Respiration and excretion rates of Calanus glacialis in arctic waters of the Barents Sea

The respiration and excretion rates of Calanus glacialis (Jaschnov) Copepodite Stages III, IV, V, and adult females from the drift-ice area east of Svalbard (Barents Sea) were measured in shipboard experiments during the period from 27 May to 13 June, 1983. The phytoplankton biomass and abundance varied considerably between localities, but these variations were not generally reflected in the respiration and excretion rates of the copepod. The respiration and excretion rates of C. glacialis at the ambient temperature of-1.8°C (average respiration rates of 0.95, 0.73, 0.57, and 0.60 l O₂ mg^-1 dry wt h^-1 for Copepodite Stage III, IV, V, and adult females, respectively) were similar to those previously reported for other large-sized copepods from cold or temperate areas. Average respiration and excretion rates tended to decrease with incubation time or time after capture. Measurements on ten occasions within a period of 27 h after capture revealed excretion rates of ammonium ranging between 2.9 and 16.8 for C III, 3.7 and 21.1 for C IV, 1.3 and 28.4 for C V, and 1.6 and 18.7 for adult females, all expressed as nmol mg^-1 dry wt h^-1. In all experiments, excretion rates of inorganic phosphate varied between 0.7 and 1.5 (C III), 0.5 and 1.1 (C IV), 0.2 and 0.8 (C V), and 0.3 and 1.0 (adult females) nmol mg^-1 dry wt h^-1. Ratios of O:N, O:P, and N:P indicated that much of the metabolic energy was derived from catabolism of proteins. Comparison of the turnover rate of carbon and nitrogen showed, however, that nitrogen turnover was between 2.6 and 8.9 times higher than that of carbon. This may indicate that the copepods deaminate ingested protein, with the carbon skeleton of the amino acids subsequently being used in the synthesis of lipid compounds, possibly wax esters.     

Molting rates and cohort development of Calanus finmarchicus and C. glacialis in the sea off southwest Nova Scotia

Species of Calanus were sampled at stations off southwest Nova Scotia during an 8-d cruise in June, 1982. A simple model using shipboard measurement of molting rates of predominant copepodite stages accurately predicted the changes in cohort structure that occurred at stations along a transect at the beginning and end of the cruise. The combined durations of C. finmarchicus (Gunnerus) stages CIII and CIV, calculated from the inverses of molting rates corrected for presence of diapausing C. glacialis Jaschnov, corresponded well with development times based on laboratory rearing experiments. We conclude that our molting rate measurements accurately reflected processes in nature. This methodology is useful for analyzing the dynamics and production of copepod populations.     

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.     

Geographical range and taxonomic divergence in North Atlantic Calanus (C. helgolandicus,C. finmarchicus and C. glacialis)

The known distribution of Calanus helgolandicus in the North Atlantic Drift is difficult to explain in the absence of a reproductively active population inhabiting continental waters off eastern North America. New evidece indicates that this population, overlooked in the past, does in fact exist. The species has been found in a study of zooplankton samples from a number of MARMAP (Marine Resource Monitoring, Assessment, and Prediction Program) cruises surveying ichthyoplankton between Cape Hatteras and the New York Bight. Sexual activity in these stocks of C. helgolandicus was indicated by the ripeness of ovaries, the frequency of males, the presence of sperm in the females' seminal receptacles and the appearance of a female bearing a Calanus spermatophore. The new records provide a likely origin for the presence of the species in the vicinity of the Labrador Grand Banks as well as further east in the North Atlantic Drift. C. finmarchicus, similarly sexually active, was found to be sympatric with C. helgolandicus in the MARMAP collections which were taken from a region contiguous with the southernmost, known distribution, of C. glacialis. The distribution of integumental organs (i.e., pore signature patterns) was examined in the three species to determine whether they would be taxonomically useful. Strikingly different patterns were found on the female urosome. Pore signature differences between the polar species C. glacialis and the temperate C. helgolandicus proved to be as pronounced as those between the boreal C. finmarchicus and its two neighboring species. The successively overlapping ranges and the distinctive differences in pore signature patterns suggest that divergence from the generic pattern of integumental organ distribution has been a product of selection against hybridizing among the three species. If this is in fact the case, the reproductive range of C. helgolandicus has overlapped with those of C. finmarchicus and C. glacialis for appreciable periods in the history of the three species.     

Annual population development and production by Calanus finmarchicus, C. glacialis and C. hyperboreus in Disko Bay, western Greenland

The populations of the copepod species Calanus finmarchicus, C. glacialis and C. hyperboreus were investigated in Disko Bay during a 14-month period in 1996-1997. The three species were predominant in the copepod community. The biomass reached a maximum at the beginning of June (127 mg C m^-3). From the end of July until the end of April the following year, the biomass was <1-6 mg C m^-3. All three species showed seasonal ontogenetic migration. The spring ascent for all three species was just prior to or in association with the break-up of sea ice and the development of the spring bloom, whereas descent occurred over a larger time span during summer. The main overwintering stages were CV for C. finmarchicus, CIV and CV for C. glacialis and C. hyperboreus. Peak abundance of juvenile copepodites, representing the new generation, was in August for C. finmarchicus, in July for C. glacialis and in May/June for C. hyperboreus. From the timing of reproduction and the population development, the life cycles were deduced to be 1 year for C. finmarchicus and at least 2 years for C. glacialis and C. hyperboreus. Secondary production and potential grazing impact of the Calanus community were estimated by two methods based on specific egg-production rates and temperature-dependent production. The Calanus community was not able to control the primary producers during the spring bloom but probably did during post-bloom. The estimates also indicated that grazing on ciliates and heterotrophic dinoflagellates contributes as an essential food source in the post-bloom period.     

Calanus marshallae,a new species of calanoid copepod closely allied to the sibling species C. finmarchicus and C. glacialis

Species of the copepod genus Calanus frequently dominate the marine zooplankton in boreal and arctic waters. Up to now there have been no operational means of identifying several species closely related to Calanus finmarchicus. Reanalysis of these taxa, using material from plankton samples collected throughout Northern Hemisphere polar and coreal waters, shows that there are 3 sibling species which have been previously combined under the names C. finmarchicus and C. glacialis. Several new taxonomic characters permit unequivocal identification of C. finmarchicus, C. glacialis, and a new species, C. marshallae. Claims that C. finmarchicus and C. glacialis are subspecies are refuted; there is no evidence that the two species continuously intergrade either where they co-occur or where they are allopatric, nor is there evidence that the two species hybridize. C. finmarchicus is basically restricted to the North Atlantic Ocean and C. marshallae to the North Pacific Ocean and Bering Sea; C. glacialis is primarily an Arctic species, but its geographical distribution slightly overlaps those of the other two species. Taxa closely related to C. finmarchicus and C. helgolandicus probably represent two separate, but closely linked evolutionary lineage; species of these two lineages are placed in one of two species groups, the finmarchicus group and the helgolandicus group.     

Vertical distribution of Calanus finmarchicus and C. helgolandicus in relation to the development of the seasonal thermocline in the Celtic Sea

The geographical distribution and annual mean abundance of Calanus finmarchicus (Gunnerus) and C. helgolandicus (Claus) in the northern North Atlantic Ocean were shown in relation to the seasonal and annual fluctuations of abundance of the species in the Celtic Sea from 1960 to 1981. These congeneric copepods, although showing allopatric distributions over most of their geographical range, have sympatric distributions in the Celtic Sea where they dominate the dry weight biomass of the plankton throughout the year. The two species respond differently to the development of the seasonal thermocline and halocline by taking up different vertical distributions in the water column. C. finmarchicus occurred in the colder, more saline water below the thermocline, while C. helgolandicus occurred in the warmer, less saline water above the thermocline. This behaviour is postulated as a mechanism by which these morphologically similar copepods more fully exploit the resources of their temporally and spatially heterogeneous environment and also minimise interspecific competition. The species have the same foraging techniques and are able to exploit the same size spectrum of particulates. The vertical depth strata in which the populations are found for most of the year in the Celtic Sea means that both species exploit the diatom bloom in early spring but, thereafter, C. helgolandicus grazes on the daily production of the autotrophs in the euphotic zone while C. finmarchicus, below the thermocline, has to rely for its food on sedimenting particulates (whole cells, detritus and faecal material). The isolating mechanisms whereby these two populations partition the habitat in the Celtic Sea are discussed.     

Grazing of phytoplankton by copepods in eastern Antarctic coastal waters

Chlorophyll a, primary productivity and grazing by copepods on phytoplankton were measured in the upper water column during the summer of 1994/1995 at a coastal site near Davis Station, East Antarctica. Chlorophyll a was at a maximum in mid-December, then dropped markedly as the coastal fast ice melted and broke‐out. Phytoplankton biomass increased again from mid‐ to late‐February. Copepods accounted for at least 65% of zooplankton biomass in the water column before sea ice break‐out, whereas larval polychaetes and ctenophores dominated after ice break‐out. Oncaeacurvata was the numerically dominant species throughout the study. The highest grazing rate (8.7 mg C␣m⁻³␣d⁻¹) was recorded on 21 December when O.␣curvata accounted for 64% of the total. Grazing had decreased markedly by 28 December (0.9 mg C m⁻³ d⁻¹); again O. curvata accounted for over 50% of the total ingested. Copepod grazing increased after ice break-out until the last experiment on 20 February (⋍5 mg C␣m⁻³␣d⁻¹). The main species responsible for grazing during this period were O. curvata, Oithonasimilis, Calanoidesacutus and unidentified copepod nauplii. It was estimated that copepods removed between 1 and 5% of primary productivity. Received: 11 October 1996 / Accepted: 22 October 1996     

Vertical distributions of Calanus finmarchicus and C. helgolandicus (Crustacea: Copepoda)

The vertical distributions of the spring populations of Calanus finmarchicus (Gunnerus) and C. helgolandicus Claus are described and compared. The differences we observed between the two species have probably confused the understanding of the vertical distribution and development of the populations of Calanus spp. in the shelf seas around the United Kingdom where the species occur together. The results imply that these two congeneric species have different behaviour patterns which minimise interspecific competition where the species have sympatric distributions. C. finmarchicus has its younger development stages overlying the older stages in the water column. In C. helgolandicus the converse is true; i. e., the majority of the populations of Stage I and II copepodites of the first spring generations are found below the thermocline. It is also suggested that the different behaviour patterns lead to different feeding regimes and strategies.JONSDAP Contribution No. 52     

Growth and development rates of Calanus finmarchicus nauplii during a diatom spring bloom

Growth and development rates were determined for nauplii of Calanus finmarchicus (Gunnerus) in the near-shore waters of a western Norwegian fjord from in situ mesocosm incubations. The major food source for the nauplii was diatoms, but Phaeocystis sp., dinoflagellates and ciliates were also part of the diet. At local temperatures ranging from 4.8 to 5.2 °C the cumulative median development time from hatching to Nauplius VI was 19 d. The time taken to molt to the next naupliar stage was approximately constant (3 d) from Stages IV to VI, but Stage III needed the longest development time (5 d). The instantaneous growth rate in terms of body carbon was negative from hatching to Nauplius Stage II, but as high as 0.25 to 0.30 d⁻¹ from Stage III to V. Enhancement of food resources by nutrient addition led to no significant change in specific growth rates. Additionally, the cohorts from different nutrient regimes showed almost equal development time, size and body carbon within stages. Length–weight relationships of nauplii from the two different food resources were: W _{low resources} = 4.17 × 10⁻⁶ × L ^2.03 (r ² = 0.84) and W _{high resources} = 4.29 × 10⁻⁶ × L ^2.05 (r ² = 0.92), where weight (W) is in micrograms of C and body length (L) in micrometers. The natural body morphology of naupliar stages I to VI is illustrated with digital images, including the final molt from Nauplius VI to Copepodid Stage I. In general, development of the nauplii was faster than that of the copepodids of C. finmarchicus, and structural growth was exponential from naupliar stages III to VI. This study validates our earlier results that nauplii of C. finmarchicus can obtain high growth and nearly maximal developmental rates at relatively low food levels (∼50 μg C l⁻¹), suggesting that nauplii exhibit far less dependence on food supply than copepodids. Received: 30 July 1999 / Accepted: 7 March 2000     

Seasonal variations in the densities of fecal pellets produced by Oikopleura vanhoeffeni (C. Larvacea) and Calanus finmarchicus (C. Copepoda)

Two abundant macrozooplankters, Oikopleura vanhoeffeni (Lohmann) and Calanus finmarchicus (Gunnerus) were collected from the coastal waters off Newfoundland in different seasons during 1990–1991 and incubated in natural seawater to collect freshly egested, field produced, fecal pellets. The densities of fecal pellets from O. vanhoeffeni and C. finmarchicus were measured in an isosmotic density gradient. These are the first reported seasonal measurements of fecal pellet densities from two different types of macrozooplankters, O. vanhoeffeni, a larvacean, filter feeder and C. finmarchicus, a crustacean, suspension feeder. Pellet density ranges and medians were significantly different among seasons for both species, depending primarily on the type of phytoplankton ingested and its ability to be compacted. Winter O. vanhoeffeni and fall C. finmarchicus feces filled with nanoplankters and soft bodied organisms had less open space [packing index (% open area) = 3.5 and 4% for O. vanhoeffeni and C. finmarchicus, respectively] and were more dense (1.23 and 1.19 g cm^-3) than spring feces filled with diatoms (packing index = 15 and 23%, density = 1.13 and 1.11 gcm^-3). For copepods, these results contrast with previously published density values and with the predicted copepod fecal pellet density calculated, in the present study, using the conventional mass/volume relationship. Copepod spring and summer diatom-filled feces had a calculated density of 1.12 and 1.24 gcm^-3 vs a measured median density of 1.11 gcm^-3 for both spring and summer feces; the fall feces containing nanoplankters had a calculated density of 1.08 gcm^-3 vs a measured median density of 1.19 gcm^-3. Knowledge of the seasonal variations in fecal pellet densities is important for the development of flux models.     

Grazing,defecation and excretion rates of copepods from inter-island channels of the Canadian Arctic archipelago

Zooplankton (>800 m) and water samples were collected at night at eleven stations in Parry Channel and adjacent waters, between 28 August and 14 September 1986. Chlorophyll concentrations varied between 17 g l^-1 at the surface at one station in Wellington Channel, Canada, and 1.5 g l^-1 throughout the top 30 m at one station in Byam Martin Channel, Canada. In tows from 0 to 50 m the zooplankton community at all stations was dominated by varying proportions of three species,Calanus hyperboreus, C. glacialis andMetridia longa. Levels of chlorophyllderived pigments inC. hyperboreus Stages V, IV and III were correlated with the concentration of chlorophyll at the chlorophyll maximum. Defecation rate constants, measured for the different stages where present, were variable but not correlated with ambient chlorophyll concentrations. Ammonia excretion was measured simultaneously for the communities in which defecation was being measured in the invividual species and stages. In these experiments the rate of ammonia accumulation decreased significantly over the period during which the copepods were actively defecating (usually the first 3 h) and then tended to a constant level (over the next 18 to 20 h). The time courses of ammonia accumulation could be described by a model comprised of the sum of a straight line and a saturating curve. For seven experiments the ammonia release given by the asymptote of the saturating component was correlated with the estimate of community defecation, obtained by summing the individual defecations, suggesting that the two processes were closely associated. Weight specific defecation and weight specific defecation-associated ammonia excretion were both correlated with ambient chlorophyll concentration. The ratio of initial to basal ammonia excretion rate varied between 2 and 20, so that defecation-associated ammonia release may be important in the estimation of in vivo nitrogen excretion or regeneration rates.     

Egg production of the Arctic copepod Calanus glacialis: laboratory experiments

Egg production of single female Calanus glacialis Jaschnov fed with Thalassiosira antarctica at super-abundant concentrations (>300 g C l^-1) was determined over several weeks. Experiments were performed directly after collection from the East Greeland Current in June 1987 and 1988, and during resumed feeding after long-term starvation over 4 (October 1988), 4.5 (October 1987) and 6.5 (January 1988) mo. In addition, in June 1987, short-term starvation experiments of 5 and 15 d were conducted. Egg production was closely related to feeding in all experiments. While directly after collection eggs were produced within a few days, it took 2 wk (October 1987 and 1988) and 10 d (January 1988), respectively, to resume egg production after long-term starvation. During long-term starvation periods eggs were not laid. The decrease in total egg production with duration of starvation was due to decreasing clutch size and increasing spawning interval. In contrast, short-term starvation experiments only affected spawning interval. Interannual variability in egg production was high, with much higher clutch sizes in 1988. Average production rates in June 1988 correponded to 5% body C female^-1 d^-1, the maximum was 7.4% (1 274 eggs in 23 d). Carbon content of eggs was 0.40 g egg^-1. C. glacialis is well adapted to pulsed food events in the Arctic by its longevity; its ability to preserve its reproductive potential over several months; its rapid mobilization of ovaries; and by its high egg production rates. The implication of prolonged spawning capacity on life cycle studies is discussed.     

Egg production of Calanus finmarchicus : effect of temperature, food and season

The use of the egg production rate of herbivorous copepods as an important parameter for understanding population dynamics and as an index of secondary production requires knowledge of the regulatory mechanisms involved and of the response to changes in food concentrations and temperature. Furthermore, the effects of season and generation on egg production have to be studied. In this context data are presented for Calanus finmarchicus from the northern North Atlantic. Prefed and prestarved females were exposed to different concentrations of the diatom Thalassiosira antarctica over 1 to 2 wk at 0 or 5 °C, and egg deposition was controlled daily. Egg production increased with higher food concentrations, but much less when prestarved. The effect of temperatures between −1.5 and 8 °C on egg production was studied in females maintained at optimum feeding conditions. Egg production rate increased exponentially over the whole temperature range by a factor of 5.2, from 14.2 to 73.4 eggs female⁻¹ d⁻¹, and carbon-specific egg production by 4, from 2.1 to 8.5% body C d⁻¹. The response to starvation was also temperature dependent. In both the temperature and feeding experiments egg production rate was regulated mainly by changes of the spawning interval, while changes of clutch size were independent of experimental conditions. Different responses to optimum feeding conditions were observed in females collected in monthly intervals on three occasions between March and May. The March females deposited more clutches than the April and May females. In May, >50% of the females did not spawn at all. Maximum egg production rates were never >25% of the rate expected at 5 °C, indicating endogenous control of egg production in addition to food and temperature effects. Received: 4 August 1996 / Accepted: 11 September 1996     

Diel variations in the nutritional physiology of Calanus glacialis from Lat. 78°N in the summer

Calanus glacialis (Jaschnov) from Isfjord, western Spitzbergen, showed no tendency to pronounced diurnal vertical migration or cyclic variations in feeding activity. The samples, taken regularly over a 24-h period and at three different depths in the beginning of August, did however show considerable individual variations in feeding activity. These variations, reflected in the results of gut analyses, were not related to variations in the concentrations of particulate protein, carbohydrate or algal pigments in the water. Analyses of the activities of two digestive enzymes and the electron-transport system (ETS) indicated that those few individuals always present below the photic zone were there temporarily, and showed no signs of slowing down their metabolism. The results suggest that the population of C. glacialis, during the Arctic summer, feeds continuously in the upper water layer, but that intermittent, non-synchronous individual feeding rhythms are the rule.     

Changes in lipids during the development of Calanus finmarchicus s.l. from Copepodid I to adult

The total lipid and wax ester content as well as the fatty acid and alcohol composition of all copepodid stages and adults of Calanus finmarchicus s.l. were investigated at different locations in the North Sea in 1983 and 1984. Total lipid and the wax ester proportion increased exponentially until Copepodid V. The females were sometimes lower in lipids than the Stage V. The wax ester proportion reached about 90% of total lipids in males and Copepodid V and up to 40% in Copepodid I. The major fatty acids were 16:0, 20:5, and 22:6 and the major fatty alcohols were 16:0, 20:1 and 22:1. At one station the 18:4 acid became one of the dominant acids, because of a Phaeocystis sp. bloom, indicating that the fatty acids of the diet are incorporated mostly unchanged into the lipids of the copepods. The other main fatty acids 20:1 and 22:1 are probably synthesized de novo, serving as precursors for the principal alcohols 20:1 and 22:1. Their levels decreased in the younger stages due to increases in 16:0 alcohol. The fatty alcohol-forming enzyme seems to be specific for saturated and monounsaturated acids, which may be synthesized de novo or derived from diet.     

Microphagous ciliates in mesohaline Chesapeake Bay waters: Estimates of growth rates and consumption by copepods

Growth rates of microphagous ciliates (forms which feed primarily on picoplankton-sized prey) were estimated, along with rates of their consumption by copepods, in shipboard experiments conducted in the mesohaline portion of Chesapeake Bay, USA, under contrasting water column conditions in April, June, and August 1987. Estimates were based on temporal changes in cell densities in size-fractionated water samples incubated under in situ conditions. In April, at low temperatures (7 to 10°C) and with oxygen present throughout the water column, similar generation times of ca. 1 to 1.5 d were estimated for surface and deep water (24 m) ciliate populations. In June, water was anoxic below 12 m and a distinct anoxic microphage community grew at about twice the rate of the surface community with generation times of ca. 7 and 14 h, respectively. In August, bottom water was again anoxic, but the sameStrobilidium sp. dominated both surface and deep waters with low or no growth apparent in anoxic waters and a generation time of ca. 8 h in surface waters. Copepod (primarilyAcartia tonsa Dana nauplii) clearance rates for microphagous ciliates in surface waters were 0.11, 0.56, and 0.53 ml h^–1 copepod^–1 for April, June and August, respectively. Calculation of removal rates, based on average densities, indicated that from 34 to 200% of surface waters were cleared d^–1 of microphagous ciliates by copepods.     

Grazing in juvenile stages of some estuarine calanoid copepods

The grazing of juvenile Eurytemora affinis, Acartia tonsa and A. clausi from the Chesapeake Bay (USA) was investigated using natural particle distributions and freshly caught copepods, live-sorted into stages. Data were analyzed in 110 size channels using an electronic particle counter, and filtering rates (FR) were estimated based on total particle removal (mean FR), and that for each size channel (giving maximum FR). Mean and maximum filtering rates increased from NVI (Nauplius Stage VI) through CVI (Copepodid Stage VI). Both rates plotted against weight satisfied a log fit best for A. tonsa, and a linear fit best for E. affinis. Results for A. tonsa were quite variable, apparently due to differences in temperature between experiments. Particle selection was investigated from the shape of the filtering rate curve over particle size. We define selective feeding by a FR curve which is higher in some size categories, and non-selective feeding by a flat FR curve. The general pattern was one of selective feeding in all copepodid stages of the three calanoid copepods investigated. E. affinis tended to track biomass peaks while Acartia spp.'s feeding was more variable, including feeding in size ranges of greatest particle concentration, on larger particles, and in other size categories as well. Experiments with nauplii tended to yield flat FR curves, and it may be that selective grazing appears with, or is greatly accentuated by, metamorphosis from NVI to CI (Copepodid Stage I).University of Maryland, Center for Environmental and Estuarine Studies Contribution No. 762.     

Life cycle, seasonal vertical distribution and feeding of Calanus finmarchicus in Skagerrak coastal water

 Day/night zooplankton sampling in Kosterfjorden, Sweden, gave information on population succession, vertical distribution and feeding of Calanus finmarchicus over 17 months. Copepodid Stage 1 and 2 (C-I, C-II) were present from December to August, indicating reproduction during most of the year. Mating and breeding for the overwintering generation mainly took place in February/March, resulting in peak abundance of C-I in March/April. Secondary breeding periods were in April and July/August, but the resulting recruitment from these were low. The relative recruitment success in the first spring was around 13 times higher than during the second spring. Low temperature in the surface water early in the year and depression of the spring phytoplankton bloom may have caused the failure in the second year. Population biomass peaked at >15 g dry weight m⁻² during the numeric peak of the youngest stages in March/April. A more sustainable level of high biomass of 8 to 10 g dw m⁻² was gradually built up during summer, mainly due to a continuous accumulation of C-Vs. Adults and C-Vs comprised the overwintering population, with 7 to 14% and 85 to 93%, respectively, for the 2 years, but only C-Vs staying in the deep water were in a resting state. Adult males showed a strong diurnal vertical migration (DVM) of the usual type from spring to early autumn and a reversed DVM during the cold season. They were in a feeding state throughout the year. Adult females showed the usual type of DVM during summer to autumn, but commonly a reversed DVM during winter to spring. They were usually in a feeding state, with no pronounced differences between surface and deep water or between day and night. C-Vs aggregated in the deep water from October to March and performed DVM in April to June. They were commonly feeding in the deep water between March and June but showed no or very low feeding activity there from July to February. C-Vs in the surface water were commonly feeding and showed the highest proportion of feeding in autumn, when the population in the deep water was inactive. C-V constituted up to nearly 100% of the population biomass, and therefore must be of profound ecological importance. Defined by this dominant role, the population of C. finmarchicus can be characterised as having an active period of feeding, reproduction and development from February to July with a following 6 to 7 months of resting in the deep water, when development is arrested and no feeding occurs. Received: 1 October 1999 / Accepted: 27 April 2000     

Grazing by adult estuarine calanoid copepods of the Chesapeake Bay

Grazing by adult female Eurytemora affinis, Acartia tonsa and A. clausi on natural distributions of particles from the Chesapeake Bay has been investigated. During the course of a year's sampling, a wide variety of particle size-biomass distributions were observed as seasonal shifts in detritus, and over 150 algal species occurred. These distributions were grouped into 5 basic types in the analyses of feeding. All three species demonstrated similar capabilities for feeding over a broad range of particle size with selection (higher filtering rates) on larger particles and on biomass peaks. Feeding on multiple-peak distributions resulted in strong selection or tracking of each biomass peak with reduced filtering rates between peaks. Evidence is presented which suggests that the copepods first feed on large particles and then successively switch to biomass peaks of the smaller size categories. Comparisons of the feeding behavior of Eurytemora affinis and the Acartia species showing that the Acartia species have greater capabilities for taking large particles may be associated with modifications of their mouth parts for raptorial feeding. The results suggest considerable flexibility in copepod feeding behavior which cannot be explained solely by the mechanism of a fixed sieve.