak signs of seasonality when examining each station separately, especially at a station close to the Polar Front, with high seasonal fluctuations in abiotic drivers indicating a stronger pelagic-benthic coupling. The lack of seasonality found both at the shelf stations south and north of the Polar Front could be related to organic matter stored in the sediments, reflected in constant levels of total organic carbon in surface sediment across time for all stations. We did observe, as expected, highly spatially structured environmental regimes and macrofauna communities associated to them from shelf to slope and basin locations. Understanding the underlying spatiotemporal mechanisms by which soft-bottom benthic communities are structured along environmental gradients is necessary to predict future impacts of climate change in this area. Our results indicate that short-term climate driven changes in the phenology of pelagic ecosystem components might not be directly reflected in the Arctic benthic system, as seafloor processes seem to be partially decoupled from those in the overlying water

Meroplankton Diversity, Seasonality and Life-History Traits Across the Barents Sea Polar Front Revealed by High-Throughput DNA Barcoding

In many species of marine benthic invertebrates, a planktonic larval phase plays a critical role in dispersal. Very little is known about the larval biology of most species, however, in part because species identification has historically been hindered by the microscopic size and morphological similarity among related taxa. This study aimed to determine the taxonomic composition and seasonal distribution of meroplankton in the Barents Sea, across the Polar Front. We collected meroplankton during five time points seasonally and used high-throughput DNA barcoding of individual larvae to obtain species-level information on larval seasonality. We found that meroplankton was highly diverse (72 taxa from eight phyla) and present in the Barents Sea year-round with a peak in abundance in August and November, defying the conventional wisdom that peak abundance would coincide with the spring phytoplankton bloom. Ophiuroids, bivalves, and polychaetes dominated larval abundance while gastropods and polychaetes accounted for the bulk of the taxon diversity. Community structure varied seasonally and total abundance was generally higher south of the Polar Front while taxon richness was overall greater to the north. Of the species identified, most were known inhabitants of the Barents Sea. However, the nemertean Cephalothrix iwatai and the brittle star Ophiocten gracilis were abundant in the meroplankton despite never having been previously recorded in the northern Barents Sea. The new knowledge on seasonal patterns of individual meroplanktonic species has implications for understanding environment-biotic interactions in a changing Arctic and provides a framework for early detection of potential newcomers to the system

Horizon scanning of potential threats to high-Arctic biodiversity, human health and the economy from marine invasive alien species: A Svalbard case study

The high Arctic is considered a pristine environment compared with many other regions in the northern hemisphere. It is becoming increasingly vulnerable to invasion by invasive alien species (IAS), however, as climate change leads to rapid loss of sea ice, changes in ocean temperature and salinity, and enhanced human activities. These changes are likely to increase the incidence of arrival and the potential for establishment of IAS in the region. To predict the impact of IAS, a group of experts in taxonomy, invasion biology and Arctic ecology carried out a horizon scanning exercise using the Svalbard archipelago as a case study, to identify the species that present the highest risk to biodiversity, human health and the economy within the next 10 years. A total of 114 species, currently absent from Svalbard, recorded once and/or identified only from environmental DNA samples, were initially identified as relevant for review. Seven species were found to present a high invasion risk and to potentially cause a significant negative impact on biodiversity and five species had the potential to have an economic impact on Svalbard. Decapod crabs, ascidians and barnacles dominated the list of highest risk marine IAS. Potential pathways of invasion were also researched, the most common were found associated with vessel traffic. We recommend (i) use of this approach as a key tool within the application of biosecurity measures in the wider high Arctic, (ii) the addition of this tool to early warning systems for strengthening existing surveillance measures; and (iii) that this approach is used to identify high-risk terrestrial and freshwater IAS to understand the overall threat facing the high Arctic. Without the application of biosecurity measures, including horizon scanning, there is a greater risk that marine IAS invasions will increase, leading to unforeseen changes in the environment and economy of the high Arctic

Seafloor warm water temperature anomalies impact benthic macrofauna communities of a high-Arctic cold-water fjord

Amid the alarming atmospheric and oceanic warming rates taking place in the Arctic, western fjords around the Svalbard archipelago are experiencing an increased frequency of warm water intrusions in recent decades, causing ecological shifts in their ecosystems. However, hardly anything is known about their potential impacts on the until recently considered stable and colder northern fjords. We analyzed macrobenthic fauna from four locations in Rijpfjorden (a high-Arctic fjord in the north of Svalbard) along its axis, sampled intermittently in the years 2003, 2007, 2010, 2013 and 2017. After a strong seafloor warm water temperature anomaly (SfWWTA) in 2006, the abundance of individuals and species richness dropped significantly across the entire fjord in 2007, together with diversity declines at the outer parts (reflected in Shannon index drops) and increases in beta diversity between inner and outer parts of the fjord. After a period of three years with stable water temperatures and higher sea-ice cover, communities recovered through recolonization processes by 2010, leading to homogenization in community composition across the fjord and less beta diversity. For the last two periods (2010-2013 and 2013-2017), beta diversity between the inner and outer parts gradually increased again, and both the inner and outer sites started to re-assemble in different directions. A few taxa began to dominate the fjord from 2010 onwards at the outer parts, translating into evenness and diversity drops. The inner basin, however, although experiencing strong shifts in abundances, was partially protected by a fjordic sill from impacts of these temperature anomalies and remained comparatively more stable regarding community diversity after the disturbance event. Our results indicate that although shifts in abundances were behind important spatio-temporal community fluctuations, beta diversity variations were also driven by the occurrence-based macrofauna data, suggesting an important role of rare taxa. This is the first multidecadal time series of soft-bottom macrobenthic communities for a high-Arctic fjord, indicating that potential periodic marine heatwaves might drive shifts in community structure, either through direct effects from thermal stress on the communities or through changes in environmental regimes led by temperature fluctuations (i.e. sea ice cover and glacial runoff, which could lead to shifts in primary production and food supply to the benthos). Although high-Arctic macrobenthic communities might be resilient to some extent, sustained warm water anomalies could lead to permanent changes in cold-water fjordic benthic systems

Uptake of sympagic organic carbon by the Barents Sea benthos linked to sea ice seasonality

On Arctic shelves, where primary production occurs in both the pelagic and sympagic (ice-associated) habitats, sympagic organic material (OM) can constitute a disproportionate fraction of benthic diets due to higher sinking rates and lower grazing pressure than pelagic OM. Less documented is how sympagic OM assimilation across feeding guilds varies seasonally and in relation to sea ice duation. We therefore investigated the relative abundance of sympagic vs pelagic OM in Barents Sea shelf megabenthos in the summer and winter of 2018 and 2019, from 10 stations where sea ice duration ranged from 0 to 245 days per year. We use highly branched isoprenoids, which are lipid biomarkers produced with distinct molecular structures by diatoms in sea ice and the water column, to determine the ratio of sympagic-to-pelagic OM assimilated by benthic organisms. From 114 samples of 25 taxa analysed, we found that the proportion of sympagic OM assimilated ranged from 0.4% to 95.8% and correlated strongly (r2 = 0.754) with the duration of sea ice cover. The effect of sea ice duration was more evident in fauna collected during summer than winter, indicating that sympagic signals are more evident in the summer than in the winter at higher latitudes. Our data show that sympagic production can supply a high fraction of carbon for Barents Sea benthos, although this is highly variable and likely dependent on availability and patchiness of sympagic OM deposition. These results are comparable to similar studies conducted on benthos in the Pacific Arctic and highlight the variable importance of sympagic OM in the seasonal ice zone of Arctic inflow shelves, which are the Arctic regions with highest rates of sea ice loss.publishedVersio