Structure,biomass distribution,and energetics of the pelagic ecosystem in the Barents Sea: A synopsis

Biomass distribution and energetics of trophic levels in the pelagic ecosystem of the Barents Sea are presented as averages over several years for the whole Barents Sea using data from the research programme Pro Mare in 1984–1989 and mathematical ecosystem models. Average biomasses range from more than 3 tonnes carbon km^–2 (zooplankton) to 0.1 kg C km^–2 (polar bears) and P/B ratios from 300 (bacteria) to 0.035 (minke whales). However, the Barents Sea ecosystem is in a far from steady state with, for instance, capelin stocks ranging from 30–700 kg C km^–2 between years and cod stocks from 150–700 kg C km^–2. As a general rule, the various fish stocks grow adequately, albeit at different rates, in warm years characterized by large influxes of Atlantic water and high zooplankton productivity. The skewed populations distribution which arises in warm years may lead to grave imbalances in cold years and even to the collapses of stocks, such as of capelin in the eighties. The food requirements of average-sized stocks of cod, seabirds and marine mammals correspond to more than twice the average productivity of capelin. Thus other species of pelagic fish (herring, polar cod) and zooplankton obviously play major roles as prey for these animals.     

Beiträge zum Mesozooplankton des Nordsylter Wattenmeers

Contributions to the mesozooplankton of the northern Wadden Sea of Sylt. From June 1975 to June 1976 temperature, salinity, mesozooplankton (>76 µm), phytoplankton and seston (dry weight, particulate organic carbon and nitrogen) were measured at different stations in the northern Wadden Sea of Sylt (German Bight, North Sea). Maxima of the planktonic copepods, which form the biggest part of the zooplankton, occurred in the summer months from June to September. Larval development from nauplii to adults was observed inAcartia clausi, Acartia discaudata, Centropages hamatus andTemora longicornis; generation times ranged from 3 (Temora longicornis) to 7 1/2 weeks (Centropages hamatus) at ca. 20 °C. Organic carbon produced by zooplankton was about 0.4 g C m^–3 year^–1 and zooplanktonic carbon decomposed in the area studied amounted to about 1.4 g C m^–3 year^–1. Meroplanktonic larvae made up ca. 60 % of the organic carbon produced by zooplankton, indicating great influence of the benthos on the water column in this very shallow part of the German Bight.     

Photoadaptation of sea-ice microalgae in the Barents Sea

Summary Variations in under-ice scalar irradiance, P vs I parameters and the CHLa C^–1 ratio of natural assemblages of sea-ice microalgae from the Barents Sea growing at -1.8°C in May and September 1988 are described, including one diurnal station. CHLa C^–1 ratios of 0.031–0.071 mg mg^–1 indicate shade adaptated assemblages both in May and September. Values for ^B (photosynthetic efficiency) were generally low, e.g. 0.0025–0.0078 mg C (mg CHLa)^–1 h^–1 (mol m^–2 s^–1)^–1, and should be typical for self-shaded algae in mats or aggregates of about 4 mm thickness. Provided no self shading and the typical spectral distribution of light under ice without algae, ^B would, however, be about 2.5 times higher. Photoinhibition of the photosynthetic response was negligible. Maximum carbon uptake P _m ^B was 0.15–0.24 and 0.032–0.088 mg C (mg CHLa)^–1 h^–1 in May and September, respectively. Diurnal variations were small, particularly for P _m ^B . Calculations of the maximum specific gross growth rate yielded an upper limit of 0.20–0.24 and 0.01–0.04 d^–1 for assemblages in May and September, respectively; the latter may have been in a resting stage.Contribution No. 245, Trondhjem Biological Station     

Spatial distribution of Pleuragramma antarcticum (Pisces: Nototheniidae) near the Filchner- and Larsen ice shelves (Weddell sea/antarctica)

Summary In the zooplankton collected during three German Antarctic expeditions to the southern Weddell Sea in 1979/80, 1980/81 and 1983, the post-larvae of the nototheniid fish Pleuragramma antarcticum were found in 71–94% of the samples and represented 85–98% in numbers of all fish caught. In 1983, the abundance of post-larvae was up to 88 ind./m² (corresponding to 3 ind./m³). Highest concentrations were generally found over the continental slope and innershelf depressions. More than 70% of the post-larvae were caught in the upper 50–100 m water layer, in the well stratified Summer Water of -1.3 to -0.5°C. The Summer Water is shifted towards the outer edge of the shelf by Ekman transport and accumulation of post-larvae in the slope front and eddies can be explained by this drift. Older Pleuragramma antarcticum of age 1 resemble in their vertical distribution the juveniles and adults, 40–60% of which were caught on the shallower parts of the shelf in cold Ice Shelf Water of -1.8 to -2.1°C in depths below 200 m. Mean abundances of the yearclasses varied by a factor of 16 in age 0 fish and by factor of 10 in age 1. The size of a yearclass may be related to the varying appearance and persistence of an ice free coastal polynya.     

Microbiological Explorations in the Northern Part of the Barents Sea in Early Winter

The total number of microorganisms and rates of microbial processes of the carbon cycle were determined in snow, sea ice, water, and seafloor sediments of the northern part of the Barents Sea from September to October, 1998. The explorations were carried out in two areas: along the transection from Franz Josef Land to Victoria Island and along the continental slope region covered with solid ice at latitude 81°–82° N and longitude 37°–39° E. At the time of study, the ice cover was represented by thick one-year old ice (up to 1.2 m), perennial ice (up to 1.85 m), and pack ice. The number of bacteria in the snow cover, sea ice, and seawater was 12 to 14, 50 to 110, and 10 to 240 × 10³ cells/ml, respectively. Rates of dark CO₂ assimilation, glucose utilization, and methane oxidation by bacteria were determined. The highest rate of microbial processes was found in samples of the lowermost newly formed sea ice. The lowest level of activity for all processes was observed in melted snow water. A direct relation was shown between the concentration of C_org, the bacterial biomass, and the values of ¹³C_org in mixtures of melted snow and ice. The number of microorganisms and rates of microbial processes in seafloor sediments measured at the stations on the continental slope are comparable to those in the central part of the Barents Sea and the northern part of the Kara Sea.     

Annual primary productivity of an antarctic continental lake: Phytoplankton and benthic algal mat production strategies

Primary production in Watts Lake, Vestfold Hills, Antarctica (68°36S, 78°13E), was measured from March 1981 to February 1982. Phytoplankton production peaked in autumn and spring, with a September maximum (340 mgC m^–2 d^–1), then declined in summer and was not detectable in winter. Benthic algal production peaked in summer at 74 mgC m^–2 d^–1), Production strategies differed, with the more efficient phytoplankton adapted to growth at low light, while benthic production increased with increasing light in summer. Estimation of annual production was 10.1 gC m^–2 and 5.5 gC m^–2 for the phytoplankton and benthos respectively.     

Phosphorus and nitrogen excretion rates of zooplankton from the eutrophic Loosdrecht lakes,with notes on other P sources for phytoplankton requirements

Phosphorus and nitrogen excretion rates by zooplankton communities from two eutrophic and shallow Dutch lakes were measured in laboratory. The variations in excretion rates in the lakes (May–October) were caused mainly by fluctuation in zooplankton biomass. Mean summer excretion rates (June–September) were 2.4 and 0.9 µg PO₄P·1^–1·d^–1 in Lake Loosdercht and Lake Breukeleveen, respectively. This difference between the lakes was caused mainly by the lower zooplankton biomass in Lake Breukeleveen. The excretion of 2.4 µg PO₄P·1^–1·d^– compared with the calculated P-demand of phytoplankton of 8.0 µg PO₄P·1^–1·d^–1 is substantial in the summer (June–September) and far more important than the external P-supply of 0.4 µg P·1^–1·d^–1 and sediment release of 0.5 µg P·1^–1·d^–1. Both temperature and composition of zooplankton affected the weight specific excretion rates of the zooplankton community. The weight specific community excretion rates of P and N increased with temperature (exponential model); 1–8 g PO₄P·mg^–1 zooplankton-C·d^–1 and 5–42 µg NH₃N·mg^–1 zooplankton-C·d^–1 (10°C–20°C).     

Latitudinal range of epiplanktonic chaetognatha and ostracoda in the western tropical Indian Ocean

An account of the chaetognath and ostracod species obtained from zooplankton samples collected along a transect in the Western Indian Ocean between 9° N–20° S and 57° 18–68° 43E in January–February 1981 is given. Species richness was maximum north of 10° S. The latitudinal variation of thirteen species of chaetognaths indicated that many of the typical Indo-Pacific species are restricted to the tropical zone with limited penetration into the subtropical region. All the sixteen species of planktonic ostracods were cosmopolitan. Based on the available information on the distribution of chaetognaths and ostracods, the species assemblages in the Indian Ocean are discussed.     

The distribution and abundance of krill faecal material and oval pellets in the Scotia and Weddell Seas (Antarctica) and their role in particle flux

Summary The abundance and depth distribution of zooplankton faeces in spring to early summer were investigated along meridional transects (47°W and 49°W) that extended from the Scotia Sea (57°S) across the Weddell-Scotia Confluence and into the Weddell Gyre (62°S). The sea ice edge retreated from 59°30S to 61°S during the study. Faeces were sampled with nets, Niskin bottles and sediment traps and subsequently analysed by light and electron (SEM) microscopy. Krill faecal strings and oval faecal pellets of unknown origin were by far the most important zooplankton faeces and highest concentrations were always found in the Confluence often close to the ice border. Krill faeces were usually more abundant in the uppermost layer (0–50m) where they contributed an average of 130 g dry weight m^–3. There was an exponential decrease with depth, with a minimum of 0.6 g dry weight m^–3 in the 500–1000 m stratum. Oval pellets were more evenly distributed in the upper 1000 m of the water column, with an average of 9 g dry weight m ^–3, although there was a small peak (20 g dry weight m^–3) in the subsurface layer (50–150 m depth). Consecutive collections (day-night) of krill faeces using drifting sediment traps showed that only the larger strings sank from 50 to 150 m depth. Peritrophic membranes appeared to deteriorate during sinking. Diatoms (in particular Nitzschia and Thalassiosira spp.) contributed by far the bulk of material in krill and oval faeces. In samples collected near or under the pack ice, remains of crustaceans in both krill- and oval faeces were also found.Data presented here were collected during the European Polarstern Study (EPOS) sponsored by the European Science Foundation     

The impact of zooplankton status on the management of Lake Kinneret (Israel)

Monthly averages of standing stock wet biomass of zooplankton in Lake Kinneret (Israel) varied between 11 and 76 g m^–2 during 1969–1981, with the exception of two months. Averaged contributions of different groups were: Cladocera 58%, Copepoda 35% and Rotifera 7%. Total standing crop wet biomass is highest during January–June, averages varied between 35 and 50 g m^–2, and decreases during summer–fall (23–36 g m^–2). The winter biomass of Cladocera fluctuated between 22 and 35 g m^–2 and dropped to a range of 9–23 g m^–2 in summer, whereas copepod biomass varied very little around an average of 18 g (ww) m^–2 with the exception of low values from April to June. The stock biomass of Rotifera is relatively high during winter floods season (December-March) whilst in summer it is very low.Young stages of fish in Lake Kinneret feed mostly on zooplankton and zoobenthic forms. The most abundant fish in the Kinneret ecosystem, Mirogrex terraesanctae terraesanctae, also feed on zooplankton at the adult stage throughout the year, and herbivorous fish consume zooplankton during the summer when lake plankton resources are limited.The summer ecosystem of Lake Kinneret is characterised as a steady state type, in which the impact of the zooplankton-chain is of great importance. Increase of predation pressure on zooplankton by fish can disequilibrate the balanced trophic relations existing between nannoplankton production and zooplankton grazing capacity. Such a situation can lead to organics accumulation as nannoplankton blooms, resulting in water quality deterioration. Management options aimed at preventing collapse of zooplankton populations are discussed.     

Phytoplankton and zooplankton (Cladocera,Copepoda) relationship in the eutrophicated River Danube (Danubialia Hungarica,CXI)

The seasonal variation in primary production, individual numbers, and biomass of phyto- and zooplankton was studied in the River Danube in 1981. The secondary production of two dominant zooplankton species (Bosmina longirostris and Acanthocyclops robustus) was also estimated. In the growing season (April–Sept.) individual numbers dry weights and chlorophyll a contents of phytoplankton ranged between 30–90 × 10⁶ individuals, l^–1, 3–12 mg l^–1, and 50–170 µg l^–1, respectively. Species of Thalassiosiraceae (Bacillariophyta) dominated in the phytoplankton with a subdominance of Chlorococcales in summer. Individual numbers and dry weights of crustacean zooplankton ranged between 1400–6500 individuals m^–3, and 1.2–12 mg m^–3, respectively. The daily mean gross primary production was 970 mg C m^–3 d^–1, and the net production was 660 mg C m^–3 d^–1. Acanthocyclops robustus populations produced 0.2 mg C m^–3 d^–1 as an average, and Bosmina longirostris populations 0.07 mg C m^–3 d^–1. The ecological efficiency between phytoplankton and crustacean zooplankton was 0.03%.     

Growth, Natural Mortality, Length–weight Relationship, Maximum Length and Length-at-first-maturity of the Coelacanth Latimeria chalumnae

Based on the re-interpretation of published data, the von Bertalanffy growth function parameters of the coelacanth, Latimeria chalumnae, are estimated as L=218cm total length, s.e. 23; K=0.059 (year^–1), s.e. 0.012; t₀=–3.3 (year), s.e. 0.5, corresponding to a life span of 48 years. The length–weight relationship of the form W=a*TLb, with wet weight (W) in g and total length (TL) in cm, is estimated as a=0.0278, b=2.89, r²=0.893, n=87, range=42.5–183cm TL. Using extreme value theory, the maximum length for female coelacanths is estimated as 199cm TL (95% confidence interval=175–223cm TL) and for males as 168cm TL (95% confidence interval 155–180cm TL). Based on data from seven females with embryos or mature eggs, the length-at-first-maturity for females is estimated to be about 150cm TL, corresponding to an age of about 16 years. Based on the value of t₀=–3.3 years and on the presence of three scale rings found in a newborn coelacanth, the period of embryogenesis lasts for about three years, the longest known in vertebrates. The natural mortality rate is estimated at M=0.12. Population food consumption is found to be 1.4 times the existing biomass per year, and gross food conversion efficiency indicates that only 10% of the consumed food is utilized for somatic growth.     

Water flux through a semi-deciduous forest grove of the Orinoco savannas

Water relations were analysed in a semi-deciduous forest grove occurring in the oxisols of the Orinoco savannas. This grove has a shallow unconsolidated ironstone cuirass, which is overlaid by a sandy loam layer (0.0–0.5 m) that contains more than 90% of the grove forest root phytomass. Evapotranspiration and through drainage were calculated by using data from the soil profile as related to physical characteristics of the site root zone, hydraulic conductivity, volumetric water content and potential hydraulic gradient. Mean annual evapotranspiration was 783 mm year^–1 and annual through drainage below the root zone was 14% (162 mm year^–1) of the gross rainfall. This drainage recharged the 42% of the annual saturation deficit of the water table. Similar mean annual evapotranspiration (770 mm year^–1) was also calculated by using the water balance components. The mean daily coupling omega factor () between the grove canopy and the surrounding atmosphere indicated that a high degree of coupling (=0.14±0.16) occurs in the grove and evapotranspiration was mainly controlled by surface conductance. As the dry season proceeded, the soil saturation deficit () increased rapidly resulting in a threshold surface conductance (0.030–0.005 m s^–1) for ranging from 0.05 to 0.10. Hypotheses to explain the omnipresence of perennial species in the wide range of physical conditions in neotropical savannas are discussed.     

Zooplankton as a compound mineralising and synthesizing system: phosphorus excretion

Data on phosphate excretion rates of zooplankton are based on measurements using the pelagic crustacean zooplankton of Lake Vechten and laboratory-cultured Daphnia galeata. In case of Daphnia sp we measured the effects of feeding on P-rich algae and P-poor algae (Scenedesmus) as food on the P-excretion rates at 20°C. The excretion rates of the natural zooplankton community, irrespective of the influence of the factors mentioned, varied by an order of magnitude: 0.025–0.275µg PO₄-Pmg^–1C in zooplankton (C _zp ) h^–1. The temperature accounted for about half the observed variation in excretion rates. The mean excretion rates in the lake, computed for 20°C, varied between 0.141 and 0.260 µg Pmg^–1C _zp h^–1. Based on data of zooplankton biomass in the lake the P-regeneration rates by zooplankton covered between 22 and 239% of the P-demand of phytoplankton during the different months of the study period.In D. galeata, whereas the C/P ratios of the Scenedesmus used as food differed by a factor 5 in the experiments, the excretion rates differed by factor 3 only. Despite the higher P-excretion rates (0.258± 0.022 µg PO₄-P mg^–1 C h^–1) of the daphnids fed with P-rich food than those fed with P-poor food (0.105 ± 0.047 µg PO₄-P mg^–1 C h^p–1), both the categories of the animals were apparently conserving P. A survey of the literature on zooplankton excretion shows that in Daphnia the excretion rates vary by a factor 30, irrespective of the species and size of animals and method of estimation and temperature used.About two-thirds of this variation can be explained by size and temperature. A major problem of comparability of studies on P-regeneration by zooplankton relates to the existing techniques of P determination, which necessitates concentrating the animals several times above the in situ concentration (crowding) and prolonged experimental duration (starving), both of which manifest in marked changes that probably lead to underestimation of the real rates.     

Plankton and hydrochemistry of Lake Futalaufquen (Patagonia,Argentina) during the growing season

Plankton communities and hydrochemistry of an oligotrophic lake occupying a glacial valley in Argentinian Patagonia (42 °49S; 71 °43W) were studied. Monthly samples at three stations integrated from 0 to 50 m and stratified samples at the site of maximum depth, were taken during the growing season. Transparency was always controlled by glacial silt, and not by phytoplankton. Lake water belongs to the calcium-bicarbonate type, with low conductivity (24 µS cm^–1), and poor buffering capacity. Forty-five phytoplankton taxa were found. Mean phytoplankton density was 49 cells ml^–1 and mean biomass 69 µg l^–1. N:P relationships, inorganic nitrogen exhaustion in the photic layer, and correlations between nutrients and phytoplankton density suggests nitrogen as the main limiting factor. Fifteen zooplankton species were found. Mean zooplankton density was 12.2 ind. l^–1 and mean biomass 22.9 µg l^–1. Diatoms and Boeckellidae were the dominant planktonic groups. Morphometry and hydrological factors were responsible for horizontal heterogeneity in phytoplankton and chemical variables.     

The relative importance of different ciliate taxa in the pelagic food web of lake constance

Abundance, biovolume, and species composition of pelagic ciliates in Lake Constance were recorded over two annual cycles (1987/88). Production was estimated from mean annual biovolumes and size-specific growth rates obtained from the literature. Cell concentrations and biovolumes ranged from 0.1 to 120 cells ml^–1 and from 3 to 1,200 mm³ m^–3, respectively. Mean annual values were, respectively, 6.8 cells ml^–1 and 94 mm³ m^–3 in 1987, and 12.0 cells ml^–1 and 130 mm³ m^–3 in 1988. In both years, prostome nanociliates (<20m) dominated numerically, while strobiliids in the size range 20–35m contributed most significantly to ciliate production. Ciliate community production, according to a crude calculation, yielded approximately 10–15 g C m^–2 year^–1.     

Living conditions, abundance and biomass of under-ice fauna in the Storfjord area (western Barents Sea, Arctic) in late winter (March 2003)

The under-ice habitat and fauna were studied during a typical winter situation at three stations in the western Barents Sea. Dense pack ice (7–10/10) prevailed and ice thickness ranged over <0.1–1.6 m covered by <0.1–0.6 m of snow. Air temperatures ranged between –1.8 and –27.5°C. The ice undersides were level, white and smooth. Temperature and salinity profiles in the under-ice water (0–5 m depth) were not stratified (T=–1.9 to –2.0°C and S=34.2–34.7). Concentrations of inorganic nutrients were high and concentrations of algal pigments were very low (0.02 g chlorophyll a l^–1), indicating the state of biological winter. Contents of particulate organic carbon and nitrogen ranged over 84.2–241.3 and 5.3–16.4 g l^–1, respectively, the C/N ratio over 11.2–15.5 pointing to the dominance of detritus in the under-ice water. Abundances of amphipods at the ice underside were lower than in other seasons: 0–1.8 ind. m^–2 for Apherusa glacialis, 0–0.7 ind. m^–2 for Onisimus spp., and 0–0.8 ind. m^–2 for Gammarus wilkitzkii. A total of 22 metazoan taxa were found in the under-ice water, with copepods as the most diverse and numerous group. Total abundances ranged over 181–2,487 ind. m^–3 (biomass: 70–2,439 g C m^–3), showing lower values than in spring, summer and autumn. The dominant species was the calanoid copepod Pseudocalanus minutus (34–1,485 ind. m^–3), contributing 19–65% to total abundances, followed by copepod nauplii (85–548 ind. m^–3) and the cyclopoid copepod Oithona similis (44–262 ind. m^–3). Sympagic (ice-associated) organisms occurred only rarely in the under-ice water layer.     

Composition,density and feeding of crustacean zooplankton community in a shallow,temperate lake (Lake Balaton,Hungary)

Composition density and filtering rates of crustacean zooplankton were studied in the open water and among the macrophytes of the oligo-mesotrophic part of Lake Balaton from 1981 to 1983. From the individual filtering rates of the different populations and the densities community grazing rates were derived.Copepoda made up 79–90% of crustacean plankton community in the open lake and 95–97% of it in littoral zone. Among them the nauplii dominated. At the end of summer 1982, when Anabaenopsis was in bloom, the filter-feeding species (Eudiaptomus, Daphnia) practically disappeared, being replaced by cyclopoids. Daphnia had the highest filtering rates followed by those of Eudiaptomus and Diaphanosoma. Among copepods, the filtering rates in decreasing order were: ovigerous > all adults > copepodites > nauplii. > . The filtering rates of the different species varied both seasonally as well as from year to year. In 1983, when the concentration of organic seston decreased, filtering rate increased compared with those in the earlier years. During the water bloom in 1982, the rates decreased by 70% on the average.The community grazing rate was very low (3% per day) in the open lake and among macrophytes, both in 1981 and 1982; also the share of crustacean zooplankton in grazing was very low. In 1983, together with the improving of water quality, the community grazing rate increased 4-fold. In 1981 and 1983 the rates were influenced by water temperature but in 1982 by seston concentration.     

Feeding of Burbot, Lota lota, at Different Temperatures

Daily food intake of adult burbot, Lota lota, fed on vendace, Coregonus albula, were estimated experimentally at four different water temperatures (2.4, 5.1, 10.8 and 23.4°C). Mean daily food intake (MDI; g d^–1) and relative daily food intake (RDI; g g^–1 d^–1) increased with temperature from 2.4 to 10.8°C and decreased at 23.4°C. Temperatures of maximum daily food intake values were 13.6°C for MDI and 14.4°C for RDI. No correlation between food intake values and burbot weight was observed. RDI values were used to estimate annual food consumption of burbot population. Annual food consumption estimates were 9.7kg ha^–1 and 24.3kg ha^–1 when burbot biomass was 2.0 or 5.0kg ha^–1, respectively.     

Zooplankton community response to reservoir aging

Changes in zooplankton diversity and density in response to reservoir aging in Pawnee Reservoir were investigated. Zooplankton samples collected from April 1992 through April 1993, were compared to a similar study conducted after initial impoundment by Helzer (1971), in 1968–1970. Since this initial study, increases in turbidity and resulting changes in biotic interactions significantly altered the zooplankton community. A significant increase in total zooplankton density and a decrease in species richness were observed between study periods. Density increased from 24.6 to 95.4 individuals L^–1, while the number of taxa declined from fourteen to ten. During this time period, Cyclops vernalis became the dominant zooplankter in the reservoir. The density of this predatory copepod increased significantly, from 0.1 l^–1 in 1968–1970, to 44.3 l^–1 in 1992–1993, which accounted for most of the increase in total zooplankton density. Though a greater spring maximum of another dominant, Bosmina spp. was found during the 1992–1993 study period, the annual density of this cladoceran was not significantly different between study periods. Similar trends for Daphnia ambigua and D. parvula were also observed, as greater spring maxima levels were attained, however overall annual densities were not significantly different. The dominance of C. vernalis (46% of annual density) and Bosmina spp. (33%), indicate that these two zooplankters were tolerant of changes in physical conditions resulting from reservoir aging and biotic interactions that followed in the reservoir during the 22 years between study periods.