Microbial processes in a high-latitude fjord (Kongsfjorden, Svalbard): II. Ciliates and dinoflagellates

The composition and ecological role of ciliates and dinoflagellates were investigated at one station in Kongsfjorden, Svalbard, during six consecutive field campaigns between March and December 2006. Total ciliate and dinoflagellate abundance mirrored the seasonal progression of phytoplankton, peaking with 5.8 × 10⁴ cells l⁻¹ in April at an average chlorophyll a concentration of 10 μg l⁻¹. Dinoflagellates were more abundant than ciliates, dominated by small athecates. Among ciliates, aloricate oligotrichs dominated the assemblage. A large fraction (>60%) of ciliates and dinoflagellates contained chloroplasts in spring and summer. The biomass of the purely heterotrophic fraction of the ciliate and dinoflagellate community (protozooplankton) was with 14 μg C l⁻¹ highest in conjunction with the phytoplankton spring bloom in April. Growth experiments revealed similar specific growth rates for heterotrophic ciliates and dinoflagellates (<0–0.8 d⁻¹). Food availability may have controlled the protozooplankton assemblage in winter, while copepods may have exerted a strong control during the post-bloom period. Calculations of the potential grazing rates of the protozooplankton indicated its ability to control or heavily impact the phytoplankton stocks at most times. The results show that ciliates and dinoflagellates were an important component of the pelagic food web in Kongsfjorden and need to be taken into account when discussing the fate of phytoplankton and biogeochemical cycling in Arctic marine ecosystems.     

Intertidal meiofauna of a high-latitude glacial Arctic fjord (Kongsfjorden, Svalbard) with emphasis on the structure of free-living nematode communities

Meiofauna communities of four intertidal sites, two sheltered and two more exposed, in Kongsfjorden (Svalbard) were investigated in summer 2001 at two different tidal levels (i.e. the low-water line and close below the driftline, referred to as mid-water (MW) level). A total of seven meiofaunal higher taxa were recorded with nematodes, oligochaetes and turbellarians being numerically dominant. Mean total meiofaunal densities ranged between 50 ind. 10 cm⁻² and 903 ind. 10 cm⁻². Our data showed a clear decrease in total meiofaunal densities with increasing coarseness of the sediment. Total meiofaunal biomass varied from 0.2 g dwt m⁻² to 2 g dwt m⁻² and, in general, was high even at low meiofaunal densities, i.e. larger interstitial spaces in coarser sediments supported larger meiofauna, especially turbellarians. The results on the vertical distribution of meiofauna contrasted sharply with typical meiobenthic depth profiles on other beaches, probably in response to ice-scouring and concomitant salinity fluctuations. Oligochaetes were the most abundant taxon, with a peak density of 641 ind. 10 cm⁻² at Breoyane Island. They were mainly comprised of juvenile Enchytraeidae, which prohibited identification to species/genus level. Nematode densities ranged between 4 ind. 10 cm⁻² and 327 ind. 10 cm⁻². Nematodes were identified up to genus level and assigned to trophic guilds. In total, 28 nematode genera were identified. Oncholaimus and Theristus were the most abundant genera. The composition of the nematode community and a dominance of predators and deposit feeders were in agreement with results from other arctic and temperate beaches. Nematode genus diversity was higher at the more sheltered beaches than at the more exposed ones. Low-water level stations also tended to harbour a more diverse nematode communities than stations at the MW level. Differences in nematode community structure between low- and MW stations of single beaches were more pronounced than community differences between different beaches and were mainly related to resources quality and availability.     

Distribution of Calanus species in Kongsfjorden, a glacial fjord in Svalbard

The distribution of Calanus species was investigated in Kongsfjordenin summer of 1996 and 1997. In both years Calanus finmarchicusand Calanus glacialis dominated, although the boreal C. finmarchicuswas more abundant than the Arctic C. glacialis in 1997. Thiscoincided with a 2°C higher water temperature at 50 m in1997, indicating stronger influence of Atlantic origin waterthat year. Advected Calanus finmarchicus occurred in deep andsubsurface layers of the outer fjord in 1996 (200 ind. m^-3,mainly CIII). A less abundant local population aggregated insurface layers of the inner fjord (100 ind. m^-3). Similarly,advected C. finmarchicus occurred in subsurface layers in 1997(446 ind. m^-3, mainly CIII and CIV) and a local population insurface layers (183 ind. m^-3, mainly CI). Calanus glacialisin 1996 aggregated as CII and CIII in the deep layers of theouter fjord (272 ind. m^-3), whereas CIII–CV were abundant(216 ind. m^-3) in cold surface waters of the inner fjord. In1997 C. glacialis (mostly CIII–CV) was more abundant inthe outer than in the inner part of the fjord (40 and 192 ind.m^-3, respectively). Within Kongsfjorden, Calanus finmarchicusneeds one year to complete its life cycle, whereas Calanus glacialisneeds two. Calanus hyperboreus seems to be an expatriate inthe fjord system.     

Decadal change in macrobenthic soft-bottom community structure in a high Arctic fjord (Kongsfjorden, Svalbard)

Marine benthic macrofauna communities are considered a good indicator of subtle environmental long-term changes in an ecosystem. In 1997/1998 and 2006, soft-bottom fauna of an Arctic glacial fjord Kongsfjorden was extensively sampled and major communities were identified along the fjord axis, which were related to the diminishing influence of glacial activity. Spatial patterns in community structure and species diversity were significantly different in the central basin of Kongsfjorden between periods while there was no change in the inner part of the fjord. In 1997/98, three faunal associations were distinguished with significant differences in species richness and diversity (H′) while in 2006 only two faunal associations were identified and there were no differences any more between the two formerly distinct associations in the central fjord. The increased input of Atlantic water due to a stronger West Spitsbergen Current may be the reason for unification of previous clear faunal division. The faunal association in the inner, well separated glacial part of the fjord, characterized by strong glacier influence, was protected from Atlantic water inflow and, hence, the macrobenthic fauna essentially remained unaffected. Reduced abundance of species typical for glacial bays in the central part of the fjord in 2006 may result from the decreasing effect of Blomstrandbreen glacier, strong increase of input of Atlantic water into the fjord and increased temperature of West Spitsbergen Current. Higher values of POC in 2006 than in 1998 are likely the effect of increased primary production resulting from warmer water temperatures.     

The distribution of picoplankton and nanoplankton in Kongsfjorden,Svalbard during late summer 2006

Abundance and biomass of pico- (<2 μm) and nanoplankton (2–20 μm) were investigated in relation to hydrography in Kongsfjorden, Svalbard (79°N, 12°E) during late summer 2006. Autotrophic and heterotrophic picoplankton abundance ranged from 0.1 × 10⁶ to 35.2 × 10⁶ cells L⁻¹ and from 0.4 × 10⁶ to 20.3 × 10⁶ cells L⁻¹, respectively. The highest number of bacteria in the entire water column was recorded at station 2 at 10 m (22.3 × 10⁸ cells L⁻¹); the lowest concentration was observed at station 1 (6.0 × 10⁸ cells L⁻¹). The abundance of autotrophic and heterotrophic nanoplankton varied from 0.4 × 10⁵ cells L⁻¹ to 46 × 10⁵ cells L⁻¹ and from 0.3 × 10⁶ to 9.1 × 10⁶ cells L⁻¹, respectively. Our results demonstrated that heterotrophic nanoflagellates and bacteria in Kongsfjorden microbial community were relatively important. The structure of plankton communities integrated with environmental variables could act as indicators of the variability of the inflow of Atlantic Water into Kongsfjorden.     

Temporal patterns of benthic community development in an Arctic fjord (Kongsfjorden, Svalbard): results of a 24-year manipulation study

We investigated temporal patterns of recolonisation and disturbance in a benthic hard bottom community in high-arctic Kongsfjorden from 1980 to 2003 through annual photographic surveys. A manipulative sampling design was applied, where half of the study area (treatment areas) was cleared at the beginning of the study. Twenty-three different taxa and groups of benthic epifauna were found in the photographs. The benthic community structures of treatments and controls converged within the first decade, but significant differences prevailed until ≤13 years after the start of the study. We could distinguish between three different time intervals with increased inter-annual changes. While the observed differences during the first two intervals could be attributed to recolonisation and succession, the changes in interval 3 were mostly due to increased external forcing and characterised by low inter-group and high inter-annual differences. During this interval, brown algae (mainly Desmarestia) and the sea urchin Strongylocentrotus droebachiensis emerged in high densities, while sea anemone populations declined. Different recolonisation patterns for individual species were related to life span, rate of maturity, predators and larval settlement. We could not find a climax stage in the succession for the benthic community at Kvadehuken, presumably due to the constant level of disturbances at the site.     

Seasonal dynamics of crustacean zooplankton, heterotrophic nanoflagellates and bacteria in a shallow, eutrophic lake

1. The seasonal development of crustacean zooplankton, heterotrophic nanoflagellates (HNF) and bacteria was examined in Grosser Binnensee, a shallow, eutrophic lake in northern Germany. The grazing impact of Daphnia on bacteria and nanoflagellates was estimated from field data on population abundances and from clearance rates obtained in laboratory experiments. 2. The seasonal succession of zooplankton showed distinct peaks of Daphnia magna, cyclopopid copepods, Bosmina longirostris and Daphnia galeata and D. hynlina. The population dynamics of Dapfinia had the strongest impact on all sestonic components. Daphnia maxima coincided with clearwater phases, and were negatively correlated with particulate organic carbon (POC), HNF and phytoplankton. Bacterial abundance was only slightly affected although daphnids were at times more important as bacterial consumers than HNF, as estimated from measured bacterial clearance rates. Other crustaceans (copepods, Bosmina) were probably of minor importance as grazers of bacteria and nanoplankton. 3. HNF abundance varied from 550 ml^?1 to more than 30000 ml^?1. HNF appeared to be suppressed by daphnids and reached highest densities when copepods dominated the metazooplankton. The variation in HNF abundance was not reflected in the concentration of heterotrophic bacteria, which fluctuated rather irregularly between 5 and 20 ± 10⁶ ml^?1. Long filamentous bacteria which were probably resistant to protozoan grazing, however, appeared parallel to the development of HNF. These bacterial cells, although small in number, could comprise more than 30% of the total bacterial biomass.     

Comparison of productivity and phytoplankton in a warm (Kongsfjorden) and a cold (Hornsund) Spitsbergen fjord in mid-summer 2002

Kongsfjorden and Hornsund are two glacial fjords without sills on the West Spitsbergen coast. Both sites are under the influence of relatively warm Atlantic-derived water, although Hornsund is more influenced by cold water from the Barents Sea. In this study, we compared the impacts of cold Arctic and warmer Atlantic waters on the pelagic ecosystems of Kongsfjorden and Hornsund. Both fjords were strongly influenced by Atlantic-derived waters during summer (2002). Diatoms were the most substantial contributors to phytoplankton biomass, especially in outer basins of both fjords, whereas the second most important contributors were autotrophic dinoflagellates in Kongsfjorden and nanoflagellates in Hornsund. Total phytoplankton biomass was highest in Hornsund. Primary production rates were an order of magnitude lower in Kongsfjorden than in Hornsund, and increased from inner to outer fjord (from 2.47 to 4.48 mg C m⁻² h⁻¹ in Kongsfjorden and from 14.00 to 86.65 mg C m⁻² h⁻¹ in Hornsund). Chlorophyll-a concentration was also substantially lower in Kongsfjorden. Zooplankton was dominated by omnivorous species in Kongsfjorden and herbivorous in Hornsund. Observed differences between the fjords may originate from (1) advection of different waters into the fjords; (2) differences in freshwater runoff and turbidity, and (3) timing of the phytoplankton bloom. Climate warming will likely increase the Atlantic water influence, and result in reduced production of diatoms and increase in flagellates.     

Microphytobenthos of Arctic Kongsfjorden (Svalbard, Norway): biomass and potential primary production along the shore line

During summer 2007, Arctic microphytobenthic potential primary production was measured at several stations around the coastline of Kongsfjorden (Svalbard, Norway) at ≤5 m water depth and at two stations at five different water depths (5, 10, 15, 20, 30 m). Oxygen planar optode sensor spots were used ex situ to determine oxygen exchange in the overlying water of intact sediment cores under controlled light (ca. 100 μmol photons m⁻² s⁻¹) and temperature (2–4°C) conditions. Patches of microalgae (mainly diatoms) covering sandy sediments at water depths down to 30 m showed high biomass of up to 317 mg chl a m⁻². In spite of increasing water depth, no significant trend in “photoautotrophic active biomass” (chl a, ratio living/dead cells, cell sizes) and, thus, in primary production was measured at both stations. All sites from ≤5 to 30 m water depth exhibited variable rates of net production from −19 to +40 mg O₂ m⁻² h⁻¹ (−168 to +360 mg C m⁻² day⁻¹) and gross production of about 2–62 mg O₂ m⁻² h⁻¹ (17–554 mg C m⁻² day⁻¹), which is comparable to other polar as well as temperate regions. No relation between photoautotrophic biomass and gross/net production values was found. Microphytobenthos demonstrated significant rates of primary production that is comparable to pelagic production of Kongsfjorden and, hence, emphasised the importance as C source for the zoobenthos.     

Spatial and temporal distribution of tintinnid (Ciliophora: Tintinnida) communities in Kongsfjorden, Svalbard (Arctic), during summer

Information of tintinnid community, as an important component of plankton, was limited in Kongsfjorden, Svalbard (Arctic), which is an important Arctic monitoring site for global change. In this paper, we studied the species composition, spatial, and vertical distribution of tintinnids in Kongsfjorden in July 2012. Water samples for tintinnid study were taken in different depths at eight grid stations in the fjord and time series (5–23 July) at one station at the pier (St. C). Eight species (morphotypes) were found: Acanthostomella norvegica, Leprotintinnus pellucidus, Parafavella edentata, Parafavella gigantea, Ptychocylis obtusa, Stenosemella spp., Tintinnopsis parvula, and Tintinnopsis spp. Total tintinnid abundance and biomass ranged from undetectable (<1 ind L^?1) to 2,435 ind L^?1 and 0–6.33 μg C L^?1, respectively. Cluster analysis of the tintinnid community in the grid stations and St. C showed that tintinnid community was uniform throughout the fjord. Four species (morphotypes) showed distinct vertical distributions: T. parvula occurred throughout the water column but showed a peak at the surface; Tintinnopsis spp. occurred only in the upper 30 m; A. norvegica occurred in low abundance throughout the water column; and Stenosemella spp. was not detected in surface waters. There was variation in abundance of different species in St. C in the 18-day period. However, the tintinnid community did not change according to cluster analysis. These results are helpful in understanding the distribution of other plankton taxon and in long-term monitoring of the Kongsfjorden marine ecosystem.     

Feeding of <Emphasis Type="Italic">Themisto libellula</Emphasis> (Amphipoda Crustacea) on natural copepods assemblages in an Arctic fjord (Kongsfjorden,Svalbard)

Ingestion rates and selectivity of the Arctic pelagic amphipod Themisto libellula were studied experimentally in Kongsfjorden (Svalbard, 78°N) during the summer period. Feeding incubations were conducted on naturally occurring copepod communities at different concentrations ranging from 25 to 250 preys L⁻¹. The ingestion rates increased with food availability from 1.3 to 17.7 preys ind⁻¹ day⁻¹, which corresponded to 0.3–11% of body carbon day⁻¹. Despite the high prey concentration used in the experiments the satiation level was not reached. We suggested that T. libellula is able to take the maximum benefit from dense patches of preys, which represent a good adaptation to the high variability in food supply characteristic of polar environment. Copepodids stage III of Calanus spp. appeared to be the preferred preys of T. libellula. Smaller copepods such as Oithona similis and Pseudocalanus spp., were also selected but only when their relative abundance exceeded 25% of the total prey available. The potential predation impact of T. libellula is discussed in relation to the mesozooplankton small-scale patchiness and predator abundance.     

Spring bloom dynamics in Kongsfjorden,Svalbard: nutrients,phytoplankton, protozoans and primary production

The marine ecosystem in Kongsfjorden (79°N), a glacial fjord in Svalbard, is to a large extent well known with regard to hydrography, mesozooplankton and higher trophic levels. Research on primary production and lower trophic levels is still scare and especially investigations from winter and spring periods. The spring bloom dynamics in Kongsfjorden were investigated in 2002. The development in nutrient conditions, phytoplankton, protozoans and primary production were followed from 15 April until 22 May. The winter/spring in 2002 was categorized as a cold year with sea ice cover and water masses dominated by local winter-cooled water. The spring bloom started around 18 April and lasted until the middle of May. The bloom probably peaked in late April, but break-up of sea ice made it impossible to sample frequently in this period. Diatoms dominated the phytoplankton assemblage. We estimated the total primary production during the spring bloom in 2002 to range 27–35 g C m⁻². There was a mismatch situation between the mesozooplankton and the phytoplankton spring bloom in 2002.     

Life histories of the copepods Pseudocalanus minutus, P. acuspes (Calanoida) and Oithona similis (Cyclopoida) in the Arctic Kongsfjorden (Svalbard)

The year-round variation in abundance and stage-specific (vertical) distribution of Pseudocalanus minutus and Oithona similis was studied in the Arctic Kongsfjorden, Svalbard. Maxima of vertically integrated abundance were found in November with 111,297 ind m⁻² for P. minutus and 704,633 ind m⁻² for O. similis. Minimum abundances comprised 1,088 ind m⁻² and 4,483 ind m⁻² in June for P. minutus and O. similis, respectively. The congener P. acuspes only occurred in low numbers (15–213 ind m⁻²), and successful reproduction was debatable. Reproduction of P. minutus took place in May/June, and stage distribution revealed a 1-year life cycle with copepodids CIII, CIV, and CV as the overwintering stages. Oithona similis exhibited two main reproductive peaks in June and August/September, respectively. Moreover, it reproduced more or less continuously throughout the whole year with all stages occurring during the entire sampling period, suggesting two generations per year. Both species migrated towards greater depth in November, but O. similis preferred to stay longer in the upper 100 m as compared to Pseudocalanus. The reproduction of the two species in Kongsfjorden seemed to be linked to phytoplankton dynamics.     

Heterotrophic nanoflagellates in water column and bottom sediments of the Rybinsk Reservoir: Species composition,abundance, biomass and their grazing impact on bacteria

Abundance, biomass, and taxonomic composition of heterotrophic nanoflagellates (HNFs) have been determined in the water column and bottom sediments of the large lowland meso-eutrophic reservoir (Rybinsk Reservoir, Upper Volga) in summer. The role of HNFs in the consumption of the bacterial production is estimated. In the reservoir, 55 species from 15 large taxa, including 35 species from the plankton, are identified and 45 species are from benthos samples. The orders Kinetoplastida, Choanomonada, and Chrysomonadida are distinguished by the highest species diversity. Abundance and biomass of HNFs in the water column average 991 ± 326 cells/mL and 41.4 ± 14.1 mg/m³, while in the bottom sediments they are (236 ± 61) × 10³ cells/mL and 10.7 ± 4.0 μg/mL, respectively. The biomass of HNFs average 11.2% of the bacterial biomass in the water column and only 0.8% of that in the sediments. Flagellates are found to be a major factor which control the development of bacterioplankton grazing, on average, 32.3% of its daily production, whereas their impact on bacteriobenthos is insignificant, as they consume, on average, only 2.0% of its production.     

Soft bottom species richness and diversity as a function of depth and iceberg scour in Arctic glacial Kongsfjorden (Svalbard)

Macrozoobenthic soft-sediment communities of central Arctic Kongsfjorden inhabiting six depth zones between 5 and 30 m were sampled using SCUBA-diving during June–August 2003 and analysed comparatively. About 63 taxa were found, nine of which had not been reported for Kongsfjorden and four for Svalbard. Suspension feeding or surface and sub-surface detritivorous polychaetes and deposit-feeding amphipods were dominant. Only 11 of the 63 taxa (45 species and additional 18 families not further identified) inhabited the complete depth range. Biomass ranged from 3.5 to 25.0 g ash free dry mass m⁻² and mean Shannon diversity (Log e) was 2.06. Similarity clustering from abundance and biomass data showed a significant difference between the shallow station (5 m) and the rest. The latter formed two sub-groups (10–20 and 25–30 m). Depth is irrevocably correlated with ice-scouring. Thus the differences in diversity together with the predicted iceberg scour intensity support the ‘intermediate disturbance hypothesis’ indicating that habitats impacted by moderate iceberg scouring enable higher diversity. In contrast, biotopes frequently affected only host pioneer communities, while mature, less diverse assemblages dominate depths of low impact.     

Trophic relationships between planktonic bacteria,heterotrophic nanoflagellates and viruses in a mesoeutrophic reservoir

The abundance, biomass, and production (Р _В) of bacrerioplankton; the taxonomic composition, abundance, biomass of heterotrophic nanoflagellates (HNF) and the rate of consumption of bacteria by HNFs; and the abundance of virioplankton, frequency of visibly infected bacterial cells, virus-induced mortality of bacterioplankton, and viral production were estimated in the mesoeutrophic Rybinsk Reservoir. The rate of bacterial mortality due to viral lysis (7.8–34.1%, on average 17.2 ± 2.0% of daily Р _В) was lower than the consumption of bacteria by the HNF community (15.4–61.3%, on average 32.0 ± 4.2% of daily Р _В). While consuming bacteria, HNFs simultaneously absorbed a significant number of viruses residing on the surface and inside the bacterial cells.     

Seasonal variability of meio- and macrobenthic standing stocks and diversity in an Arctic fjord (Adventfjorden, Spitsbergen)

Strong environmental seasonality is a basic feature of the Arctic system, still there are few published records of the seasonal variability of the Arctic marine biota. This study examined the year-round seasonal changes of soft bottom macro- and meiobenthic standing stocks and diversity on a station located in an Arctic fjord (Adventfjorden, Spitsbergen). The seasonality observed in benthic biota was related to the pelagic processes, primarily the seasonal fluxes of organic and inorganic particles. The highest abundance, biomass and richness of benthic fauna occurred in the spring after the phytoplankton bloom. During the summer, when a high load of glacial mineral material was transported to the fiord, the number of both meio- and macrobenthic individuals decreased remarkably. The strong inorganic sedimentation in summer was accompanied by a decline in macrobenthic species richness, but had no effects on evenness. Redundancy analysis (RDA) pointed to granulometric composition of sediments (depended on mineral sedimentation) and organic fluxes as factors best related to meio- and macrobenthic taxonomic composition, but no clear seasonal trend could be observed on the nMDS plots based on meiobenthic higher taxa or macrobenthic species abundances in the samples. This study addresses the possible effects of changes in the winter ice cover on the fjordic benthic systems because it was performed in a year with no ice cover on the fjord.     

Sea ice microbial communities. III. Seasonal abundance of microalgae and associated bacteria,Mcmurdo Sound,Antarctica

Numbers of bacteria in annual sea ice increased directly with numbers of algae during the 1981 spring ice diatom bloom in McMurdo Sound, Antarctica. Algae and bacteria in a control site grew at rates of 0.10 and 0.05 day^–1, respectively, whereas in an experimentally darkened area neither increased after six weeks. Epiphytic bacteria grew at a rate twice that of the nonattached bacteria and were significantly larger, contributing approximately 30% of the total bacterial biomass after October. The microalgal assemblage was dominated by two species of pennate diatoms, anAmphiprora sp. andNitzschia stellata. Greater than 65% of epiphytic bacteria were associated withAmphiprora sp. after October.N. stellata, however, remained largely uncolonized throughout the study. We hypothesize that microalgae stimulate bacterial growth in sea ice, possibly by providing the bacteria with organic substrates.     

The densities of bacteria, picophytoplankton, heterotrophic nanoflagellates and ciliates in Lake Kasumigaura (Japan) monitored monthly since 1996

This data paper describes the densities of the bacterioplankton, picocyanobacteria, eukaryotic picoplankton, heterotrophic nanoflagellates and ciliates in the water of Lake Kasumigaura??a shallow, eutrophic lake that is the second largest in Japan. All of these planktonic organisms are components of a microbial loop and are countable using an epifluorescence microscope. These data represent the results of monthly collections from April 1996 through March 2010 at two sites on the lake, and this data set is unique among the available published data papers concerning lakes or plankton and continues to be freely available. The monitoring was performed as a component of the Lake Kasumigaura Long-term Environmental Monitoring program conducted by National Institute for Environmental Studies since 1977 and includes water quality, plankton, and benthos. The data have been used for ecological studies and for studies on the management of water quality.