Structure and resilience of overwintering habitats of Calanus finmarchicus in the Eastern Norwegian Sea

Very high concentrations of overwintering Calanus finmarchicus were found in the eastern Lofoten Basin of the Norwegian Sea close to the shelf break in January 2001–2002. A coupled 3D hydrodynamic and ecological model was used to study the formation of this deep overwintering aggregation and its stability. The ecological model includes nutrients, phytoplankton and microzooplankton in addition to a stage-structured model of C. finmarchicus. Using a Eulerian approach, the model was initiated with an overwintering stock evenly distributed in the oceanic regions of the Norwegian Sea, i.e. where depths>600 m. Spawning and development of the new generation take place in response to vertical mixing and phytoplankton development. Animals are assumed to begin their descent to overwintering depths of 700–1000 m as late stage Vs. Model results show that, in late summer, high concentrations of animals were found at overwintering depths near the shelf break north of the North Sea, off the northeastern Vøring Plateau and in the eastern Lofoten Basin along the slope of the Barents Sea shelf. They remained there for months due to deep eddies and southward, deep currents along the Norwegian shelf. The simulation experiments indicate that the combined effect of deep anticyclonic circulation and vertical migration behavior of the animals may explain the high concentrations of overwintering C. finmarchicus found in field surveys in the Eastern Lofoten Basin, close to the shelf break.     

Oceanic distribution and life cycle of Calanus species in the Norwegian Sea and adjacent waters

The distribution and demography of Calanus finmarchicus, C. glacialis and C. hyperboreus were studied throughout their growth season on a basin scale in the Norwegian Sea using ordination techniques and generalized additive models. The distribution and demographic data were related to the seasonal development of the phytoplankton bloom and physical characteristics of water masses. The resulting quantified relationships were related to knowledge on life cycle and adaptations of Calanus species. C. finmarchicus was the numerically dominant Calanus species in Coastal, Atlantic and Arctic waters, showing strong association with both Atlantic and Arctic waters. C. hyperboreus and C. glacialis were associated with Arctic water; however, C. glacialis was occasionally observed in the Norwegian Sea and is probably an expatriate advected into the area from various origins. Demography indicated one generation per year of C. finmarchicus, a two-year life cycle of C. hyperboreus, and both one- and two-year life cycles for C. glacialis in the water masses where they were most abundant. For the examined Calanus species, young copepodites of the new generation seemed to be tuned to the phytoplankton bloom in their main water mass. The development of C. finmarchicus was delayed in Arctic water, and mis-match between feeding stages and the phytoplankton bloom may reduce survival and reproductive success of C. finmarchicus in Arctic water. Based on low abundances of C. hyperboreus CI–III in Atlantic water and main recruitment to CI prior to the phytoplankton bloom, we suggest that reproduction of C. hyperboreus in Atlantic water is not successful.     

Egg production of Calanus finmarchicus—A basin-scale study

Egg production of Calanus finmarchicus was studied during joint basin-scale surveys in April–June 2003 in the Norwegian Sea. Surveys covered the whole Norwegian Sea and were conducted from Norwegian, Icelandic and Faroese research vessels. Stations were classified as being in pre-bloom, bloom or post-bloom phase according to levels of chlorophyll a and nitrate. Individual egg production rates and population egg production rates were calculated and compared between areas. Both individual egg production rates (eggs female⁻¹ day⁻¹) and population egg production rates (eggs m⁻² day⁻¹) were significantly higher in bloom areas compared with pre-bloom and post-bloom areas. However, when integrated over an estimated duration of the three phases, the time-integrated egg production (eggs m⁻²) in most years was highest in the pre-bloom phase, and this was explained by the longer duration of this phase compared with the two other phases.     

Trans-Atlantic responses of Calanus finmarchicus populations to basin-scale forcing associated with the North Atlantic Oscillation

Populations of the copepod species Calanus finmarchicus often dominate the springtime biomass and secondary production of shelf ecosystems throughout the North Atlantic Ocean. Recently, it has been hypothesised that interannual to interdecadal fluctuations observed in such populations are driven primarily by climate-associated changes in ocean circulation. Here, we compare evidence from the North Sea and Gulf of Maine/Western Scotian Shelf (GoM/WSS) linking fluctuations in C. finmarchicus abundance to changes in ocean circulation associated with the North Atlantic Oscillation (NAO). A particularly striking contrast emerges from this Trans-Atlantic comparison: whereas the North Sea C. finmarchicus population exhibits a negative correlation with the NAO index, the GoM/WSS population exhibits a more complex, positive association with the index. The physical processes underlying these contrasting population responses are discussed in the context of regional- to basin-scale circulation changes associated with the NAO.     

Distribution of Calanus finmarchicus in the northern North Atlantic and Arctic Ocean—Expatriation and potential colonization

The distribution of Calanus finmarchicus was studied on a transect across the central Greenland Sea, and on five transects from the Eurasian shelves across the Atlantic Inflow in the Arctic Ocean. Stage composition was used as an indicator for successful growth; gonad maturity and egg production were taken as indicators for reproductive activity. On the Arctic Ocean transects, these parameters were measured simultaneously from the sibling species Calanus glacialis. Response of egg production rate to different temperatures at optimal food conditions was very similar between both species in the laboratory. C. finmarchicus was present at all stations studied, but young developmental stages were only present close to the regions of submergence of Atlantic water under the Polar water. This together with a decreasing abundance and biomass from west to east along the Atlantic Inflow in the Arctic Ocean and reproductive failure indicates that C. finmarchicus is expatriated in the Arctic Ocean. We hypothesize that the late availability of food in the Arctic Ocean, rather than low temperature per se, limits reproductive success. Better reproductive success in the very low temperature regions of the Return Atlantic Current and the marginal ice zone in the Greenland Sea supports this hypothesis. The possibility for a replacement of C. glacialis by C. finmarchicus and consequences for the ecosystem after increasing warming of the Arctic are discussed.     

The dynamics of the carbon cycle in the surface water of the Norwegian Sea

Historical data of total dissolved inorganic carbon (C_T), together with nitrate and phosphate, have been used to model the evolution of these constituents over the year in the Atlantic water of the Norwegian Sea. Changes in nutrient concentration in the upper layer of the ocean are largely related to biological activity, but vertical mixing with the underlying water will also have an impact. A mixing factor is estimated and used to compute the entrainment of these constituents into the surface water from below. After taking the mixing contribution into account, the resulting nutrient concentration changes are attributed to biological production or decay. The results of the model show that the change in C_T by vertical mixing and by biological activity based on nutrient equivalents needs another sink to balance the carbon budget. It cannot be the atmosphere as the surface water is undersaturated with respect to carbon dioxide and is, thus, a source of C_T in this region. Inasmuch as the peak deficit of carbon is more than a month later than for the nutrients, the most plausible explanation is that other nitrogen and phosphate sources than the inorganic salts are used together with dissolved inorganic carbon during this period. As nitrate and phosphate show a similar trend, it is unlikely that the explanation is the use of ammonia or nitrogen fixation but rather dissolved organic nitrogen and phosphate, while dissolved organic carbon is accumulating in the water.     

Changes in the population structure of Calanus sinicus during summer–autumn in the southern Yellow Sea

Calanus sinicus is a calanoid copepod widely distributed in coastal waters of China and Japan, and oversummering strategies may have major impacts on their population dynamics which in turn affect local marine food web structure. The abundance, stage composition, and sex composition of the planktonic copepod C. sinicus were studied from August to October 2002 in the southern Yellow Sea to understand how its population recovers from the over-summering state. Results showed that C. sinicus had low reproduction in August due to high temperature, except in waters near the Cheju Island with rich food and moderate bottom temperature, but the reproduction rates here decreased in September–October as food availability declined. When temperature dropped in September–October, C. sinicus actively propagated in coastal shallow waters. However, reproduction rates of C. sinicus individuals inhabiting the Yellow Sea Cold Water Mass(YSCWM) remained low during the three months of the study. The percentage of C. sinicus females was high during the reproductive period, which suggests that the sex composition of adult C. sinicus may reflect whether or not the population is in the reproductive mode.Numerous fifth copepodite stage(CV) C. sinicus aggregated in the YSCWM in a suspended developmental stage during the three months of this study, and they potentially served as the parental individuals for population development when conditions became optimal for reproduction later in the year.     

Seasonal distribution of Calanus sinicus (Copepoda, Crustacea) in the East China Sea

On the basis of the four-season investigation in 23°30′~33°N and 118°30′~128°E of the East China Sea from 1997 to 2000, the seasonal distribution of Calanus sinicus was studied with aggregation intensity, regression contribution and other statistical methods. It was inferred that C. sinicus’s predominance presented from winter to summer, especially in spring and summer, because its dominance amounted to 0.62 and 0.29 respectively. The percent of its abundance in copepod abundance was 76.71% in summer, greater than 66.60% in spring, greater than 19.02% in winter, greater than 4.02% in autumn. The occurrence frequency in winter and spring was 83.08% and 93.89%, higher than that in summer and autumn, 76.71% and 73.87%. Compared with other dominant species of copepods, C. sinicus’s contribution to the copepod abundance was obviously greater than that of the other species in winter, summer and spring, but smaller in autumn. C. sinicus tended to have an aggregated distribution. The clumping index peaked in summer (50.19), followed in spring (19.60), declined in autumn (13.18) and was the lowest in winter (3.04). The abundance changed in different seasons and areas, relating to temperature but not salinity in spring and autumn, to salinity but not temperature in summer; to neither temperature nor salinity in winter. In spring and summer, its high abundance area was often located in the mixed water mass formed by the Taiwan Warm Current, the Huanghai Sea Cold Water Mass, the coastal water masses and the Changjiang Dilute Water. In spring and autumn, its abundance was affected by the warm current, as well as the runoff from continental rivers affected it in summer. It can be inferred that C. sinicus was adapted to wide salinity and temperature, as a euryhalinous and eurythermous species in the East China Sea.     

春季中华哲水蚤在不同海湾或河口对环境的适应性

为探究南黄海和东海不同海域春季中华哲水蚤(Calanus sinicus)分布主要受何种环境因子影响及对各环境因子的响应差异,利用2020年春季南黄海海州湾至长江口以北海域和东海长江口海域、三门湾邻近海域、福建中部近岸海域调查数据,基于广义可加非线性模型(GAMs)分析了中华哲水蚤与环境因子的关系。结果表明：海州湾至长江口以北海域中华哲水蚤主要受温度、溶解氧、化学需氧量、Chl a、pH和氨氮的影响。其密度随着温度的增加呈一个波谷曲线变动,温度在14℃时密度最低;随着溶解氧的增加其密度呈一个波谷曲线变动,溶解氧含量在10 mg/L时中华哲水蚤密度最低;随化学需氧量增加其密度波动变化;随着Chl a浓度的增加其密度先下降再呈波动上升;随着pH的增加中华哲水蚤密度逐渐降低;随着氨氮浓度的增加中华哲水蚤密度逐渐增加。长江口海域中华哲水蚤主要受盐度、水深、化学需氧量和活性磷酸盐影响,随盐度和水深的增加其密度逐渐增加;随化学需氧量和活性磷酸盐浓度增加其密度逐渐降低。三门湾邻近海域中华哲水蚤主要受盐度、pH和透明度影响,随盐度的增加其密度逐渐增加;随pH的增加其密度逐渐降低;随透明度的增加其密度先...     

Trophic position of Calanus finmarchicus (Copepoda, Calanoida) in the Trondheim Fjord

The trophic position of Calanus finmarchicus in the Trondheim Fjord in 2004 was determined through stable isotope analyses. Wild specimens were sampled monthly in the fjord and δ¹³C and δ¹⁵N signatures of the developmental stages from CIII to adults were measured. There were statistically significant differences in the δ¹³C and δ¹⁵N signatures of three identified groups: overwintered parental generation, developing new generation and new generation preparing for overwintering. C. finmarchicus individuals raised in a laboratory on a pure algal diet (Dunaliella tertiolecta and Isochrysis galbana) provided stable isotope signatures for purely herbivorous copepods. With these signatures as comparison, the trophic position of C. finmarchicus in the Trondheim Fjord in 2004 was determined as trophic level 2.4, thus indicating omnivory under natural conditions. Additionally, our data suggest that seasonal differences in the δ¹³C signatures of C. finmarchicus are due to the varying lipid content of the different developmental stages.     

Diel vertical migration of the copepod Calanus sinicus before and during formation of the Yellow Sea Cold Bottom Water in the Yellow Sea

To understand the effects of the Yellow Sea Cold Bottom Water(YSCBW)on the diel vertical migration(DVM)of the copepod Calanus sinicus,we surveyed vertical distribution of C.sinicus at a fixed station in the Yellow Sea before(spring)and during(summer)formation of the YSCBW.Cold water(<10 C)was observed in the bottom layer when the water column was thermally stratified in summer,but the water column was thermally well-mixed in spring 2010.Samples were collected from five different layers at 3-h intervals using an opening-closing net.Adult females(1–155 ind./m3)showed a clear normal DVM pattern throughout the entire water column in spring,whereas adult males did not migrate.DVM of copepodite V(CV)individuals was not clear,but the maximum abundance of CI–CIV occurred consistently in the upper 10–20 m layer,where there was a high concentration of chlorophyll-a(Chl-a)(0.49–1.19μg/L).In summer,weak DVM was limited to cold waters beneath the thermocline for adult females(<30 ind./m3),but not for adult males.The maximum abundance of CI–CIV also occurred consistently in the subsurface layer(20–40 m)together with high concentrations of Chl-a(0.81–2.36μg/L).CV individuals(1–272 ind./m3)moved slightly upward nocturnally to the near-surface layer(10–20 m),where the average temperature was 25.74 C,but they were not found in the surface layer(0–10 m;28.31 C).These results indicate that the existence of the YSBCW affected food availability at depth and the vertical temperature distribution,leading to variation in the amplitude and shape of stage-specific vertical distributions(CI to adults)in C.sinicus before and during the formation of cold waters in the Yellow Sea during the study period.     

低氧对中华哲水蚤存活和繁殖影响的实验研究

为研究低氧对海洋浮游生物关键种存活、繁殖等的影响,作者采用了实验室模拟低氧环境的方法,研究了低溶解氧(dissolved oxygen, DO)浓度对中华哲水蚤(Calanus sinicus)致死率、产卵和孵化的影响.研究发现,中华哲水蚤对低氧环境比较敏感,当DO质量浓度为2 mg/L左右时,中华哲水蚤会在1 h内全部死亡;当DO质量浓度为3 mg/L左右时,中华哲水蚤会在96 h内全部死亡;当DO质量浓度为4~6 mg/L 时,基本无死亡,但对中华哲水蚤的产卵有明显抑制作用,首次产卵时间推迟,产卵周期缩短.当DO质量浓度为4 mg/L时,基本不产卵,当DO质量浓度为6 mg/L时,产卵量也仅为常氧组的15%左右.当DO质量浓度为4~6 mg/L时,中华哲水蚤所产卵的孵化率为常氧组的60%左右,但差异不显著.可见,低氧对中华哲水蚤的存活和繁殖均有抑制作用,长时间低氧很可能会降低中华哲水蚤的种群数量,甚至改变海洋生态系统结构.     

黄海中部太平洋磷虾的大小组成和丰度分布的季节变化及其与环境因子的关系

The seasonal size structure and spatial abundance distributions of Euphausia pacifica populations were investigated in the central part of southern Yellow Sea from August 2009 to May 2010.The abundance and biomass of E.pacifica were higher in spring and summer,and lower in autumn and winter.The mean abundance and biomass(calculated by carbon)were 74.94 ind./m~3 and 8.23 mg/m~3,respectively.Females with total length(TL)ranging between 10 and 19 mm in summer had a substantial contribution to the population biomass,whereas larvae of TL of 3–7 mm in spring were the main contributor to the population abundance.The sex ratio(female:male)showed a female bias in four seasons.Its value peaked in summer,and then decreased in autumn,spring,and winter successively.Cohort analysis revealed that the length-frequency distribution of E.pacifica could be characterized as one group with large animals(mean TL12 mm)accompanied by one or two subgroups of small individuals(mean TL7 mm).Regarding the spatial distribution,juveniles and adults of E.pacifica tend to concentrate in relatively deep water with low temperature(～11℃)and high salinity(32),whereas its larvae showed more abundance in inshore water with rich chlorophyll a,low salinity(32),and warm temperature(11℃),especially in summer and autumn.Associations changed seasonally between stage-specific abundance and environmental factors.     

中华哲水蚤抗氧化系统对海水酸化的响应

海洋浮游动物可以对因二氧化碳浓度升高导致的海水酸化做出敏感的响应,然而关于海水酸化对其生理机能的研究却较少。因此,我们研究了暴露于不同二氧化碳浓度（0.08,0.20,0.50 and 1.00%）的酸化条件下中华哲水蚤主要抗氧化酶以及2种解毒酶的活性变化情况。结果表明,暴露于酸化海水中的中华哲水蚤的GPx酶活性显著高于对照组,然而,其它抗氧化组分,包括GST、SOD活性,GSH水平和GSH/GSSG比值均被显著抑制。中华哲水蚤的ATPase活性被海水酸化显著刺激,而AchE活性却显著被抑制。此外,主成分分析结果表明,各指标变化的75.93%可以用第一和第二主成分解释。因二氧化碳浓度升高导致的海水酸化可以通过影响桡足类体内一些酶类的活性来影响其新陈代谢及存活。在未来的研究工作中,需要对海水酸化和其它环境因素的协同作用对桡足类的影响作进一步研究。     

The Norwegian system and the distribution of claims to redfeed

Identifying competing interests and potential claims to a new marine resource, the plankton redfeed, which may be central as a solution to diminishing supplies of marine oils world wide, is critical prior to a commercial harvest thereof. The competing interests are found to be between those Norwegian fisheries organization taking an encompassing view of fisheries interests and those wanting more focus on the well-being of the coastal fishermen. Working within a corporatist setting, it is found, that interest organizations in Norway are included comprehensively by the national government in the decision-making process, but that the main group having the most power is Norges Fiskarlag, which will draw the quota distribution of a future redfeed harvest off shore and away from the coastal fishermen.     

Modelling phytoplankton succession on the Bering Sea shelf: role of climate influences and trophic interactions in generating Emiliania huxleyi blooms 1997–2000

Several years of continuous physical and biological anomalies have been affecting the Bering Sea shelf ecosystem starting from 1997. Such anomalies reached their peak in a striking visual phenomenon: the first appearance in the area of bright waters caused by massive blooms of the coccolithophore Emiliania huxleyi (E. huxleyi). This study is intended to provide an insight into the mechanisms of phytoplankton succession in the south-eastern part of the shelf during such years and addresses the causes of E. huxleyi success by means of a 2-layer ecosystem model, field data and satellite-derived information. A number of potential hypotheses are delineated based on observations conducted in the area and on previous knowledge of E. huxleyi general ecology. Some of these hypotheses are then considered as causative factors and explored with the model. The unusual climatic conditions of 1997 resulted most notably in a particularly shallow mixed layer depth and high sea surface temperature (about 4 °C above climatological mean). Despite the fact that the model could not reproduce for E. huxleyi a clear non-bloom to bloom transition (pre- vs. post-1997), several tests suggest that this species was favoured by the shallow mixed layer depth in conjunction with a lack of photoinhibition. A top-down control by microzooplankton selectively grazing phytoplankton other than E. huxleyi appears to be responsible for the long persistence of the blooms. Interestingly, observations reveal that the high N:P ratio hypothesis, regarded as crucial in the formation of blooms of this species in previous studies, does not hold on the Bering Sea shelf.     

The hydrography and biology of a bloom of the coccolithophorid Emiliania huxleyi in the northern North Sea

In June/July 1994 a study was made of a small bloom of the coccolithophorid Emiliania huxleyi in an area of the North Sea to the east of the Shetland Islands. Observations on the hydrography of the study area indicated the bloom was associated with Atlantic water and was confined to an area in which a stable shallow mixed layer had formed. There was no evidence to suggest association of horizontal physical structure with the bloom development. High cell densities of >1– cells dm⁻³, together with low concentrations of PIC (<50 μg dm⁻³) and detached liths (2– liths cm⁻³) indicated that the bloom was studied at an early stage of development. Biochemical and physiological observations indicated active growth was taking place. The results presented are discussed in comparison with previous studies carried out in both oceanic and shelf seas.     

Reproductive success of Calanus pacificus during diatom blooms in Dabob Bay, Washington

While numerous laboratory studies in the last decade have shown that diatoms can induce reproductive failure in copepods, field evidence for a negative diatom effect is equivocal. To unambiguously elucidate the effects diatoms have on copepod reproduction in situ, we undertook a study of the abundance, distribution, grazing rates, and reproductive success of Calanus pacificus in Dabob Bay, Washington, USA, during two spring bloom periods. We simultaneously measured the phytoplankton composition, abundance, and distribution. Here we present results for the reproductive success of C. pacificus using four measurements: egg production rate, clutch size, egg hatching success, and naupliar survival (to the first feeding stage). Egg production rate was positively correlated with chlorophyll a concentration, and egg hatching success and naupliar survival were usually greater than 80%. However, in February 2002 and 2003 – during blooms of diatoms of the genus Thalassiosira – either egg hatching success, naupliar survival, or both were significantly depressed compared to other times in spring and summer. These effects, combined with evidence that C. pacificus was grazing aldehyde-producing Thalassiosira at the time of their blooms, indicate that diatoms can negatively affect copepod reproduction in the field, albeit only under specific circumstances and for brief periods.