Kurt S. Tande

Ecological investigations on the zooplankton community of Balsfjorden, northern Norway: Generation cycles, and variations in body weight and body content of carbon and nitrogen related to overwintering and reproduction in the copepod Calanus finmarchicus (Gunnerus)

The generation cycles of Calanus finmarchicus (Gunnerus) are described together with the seasonal variations in length, wet wt, dry wt, carbon content, nitrogen content and CN ratio in copepodite stage IV, V and stage VI males and females from Balsfjorden (69°21′N: 19°06′E), a subarctic fjord in northern Norway. C. finmarchicus overwinters in copepodite stage IV (≈ 20%) or V (≈80%) and produces one generation a year. Variations in body weight and body content of carbon and nitrogen in the different copepodite stages showed a pronounced seasonal pattern. For instance, the CN ratio was lowest (4.9) in adult females during the spawning period. Copepodite stage IV and V had higher CN values in summer and autumn (12 to 14) than in spring (8 to 10). Variations in length, weight and chemical composition revealed that the overwintering stock of C. finmarchicus went through two growth phases during this period. From September to January no significant changes in the measured variables were detected. During the second phase of the overwintering period, January to April, the different stages showed a profound decrease in weight and change in chemical composition. This seemed to be connected with the onset of sexual differentiation in stage V starting in January, subsequent moulting into adults and gonad maturation in these adults. These results are further discussed in relation to the different prevailing hypotheses concerning overwintering strategy in Calanus species.

Lipid Composition of Zooplankton in Relation to the Sub-Arctic Food Web

SummarySeasonal changes in the lipid class composition and fatty acid and fatty alcohol composition of neutral lipids were determined for Calanus finmarchicus, Metridia longa and Sagitta sp. in Balsfjord, northern Norway. Similar analyses were obtained for C. hyperboreus and Parathemisto abyssorum in an adjacent fjord, Ullsfjord, in spring. C. finmarchicus, C. hyperboreus, M. longa, and Parathemisto abyssorum all contained large amounts of wax esters whereas Sagitta sp. contained small amounts of triacylglycerols and traces of wax esters. the levels of wax ester in C. finmarchicus and M. longa were highest in late autumn (respectively 88% and 84% of total lipid) and lowest in early spring (respectively 85% and 27% of total lipid). The accumulation of these neutral lipids in spring and summer is related to the feeding activity during the primary production period, while their decline in late winter is associated with the mobilisation of metabolic energy for production of gonads. The major fatty alcohols in the wax esters of C. finmarchicus and C. hyperboreus and Parathemisto abyssorum were 20:1 and 22:1 while those in the wax esters of M. longa were 14:0 and 16:0. The traces of wax esters in Saqitta were rich in 20:1 and 22:1 fatty alcohols. These analyses are consistent with C. finmarchicus and C. hyperboreus being strictly herbivorous, M. longa being more carnivorous and both Sagitta sp. and Parathemisto being highly carnivorous, probably ingesting substantial amounts of calanoid copepods.

The trophic significance of Phaeocystis blooms

Abstract Both colonies and solitary cells of the prymnesiophyte Phaeocystis are ingested by a wide array of marine vertebrates. Grazers include protozoa, bivalves, amphipods, euphausiids and many copepod species. Common fish species such as mackerel (Scomber scombrus) and flounder (Pleuronectus flesus) have also been reported to feed upon Phaeocystis. While qualitative observations on the potential trophic significance of Phaeocystis exist for a long time, attempts to quantify feeding rates on Phaeocystis have begun only during the last decade. In feeding experiments, copepods have received most attention. Many copepod species appear to consume Phaeocystis colonies and solitary cells in the North Atlantic and in polar seas. Ingestion rates are highly variable depending on size and physiological state of Phaeocystis as well as on copepod species. Within the same species, large variations in individual feeding rates have been reported from different areas and investigators. When Phaeocystis co-occurs with larger amounts of diatoms, the latter seem to be preferred by some copepod species while others do not select against Phaeocystis. At present it is unclear whether this is primarily due to unsuitable size of Phaeocystis or because it is poor quality food. The relative nutritive value of Phaeocystis single cells and colonies should be investigated in more detail. Feeding of protozoa on Phaeocystis has been little studied. Ciliates and the giant dinoflagellate Noctiluca are known to consume Phaeocystis solitary cells. Protozoa might efficiently control Phaeocystis blooms during their initial phases when the share of solitary cells relative to total Phaeocystis biomass is higher than during later stages of the bloom. By switching their food preference towards heterotrophic food, copepods might benefit from enhanced protozoan biomass during Phaeocystis blooms. The potential succession of protozoan and metazoan grazers in the course of a Phaeocystis bloom awaits further research.

Does the spawning of Calanus finmarchicus in high latitudes follow a reproducible pattern

In the spring of 1989, an experimental study of the spawning behaviour of Calanus finmarchicus was carried out in Malangen, northern Norway. Here, a single cohort of females reproduce from mid-March to May, approximately coinciding with the wax and wane of the spring phytoplankton bloom. An evaluation of population characteristics such as the proportion of adults, sex ratio, as well as gonad maturation and daily productivity of the females clearly reveals three phases within the populations reproductive period. In between incline and decline, the highest spawning rates (on average >20 eggs female-1 d-1, equivalent to 5.7% body C d-1) occur after the males have disappeared from the population and almost all females have mature gonads. During this period, the ratio of adults to copepodid Stage Vs changes from dominance of adults to that of CVs. Although first egg production was observed prior to the phytoplankton increase, it is suggested that the onset of the phytoplankton spring bloom in the first few days of April enhances the final maturation of ovaries in the females and therefore triggers the onset of the main spawning period. The clutch sizes (max. 95 eggs clutch-1) vary with the “age” of the females, while the spawning frequencies depend on the available food quantities. The overlap of an estimated minimal 4 wk spawning period for the individuals leads to a main reproductive phase for the population of ca. 3 wk, during which time mean clutch sizes and spawning frequencies are maximal (highest average clutch size: 70 eggs female-1 clutch-1, 100 to 60% of the females spawning). This period ends before the end of the phytoplankton bloom. Calculated by stepwise interpolation and summation of the mean daily egg production in the population, an average female produced ca. 600 eggs during the spring bloom in Malangen 1989. We suggest that reproduction and population development of C. finmarchicus in spring follows a reproducible pattern for a given temperature regime and non-limiting food conditions. In the case of clearly identifiable cohorts, it seems possible to trace the state of reproduction by evaluating population parameters.

Ecological investigations of the zooplankton community of balsfjorden, Northern Norway: An analysis of growth and overwintering tactics in relation to niche and environment in Metridia longa (Lubbock), Calanusfinmarchicus (Gunnerus), Thysanoessa inermis (Krøyer) and T. raschi (M. Sars)

Abstract The copepods Calanus finmarchicus (Gunnerus) and Metridia longa (Lubbock) together with the euphausiids Thysanoessa inermis (Kroyer) and T. raschi (M. Sars) are the dominant biomass of the zooplankton in Balsfjorden. Although they all spawn during the phytoplankton production season, a significant portion of their gonad maturation and development occurs during the winter when phytoplankton is absent. Nevertheless, there is a tendency for off-set spawning times and growth patterns in these species. Maximum growth in body weight, favouring increases in lipid and carbon rather than protein or nitrogen, occurs during the primary production season (April to October). In all species overwintering (November to March) results in marked decreases in dry body weight, chiefly as a result of lipid and carbon catabolism. Decreases in dry, carbon, and lipid body weights during winter in both copepods and euphausiids accounted for 40–70% of that present at the end of the primary production season. Although Metridia longa and Thysanoessa raschi take some non-phytoplankton food during winter, metabolism during winter is chiefly sustained on reserves elaborated during spring and summer. There is, however, a trend for the more active copepod ( Metridia longa ) and euphausiid ( Thysanoessa raschi ) to utilize proportionately less of their carbon and lipid than the less trophically active copepod ( Calanus finmarchicus ) and euphausiid ( Thysanoessa inermis ). The last two also rely more on wax esters and less on triacylglycerol as their depot lipid. Such a range of life-cycle tactics is likely to facilitate coexistence of these chiefly herbivorous species in environments where primary production provides short periods with plentiful food followed by long periods of food shortage.

Ecological investigations of the zooplankton community of balsfjorden, northern norway: The genital system in Calanus finmarchicus and the role of gonad development in overwintering strategy

The structure of the reproductive system in Copepodite Stages IV and V of the marine copepod Calanus finmarchicus (Gunnerus) is described, together with seasonal variations in gonad development and sex ratio, from Balsfjorden (69°21′N; 19°06′E), a subarctic fjord in northern Norway. The genital system in Copepodite Stage IV consists of a small gonad with two genital ducts. The gonad of Copepodite Stage V in its most immature condition resembles that seen in Copepodite Stage IV, whereas the sex of Stage V copepodites with mature gonads can be easily and reliably determined. The genital ducts are the key characters for separating males and females in Copepodite Stage V; potential males have one and potential females two genital ducts connecting the gonad to the genital opening on the first urosome segment. Adults males were rare compared to females, and were only in the majority at the end of January and beginning of February. The sex-ratio also favoured females in Copepodite Stage V. Seasonal peaks in the proportion of adult males reflected peaks of males in Copepodite Stage V, suggesting that seasonal variations in the adult sex-ratio are a direct result of a situation which has already been determined in Copepodite Stage V. The seasonal variation in gonad development in Copepodite Stages IV and V of C. finmarchicus reveals that the size of maturity of the gonad varies throughout the year. This study indicates that visible sex-differentiation in Stage V and moulting into adults occurs in Balsfjorden at least 2 months before spawning in April. Phytoplankton levels are immeasurable before the spring diatom increase starts at the end of March, implying that development and maturation of gonads are dependent upon internal energy resources, whereas the final act of spawning seems to require energy input from phytoplankton.

Grazing rates of the copepods Calanus glacialis and C. finmarchicus in arctic waters of the Barents Sea

Natural feeding rates of Copepodite Stages IV and V, and adult female Calanus glacialis (Jaschnov) and Copepodite Stage V and adult female C. finmarchicus (Gunnerus) were estimated using fluorescence analysis of gut contents. Measurements were made on copepods sampled from arctic waters east of Svalbard (Barents Sea) during the spring phytoplankton increase, in the period from 27 May to 13 June, 1983. Observations on Copepodite Stages IV and V and adult female C. glacialis suggest that the gastric evacuation rate is independent of developmental stage, whereas C. finmarchicus Copepodite Stage V showed a lower gastric evacuation rate than adult females. Gut fullness displayed a low correlation with the ambient chlorophyll concentrations. Ingestion rates calculated for C. glacialis were 0.3, 2.3, and 11.0 μg C h-1 for Copepodite Stages IV and V and adult females, respectively. Copepodite Stage V and adult female C. finmarchicus ingested 0.9 and 1.1 μg C h-1 at a temperature of ca.-1.0°C. The maximum ingestion rate in terms of percent body carbon d-1 was higher for adult female C. glacialis and C. finmarchicus than for the respective Copepodite Stage Vs. The results are discussed both in relation to the physiological state of the species and to the environmental conditions.

On the trophic fate of Phaeocystis pouchetii (Hariot). II. Grazing rates of Calanus hyperboreus (Krøyer) on diatoms and different size categories of Phaeocystis pouchetii

Experiments were conducted with CIV and C V copepodites of Calanus hyperboreus (Kroyer) to determine if they would feed on the prymnesiophyte Phaeocystis pouchetii (Hariot). We used analysis of gut pigment to estimate ingestion and clearance rates. In applying this methodology we have demonstrated that pigments can be completely extracted from whole animals within 90 min, and that laborious procedures of tissue homogenization and centrifugation are not required. We conducted two experiments. In the first experiment Stage IV copepodites were exposed to ≈1 mg C·1−1 of either P. pouchetii flagellates, small colonies (25–200 μm), large colonies (> 200 μm) or mixed diatoms > 25 μm (primarily Chaetoceros socialis Lauder and Nitzschia grunowii Hasle). Ingestion rates and daily rations were almost four times greater on both sizes of colonies than on either Phaeocystis pouchetii flagellates or mixed diatoms. Daily rations of copepodites feeding on colonies ranged from 8.1 to 12.4% · day−1, well within the range previously reported for Calanus hyperboreus or sympatric copepods of similar size. From the second experiment we determined that Stage V copepodites obtained a daily ration of 6.2 to 10.8% · day−1 when feeding on small colonies of Phaeocystis pouchetii. We conclude that a diet of P. pouchetii colonies should sustain the metabolic and growth requirements of Calanus hyperboreus copepodites.

Ecological investigations on the plankton community of Balsfjorden, Northern Norway

Abstract The ‘Balsfjord Project’, is an inter-disciplinary research project which concentrates on fish, shell-fish, plankton, and hydrography. In this paper aspects of physical environment relevant to the ecology of phytoplankton and Zooplankton communities in Balsfjorden are presented. Balsfjorden is a single basin (max. depth c. 195 m) fjord separated from the surrounding coastal waters and sounds by shallow sills. It is a cold-water fjord with temperatures from c. 1–7° C during most of the year. Salinity varies from c. 32.80–34.00 %. Temperature and salinity tend to increase slightly with increasing depth. Variations in density are closely related to salinity. Conditions are aerobic, with an oxygen content of 70-120 % saturation. Stratification starts in May and lasts until September. Fresh-water run-off, due to melt water, peaks in June. The pycnocline oscillates around 10 m. A yearly inflow of heavier and warmer Atlantic water occurs, starting in the spring, resulting in deep-water renewal in the bas...

Post-bloom grazing byCalanus glacialis, C. finmarchicus andC. hyperboreus in the region of the Polar Front, Barents Sea

The plankton community in the Polar Front area of the Barents Sea was investigated during a cruise from 14 to 28 July 1987. The colonial algaePhaeocystis pouchetii andDinobryon pellucidum dominated the phytoplankton. Depth integrated carbon assimilation rates varied from 190 to 810 mg C m−2 d−1. A high carbon:chlorophyll ratio (which varied from 123 to 352) prevailed at the three stations investigated, which may relate to facultative heterotrophic behaviour byD. pellucidum. The herbivorous zooplankton community was dominated byCalanus glacialis, C. finmarchicus, andC. hyperboreus. Maximum zooplankton biomass was found in the same depth strata as phytoplankton chlorophyll maximum. The herbivorous copepod populations did not display consistent day-night vertical migration patterns. Phytoplankton consumption rates of the various life stages were estimated from the turnover rate of plant pigments in the gut. The gut defecation rate constant (R) varied from 0.014 to 0.027 min−1 at 0°C in copepodites (Stage II to adult female) ofC. glacialis, independent of developmental stage.Calanus spp. community carbon ingestion rates calculated from particulate carbon:chlorophyll ratios, were 10, 65 and 400% of daily phytoplankton carbon fixation rates at Stations 1, 2 and 3, respectively.