Multidecadal stability of benthic community structure in a high-Arctic glacial fjord (van Mijenfjord,Spitsbergen)

Long-term change in benthic community structure may have significant impact on ecosystem functions. Accelerating climate change and increased human activity in the Arctic suggest that benthic communities in this region may be expected to exhibit change over time scales coinciding with these potential stressors. In 2000 and 2001, we resampled the soft-sediment communities of van Mijenfjord, a semi-closed (silled) fjord system on the west coast of Spitsbergen, following initial surveys in 1980. Multivariate community analyses and biodiversity indices identified distinct regions within the fjord. The communities characteristic of two regions were very similar to those sampled 20 years earlier. Regions corresponded with fjord basins and to community patterns and diversity gradients identified for many other Arctic fjords. Benthic communities in open (unsilled) fjords in the area have recently been shown to respond to decadal scale climatic fluctuation. We suggest that semi-closed fjords may be less susceptible to this type of environmental variability, and that communities are shaped by an interaction of impacts from local topography, glacial runoff, local circulation patterns, and faunal life-history traits. Open and closed fjords may respond to climatic warming trends in different ways, resulting in a subsequent divergence in spatial patterns of resident communities.     

Distinct or similar? Soft bottom polychaete diversity in Arctic and Antarctic glacial fjords

The main aim of this study was to compare the polychaete communities in two similar polar areas: an Arctic fjord, Hornsund (Svalbard) and an Antarctic fjord, Ezcurra Inlet (South Shetlands). This is the first attempt to compare Arctic and Antarctic diversity based on fully comparable datasets. Forty van Veen grab samples were collected in each fjord: twenty replicates were taken in each of two fjord areas characterized by a different level of glacial disturbance—in the inner (glacial bay) and outer (fjord mouth) region of both fjords, from depths of about 100 m in 2005 (Hornsund) and in 2007 (Ezcurra Inlet). In the glacial bays, species richness and diversity were significantly higher in Ezcurra Inlet than in Hornsund due to higher rate of glacial disturbance in the latter one. In the outer areas, species richness was similar in both fjords, although diversity values were higher in Ezcurra Inlet. Polychaete species richness in the habitats characterized by similar level of disturbance (outer areas of the fjords) was the same in both polar regions. At this small scale, where community drivers are very similar, the species richness seems to be independent from the local or regional species pool.     

Blue carbon gains from glacial retreat along Antarctic fjords: What should we expect?

Rising atmospheric CO₂ is intensifying climate change but it is also driving global and particularly polar greening. However, most blue carbon sinks (that held by marine organisms) are shrinking, which is important as these are hotspots of genuine carbon sequestration. Polar blue carbon increases with losses of marine ice over high latitude continental shelf areas. Marine ice (sea ice, ice shelf and glacier retreat) losses generate a valuable negative feedback on climate change. Blue carbon change with sea ice and ice shelf losses has been estimated, but not how blue carbon responds to glacier retreat along fjords. We derive a testable estimate of glacier retreat driven blue carbon gains by investigating three fjords in the West Antarctic Peninsula (WAP). We started by multiplying ~40 year mean glacier retreat rates by the number of retreating WAP fjords and their time of exposure. We multiplied this area by regional zoobenthic carbon means from existing datasets to suggest that WAP fjords generate 3,130 tonnes of new zoobenthic carbon per year (t zC/year) and sequester >780 t zC/year. We tested this by capture and analysis of 204 high resolution seabed images along emerging WAP fjords. Biota within these images were identified to density per 13 functional groups. Mean stored carbon per individual was assigned from literature values to give a stored zoobenthic Carbon per area, which was multiplied up by area of fjord exposed over time, which increased the estimate to 4,536 t zC/year. The purpose of this study was to establish a testable estimate of blue carbon change caused by glacier retreat along Antarctic fjords and thus to establish its relative importance compared to polar and other carbon sinks.     

Environmental drivers of macroinvertebrate communities in high Arctic rivers (Svalbard)

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From the worm’s point of view. I: Environmental settings of benthic ecosystems in Arctic fjord (Hornsund,Spitsbergen)

As a consequence of ongoing climate warming, nearly all tidal glaciers in Arctic are retreating; hence, the seascape of glacial fjords is changing in many aspects. We took the example of Hornsund, the well-studied Svalbard fjord, with over 30 years of almost continuous observations of marine system. Recent data were collected during summer oceanographic surveys between 2001 and 2013 and compared with archival data from 1980s. As most of the phenomena connected with the warming happen at the sea surface (ice, wind, waves, surface currents, brackish water), we were interested, how the presumably stable, near-bottom waters in fjords behave, what are the environmental changes that are experienced directly by the worms living in the sediment. We have found that both the inner fjord basins (usually regarded as stable) and the outer fjord parts (exposed to the direct influence of shelf waters) has changed. Warming was documented in the inner basins, while cooling and warming episodes were recorded in the outer parts of the fjord. We demonstrate that following the increase melting and retreat of the glaciers, the area of shallows increased, salinity decreased and temperature increased—partly due to the advection of Atlantic waters from the shelf. Observed changes are in accordance with the model of arctic fjords evolution towards boreal ones associated with increased organic matter turnover. The observed changes are most likely typical for all cold water, glaciated fjords that are exposed to climate warming.     

Benthic infaunal community variability on the northern Svalbard shelf

Marine benthic communities are effective indicators of environmental change. Yet in the Arctic, there are few empirical tests of how sustained climatic change may influence community structure. Northern Svalbard is influenced by both warm Atlantic and cold Arctic water masses, providing an opportunity to assess potential effects of long-term environmental changes by examining spatial variation in community structure. We examined benthic macroinfaunal communities and sediment pigments under Atlantic and Arctic water masses on the northern shelf and fjords of Svalbard. We report on infaunal biomass, abundance, species composition, and diversity at 10 stations spanning 79°–81°N and ranging in depth from 200 to 500?m. Benthic biomass averaged 128?g?WW?m^?2 (48–253?g?WW?m^?2), mean density was 3,635?ind.?m^?2 (780–7,660?ind.?m^?2), and species richness varied from 45 to 136?taxa?stn.^?1. Abundance-based community structure clustered stations in groups related to water mass characteristics, with Atlantic and Arctic shelf stations being well distinguished from each other. Dominant taxa were different in Atlantic- and Arctic-influenced locations. Faunal biomass was highest in the Atlantic-influenced fjords, followed by Arctic fjords and Arctic shelf stations, with Atlantic shelf stations having the lowest biomass. Species richness and diversity were inversely related to biomass. Patterns in faunal biomass were strongly correlated with sedimentary pigments (R ²?=?0.74 for chl a and R ²?=?0.77 for phaeopigments), with large differences in sedimentary pigment concentration among stations. These relationships suggest that benthic fauna on the northern Svalbard shelf are food limited and dependent on predictable, albeit episodic, delivery of organic matter from the water column.     

Distribution patterns of polychaete fauna in an Arctic fjord (Hornsund, Spitsbergen)

Polychaetes are one of the most important groups of benthic organisms in marine ecosystems. They dominate on the Arctic shelf and play an important role in ecosystem functioning. This study focuses on the polychaete biodiversity and their distribution patterns in Hornsund, an open glacial fjord, in western Spitsbergen and provides important baseline data for future studies of temporal fluctuations in benthic fauna. The main aim of this study was to assess how the polychaete abundance, biomass, diversity, community structure, and function vary along the Hornsund fjord’s axis, in relation to the environmental factors. Eighty-eight polychaete taxa were identified; an average density was 457 ind. m^?2 ± 237.5 SD. Three assemblages were distinguished (INNER, MIDDLE, and OUTER) along the fjord axis, reflecting the intensity of glacial disturbance. A clear division between the highly disturbed inner part of the fjord (Brepollen) and the less impacted middle and outer parts was observed. Continuous gradients in abundance, biomass, and diversity were found with all those values diminishing toward the inner region of the fjord. The polychaete assemblages' indices were significantly correlated with bottom temperature, sediment characteristics (grain size), and distance to the glacier (longitude). No significant correlations were found with depth or total organic carbon content. Carnivore and motile surface deposit feeding polychaete species dominated in the areas close to the glaciers, while the OUTER community was dominated by carnivores and surface sessile and discretely motile species, and had more complex trophic structure, with multiple species representing different functional groups including carnivores, sessile, discretely motile, and motile surface deposit feeders and motile burrowers.     

Polar zoobenthos blue carbon storage increases with sea ice losses,because across‐shelf growth gains from longer algal blooms outweigh ice scour mortality in the shallows

One of the major climate‐forced global changes has been white to blue to green; losses of sea ice extent in time and space around Arctic and West Antarctic seas has increased open water and the duration (though not magnitude) of phytoplankton blooms. Blueing of the poles has increases potential for heat absorption for positive feedback but conversely the longer phytoplankton blooms have increased carbon export to storage and sequestration by shelf benthos. However, ice shelf collapses and glacier retreat can calve more icebergs, and the increased open water allows icebergs more opportunities to scour the seabed, reducing zoobenthic blue carbon capture and storage. Here the size and variability in benthic blue carbon in mega and macrobenthos was assessed in time and space at Ryder and Marguerite bays of the West Antarctic Peninsula (WAP). In particular the influence of the duration of primary productivity and ice scour are investigated from the shallows to typical shelf depths of 500 m. Ice scour frequency dominated influence on benthic blue carbon at 5 m, to comparable with phytoplankton duration by 25 m depth. At 500 m only phytoplankton duration was significant and influential. WAP zoobenthos was calculated to generate ~10⁷, 4.5 × 10⁶ and 1.6 × 10⁶ tonnes per year (between 2002 and 2015) in terms of production, immobilization and sequestration of carbon respectively. Thus about 1% of annual primary productivity has sequestration potential at the end of the trophic cascade. Polar zoobenthic blue carbon capture and storage responses to sea ice losses, the largest negative feedback on climate change, has been underestimated despite some offsetting of gain by increased ice scouring with more open water. Equivalent survey of Arctic and sub‐Antarctic shelves, for which new projects have started, should reveal the true extent of this feedback and how much its variability contributes to uncertainty in climate models.     

Marine‐terminating glaciers sustain high productivity in Greenland fjords

Accelerated mass loss from the Greenland ice sheet leads to glacier retreat and an increasing input of glacial meltwater to the fjords and coastal waters around Greenland. These high latitude ecosystems are highly productive and sustain important fisheries, yet it remains uncertain how they will respond to future changes in the Arctic cryosphere. Here we show that marine‐terminating glaciers play a crucial role in sustaining high productivity of the fjord ecosystems. Hydrographic and biogeochemical data from two fjord systems adjacent to the Greenland ice sheet, suggest that marine ecosystem productivity is very differently regulated in fjords influenced by either land‐terminating or marine‐terminating glaciers. Rising subsurface meltwater plumes originating from marine‐terminating glaciers entrain large volumes of ambient deep water to the surface. The resulting upwelling of nutrient‐rich deep water sustains a high phytoplankton productivity throughout summer in the fjord with marine‐terminating glaciers. In contrast, the fjord with only land‐terminating glaciers lack this upwelling mechanism, and is characterized by lower productivity. Data on commercial halibut landings support that coastal regions influenced by large marine‐terminating glaciers have substantially higher marine productivity. These results suggest that a switch from marine‐terminating to land‐terminating glaciers can substantially alter the productivity in the coastal zone around Greenland with potentially large ecological and socio‐economic implications.     

Main differences in macrobenthos and benthic communities of the arctic and antarctic,as illustrated by comparison of the Laptev and Weddell sea faunas

A comparison of the benthic macrofaunas of the Laptev and Weddell seas revealed considerable differences in the species composition of the Arctic and Antarctic shelf faunas. In the Arctic, the infauna has the highest species diversity and plays the main role in the benthic communities, whereas in the Antarctic the epifauna predominates. The main reasons for these essential differences are (1) the different sediment composition at the time of formation of cold-water faunas, (2) the different productivity of the ecosystems, (3) the different extent of exchange with the adjacent oceans, and (4) the different history of the origin of both faunas.     

Tanaidacea (Crustacea,Malacostraca) of two polar fjords: Kongsfjorden (Arctic) and Admiralty Bay (Antarctic)

The tanaidacean faunas of two polar fjords, Kongsfjorden (Arctic) and Admiralty Bay (Antarctic), were compared. The results show that Tanaidacea in the southern fjord are more diversified than those in the northern one. This difference is especially evident in species richness (12 species vs 3 species, respectively), but is not too significant in terms of diversity. In both polar fjords, the highest densities of tanaids were noted in sites where mud was swept off by the water current or eroded off steep bottom expoing coarser mineral particles used for tube building. The distribution of tanaids is suggested to be based on the joint action of inter alia factors such as the quality of bottom sediments, sedimentation ratio, accessibility of food, predation pressure, behaviour and heterogeneity of habitats.An erratum to this article can be found at     

ROV observations of benthic fishes in the Northwind and Canada Basins, Arctic Ocean

Abyssal and midslope Arctic benthic fishes were sampled nonquantitatively by still photography and videography from a remotely operated vehicle (ROV) between 13 August and 7 September 2002. Species diversity was low: only six putative species were seen, including Lycodes frigidus Collett 1879, Lycodes sp., Rhodichthys regina Collett 1879, Paraliparis bathybius (Collett 1879), Raja (Amblyraja) hyperborea Collett 1879, and Cottunculus species diversity varied among stations sampled. An analysis of 1-min segments of videotape from one long ROV dive suggested that habitat selection by demersal nekton in these environments may be associated with the presence, absence, or density of other benthic animals.     

Variability of protistan and bacterial communities in two Arctic fjords (Spitsbergen)

Krossfjorden and Kongsfjorden are Arctic fjords on the western side of Spitsbergen. These fjords share a common mouth to the open sea, and both are influenced by the input of sediment-rich glacial meltwater leading to decreased surface salinity, increased turbidity and decreased light penetration during summer. Earlier classical taxonomic studies had described the pelagic protistan composition of the Kongsfjorden during summer, revealing the dominance of flagellates of often unresolved taxonomic origin. Only little information existed on microbial eukaryote composition of the Krossfjorden as well as the bacterial composition of both fjords. The aim of the present study was to analyze and compare surface summertime protistan and bacterial communities in both fjords, using molecular approaches (16S and 18S rRNA DGGE, sequencing). Samples were collected three times a week from the central Kongsfjorden over a 1-month period. Additionally, 10 marine and 2 freshwater sites were sampled within a 1-week period in both Kongsfjorden and Krossfjorden. The central Kongsfjorden revealed a relatively stable protistan community over time with dinoflagellates, chlorophytes and small heterotrophs dominating. In contrast, the bacterial community varied over time and appeared to be correlated with the inflow of glacial meltwater. The Kongsfjorden and Krossfjorden were found to harbor distinctive bacterial and eukaryotic communities. We speculate that differences in glacial meltwater composition and fjord bathymetry affect the surface water properties and therefore the observed spatial variability in the community fingerprints.     

Explaining bathymetric diversity patterns in marine benthic invertebrates and demersal fishes: physiological contributions to adaptation of life at depth

Bathymetric biodiversity patterns of marine benthic invertebrates and demersal fishes have been identified in the extant fauna of the deep continental margins. Depth zonation is widespread and evident through a transition between shelf and slope fauna from the shelf break to 1000 m, and a transition between slope and abyssal fauna from 2000 to 3000 m; these transitions are characterised by high species turnover. A unimodal pattern of diversity with depth peaks between 1000 and 3000 m, despite the relatively low area represented by these depths. Zonation is thought to result from the colonisation of the deep sea by shallow‐water organisms following multiple mass extinction events throughout the Phanerozoic. The effects of low temperature and high pressure act across hierarchical levels of biological organisation and appear sufficient to limit the distributions of such shallow‐water species. Hydrostatic pressures of bathyal depths have consistently been identified experimentally as the maximum tolerated by shallow‐water and upper bathyal benthic invertebrates at in situ temperatures, and adaptation appears required for passage to deeper water in both benthic invertebrates and demersal fishes. Together, this suggests that a hyperbaric and thermal physiological bottleneck at bathyal depths contributes to bathymetric zonation. The peak of the unimodal diversity–depth pattern typically occurs at these depths even though the area represented by these depths is relatively low. Although it is recognised that, over long evolutionary time scales, shallow‐water diversity patterns are driven by speciation, little consideration has been given to the potential implications for species distribution patterns with depth. Molecular and morphological evidence indicates that cool bathyal waters are the primary site of adaptive radiation in the deep sea, and we hypothesise that bathymetric variation in speciation rates could drive the unimodal diversity–depth pattern over time. Thermal effects on metabolic‐rate‐dependent mutation and on generation times have been proposed to drive differences in speciation rates, which result in modern latitudinal biodiversity patterns over time. Clearly, this thermal mechanism alone cannot explain bathymetric patterns since temperature generally decreases with depth. We hypothesise that demonstrated physiological effects of high hydrostatic pressure and low temperature at bathyal depths, acting on shallow‐water taxa invading the deep sea, may invoke a stress–evolution mechanism by increasing mutagenic activity in germ cells, by inactivating canalisation during embryonic or larval development, by releasing hidden variation or mutagenic activity, or by activating or releasing transposable elements in larvae or adults. In this scenario, increased variation at a physiological bottleneck at bathyal depths results in elevated speciation rate. Adaptation that increases tolerance to high hydrostatic pressure and low temperature allows colonisation of abyssal depths and reduces the stress–evolution response, consequently returning speciation of deeper taxa to the background rate. Over time this mechanism could contribute to the unimodal diversity–depth pattern.     

Diversity,Abundance and Community Structure of Benthic Macro- and Megafauna on the Beaufort Shelf and Slope

Diversity and community patterns of macro- and megafauna were compared on the Canadian Beaufort shelf and slope. Faunal sampling collected 247 taxa from 48 stations with box core and trawl gear over the summers of 2009–2011 between 50 and 1,000 m in depth. Of the 80 macrofaunal and 167 megafaunal taxa, 23% were uniques, present at only one station. Rare taxa were found to increase proportional to total taxa richness and differ between the shelf ( 100 m) where they tended to be sparse and the slope where they were relatively abundant. The macrofauna principally comprised polychaetes with nephtyid polychaetes dominant on the shelf and maldanid polychaetes (up to 92% in relative abundance/station) dominant on the slope. The megafauna principally comprised echinoderms with Ophiocten sp. (up to 90% in relative abundance/station) dominant on the shelf and Ophiopleura sp. dominant on the slope. Macro- and megafauna had divergent patterns of abundance, taxa richness ( diversity) and diversity. A greater degree of macrofaunal than megafaunal variation in abundance, richness and diversity was explained by confounding factors: location (east-west), sampling year and the timing of sampling with respect to sea-ice conditions. Change in megafaunal abundance, richness and diversity was greatest across the depth gradient, with total abundance and richness elevated on the shelf compared to the slope. We conclude that megafaunal slope taxa were differentiated from shelf taxa, as faunal replacement not nestedness appears to be the main driver of megafaunal diversity across the depth gradient.     

The biodiversity of the deep Southern Ocean benthos

Our knowledge of the biodiversity of the Southern Ocean (SO) deep benthos is scarce. In this review, we describe the general biodiversity patterns of meio-, macro- and megafaunal taxa, based on historical and recent expeditions, and against the background of the geological events and phylogenetic relationships that have influenced the biodiversity and evolution of the investigated taxa. The relationship of the fauna to environmental parameters, such as water depth, sediment type, food availability and carbonate solubility, as well as species interrelationships, probably have shaped present-day biodiversity patterns as much as evolution. However, different taxa exhibit different large-scale biodiversity and biogeographic patterns. Moreover, there is rarely any clear relationship of biodiversity pattern with depth, latitude or environmental parameters, such as sediment composition or grain size. Similarities and differences between the SO biodiversity and biodiversity of global oceans are outlined. The high percentage (often more than 90%) of new species in almost all taxa, as well as the high degree of endemism of many groups, may reflect undersampling of the area, and it is likely to decrease as more information is gathered about SO deep-sea biodiversity by future expeditions. Indeed, among certain taxa such as the Foraminifera, close links at the species level are already apparent between deep Weddell Sea faunas and those from similar depths in the North Atlantic and Arctic. With regard to the vertical zonation from the shelf edge into deep water, biodiversity patterns among some taxa in the SO might differ from those in other deep-sea areas, due to the deep Antarctic shelf and the evolution of eurybathy in many species, as well as to deep-water production that can fuel the SO deep sea with freshly produced organic matter derived not only from phytoplankton, but also from ice algae.     

Distribution of meiofauna in Kongsfjorden,Spitsbergen

Kongsfjorden, a glacial fjord, is a typical fjord in the Spitsbergen (Svalbard archipelago) in the Arctic. The study supports a hypothesis that meiofauna and macrofauna are affected by natural environmental disturbances. Therefore, meiofaunal and macrofaunal analyses can be used to assess the effects of natural environmental disturbances in similar fjords in the Spitsbergen. Inputs from tidal glaciers create steep environmental gradients in sedimentation and salinity along the fjord. The magnitude of the glacial outflow diminishes towards the outer part of the fjord. Glacial-related physical stress causes reduced abundance, biomass and diversity among the meiofaunal assemblages in the inner part of the fjord. Based on quantitative and qualitative analyses of the composition of collected samples, three groups of meiofauna have been distinguished: one outer basin association and two in the inner, glacial bay. The presented results demonstrate that both the meiofauna and the macrofauna are affected on a similar scale by natural environmental disturbances. Therefore, as for macrofauna, meiofaunal analysis can be used to assess the effect of natural environmental disturbances.     

Genetic differentiation between Arctic and Antarctic monothalamous foraminiferans

Monothalamous (single-chambered) foraminifers are a major component of the benthic meiofauna in high latitude regions. Several morphologically similar species are common in the Arctic and Antarctic. However, it is uncertain whether these morphospecies are genetically identical, or whether their accurate identification is compromised by a lack of distinctive morphological features. To determine the relationship between Arctic and Antarctic species, we have compared SSU rDNA sequences of specimens belonging to four morphotaxa: Micrometula, Psammophaga, Gloiogullmia, and one morphospecies Hippocrepinella hirudinea from western Svalbard (Arctic) and McMurdo Sound (Antarctic). Wherever possible, we include in our analyses representatives of these taxa from the deep Arctic and Southern Oceans, as well as from Northern European fjords. We found that in all cases, the bipolar populations were clearly distinct genetically. As expected, Arctic specimens were usually more closely related to those from Northern Europe than to their Antarctic representatives. The deep-sea specimens from Weddell Sea branched as a sister to the McMurdo Sound population, while those from the Arctic Ocean clustered with ones from Norwegian fjords. Our study has revealed a high number of cryptic species within each of the examined genera, and demonstrates the unexplored potential of monothalamous foraminifers for use as a tool to evaluate the origin and biogeography of polar meiofauna.     

Habitat specialization in tropical continental shelf demersal fish assemblages

The implications of shallow water impacts such as fishing and climate change on fish assemblages are generally considered in isolation from the distribution and abundance of these fish assemblages in adjacent deeper waters. We investigate the abundance and length of demersal fish assemblages across a section of tropical continental shelf at Ningaloo Reef, Western Australia, to identify fish and fish habitat relationships across steep gradients in depth and in different benthic habitat types. The assemblage composition of demersal fish were assessed from baited remote underwater stereo-video samples (n = 304) collected from 16 depth and habitat combinations. Samples were collected across a depth range poorly represented in the literature from the fringing reef lagoon (1-10 m depth), down the fore reef slope to the reef base (10-30 m depth) then across the adjacent continental shelf (30-110 m depth). Multivariate analyses showed that there were distinctive fish assemblages and different sized fish were associated with each habitat/depth category. Species richness, MaxN and diversity declined with depth, while average length and trophic level increased. The assemblage structure, diversity, size and trophic structure of demersal fishes changes from shallow inshore habitats to deeper water habitats. More habitat specialists (unique species per habitat/depth category) were associated with the reef slope and reef base than other habitats, but offshore sponge-dominated habitats and inshore coral-dominated reef also supported unique species. This suggests that marine protected areas in shallow coral-dominated reef habitats may not adequately protect those species whose depth distribution extends beyond shallow habitats, or other significant elements of demersal fish biodiversity. The ontogenetic habitat partitioning which is characteristic of many species, suggests that to maintain entire species life histories it is necessary to protect corridors of connected habitats through which fish can migrate.