Switching between herbivory and carnivory by the planktonic marine copepod Calanus pacificus

Adult females of the omnivorous copepod Calanus pacificus, collected from the plankton off La Jolla, California, USA (June, 1978), fed disproportionately on the prey in greatest relative abundance when given mixtures of diatoms (Thalassiosira fluviatilis) and copepod (C. pacificus) nauplii as food. This switch from herbivorous to carnivorous behavior may be significant in nature during the decline of phytoplankton blooms. More generally, the widespread omnivorous habit among pelagic animals suggests a responsive and flexible trophic organization which contributes to the resiliency of planktonic communities in a dynamic physical environment.Contribution No. 1223 from the Department of Oceanography, University of Washington, Seattle, Washington 98195, USA     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in November 1988 and March 1989, when the water temperature was weakly stratified in a reversed manner. In both investigations a pronounced ontogenetic difference in vertical distribution was found. Spawning always occurred during nighttime, being confined to the upper 40 m water column in November but to the layer below 35 m in March. The distribution of pre-feeding nauplius stages, NI and NII, was more or less similar to that of the eggs. The first-feeding NIII performed a marked upward migration, and late nauplius stages (NIV to NVI) and early copepodite stages (CI and CII) continuously aggregated in the upper water column where phytoplankton was abundant. CIII to CVI (adult female and male) tended to disperse in the whole water column. In November, however, they avoided the upper 10 m strate during daytime and some individuals migrated upward to the surface during nighttime. In March, CV and CVI aggregated in the layer between 5 and 15 m deep in the daytime and migrated both upward and downward at dusk, resulting in homogeneous distributions during the nighttime.     

Ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus in the Inland Sea of Japan

The ontogenetic diel vertical migration of the planktonic copepod Calanus sinicus was investigated in the Inland Sea of Japan in June 1989, when the water column was thermally weakly stratified. Because of fewer eggs and less variation in their abundance, nocturnal spawning was not apparent. A pronounced upward migration occurred in NIII. NIII to CIII resided in the upper 20 m layer throughout the day, and from CIV on their median depths descended. CV and adult females underwent significant diel vertical migration, whereas adult males did not migrate. By integrating the results from the present study and those from our previous investigations (in August–September 1988, November 1988 and March 1989), we review seasonal variation in the ontogenetic diel vertical migration of C. sinicus. Spawning was largely nocturnal, reaching its maximum level around dawn, but spawning depth and fecundity changed seasonally. The distribution of pre-feeding stages, NI and NII, was similar to that of eggs. A pronounced upward migration always occurred in the first feeding stage, NIII, and late nauplii and early copepodites always resided in the food-rich upper layer, indicating that upward migration by NIII is feeding migration. As the stages progressed, they extended their vertical distribution range, and CV and adult females usually underwent diel vertical migration. However, the pattern and strength of this migration differed seasonally. Their day depths increased with the increase of relative biomass of planktivorous fish, indicating that predator avoidance induces their diurnal downward migration. High chlorophyll a concentrations in the upper layer (<15 m deep) relative to the lower layer (>20 m deep) amplified their diel vertical migrations. Diel vertical migration of C. sinicus is a phenotypic behavior.     

Grazing and ingestion rates of nauplii,copepodids and adults of the marine planktonic copepod Calanus helgolandicus

The average grazing and ingestion rates of all stages of the marine planktonic copepod Calanus helgolandicus (Calanoida) from nauplius stage IV to adults were measured experimentally at 15°C in agitated cultures. The chain-forming diatom Lauderia borealis and the unarmoured dinoflagellate Gymnodinium splendens were offered as food. The food concentrations were close to natural conditions and ranged from 36 to 101 g of organic carbon per liter. The medium body weights expressed in g of organic carbon of almost all larval stages raised at 49 g C/1 were identical with the weight of the same stages caught in the Pacific Ocean off La Jolla, California, USA. In a log-log system, grazing and ingestion rates increased almost linearly with increasing body weight. Grazing rates ranged from 4 to 21 ml/day/nauplius stage IV to 286 ml to 773 ml/day/female. Ingestion rates increased from 0.2 g to 0.8 g C/day/nauplius stage IV to 18 g to 69 g C/day/female. Grazing and ingestion rates per unit body weight decreased gradually with increasing body weight. The daily ingested amount of food decreased from 292 to 481% of the body weight (g C) of nauplius stage V to 28–85% of the body weight of adult females. Grazing and ingestion performances of all stages increased with increasing particle size. Grazing rates decreased and ingestion rates increased with increasing food concentrations. The published data on food intake of the different age groups of C. helgolandicus show that the young stages of herbivorous planktonic copepods can play a major part in the consumption of phytoplankton in the sea due to their high grazing and ingestion rates.     

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     

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.     

Potential impact of a toxic dinoflagellate (Alexandrium excavatum) bloom on survival of fish and crustacean larvae

During the last 20 yr the western half of the Dutch Wadden Sea has undergone significant eutrophication: concentrations of P and N compounds and planktonic algae have roughly doubled, as has primary production. Though oxygen levels are often low in summer, anoxic areas are small and rare due to strong tidal mixing. During the 1970 to 1990 period, macrozoobenthos was sampled annually at 15 stations at Balgzand, a 50-km² tidal-flat area in the westernmost part of the Wadden Sea. Not only did the estimates of total numbers, biomass, and production double during these two decades, but significant changes in the composition of the benthic community were observed, too: (1) the numerical proportion of polychaetes increased at the expense of molluscs and crustaceans, (2) the overall mean weight per individual of the macrozoobenthos decreased (numbers of individuals of small-sized species increased more rapidly than those of large-sized species), and (3) though absolute numbers and biomass of all feeding types increased, the share of carnivores declined and that of deposit feeders increased; the proportion of suspension feeders showed little change. This study refers to true macrobenthos only (1-mm sieve) and further excludes two taxa (Corophium spp. andHydrobia ulvae) which occasionally exercised an undue influence on numbers. Mass mortalities caused by low oxygen concentrations were of a small-scale nature only. Total number of species fluctuated without a clear trend. As a consequence of the increasing numerical densities, trends in species numbers were slightly increasing when expressed per unit of area and slightly decreasing when estimated per 100 individuals (by rarefaction).     

Importance of food quantity to structural growth rate and neutral lipid reserves accumulated in Calanus finmarchicus

 Growth and developmental rates were determined for copepodids of Calanus finmarchicus (Gunnerus) from experimental seawater mesocosms in a western Norwegian fjord. The instantaneous growth rates (g) from copepodid stage I (CI) to adult ranged from 0.08 to 0.10 d⁻¹. Daily per capita mortality rate of the cohorts was as low as 0.012 d⁻¹ (1.2% d⁻¹). At local increasing temperatures (5.1 to 8.3 °C), development was equiproportional, and the cumulative median development time from egg to CV was approximately 65 d. CV moulted to males and females, and egg production was initiated. Enhancement of food resources by nutrient addition caused a 23.4% increase in growth rates from CI to adult. Additionally, copepodid stages showed a generally larger body size, carbon and nitrogen content and total storage lipid content (wax esters + triacylglycerols) in response to enhanced resources. Our data support an elsewhere proposed exponential-growth hypothesis; growth of the structural compartments and store lipids (mostly wax esters) was exponential during the copepodid stages. However, a sigmoidal pattern of growth best described growth of adult stages if reared at high resources, and depot lipid accumulation in late CVs and adults at high resources. Body nitrogen growth increased exponentially, however, no significant changes in nitrogen specific growth rates were found between individuals from low and high resources. CV and adults seem to have reached near-maximal weights at high resources, whereas structural weight continued to increase at low resources. Despite the differences in structural growth dynamics, cohort development was similar until the end of CV. During the onset of sexual differentiation, the male:female ratio and the adult:CV ratio were highest at high food resources, suggesting that the time used for the final moult depends on the feeding history of the copepods in relation to food quality and quantity. It appears that relatively small changes in food availability strongly influence the biochemical composition of C. finmarchicus copepodids. Received: 28 May 1999 / Accepted: 10 February 2000     

Cyclical contributions of the digestive epithelium to faecal pellet formation by the copepod Calanus helgolandicus

Faecal pellet formation within the gut of Stage V and adult females of the copepod Calanus helgolandicus Claus involves (1) cyclical processes of digestion and (2) the contribution of parts of the gut epithelium to the pellets. During an experimental regime in which dim lighting was restricted to day-time and feeding to night-time (17.00 to 09.00 hrs), the copepods responded with cyclical changes in both the quantity of pellets they produced and the fine structure of the contents. During the feeding period, the contents showed changes in the relative amounts of materials originating from disintegrated cells of the digestive epithelium and those derived directly from the ingested food. The vacuolar B-cells of the gut contribute to the content of the pellets and the distal, necrotic N-cells appear to be involved in forming the peritrophic membrane which encloses each pellet. Cells of the gut epithelium which are broken down during feeding are all replaced during the non-feeding period. Other individuals were taken directly from the sea and in these, also, the cells of the gut broke down during feeding and contributed to the faecal pellets. The supply of epithelial cells may limit the duration of the feeding period.     

Sensitivity of marine fishes to toxins from the red-tide dinoflagellate Gonyaulax excavata and implications for fish kills

Marine fishes (Atlantic herring, American pollock, winter flounder, Atlantic salmon, and cod) were dosed orally and intraperitoneally (i.p.) with paralytic shellfish toxins extracted from Bay of Fundy Gonyaulax excavata (tamarensis) cells. The toxins are lethal to these fishes in low oral doses, and in extremely low i.p. doses. Symptoms are the same among these fishes, both for oral and i.p. administrations, including loss of equilibrium within 5 to 15 min, followed by immobilization and shallow, arrhythmic breathing. Death generally occurs within 20 to 60 min of toxin administration. Dose responses are also similar among these fishes. Oral LD50 values are 400 to 750 g saxitoxin (STX) equivalent kg^-1 body weight. Intraperitoneal LD50 values are 4 to 12 g STX equivalent kg^-1. Toxins are undetectable in fish muscle tissue following lethal oral doses. The similarity of symptoms and dose responses suggest that fish as a group are sensitive to G. excavata toxins. Results, in combination with reports implicating these toxins in herring, sand lance, and menhaden kills, show the plausibility that the nearly worldwide blooms and red tides of G. excavata and its relatives may cause kills of a variety of fishes.     

Population structure of the planktonic copepod <Emphasis Type="Italic">Calanus pacificus</Emphasis> in the North Pacific Ocean

The pelagic copepod Calanus pacificus ranges nearly continuously across temperate-boreal regions of the North Pacific Ocean and is currently divided into three subspecies—C. pacificus oceanicus, C. pacificus californicus, C. pacificus pacificus—based on subtle morphological differences and geographic location. The relation between geography and genetic differentiation was examined for 398 C. pacificus individuals sampled from six widely distributed locations across the North Pacific, including an open ocean site and coastal sites on both sides of the North Pacific basin. For each individual copepod, the DNA sequence was determined for a 421-bp region of the mitochondrial coxI gene (mtCOI). A total of sixty-three different mtCOI sequences, or haplotypes, were detected, with a sequence divergence between haplotypes of 0.2–3.1%. The number and distribution of haplotypes varied with sampling location; 12 haplotypes were distributed across multiple sampling locations, and 51 occurred at only one location. Five genetically distinct populations were detected based on F _ST values. Haplotype minimum spanning networks, nucleotide divergence and F _ST values indicated that individuals from coastal sites in the North Pacific Ocean were more closely related to each other than to individuals from the open ocean site at Station P. These results provide genetic support for the designation of two subspecies—a coastal subspecies that consists of what is currently referred to as C. p. pacificus and C. p. californicus and an open ocean subspecies C. p. oceanicus. This work also indicates that planktonic copepods with potentially high dispersal capacity can develop genetically structured populations in the absence of obvious geographic barriers between proximate locales within an ocean basin.     

Diet and respiration of the small planktonic marine copepod Oithona nana

The functional responses of Oithona nana (Giesbr.) to various phytoplankton and zooplankton food species are described. The food species were divided into three size categories, the seasonal abundances of which were measured in Loch Turnaig, a Scottish sea loch in 1977. The seasonal variations in feeding rates in the sea for each size class were derived. The seasonal variation in respiration rate of O. nana was measured, and metabolic requirements were claculated as between 6 and 40% of the food material estimated as being eaten. O. nana differs from other common copepods in having a wide food-particle size spectrum and a low metabolic rate. It is suggested that these adaptations constitute the strategy whereby O. nana maintains its population levels throughout the year.     

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.     

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.     

Large-scale oceanic distribution and population structure of Calanus finmarchicus, in relation to physical environment, food and predators

The investigation was carried out from 62°N to 73°N and from 14°E to 11°W in the Norwegian Sea during 19 June-12 July 1997. Regional differences in the phase of the seasonal development of the plankton community were evident, most pronounced across the Arctic front. In the Coastal and eastern Atlantic domains, post-bloom conditions prevailed, characterised by low chlorophyll a (chl a) levels and a phytoplankton assemblage dominated by coccolithophorids and small flagellates. During the study period, egg production rates of Calanus finmarchicus were low (<10 eggs female^-1 day^-1), older copepodite stages dominated, and the seasonal descent to deeper waters had started. In the Arctic domain, bloom conditions were evident by high chl a levels and a high abundance of large diatoms. Egg production rates were higher (a maximum of 29 eggs female^-1 day^-1), but the dominance of stages CI-CIII indicated that considerable spawning had already occurred prior to the spring bloom. The seasonal descent had barely started. Both invertebrate and fish predators were most abundant in the Coastal and eastern Atlantic domains, with abundance strongly decreasing north-westwards. No tight relationship between total abundance of invertebrate or fish predators and that of C. finmarchicus was apparent. However, a weak, but significant, relationship between abundance of young stages of chaetognaths and Euchaeta spp. versus young stages of C. finmarchicus was found, indicating that these invertebrate predators develop parallel to the development of the new cohort of C. finmarchicus. In early summer, C. finmarchicus had reached overwintering stages, and had started to accumulate in deeper waters in areas with the highest abundance of horizontally migratory planktivorous fish.     

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 g 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 g 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 V's. The results are discussed both in relation to the physiological state of the species and to the environmental conditions.     

Food ingestion by the marine planktonic copepod Paracalanus in relation to abundance and size distribution of food

Juveniles and adult females were presented a food spectrum of three algae of different sizes (4.5, 12 and 20 m cell width). The increase in rate of ingestion of the medium-sized alga with an increase in copepod size was significantly greater than the increase in rate of ingestion of the small alga. It is hypothesized that the perception of chemical signals from the small alga by a copepod decreases as the copepod moults from stage to stage. The rate of ingestion of the large alga by copepod stage V (CV) and adult females was lower than the rate of ingestion of the medium-sized alga at mid- and high phytoplankton concentrations. The amount of nitrogen ingested when the medium-sized alga alone was offered was either higher than (stage C II) or not significantly different from that when the three algae were offered together (stage C IV). Ingestion rates are reduced when there is a multialgal food source. This implies that there is increased stability in the ocean because multiparticle food sources are more slowly depleted than unialgal foods. Weight-specific ingestion rates of copepods fed the three algae simultaneously increased from nauplius to stage C III and then decreased as adulthood approached. The contribution of the small alga to the total amount of nitrogen ingesied was greatest for naupliar stages while the contribution of the medium-sized cells was greater for later stages. The largest alga was readily ingested by stage C V and adult females but never contributed more than 25% of the nitrogen ingested. Eight to 12% of the nitrogen ingested by adult females was from the small alga. It is hypothesized that the algal cell size for maximum nitrogen ingestion in upwelled waters is relatively small, round or square and close to the size threshold below which adult females do not sense individual cells.     

Seasonal observations on the biometry and development in Copepodite Stage V of Calanus finmarchicus from the Firth of Clyde

Relationships between linear dimensions of casts produced at the final moult in the laboratory and definitive sex confirm that Calanus finmarchicus (Gunn.) is sexually dimorphic in Copepodite Stage V. Two forms of Stage V copepodite were present at all times of the year in the Firth of Clyde, Scotland. Except in early spring, when size was similar, these were consistently distinguished by the prosome length. The small form moulted to adult females and the large form to adults of either sex. Relative to the metasome, the length of the cephalosome was greater in the large than small form of Stage V copepodite. This, to an increased extent, distinguished males from all females in the adult. Differences between females of the large and those of the small form were similar in the Stage V copepodite and adult. Feminization in the large form entailed an arrest in the differentiation of secondary male sex-characters. Most morphological females moulting from both forms of Stage V copepodite possessed an apparently normal ovary, with eggs extending into oviducal diverticula. In the case of the large form, however, eggs were occasionally absent from diverticula. The gonad in two specimens resembled an immature testis. In males, a normal testis was always present. Differentiation into the adult male entailed the internal suppression of feminizing factors. On average, within collections, females of the large form moulted later than the small form and definitive males. A similarly consistent difference between females of the small form and males was absent. Moulting times in the large form of Stage V copepodite depended on the presence or not of feminization. Differentiation into the adult male required the environmental stimulation of development. Comparisons with moulting patterns in the sea suggest a mechanism that regulates the recruitment of males to the adult.     

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.     

Lipid composition of the copepod Calanus hyperboreas from the Arctic Ocean. Changes with depth and season

A build-up of reserve lipid, predominantly wax esters, occurred during the summer in the copepod Calanus hyperboreas, collected off an Arctic ice-island. This lipid storage was correlated with a phytoplankton bloom and was followed by a progressive decrease of lipid from 2.1 mg per individual in September to 0.4 mg in June. There was a rapid decrease in lipid utilization between October and December, followed by much slower decreases until June. Lowered respiration rates or the availability of different types or quantities of food in the winter and spring are possible explantions for the slower rate of lipid use. Laboratory starvation experiments for up to 90 days correlated with results from the field. Gas-liquid chromatographic studies of the lipids showed that both the alcohols and fatty acids of the wax esters were highly variable with season and depth, whereas the phospholipid fatty acids were not affected by changes in these parameters. Only summer samples bad wax esters with a phytoplankton-like fatty acid composition, and upper water winter copepods had wax esters with little polyunsaturation. The deep-water winter copepods had a very different wax-ester composition from the upper water samples, with a predominance of hexadecanol (all other copepod samples had 20:1 and 22:1¹ as the principal alcohols) and a high content of polyunsaturated acids. Deep-water C. hyperboreas may differ in food habits and life history from those in the upper water community.