2. National Centre for Antarctic and Ocean Research, Ministry of Earth Sciences, Government of India, Goa 403 804
Author Correspondence author
International Journal of Marine Science, 2016, Vol. 6, No. 54 doi: 10.5376/ijms.2016.06.0054
Received: 28 Oct., 2016 Accepted: 13 Dec., 2016 Published: 15 Dec., 2016
Bijoy Nandan S., Krishnapriya P.P., Akhilesh V., Asha C.V., Jayachandran P.R., and Krishnan K.P., 2016, Benthic Faunal Assemblage of the Arctic Kongsfjorden System, Norway, International Journal of Marine Science, 6(54): 1-8 (doi: 10.5376/ijms.2016.06.0054)
Kongsfjord is an open glacial fjord on the west coast of Svalbard, located northwards to Norway. Benthic samples were collected from seven selected locations as a part of summer Phase III group 2 – Indian Arctic Expedition 2011. The characteristic macro benthic fauna observed in the soft bottom glacial community include Foraminifera, Nematoda, Polychaeta, Bivalvia, Amphipoda, Calanoid copepoda and Ostracoda, out of which polychaetes (59%) and nematodes (21%) were abundant. The meio benthic group was constituted by Nematoda, Foraminifera, Bivalvia, Polychaeta, Harpacticoid copepoda, Gastrotricha, and Kinoryncha, of which nematodes (54%) followed by foraminferans (37%) were the dominant group. A significant difference in the distribution of meio benthos between the inner and outer fjords was observed. Multivariate analysis of macro and meio fauna showed an overall similarity of ~ 80% among stations. The inner and outer fjords were similar in terms of granulomertic composition and were also silt dominated in the entire Kongsfjorden system. Total carbon (TC) and total organic carbon (TOC) were higher in the outer fjord, but the total inorganic carbon (TIC) was slightly higher in the inner fjord. The diversity and abundance of benthic organisms was strongly influenced by sediment characteristics, glacial inputs and Atlantic water mass movements making the ecosystem unpredictable.
Introduction
Fjords are semi-enclosed marine inlets that remain under strong terrestrial and oceanic influences and are considered to be strongly dependent on regional species pools of neighbouring open seas. The mixing of melt water from the glaciers and the saline water from the ocean form a suitable environment for the flourishing benthic organisms. Studies on the sediment characteristics are important for benthic studies. The works done by several researchers have got immense attention (Birgel et al., 2004; Chetelat et al., 2010; Holte et al., 1996; Urban-Malinga et al., 2005, 2009; Pawlowska et al., 2011; Wlodarska-Kowalczuk et al., 2004, 2012). Meiobenthic abundance and diversity exceed that of macrofauna and megafauna with increasing water depth (Giere, 2009). Shallow water soft bottom macro faunal communities have received attention in recent times but were not sampled effectively for quantitative studies. Hence the characterisation of benthic fauna both macro and meio have immense understanding in studying their distribution and role in environmental monitoring programs. This paper discusses the composition, abundance and diversity of macro benthic fauna with regard to sediment characteristics in the arctic Kongsfjorden system.
Materials and Methods
Study area and sampling stations
The study area was Kongsfjorden glacial fjord (79O N, 12O E) in Arctic (Svalbard) located north wards to Norway. Kongsfjorden is a high latitude glacial fjord influenced by Atlantic and Arctic water masses. The glacial outflow of fresh melt-water containing mineral materials influences the salinity, water transparency and primary production and sedimentation rates. The samples were collected from seven stations (Fig.1). The depth of the area ranged from 40 to 303 meters (Table 1). Based on depth profile, the fjord stations were divided into two subsets representing the outer and inner parts of the fjord. Stations 1, 2 and 3 were considered as inner fjord whereas 4, 5, 6 and 7 as outer in the fjord.
Fig. 1 Map of Kongsfjord showing the study stations |
Table 1 Details of sampling stations in Kongsfjorden system |
Collection and processing of samples
The samples were collected as a part of Indian Arctic Expedition from 17 July to 6 August, 2011 at the Indian research base “Himadri Station” at Spitsbergen, Svalbard, Norway; which is a part of the International Arctic Research base, Ny-Alesund. The boat “Teisten” of Kings Bay was employed for collecting samples from the selected transects. The samples, both macro and meio benthos were collected separately using van Veen grab (KC Denmark A/S) of 0.1 m2 area. Sub samples for meiofauna analysis were collected by a glass corer with cross-section area of 5cm2 that was pushed into sediment to a depth of 5 cm. (Pfannkuche and Thiel, 1988). The samples were preserved in formaldehyde – seawater solution of 4% and stained in Rose Bengal. Organisms were extracted from the sediment using decantation technique (Higgins and Thiel, 1988). Meiofauna after passing through a 1-mm sieve were retained on a 63 µm sieve, was counted. The meiobenthic and macrobenthic organisms were identified using suitable taxonomic keys (Giere, 2009; Fauvel, 1953) using, Olympus - Magnus MS 24.
Sediment characteristics
Sediment pH was determined using Systronics analyser (No.371, [accuracy ± 0.01]). Digital Eh meter (Systronics, No.318) was used for determining Oxidation reduction potential (Eh) and expressed in mV. The moisture content of the sediment was determined by drying sediment samples at a uniform temperature of 1000C. Sediment granulometric analysis were performed with a Sympatec (KFS Magic) Serial No: 1131, 41362 laser diffraction granulometer for finding the fractions of silt, clay and sand in the sediment. Total carbon (TC), total organic carbon (TOC) and total inorganic carbon (TIC) concentrations were determined on dried sediment samples by thermal combustion using a TOC – analyzer, Analytik Jena multi N/C 2100s. Samples for inorganic carbon were pre-treated with HCl to remove carbonates.
Data analysis
PRIMER (Plymouth Routines in Multivariate Ecological Research, Version 6.1.6) was used for univariate and multivariate analysis of data (Clarke and Gorley, 2006). Faunal diversity was calculated using Shannon-Wiener index (H’, log2). Variation pattern in community structure was evaluated by multivariate methods as a grouping analysis (Cluster) based on Bray-Curtis’ similarity index, ordination through non-metric Multi-dimensional Scaling (MDS) and Analysis of Similarity (ANOSIM).
Results
Sedimentology
In the summer months of August, the average sediment temperature of Kongsfjord ranged from 2.7 to 4.1OC. The mean value of pH in the inner fjord was 6.92±0.294 and that of outer fjord was 7.27±0.126. The mean value of Eh in the inner fjord was -294.33±59.28 mV and that of outer fjord was -183.75±56.68 mV. The mean value of sediment moisture in the inner fjord was 4.99% and that of outer fjord was 6.5%. The total carbon in the inner fjord was 31.25±6.698 g/kg and that of outer fjord was 35.44±0.843g/kg. The inorganic carbon varied from a lowest value of 13.01 g/kg in station 2 to a highest value of 27.1 g/kg in station 1. Inorganic carbon in the inner fjord was 21.79±7.65 g/kg and that of outer fjord was 20.17 ± 2.24g/kg. The total organic carbon varied from a lowest value of 8.69 g/kg in station 1 to a highest value of 16.49 g/kg in station 7. The total organic carbon in the inner fjord was 9.47±0.967 g/kg and that of outer fjord was 15.28±1.42 g/kg (Table 2). The sediment fraction consists of silt, sand and clay. The percentage fractions of the sediment are detailed in Table 2. The average silt fraction in the Kongsfjorden system during the study period was 66.33 %, that of inner fjord was 70.33% and that of outer fjord was 63.33. The average sand fraction in the entire system during the study period was 45.87 %, of which the inner fjord and outer fjords were 49.93% and 42.83 % respectively. The average clay fraction in the system was 20.71 % of which the inner and outer fjords were 21 % and 20.5 % respectively.
Table 2 Physio-chemical parameters of sampling stations in Kongsfjord |
Benthic macro fauna – composition, abundance and diversity
The characteristic macro benthic fauna observed in the soft bottom glacial community included Foraminifera (14.7%), Nematoda (21%), Polychaeta (59.18%), Bivalvia (1.84%), Amphipoda (1.98%), Calanoid copepoda (0.57%) and Ostracoda (0.71%). Out of which, polychaetes (835 ind. / m2) and nematodes (296 ind. / m2) were the numerically dominant taxon. Three polychaete families, Eunicidae (1.39%), Spionidae (0.90%) and Maldanidae (0.57%) were abundant in most of the study stations. Lagenidae (44.2%) and Buliminidae (Globobulimina auriculata) (27.8%) were the abundant families in Foraminifera (Fig. 2). The inner fjord fauna was dominated by three families of polychaetes, Syllidae, Ampharetidae, and Capitellidae. The outer fjord was comprised of the foraminiferan species, Virgulina complanata, Virgulina proboscidea, Bolivina sp., Reophax dentaliniformis and Cornuspira foliacea. In the inner fjord the macrofaunal abundance was higher in station 1 (344 ind./m2) followed by station 2 (225 ind./m2), having an average of 246 ind./m2 , where the outer fjord had 168 ind./m2. The Shannon Wiener index (H’) was higher in station 4 (1.895) followed by station 7(1.878) and that it was 1.36 in the inner fjord and 1.72 in the outer fjord. The multivariate analysis of macrofaunal abundance formed two major clusters with ~ 80% similarity. Stations 2, 3, 5 and 6 represented cluster 1 and stations 1, 4 and 7 in cluster 2. Highest similarity in abundance was observed in stations 4 and 7 (90%) of cluster 2 (Fig. 3). A permutation based hypothesis testing ANOSIM (one way) was done to find out whether there is any significant variation in the distribution of macro fauna between inner and outer fjord. Similarity analysis (ANOSIM) indicated that there was no significant variation in the distribution of macro fauna between inner and outer fjords (Global R = 0.074, P < 0.001).
Fig. 2 Mean percentage abundance of macro fauna in the Kongsfjorden system |
Fig. 3 Dendrogram representation of square - root transformed data of macrofaunal abundance of Kongsfjorden system-Arctic |
Benthic meiofauna- composition, abundance and diversity
Seven meiofaunal groups were recorded in the summer months of Kongsfjorden glacial system that was constituted of Nematoda, Foraminifera, Bivalvia, Polychaeta, Harpacticoid copepoda, Gastrotricha, and Kinoryncha. Maximum numerical abundance of the taxa was recorded in station 2 (2273 ind. /10 cm2) followed by station 1 (2154 ind. /10 cm2) and station 3 (1862 ind. / 10 cm2) respectively. Nematoda was the numerically dominant taxon in every station, constituting a minimum of 54% of total meiobenthic abundance. The percentage abundance of other subdominant taxa were, Foraminifera (37 %), Harpacticoid copepoda (0.2%), Bivalvia (3.1 %), Gastropoda (0.2 %), Polychaeta (3 %) and Kinoryncha (3%) (Fig. 4). Nematodes formed the abundant group both in the inner and outer fjords with a percentage abundance of 47% in the inner fjord and that of 52.7% in the outer fjord.
Fig. 4 Mean Percentage Abundance of Meiofauna in the Kongsfjorden system |
The Shannon Wiener index (H’) varied from 1.684 in station 1 to a 1.66 in station 2 with an average of 1.61 in the inner fjord and 1.28 in the outer fjord. The diversity of meio fauna was higher in the inner fjord. All meiobenthic groups were present in station 1 which was located in the inner fjord and it was dominated by nematodes. Multivariate analysis of meiofaunal abundance showed an overall similarity of 80% between stations that indicated least variability among stations, since station wise cluster analysis revealed 2 major clusters where the abundance of meiofauna was maximum between station 5 and 7 (95%) followed by stations 4, 5 and 7 (94%) (Fig.5). Hence, the similarity in abundance of fauna was more conspicuous in stations 4, 5 and 7. Cluster analysis revealed that the inner and outer fjords had an overall similarity of ~ 85% between stations (Fig. 6). Similarity analysis (ANOSIM) indicated that there was significant variation in the distribution of meio fauna between inner and outer fjords (Global R = 0.833, P < 0.001).
Fig. 5 Dendrogram representation of square - root transformed data of meiofaunal abundance of Kongsfjorden system-Arctic |
Fig. 6 Dendrogram representation of square - root transformed data of meiofaunal abundance of Kongsfjorden system in the inner and outer fjord of Kongsfjord |
Discussion
The mean value of sediment moisture in the outer fjord was higher when compared to the inner fjord. The moisture content indicates the water holding capacity of the sediment. Station seven had higher water holding capacity but the rest of the stations depicted a uniform nature. Fine sediment with higher water content was probably a more difficult barrier to the upward burrowing activity of fauna than coarse sediment (Turk, 1981). The amount of water in the sediment is an important factor determining the distribution of organisms. Total carbon in the sediment was higher in the outer fjord, than the inner fjord. The average inorganic carbon was slightly higher in the inner fjord, than the outer fjord. Among the two forms of carbon (organic and inorganic) present in the soil, total organic carbon is the major form of carbon which is available to the food web which gets transferred from one trophic level to another. The total organic carbon values showed a direct relationship within the Kongsfjorden system. The decreasing concentration of organic matter in the sediment could be the result of glacial mixing (Holte and Gulliksen, 1998). The proportion of silt, sand and clay in the sediment in all the seven stations were almost similar, hence the inner and outer fjords also was similar in terms of granulometric composition. Similar results were observed in the studies of Kotwicki et al. (2004) and Wlodarska-Kowalczuk and Pearson (2004). In the present study, it was also observed that, silt dominated in sediment fractions among the seven study stations in Kongsfjord. The granulometric characteristics, quantity of organic carbon and the stability of sediments, all reflect the glacial sedimentation gradients in the Arctic glacial fjord. Sedimentation rates of suspended particulate matter were reported to be highest in the glacier’s proximal inner basin (Svendsen et al., 2002). Salinity and temperature gradients are produced by the inflow of fresh and cold glacial melt waters in the Kongsfjorden system, which are controlled by the glacial activity. During the summer months, the high influx of terrigenous material transported by the glacial inputs produces high particle flux which affected both the benthic food availability and substrate condition while the lack of sediment stability and permanent mineral sedimentation processes appear to be much more important, influencing the meiofaunal distribution and macro faunal densities in Kongsfjord (Kotwicki et al., 2004; Wlodarska-Kowalczuk and Pearson, 2004). But, it is also important that the high levels of mineral sedimentation and sediment deposition have shown to be an acute disturbance agent causing a dramatic decrease in benthic densities and diversity. In the present study also it is noticed that macro benthos have higher diversity towards the outer fjord but their abundance was lower.
During the summer investigation of 2011 in the Kongsfjord, the macrofunal abundance showed that 33.2% families were recorded from the inner fjord whereas 66.8% from the outer fjord. From the multivariate analysis of the seven stations, a similar trend in macrofaunal abundance was observed. From the ANOSIM results significant differences were not observed in the distribution of macrobenthic communities among the inner and outer fjords. The overall diversity pattern indicated an increase in faunal diversity with increasing depth but the abundance of fauna decreased with increasing depth. The patterns of decline in species diversity across the fjord have been observed by Kotwicki et al. (2004) and Somerfield et al. (2006). The decrease in the species richness in the inner fjord has been reported for several benthic groups including mostly brooding species (crustaceans) and those with dispersing larvae (polychaetes and molluscs) (Wlodarska-Kowalczuk et al., 2012). The reason for the decline in faunal abundance and diversity in the Arctic fjord ecosystems are strongly influenced by glacial derived disturbances, mainly by the outflow of melt water produced by active tidal glaciers. Tidewater glaciers are often situated in the innermost parts of the fjords and they have a strong effect on the physical regimes of whole basins and can impact hard and soft-bottom benthic communities. Organic matter quality can play a critical role in determining macro faunal distribution in the fjord system. There were changes in both free living nematodes and macro faunal assemblages with increasing distance from the larger glacier, Kongsbreen which correlate to some extent with the organic content (C:N ratio) of the sediment (Somerfield et al., 2006). In the present study also a decreasing pattern of macrobenthic abundance with increasing distance from glaciers was noticed. Natural factors (depth of basins, presence of sills, variable terrestrial runoff, atmospheric forcing, seasonal phytoplankton blooms) and anthropogenic factors (organic enrichment, fishing activities) may be leading to community changes.
The similarity in abundance of meiofauna was more conspicuous in stations 4, 5 and 7. It could be the presence of almost similar pattern of meiobenthic distribution in these stations and it was also noticed that the three stations were located in the outer fjord. From the cluster analysis and ANOSIM, it was clear that there were significant differences in the distribution of meiofauna between inner and outer fjords. The average abundance of nematodes was higher in the inner region but foraminiferal abundance was higher in the outer fjord and this difference in distribution could be a reason for this dissimilarity. The persistence of stability of meiofaunal abundance and diversity in the inner and outer fjord of Kongsfjord revealed that meiofauna can be less sensitive than macro fauna (Austen et al., 1989; Warwick et al., 1990) to sediment disturbance caused by decreased sediment re-suspension due to glacial activity. Meiofaunal distribution is influenced by the uniform granulometry, unstable sediment and constant mineral sedimentation processes. The glacial inputs and the inflow of Atlantic water masses also contribute to the distribution of meiofauna (Gulliksen et al., 1985; Kendall, 1994; Holte et al., 1996; Holte and Gulliksen 1998). The present study on meiofauna corroborates with that by Kotwicki et al. (2004). Urban-Malinga et al. (2005) also reported that nematodes, oligochaetes and turbellarians were numerically dominant among the meio benthic taxon throughout the fjord. In the present study nematodes were the numerically abundant taxon followed by foraminiferans. Likewise the study of Pawlowska et al. (2011) revealed that in the Adventfjord, Spitsbergen (an arctic fjord) more than 90% of all individuals were represented by nematodes.
From the present study the following conclusions can be arrived, where the macro benthic community showed an increase in diversity towards the outer fjord but their abundance was higher in the inner fjord. Meiofaunal community displayed higher abundance and diversity towards the inner fjord of the Kongsfjord. Polychaetes were the dominant macro benthic taxon whereas nematodes were the dominant meio benthic taxon during the study period. There was no significant difference in the distribution of macro benthic communities among the inner and outer fjords but meio fauna had significant differences in the distribution between the two fjords. Thus, macrofauna and meiofauna provide different but complementary types of information on their occurrence and also in relation to the prevailing environmental conditions. Meiofauna can be sensitive to environmental disturbances as the sediment in the outer fjord area was always under commotion by the inflow and out flow of water masses hence they were highly concentrating to the less disturbed inner fjord. In summer months the input of terrigenous material transported by the rivers produced high particle flux which affected both the benthic food availability and substrate conditions. Hence, seasonality was evident for both macro and meio faunal distribution in the present study. Therefore, long term, spatio-temporal studies are further required to evolve at a detailed and comprehensive information on benthic fauna and its variabilities in the Arctic fjords especially in the context of emerging global climate change.
Acknowledgements
The authors are thankful to the National Centre for Antarctic and Ocean Research (NCAOR), Govt. of India and to the Head, Dept. of Marine Biology, Microbiology and Biochemistry, School of Marine Sciences, Cochin University of Science and Technology for providing the financial support and necessary facilities for undertaking the study.
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