Dynamics of coexisting Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus populations in a high-Arctic fjord

We studied the population dynamics of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in Billefjorden, Svalbard (78°40N). All three species reproduced in the fjord with different timing. The maximum abundance of Calanus spp. copepodite stages peaked on the 11th of July (29,000 ind m^–2). C. glacialis was the dominant species accounting for 60–80% of the total Calanus abundance. C. finmarchicus appear to thrive in the fjord despite the low temperatures (–1.86°C to 5°C) and accounted for 20–30% of the total population. C. hyperboreus contributed less to the total abundance (5–20%). A 1-year life cycle is suggested for C. finmarchicus and C. hyperboreus in the fjord, C. glacialis has a 1- to 2-year life cycle. Highest mortality rates were observed for copepodite stage CV in C. finmarchicus and C. glacialis (0.09 and 0.075 d^–1, respectively) and for females in C. hyperboreus (0.149 d^–1). Mortality of copepodite stages was substantially lower in C. glacialis than in the other species. This is particularly obvious in the early and numerous copepodite stages (CI + CII) during the period of recruitment to these stages. This suggests that differences in secondary production in Arctic pelagic ecosystems are controlled partly by population loss rates.     

Lipid composition of copepodite stages and adult females of Calanus glacialis in Arctic waters of the Barents Sea

Summary The composition of lipid classes and their component fatty acid are described for copepodite stages III, IV, V, and adult females of Calanus glacialis sampled from Arctic waters of the Barents Sea during summer. Was esters were the principal component of the lipid in all copepodite stages examined, averaging 73% over all the stages. The proportion of triacylglycerols varied from 1.5% to 10.5% of total lipid among copepodite stages. The lipids of adult females contained lower levels of wax esters and higher levels of triacylglycerols than copepodite stages III, IV and V. Fatty alcohols of wax esters from copepods sampled in June and July were dominated by 20:1 (n-9) and 22:1 (n-11) alcohols with the proportion of 20:1 (n-9) increasing from stages III to adult female. 14:0 and 16:0 alcohols were the principal fatty alcohols of wax esters of a sample comprising mainly of copepodites stage III taken in August. 16:1 (n-7), 20:1 (n-9) and 20:5 (n-3) were the major fatty acids in the was esters of animals captured in June and July whereas 18:1 (n-9) predominated in the August sample. The polar lipids of the copepodite stage III from August also contained lower levels of polyunsaturated fatty acids than from all stages of copepods from June and July. The data are discussed in relation to life cycle strategies and trophic aspects of Calanus glacialis in the Arctic pelagic community of the Barents Sea.     

Vertical distribution of zooplankton biomass and species in northeastern Baffin Bay related to temperature and salinity

Summary Vertical distributions of various species and stages of zooplankton at different times of the day were determined by stratified sampling with the BIONESS in northeastern Baffin Bay during early August. The water column was divided into an upper subarctic zone (>0° C, salinity < 32), a lower subarctic zone (< 0° C, salinity 32 to 34) and a deep zone of Atlantic water (>0° C, salinity 34). The upper subarctic zone was dominated by two species of pteropod molluscs; the lower subarctic water was dominated by the copepods, Calanus finmarchicus, C. glacialis and C. hyperboreus whose copepodite stages showed depth distributions that were different from one another, with the copepodite stage 5 and adult females generally shallower than the younger stages. All stages of all Calanus species were in the zone of primary production (10 to 50 m) while the copepods Pseudocalanus, Metridia and Oithona were generally found below this zone. Only C. finmarchicus and C. glacialis showed evidence of diurnal migration, migrating to the surface waters when the sun was at its lowest position on the horizon (i.e. at 0100 h).     

Reproduction of Calanus glacialis in the Laptev Sea, Arctic Ocean

Abundance and reproductive biology (gonad maturation and egg production) of the Arctic copepod Calanus glacialis were studied in the Laptev Sea and adjacent Arctic Ocean in September 1993 and from July to September 1995. Both abundance and reproductive activity were subject to strong spatial and seasonal variability, which was related to the ice cover, feeding conditions and circulation pattern. Maximum abundance of the C. glacialis population was generally confined to the outer shelf and slope with depths between 50 and 1000 m. During both cruises, highest egg production rates and largest number of young copepodite stages were observed in the eastern Laptev Sea, where the development of the C. glacialis population seems to follow the opening of the “Siberian Polynya”. In the western part, which is usually covered by pack ice, females were all immature, and no young stages were found. However, females responded quickly to a temporary opening of the ice there in 1995 and spawned. Starvation experiments showed that food-independent reproduction fuelled by internal energy resources was at least partly responsible for relatively high egg production rates at low ambient food concentrations. Egg production rates in starved females were considerably higher than those previously reported. Accepted: 14 June 2000     

The year cycle of Eudiaptomus vulgaris (Schmeil, 1896) (Copepoda,Calanoida) in a small,acid water body during 1973. Development in the natural habitat and relationships between temperature and duration of development stages

The development of a population of Eudiaptomus vulgaris (Schmeil, 1896) in the Meeuwenven, a shallow acid guanotrofic moorland pool, has been described during one year. The population hibernates as copepodite 5 stages, adults and, to a small extent as naupliar stages N1, N2 and N3 (which could not develop further at low temperatures in autumn). In spring the population development starts at temperatures above 10°C and shows 3 or 4 pulses a year. An attempt has been made to explain seasonal changes in the size of adult males and females and in the sex ratio.In order to establish the duration of the various development stages, the animals have been cultured at different temperatures under illumination with 2000 Lux at a daylength of 14 hours. An adequate quantity of food from the natural habitat was available.Total egg development and total naupliar and copepodite development have been compared with the results of other workers, especially with those from Eckstein (1964), who studied Eudiaptomus vulgaris in the deep Schluchsee. The duration curves do not differ markedly with those of Eckstein and are strongly temperature dependent.The relation between the development times of the various stages with temperature can be generally expressed as parabolic regressions of the type D = a + b₁T + b₂T², the C₅ and adult stages being the only exception at higher temperatures. Comparison of the relative duration of the stages at different temperatures did show that younger stages can take a larger share of the total development time at lower temperatures, stage N6 being the most temperature-sensitive.     

Feeding,metabolism and body composition of the Antarctic copepod Calanus propinquus Brady with special reference to its life cycle

Summary In the Weddell Sea, several biochemical and physiological characteristics of copepodite stage III to adult females of Calanus propinquus from the surface layer (0–100 m) and the deep layer (500–1,000 m) were compared at a time of high phytoplankton abundance (February) and a month later, when chlorophyll concentration was much lower and older copepodite stages had migrated to their overwintering depth. Daily rations of surface animals, estimated by the gut fluorescence method, varied from 0.8% to 7.8% body carbon (increasing with phytoplankton concentration). Respiration rate (calculated from measurements of electron-transport-system activity) and lipid and protein body content of animals inhabiting the surface layer decreased from February to April. Deep water animals (stage V and adult females) were characterized by high lipid and protein content; their respiration and excretion rates were much lower than in surface copepods. Our calculation showed that they could survive without an additional source of energy for 8–9 months. Based on our own and published data on bio-chemistry and physiology the possibility of a two-year life cycle is suggested.     

Lipids and life strategies of Calanus finmarchicus, Calanus glacialis and Calanus hyperboreus in late autumn, Kongsfjorden, Svalbard

Stage IV and V copepodites were the dominant forms of Calanus finmarchicus, C. glacialis and C. hyperboreus in Kongsfjorden in late September 1997. Stage IV and V copepodites of C. glacialis and C. hyperboreus were rich in lipid, largely wax esters, and were well fitted to overwinter. Stage IV copepodites of C. finmarchicus were also rich in wax esters, but stage V copepodites of C. finmarchicus were less wax ester-rich. Large size increments between stage IV and V copepodites and between stage V copepodites and females were noted in C. finmarchicus. A very large increment between stage IV and V copepodites was noted for C. glacialis but the size difference between stage V copepodites and females was very small in this species. Particularly large increments were noted between stage IV and V copepodites of C. hyperboreus and also between stage V copepodites and females of this species. The very large, wax ester-rich C. hyperboreus is well adapted to survive the most extreme variations in the Arctic, in Arctic basin waters, whereas the smaller, wax ester-rich C. glacialis is adapted to survive less extreme Arctic variations, as in Arctic shelf waters. The smallest of the three, C. finmarchicus, is best adapted to survive the more predictable waters of the North Atlantic and the Barents Sea. Accepted: 3 January 2000     

Postembryonic development of Drescheriella glacialis Dahms & Dieckmann (Copepoda,Harpacticoida) reared in the laboratory

Summary All 6 naupliar and 5 copepodite stages of Drescheriella glacialis, a tisbid from Antarctic fast ice, are described. A key for the identification of the naupliar stages is given. Sexual dimorphism makes its first appearance in the copepodid IV. The setae configuration of the exopod of the second antenna in copepodid I is considered to be an ancestral state, and subsequent ontogenetic reductions are proposed to be apomorphic processes.     

Lipids in copepodite stages of <Emphasis Type="Italic">Calanus glacialis</Emphasis>

Calanus glacialis is a key herbivore in Arctic shelf seas. It feeds on primary producers and accumulates large energy reserves, primarily as wax esters. Lipid classes, fatty acids (FAs) and fatty alcohols (FAlcs) from copepodite stage II (CII) to adult females (AF) from Kongsfjorden, Svalbard, were studied in May 2004. Wax esters were the dominating lipid class in all stages, ranging from 34% of total lipids in CII to 60% in CIII–CV. Triacylglycerols increased from 8% of total lipids in CII to 23% in AF. In the earlier stages, 16:1n7 and 16:0 FAs and FAlcs were the major components of the neutral lipids, whereas the later stages were mainly characterized by the long-chained FAs and FAlcs 20:1n9 and 22:1n11. C. glacialis utilizes the short spring bloom to build up lipid reserves, mainly as wax esters, and it also incorporates effectively essential polyunsaturated FAs such as 20:5n3 and 22:6n3 in its polar lipids.     

Seasonal size distribution of developmental stages of sub-antarctic copepod

Drepanopus pectinatus, the most numerous species of Copepod in the Kerguelen Archipelago completes 4 generations a year. The succession of generations follows the hydrological seasons. Each generation may be characterized by the cephalothorax length of the developmental stages C1 to C6. The mean lengths of stages C2 to C6 are shown to be significantly different (P > 0.05) between seasons (i.e. between generations). The size-classes have a unimodal distribution from stages C1 to C4. For copepodite C5 and adults, bimodal histograms reflect the differentiation of males and females.A reverse trend of the development pattern of lengths is demonstrated in this subantarctic species: a decrease with decreasing temperatures (6° to 2°) from April to September, and an increase with the rising temperatures (2° to 7°) from October to February.Two populations of females are distinguishable in spring and summer, by their small (0.950–1.299 mm length of cephalothorax, 55%) or large (1.300–1.800 mm, 45%) size, respectively.The differences in mean length cannot be explained by their relation to the mean temperature alone. Pigment content is suspected to play a role.     

Chick diet of dovekies Alle alle in Northwest Greenland

This paper reports the results of an investigation of dovekies (Alle alle) breeding at Hakluyt Island in the Thule District, Northwest Greenland. The paper presents a comprehensive examination of the chick diet of dovekies in the species' most important breeding area. Food delivered to nestlings consisted primarily of copepods Calanus hyperboreus and C. glacialis in the copepodite stages IV and V. Themisto libellula, other amphipods, decapods and fish larvae in small numbers were found in the chick diet. The zooplankton in the chick diet is presented both in terms of numbers and dry weight. The characteristic copepod of high-arctic waters, C. hyperboreus, was found to be the single most important species in the chick diet, contributing approximately 54% of the dry food weight. The food samples revealed low diversity, and the three species C. hyperboreus, C. glacialis and T. libellula made up 91.5% of the total zooplankton dry weight. The results from this paper are compared with previous dovekie-chick diet studies. Accepted: 14 June 2000     

Feeding and digestive activity of cyclopoid copepods in active diapause

Three different methods were used to compare the feeding patterns of active cyclopoids and cyclopoids in active diapause from different habitats. First, we offered ¹⁴C- labeled algal detritus to stage III, stage IV (two different physiological groups), and adult females of Acanthocyclops vernalis from temporary water-bodies of the Volga Delta. Second, we compared the functional responses of active and actively diapausing cyclopoids preying on copepod nauplii. Stage III, stage IV (various physiological groups), stage V, adult males and females of Acanthocyclops viridis, Cyclops kolensis, C. abyssorum sevani from the Volga Delta, Lake Sevan (Armenia), and Mozhaisk Reservoir (Moscow District) were tested. Finally, we examined the activity of digestive enzymes (proteases) of CIV of C. kolensis from Mozhaisk Reservoir at different temperatures and temperature acclimation regimes. All three approaches used have led to similar conclusions. Actively diapausing cyclopoids were capable of feeding on both plant and animal prey; however, their feeding rates and digestive enzyme activity were 2–8 times lower than those of active cyclopoids of similar size. At the initial stage of active diapause the feeding rates were comparable with those of active cyclopoids then declined to a low level during approximately two weeks. Transfer of cyclopoids undergoing winter diapause from ambient temperature (4 °C) to warmer water (20 °C) resulted in dramatic increase of proteases activity only if the acclimation period was relatively long (two weeks).     

Ecological investigations of the zooplankton community of Balsfjorden,northern Norway: Generation cycle,seasonal vertical distribution,and seasonal variations in body weight and carbon and nitrogen content of the copepod Metridia longa (Lubbock)

Metridia longa (Lubbock) has an annual life cycle in Balsfjorden (69°21'N:19°06'E) with spawning occurring from early to mid-May. Development through copepodite stages I-V takes place during summer. At the end of August copepodite stage V accounted for >80% of the population. Overwintering (October-March) was mainly performed by adults. At noon throughout the year copepodite stage V and adults were found within the bottom 30 m of the fjord. The majority of stage IV occurred within the bottom 80 m, while copepodite stages I-III were mainly concentrated within the surface 50 m. Body length, weight, carbon and nitrogen content, and $\text{C}\text{N}$ ratio were determined for copepodite stage V, adult males and females throughout the year. While body length was constant, weight and C and N content varied with season. Lowest $\text{C}\text{N}$ ratios (4–6) were found in copepodite stage V in July and in adults in late winter. Highest $\text{C}\text{N}$ ratios (16–20) were measured in adults from October through early winter. These results are discussed in relation to the life cycle of M. longa and the primary production cycle of the fjord. It is argued that M. longa overwinters in an active state compared with Calanus finmarchicus (Gunnerus) from the same locality.     

The reproductive cycle and life history of the Arctic copepod Calanus glacialis in the White Sea

Seasonal variations in the gonad development and sex ratio of copepodite stage V (CV) and adults were examined from February to November in order to understand the reproductive cycle and the life history of Calanus glacialis in the White Sea. Gonad maturation, sexual differentiation and moulting to adults take place during the 2nd year of development. Energy accumulation takes place in the spring and summer of the 2nd year. The following autumn/winter is the major period of CV maturation, which occurs independent of food supply. Maturation of males precedes that of females by 2–3 months. The maximum proportions of CV and adult males are found in the population in October and November. The onset of female maturation is observed in February and March, ca. 2 months prior to the spring phytoplankton bloom. Reproduction takes place between April and June. Its termination in the second half of June coincides with the warming of the surface water layer where egg laying takes place. Variations in the gonad morphology throughout the year suggest long life spans and iteroparity of females of C. glacialis in the White Sea. Many of them survive for several months after reproduction and are able to overwinter again. Therefore, females with different life histories co-occur in the population in winter: “young” females recently moulted from the overwintering CVs, and “old” females which have spawned at least once in their life, after which they return to overwintering conditions. In contrast, males have shorter life spans of 3–4 months resulting in a sex ratio skewed toward females at all seasons. Accepted: 27 April 1999     

Development of eggs of Antarctic krillEuphausia superba in relation to pressure

Summary This paper focuses upon the influence of increasing hydrostatic pressure on the development of krill eggs at 2°C. This experimental study on the embryology ofEuphausia superba was conducted at the Palmer Station, Antarctica during the 1982–1983 austral summer. The gravid females were captured from Bransfield Strait aboard theR/V Hero. The various embryological stages such as early cleavage, blastula, gastrula and limb-bud nauplius larva were defined and described. The duration for these various developmental stages at 1 atm was also established at +2°C and compared with the timing of this event at negative temperature. Krill embryonic development is inhibited at 4°C. The sinking rate of eggs and embryos was also measured at various pressure. The data suggest that pressure does not significantly influence the sinking rate. There appears to be a wide variation of sinking rates of eggs within the same brood. based on a simulated model of sinking rate, egg development was studied at increasing pressure. Pressure of 5–20 atm accelerates the rate of cleavage and therefore the 32-celled stage is attained within 5–8 h, while at 1 atm it took 13 h to reach the same stage. Pressure thus seems to have some influence on the duration of the development of different developmental stages of krill embryos.     

Winter feeding: an adaptive strategy broadening the niche of the central mudminnow,Umbra limi

Synopsis Mudminnows, Umbra limi, were euryphagic carnivores during the ice-free period consuming invertebrates of terrestrial and aquatic origins. In winter, diversity and amount of food eaten were reduced. Age 0 fish consumed aquatic invertebrates as did males aged 1 and 2+ but females aged 1 and 2+ consumed mainly fishes. In the laboratory mudminnows were able to capture small fish at temperatures of 12.6° C and less. Their rates of digestion of fish prey varied directly with temperature and estimates of time for 100%, gastric evacuation ranged from 161 h at 1.1° C to 26 h at 12.6° C. The ability to forage actively during winter and to digest food relatively rapidly at cold temperatures are seen as mechanisms to broaden the niche along dimensions of resources utilized and time, thereby increasing the chances of survival and reproduction in an environment with high temporal heterogeneity.     

Respiratory metabolism of Cyclops bicuspidatus (sensu stricta) (Claus) (Copepoda: Cyclopoida) from Esthwaite Water,Cumbria

Summary The respiration rate of all stages of Cyclops bicuspidatus (s. str.) (Claus), a benthic copepod, from Esthwaite Water in the English Lake District, was determined at a range of field temperatures, 4° C–12° C, using cartesian diver microrespirometry. The population of C. bicuspidatus in Esthwaite Water was found to be adapted to low temperatures, with an optimum rate of metabolism at 8° C. Weight varied with temperature, in general the largest weights occurred at the lower temperatures. Adult males had higher rates of respiration than adult females, which were on average twice the size of males. Gravid and non-gravid females had similar levels of metabolism. Regression coefficients (b), derived from regressions of log oxygen consumption against log dry weight were low, ranging between 0.25–0.51 according to temperature.     

Experimental studies on the life cycle ofJassa falcata (Crustacea,Amphipoda)

Jassa falcata (Leach), an unselective suspension feeder, was cultivated in standing, unaerated finger bowls, feeding on a mixture ofBrachionus plicatilis, Scrippsiella faeroense, Ulva spec., and natural detritus. This amphipod can also be maintained with diets composed of dead or live material of both plant and animal origin, but an addition of living zooplankton is necessary for longterm cultivation. Starvation resistance is higher in females than in males, and it is increased by lower temperatures. The life span ofJ. falcata increases with decreasing temperature, and it is generally higher in females (maximum: 252 days at 10°C) than in males. In laboratory culture, a bimodal mortality pattern is typical with high juvenile death rates, low mortality during the reproductive phase, and again increasingly high death rates toward the end of the life cycle. Growth rate depends on temperature, sex, and individual age. The temperature dependence of growth is particularly high in males, and it is higher in adults than in juveniles. The total number of moults is lower in males (5 to 6) than in females (7 to 9). Sexual maturity is attained at moult IV to V. The average incubation time of eggs is about 9 to 16 days (highest value at 10°C). Three to 4 broods were observed, with largest numbers of offspring at 10°C.     

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 $\text{C}\text{N}$ 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 $\text{C}\text{N}$ ratio was lowest (4.9) in adult females during the spawning period. Copepodite stage IV and V had higher $\text{C}\text{N}$ 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.     

Diapause in the life cycle of Calanoides carinatus (Kroyer), (Copepoda, Calanoida)

The distribution and stage composition of Calanoidescarinatus (Kroyer) are described together with changes in protein and lipid content, respiration rate and gonad development in copepodite stages IV and V (CV) and adults in the Northern Benguela current (17°S, 23°S and 25°S).During active upwelling the population consisted of two parts: the surface part over the shelf was represented by all development stages, while the deep part offshore was dominated (90–95%) by diapausal CVs. In the surface CVs the surplus assimilated energy was allocated to structural growth and maturation or to synthesis of reserve lipids. CVs with large oil sacs and high lipid content descended into deeper layers and formed diapausal stock; they were characterized by a dramatic decrease of respiration rate. Increase of gonad size in association with decrease of oil sac volume in diapausal CV suggests that reserve lipids were expended not only for respiration but also for gonad development. The moulting of dispausal CVs into adults took place in deep water.These results are discussed in relation to the life cycle of the C. carinatus population and the factors causing the formation and termination of the diapausal phase.