Copepod grazing and its potential impact on the phytoplankton development in the Barents Sea

Compiled data from published and unpublished sources on copepod grazing of the large-sized copepods in the Barents Sea give wide ranges in grazing rates. Approximate average values indicate daily rations of 7–18% for copepodite stages V and VI and considerably higher values for the earliest copepodite stages. It is demonstrated that individual variability in gut fullness of copepods from a given locality is typically very high and not closely related to variable food abundance or depth of occurrence. There is no diel feeding rhythm during the summer, and even when relating copepod grazing to a number of biotic and abiotic factors through stepwise linear regression analysis, much of the variability remains unexplained. It is suggested that feeding behaviour, food quality and feeding history of the copepods all play important roles as factors which regulate copepod grazing. Model simulations on the phytoplankton succession, using literature data on laboratory-determined growth characteristics for solitary cells and colonies of the prymnesiophyte Phaeocystis pouchetii and large diatoms, indicate that the extent of the mixed layer and selective grazing by zooplankton are important factors that may explain the occurrence of dense blooms of P. pouchetii colonies, frequently observed during the spring.     

Prey composition and feeding rate of Sagitta elegans var. arctica (chaetognatha) in the Barents Sea in early summer

Sagitta elegans var. arctica , the dominant and locally abundant chaetognath in the Barents sea, was collected from the upper 50 m in Arctic water masses during an ice edge bloom in early summer 1983. In situ sampling was made along a transect at discrete depths with a 375 μm mesh net mounted on a plankton pump. Prey composition and feeding rate were estimated from gut content analyses on preserved specimens combined with data on digestion times from previous studies. No diel variations were found in feeding activity. The diet reflected the composition of available prey in the zooplankton and consisted mainly of nauplii, small copepods (early stages of Calanus, Pseudocalanus, Oithona ) and appendicularians. Prey usually occurred as a single item in the gut.
Mean prey body width related to chaetognath head width yielded a power curve, with a large amount of scatter, showing that chaetognaths in the Barents Sea can use a wide spectrum of prey sizes. Similarly, maximum prey body width was related to chaetognath head width as a power curve, reflecting the existence of an upper prey size limitation due to the chaetognath mouth size. The highest abundance of S. elegans (5 specimens m⁻³), and the most intense feeding activity, were found within or beneath the maximum zooplankton biomass. Further, distribution and feeding were affected by light intensity, salinity, and the population structure of 5. elegans var. arctica.
Estimated feeding rates ranged between 0.30 and 1.05 prey items per chaetognath day⁻¹. This corresponds to an ingestion of 8-54 μg AFDW day⁻¹, and a consumption of 0.08–0.22% of the zooplankton standing stock day⁻¹. From these rates, the calculated yearly ingestion by S. elegans var. arctica was 3% of the annually secondary production.     

Calanus in North Norwegian fjords and in the Barents Sea

The Physical environment of a North Norwegian fjord and of the Atlantic and Arctic domains of the Barents Sea are described. The seasonal variation of primary production and biomass of the most important copepod species are described in order to contrast regional differences in the timing of the plankton cycles. Analysis of the seasonal variation in the biomass of six different copepod species in Balsfjorden clearly demonstrate the importance of Calanus finmarchkus as a spring and early summer form, whereas Pseudoculanus acuspes , the most important smaller form, reaches the highest biomass later during the productive season. In the Atlantic part of the Barents Sea, C. finmarchkus is the dominant herbivorous form. The next most important species, Pseudocalanus sp. and M. longa , play a less important role here than in Balsfjorden. In the Arctic domain, the smaller copepod forms appear to have been replaced in trophodynamic terms by the youngest year-group (C-CIII) of C. glacialis , which prevails during the Arctic summer and autumn periods. The coupling between primary producers and Calanus on a seasonal basis is addressed through the grazing and the vertical organisation of the plant-herbivore community. The productivity of these two Calanus species is considered in relation to the seasonal and inter-annual variation in climate; although different mechanisms are utilised, cold periods tend to lower Calanus productivity both in the Arctic and the Atlantic domains of the Barents Sea. Interannual variations in Calanus biomass and productivity are discussed in the perspective of endemic and advective processes.     

Gonad maturation as an indication of seasonal cycles for several species of small copepods in the Barents Sea

In a remote oceanic area like the Barents Sea, it is often difficult to follow the seasonal development of copepod populations in detail. Information on the gonad maturation stage of older juveniles and adults of a species will reveal the immediate state of reproduction and the expected development of juveniles into reproductively active adults. Winter "resting stages" in juveniles can also be recognised.
Zooplankton were caught during Pro Mare cruises in early March, May, July/August, mid-September and mid-October. Abundance and composition of developmental stages of small copepods were determined for several stations from each cruise. Samples consisting of Stages CIV to CV1 of Pseudocalanus acuspes (Giesbrecht 1881), P. minutus (Kröyer), Microcalanuspusillus (Sars), and M. pygmaeus Sars were stained with carmine and analysed with respect to gonad maturation stage, length, width, and area of the prosome and the area of the gonad and the oil sac. Image analyses were performed from photographs or drawings of copepods using a digitising pad.
With additional information on abundance and stage composition, and by comparing the present data set with information on Pseudocalanus spp. from Balsfjorden, northern Norway, seasonal cycles for the species could be inferred.     

Horizontal distribution of calanoid copepods in the western Arctic Ocean during the summer of 2008

The horizontal distribution of the epipelagic zooplankton communities in the western Arctic Ocean was studied during August–October 2008. Zooplankton abundance and biomass were higher in the Chukchi Sea, and ranged from 3,000 to 274,000 ind. m^?2 and 5–678 g WM m^?2, respectively. Copepods were the most dominant taxa and comprised 37?94% of zooplankton abundance. For calanoid copepods, 30 species belonging to 20 genera were identified. Based on the copepod abundance, their communities were classified into three groups using a cluster analysis. The horizontal distribution of each group was well synchronized with depth zones, defined here as Shelf, Slope and Basin. Neritic Pacific copepods were the dominant species in the Shelf zone. Arctic copepods were substantially greater in the Slope zone than the other regions. Mesopelagic copepods were greater in the Basin zone than the other regions. Stage compositions of large-sized Arctic copepods (Calanus glacialis and Metridia longa) were characterized by the dominance of late copepodid stages in the Basin. Both the abundance and stage compositions of large copepods corresponded well with Chl. a concentrations in each region, with high Chl. a in the Shelf and Slope supporting reproduction of copepods resulting in high abundance dominated by early copepodid stages.     

Photosynthesis-irradiance relationships in natural phytoplankton populations of the Barents Sea

An analysis is made of the photosynthesis-irradiance relationships in natural phytoplankton populations in the Barents Sea. The data set comprises 232 experiments carried out during a 10-year period, both in open and ice-covered waters. The variability on the P-I parameters is discussed and examined in relation to the variation in a variety of environmental conditions. The results suggest that in the Barents Sea, as in other Arctic areas, phytoplankton photosynthesis is mainly controlled by physical variables. However, control of the phytoplankton stock, i.e. by zooplankton grazing, seems also to have a considerable indirect influence on P-I parameters, especially after the spring bloom and the depletion of winter nutrients.     

The diets of common and Brünnich's guillemots Uria aalge and U. lomvia in the Barents Sea region

Guillemots Uria spp. account for ca. 70% of the total harvest of prey taken by seabirds breeding in the Barents Sea region. This paper presents guillemot chick diet data collected recently at four localities (Finnmark, Murman, Bjørnøya and Spitsbergen) and collates all the data found by the authors in the literature and in the archives of Tromsø Museum, the Norwegian Polar Institute and Kandalaksha State Nature Reserve. Guillemots consume a wide variety of prey and, in comparison to the harvest by predatory fish and marine mammals, their impact on the Barents Sea ecosystem is considered to be minimal. We point out the need for more systematic collection of data from different regions and at different times of the year before a final impact assessment can be made.     

Differential responses of zooplankton populations (Bosmina longirostris) to fish predation and nutrient-loading in an introduced and a natural sockeye salmon nursery lake on Kodiak Island, Alaska, USA

Stratigraphic changes in the remains of Bosmina longirostris from a lake with an introduced sockeye salmon population and a lake with a natural salmon run on Kodiak Island demonstrated markedly different responses to past fluctuations in salmon populations. In both lakes, there was a positive correlation between the density of Bosmina microfossils and the abundance of sockeye salmon. However, opposite size trends were observed in the two lakes. In Karluk Lake, which has a native sockeye salmon population, Bosmina mean carapace lengths were largest at high salmon densities, and mean mucro and antennule lengths were also large, suggesting strong predation pressure from cyclopoid copepods, and less intense pressure from juvenile sockeye salmon. As salmon-derived nutrients are important in driving primary productivity in this system, changes in zooplankton productivity track salmon escapement, but grazing pressure on Bosmina from juvenile salmon is less important than that from cyclopoid copepods. In Frazer Lake, a lake with an introduced salmon population, Bosmina morphologies were smallest during periods of high sockeye salmon in the lake, suggesting much stronger predation effects from sockeye salmon due to the suppression of Cyclops columbianus. Latent development of compensatory mechanisms and the delayed recovery of copepod populations to salmon introductions has resulted in zooplankton populations that are still recovering from shifts in fish populations that occurred decades earlier. The differential response of Bosmina populations between the natural and manipulated lakes suggests that care must be taken when attempting to extrapolate results from whole-lake manipulations and short-term experiments to natural systems.     

2012年夏季西北冰洋浮游动物群落特征

北冰洋夏季海冰覆盖面积在2012年达到低值。为了了解海冰变化对浮游动物群落的影响, 利用夏季西北冰洋22个站位的网采样品, 通过种类组成和丰度研究了群落的类型、结构和地理分布, 探讨了其分布特征与环境因子的关系。根据记录到的54种(类)浮游动物, 21站位可以划分成在地理上基本隔离的三个浮游动物群落: 楚科奇海南部群落, 藤壶幼体数量占优, 站位丰度百分比在56.6%—79.8%之间, 桡足类次之(18.0%—42.2%), 同时还含有少量的白令海种类;楚科奇海中北部群落以广布性桡足类占绝对优势(62.3%—96.8%), 藤壶幼体次之(0—30.9%);深海群落浮游动物的丰度极低, 组成上以桡足类为主(71.6%—89.8%), 且多数是体型较大极地种。楚科奇海陆坡边缘的M06站丰度较高但是种类组成与深海站位相似, 没有归入任何群落。两个浅水群落优势种都是北极哲水蚤(Calanus glacialis)、伪哲水蚤类(Pseudocalanus sp.)、圆胃住囊虫(Oikopleur vanhoffeni)以及藤壶幼体(Barnacle larva), 但优势度各异。深海群落优势种较多, 北极哲水蚤(Calanus glacialis)、极北哲水蚤(Calanus hyperboreus)、细长长腹水蚤(Metridia longa)以及北极拟真刺水蚤(Paraeuchaeta glacialis)等体型较大的桡足类优势度较高, 体型较小的矮小微哲水蚤(Microcalanus pygmaeus)、长腹剑水蚤(Oithona similis)优势度相对较低。与群落类型按维度和深度的变化趋势一致, 统计分析显示表层温度和表层盐度是最重要的影响因子。与海冰覆盖面积较高的2003年相比, 群落类型和地理分布没有显著变化, 但是楚科奇海浮游动物丰度增加了1—2倍, 深海群落丰度降低而组成上大型种类比例升高。     

Correlations between pelagic distribution of Common and Brünnich's Guillemots and their prey in the Barents Sea

Correlations between guillemots (including Common Guillemots Uria aalge and Brünnich's Guillemots U. lomvia) and their prey (divided into five prey categories, capelin Mallotus villosus , herring Clupea harengus , polar cod Boreogadus saida , plankton, and a mixture of other prey species) at two depths (10-100 m and 100-200m) were estimated along an extended transect of 3,060 nautical miles (5,667 km) in the Barents Sea in April/May 1986. Spatial concordance was highest during daylight hours when the largest number of birds were seen on water (presumably feeding birds). Capelin was the single prey category which was most often associated with birds but no single prey category could alone explain the distribution of birds. Although only a small fraction of guillemots could be identified to species, there was some evidence that capelin were of greater importance to Common than to Brünnich's Guillemots. Overall correlation between birds and total prey density was statistically significant at the smallest scale of 5 nautical miles (n.m.). The removal of herring from the calculations increased the strength of the correlation. The depth at which prey was located had little effect on the distribution of birds. The correlation between birds and prey was scale dependent, and reached a maximum at 90 n.m., although there seemed to be some upper threshold in the coefficient at c. 40 n.m. Numerical concordance (including only 5 n.m. periods where both prey and birds were present) was significant at the 5 n.m. scale but was higher for high density than for low density prey patches. The results are discussed in relation to the few similar studies in other oceans and in relation to the severe reduction of important prey species in the Barents Sea.     

南极沿海伯尔顿盐湖两种越冬浮游动物温度耐性的初步研究

本文报导了南极大陆沿海伯尔顿湖两种越冬浮游动物:双刺镰状水蚤和球栉水母的耐热和耐过冷能力的实验.结果表明,两种动物具有较强的温度耐性,其12小时LD_(50)的上、下限温度,雌性双刺镰状水蚤可在14.8℃和—10℃偏下,球栉水母可在13.1℃和—8℃偏下.它们忍受的温度范围大于其他一些南极沿岸海洋无脊椎动物的.这两种动物的耐温能力有明显的季节差异,表明动物具有较强的适应环境温度变化和生理调节能力,这可能是受环境温度自然驯化的结果.     

Impact of grazing from capelin (Mallotus villosus) on zooplankton: a case study in the northern Barents Sea in August 1985

The distribution of capelin was mapped in the area east of Hopen. Zooplankton was sampled with Juday net and 1 m² MOCNESS sampler, and analysed with respect to hydrography and capelin abundance. The capelin "front" coincided more or less with the physical Polar Front, and this complicated the interpretation of the results. Strong indications for a grazing impact by capelin on zooplankton were nevertheless obtained. The zooplankton biomass was significantly lower in the area with high abundance of capelin than in the area with no capelin. This effect was due to a lower biomass of relatively large zooplankton (> 1 mm size fraction) and seen most clearly in data obtained with the MOCNESS. The biomass of zooplankton in the upper 100 m was very low where capelin was present, suggesting rapid depletion of the major prey items. The biomass (m ⁻²) of capelin in the capelin front area was about three times higher than the biomass of zooplankton in areas without capelin. The capelin front would therefore have the potential to graze down the available prey in 3-4 days. Light seems to be an important factor for the predation impact by capelin, resulting in strong interactions between capelin predation and zooplankton vertical distribution.     

Spatial variability of phytoplankton, nutrients and new production estimates in the waters around Svalbard

Phytoplankton dynamics and carbon input into Arctic and sub-Arctic ecosystems were investigated around Svalbard, in summer 1991. Phytoplankton biomass, species composition and dissolved nutrient concentrations were analysed from water samples collected along seven transects. Phytoplankton biomass was low especially to the north (Chlorophyll-a mean 0.3 pg 1- '), where flagellates dominated the communities and only ice-diatoms were present. To the west, the phytoplankton composition was representative of a summer Atlantic community, in a post-bloom state. Zooplankton grazing, mainly by copepods, appeared to be the main control on biomass to the west and north of Svalbard.
In the Barents Sea (east of Svalbard), an ice edge bloom was observed (Chlorophyll-a max. 6.8 pgl-') and the ice edge receded at a rate of approximately 1 1 km day-'. The phytoplankton community was represented by marginal ice species, especially Phaeocystis poucherii and Chaeroceros socialis. South of the ice edge, Deep Chlorophyll Maxima (DCM) were observed, as surface waters became progressively nutrient-depleted. In these surface waters, the phytoplankton were predominantly auto- and heterotrophic flagellates.
Carbon production measurements revealed high net production (new and regenerated) to the north of the Barents Sea Polar Front (BSPF); it was especially high at the receding ice edge (reaching 1.44gC m-'day-'). To the south, a low level of production was maintained, mainly through regenerative processes.     

Egg production rates of two common copepods in the Barents Sea in summer

Small copepod species play important roles in the pelagic food webs of the Arctic Ocean, linking primary producers to higher trophic levels. The egg production rates (EPs) and weight-specific egg production rates (SEPs) of two common copepods, Acartia longiremis and Temora longicornis, were studied under experimental conditions in Dalnezelenetskaya Bay (southern Barents Sea) during summer. The average EP and SEP at 5–10 °C were 4.7 ± 0.4 eggs female⁻¹ day⁻¹ and 0.025 ± 0.002 day⁻¹, respectively, for A. longiremis and 13.1 ± 0.9 eggs female⁻¹ day⁻¹ and 0.075 ± 0.006 day⁻¹, respectively, for T. longicornis. EP and SEP were significantly higher at 10°C than at 5°C for both species. The mean egg diameter correlated positively and significantly with female prosome length (PL) in each species. SEP of T. longicornis correlated negatively and significantly with PL. Daily EP and SEP were similar to rates recorded for other Acartia and Temora species in temperate and warm regions. The influence of environmental factors (temperature, salinity, and phytoplankton concentration) on EP of both species is discussed. We conclude that temperature is the main factor determining the reproduction rate and timing in A. longiremis and T. longicornis in the Barents Sea.     

2012年马卡诺夫海盆与楚科奇深海平原浮游动物夏季的垂直分布及地理差异

利用2012年9月1—6日采自马卡诺夫海盆3个站位和楚科奇深海平原1个站位的分层浮游动物样品,研究了浮游动物在0—1 000 m水层的垂直分布以及地理差异。结果表明,浮游动物在上层分布密集而在深层比较稀少。4个站位在0—50 m、50—100 m和100—200 m水层的平均丰度分别为265.0、360.7和231.2 ind·m^-3,而在200—500 m和500—1 000 m的丰度只有64.4和36.9 ind·m^-3。在数量上占优势的种类中,植食性为主的拟长腹剑水蚤(Oithona similis)、北极哲水蚤(Calanus glacialis)和极北哲水蚤(Calanus hyperboreus)集中在200 m以浅的水层。虽然在200m以下杂食性种类矮小微哲水蚤(Microcalanus pygmaeus)、隆剑水蚤(Oncaea spp.)和细长长腹水蚤(Metridia longa)的丰度明显降低,但是占浮游动物总丰度的比例却明显更高。两个调查海区浮游动物种类组成相似,但是楚科奇深海平原大型桡足类极北哲水蚤的丰度较高,而小型桡足类丰度较低?垂直分布上差异主要在于500—1 000 m水层,马卡诺夫海盆站位丰度为22.7—92.6 ind·m^-3,而楚科奇深海平原只有1.6 ind·m^-3。深海区浮游动物丰度的地理差异说明生物泵的作用存在空间异质性。类似地理差异产生的原因在于楚科奇深海平原存在数量较多的极北哲水蚤,它们在春季融冰前就上升到表层摄食冰藻,显著降低了有机物的垂直通量。     

Are bacteria active in the cold pelagic ecosystem of the Barents Sea?

Bacterial biomass and activity indicators have been studied at low water temperatures (−1.9 to +4°C) in Barents Sea. Strong responses by indicators of bacterial activity, such as hydrolytic enzyme and substrate uptake potentials, were observed in association with the development of phytoplankton blooms. At late successional stages of blooms, observation by epifluorescence microscopy revealed heavy bacterial colonisation of detrital matter, in particular of senescent colonies of Phaeocystis pouchetii . Based on the retention of bacteria on filters of 1 μm pore size, up to 55% of the bacterial population was estimated to be attached to organic aggregates in some cases. Based on thymidine incorporation and a conventional conversion factor, bacterial generation times as short as one day were estimated at temperatures below zero. Changes in substrate availability governed by the successional stages of the planktonic ecosystem seem to be more important as controlling factors for bacterial growth than the low temperatures of the Barents Sea.     

Ice draft and current measurements from the north-western Barents Sea, 1993–96

From 1993 to 1996, three oceanographic moorings were deployed in the north-western Barents Sea, each with a current meter and an upward-looking sonar for measuring ice drafts. These yielded three years of currents and two years of ice draft measurements. An interannual variability of almost I m was measured in the average ice draft. Causes for this variability are explored, particularly its possible connection to changes in atmospheric circulation patterns. We found that the flow of Northern Barents Atlantic-derived Water and the transport of ice from the Central Arctic into the Barents Sea appears to be controlled by winds between Nordaustlandet and Franz Josef Land, which in turn may be influenced by larger-scale variations such as the Arctic Oscillation/North Atlantic Oscillation.     

Primary production of the northern Barents Sea

The majority of the arctic waters are only seasonally ice covered; the northern Barents Sea, where freezing starts at 80 to 81°N in September, is one such area. In March, the ice cover reaches its greatest extension (74-75°N). Melting is particularly rapid in June and July, and by August the Barents Sea may be ice free. The pelagic productive season is rather short, 3 to 3.5 months in the northern part of the Barents Sea (north of the Polar Front, 75°N), and is able to sustain an open water production during only half of this time when a substantial part of the area is free of ice. Ice algal production starts in March and terminates during the rapid melting season in June and July, thus equalling the pelagic production season in duration.
This paper presents the first in situ measurements of both pelagic and ice-related production in the northern Barents Sea: pelagic production in summer after melting has started and more open water has become accessible, and ice production in spring before the ice cover melts. Judged by the developmental stage of the plankton populations, the northern Barents Sea consists of several sub-areas with different phytoplankton situations. Estimates of both daily and annual carbon production have been based on in situ measurements. Although there are few sampling stations (6 phytoplankton stations and 8 ice-algae stations), the measurements represent both pelagic bloom and non-bloom conditions and ice algal day and night production. The annual production in ice was estimated to 5.3 g Cm^-2, compared to the pelagic production of 25 to 30 g Cm^-2 south of Kvitøya and 12 to 15 g Cm^-2 further north. According to these estimates ice production thus constitutes 16% to 22% of the total primary production of the northern Barents Sea, depending on the extent of ice-free areas.     

Insights into the lithospheric structure and tectonic setting of the Barents Sea region from isostatic considerations

We study the tectonic setting and lithospheric structure of the greater Barents Sea region by investigating its isostatic state and its gravity field. 3-D forward density modelling utilizing available information from seismic data and boreholes shows an apparent shift between the level of observed and modelled gravity anomalies. This difference cannot be solely explained by changes in crustal density. Furthermore, isostatic calculations show that the present crustal thickness of 35–37 km in the Eastern Barents Sea is greater than required to isostatically balance the deep basins of the area (>19 km). To isostatically compensate the missing masses from the thick crust and deep basins and to adequately explain the gravity field, high-density material (3300–3350 kg m⁻³) in the lithospheric mantle below the Eastern Barents Sea is needed. The distribution of mantle densities shows a regional division between the Western and Eastern Barents and Kara Seas. In addition, a band of high-densities is observed in the lower crust along the transition zone from the Eastern to Western Barents Sea. The distribution of high-density material in the crust and mantle suggests a connection to the Neoproterozoic Timanide orogen and argues against the presence of a Caledonian suture in the Eastern Barents Sea. Furthermore, the results indicate that the basins of the Western Barents Sea are mainly affected by rifting, while the Eastern Barents Sea basins are located on a stable continental platform.     

Food energy requirements of the harp seal (Phoca groenlandica) population in the Barents and White Seas

The harp seal ( Phoca groenlandica ) population of the Barents and White Seas has probably decreased from about one million individuals to half this size the last few years. Energy requirements of the population have been estimated by use of the simulation model SEAERG. In this model the energy requirements of an individual seal from each age group is multiplied with the group size and summed to provide the requirements of the population. In addition to population size and age structure the total food and energy requirement is sensitive to individual activity levels as well as metabolic levels and other specified physiological functions. The interactions between the seal population and fisheries depends on the caloric density of the prey species which varies with season and location. Realistic simulations of interactions between seals and fisheries require more information about spatial and temporal variations in the prey selection of harp seals than is available today.
Present estimates indicate average maintenance requirements of about 13,600 and ll,150kcal/day for adult female and male harp seals respectively. The high value for the females is due to the costs of pregnancy and lactation. With a mean energy density of prey of 1500 kcal/kg, the corresponding food consumption is 9 kg/day for females and 7.4 kg/day for males.