Egg production and hatching success in the peri-Antarctic copepod Calanus simillimus

In situ rates of egg production and hatching success are reportedfor Calanus simillimus, one of the most abundant calanoid speciesin peri-Antarctic regions, during the Italian ‘Italics’cruise in the Straits of Magellan in March-April 1995. Low fecundity(8.6 eggs female^–1 day^–1) and fecal pellet production(0.8 fecal pellets female^–1 day^–1) in this periodindicate that the species was feeding very poorly. Sixty-sixof the 126 females sampled did not produce eggs and 80 of thesedid not produce any fecal pellets during the 24 h period ofincubation. Striking abnormal naupliar and embryonic developmentwas recorded in 81.8% of the eggs spawned. Aberrant eggs didnot undergo normal cleavage, and failed to develop to hatching.Deformed nauplii were asymmetrical and presented strong anatomicalanomalies in the total body length and number of swimming appendages.These results are discussed in the light of recent findingson the causes of low hatching success in copepods.     

The relative importance of food and temperature to copepod egg production and somatic growth in the southern Benguela upwelling system

The fecundity and somatic growth rates of Calanus agulhensisand Calanoides carinatus, the dominant large calanoid copepodsin the southern Benguela upwelling system, as well as the fecundityof several other common copepods, were measured between Septemberand March of 1993/94 and 1994/95. Mean egg production of mostcopepods was low at >30 eggs female^-1 day^-1 {Calanoides carinatus23.7, Calanus agulhensis 19.0, Neocalanus tonsus 16.1 and Rhincalanusnasutus 26.1), whereas the mean fecundity of Centropages brachiatuswas significantly greater (83.6 eggs female^–1 day^-1).This study also presents the first comprehensive field estimatesof the fecundity of Nanno-calanus minor (mean: 26.1 eggs female^–1day^–1, range: 0.0–96.2 eggs female^–1 day^–1)and of somatic growth of N6 and all copepodite stages of Calanoidescarinatus (decreasing from 0.58 day^–1 for N6 to 0.04 day^–1for C5). Somatic growth rates of Calanus agulhensis also declinedwith age: from 0.57 day1 for N6 to 0.09 day1 for C5. Data ongrowth rates were used to assess the relative importance offood [as measured by total chlorophyll (Chi) a concentration],phytoplankton cell size (proportion of cells >10 µm)and temperature to the growth of copepods. Multiple regressionresults suggested that fecundity and somatic growth rates werepositively related to both Chi a concentration and phytoplanktoncell size, but not to temperature. Although it was not possibleto separate the effects of Chi a concentration and phytoplanktoncell size, data from previous laboratory experiments suggestthat copepod growth is not limited by small cells per se, butby the low Chi a concentrations that are associated with theseparticles in the field. Despite growth not being directly relatedto temperature, a dome-shaped relationship was evident in somespecies, with slower growth rates at cool (<13°C) andwarm (>18°C) temperatures. The shape of this relationshipmirrors that of Chi a versus temperature, where poor Chi a concentrationsare associated with cool and warm temperatures. It is concludedthat the effect of food limitation on growth of copepods outweighsthat of temperature in the southern Benguela region. Sourcesof variability in relationships between growth and Chi a concentrationare discussed.     

Seasonal variation of reproduction rates and body size of Calanus sinicus in the Southern Yellow Sea, China

Annual variations of egg production rate (EPR) and clutch sizeof Calanus sinicus, as well as body size of females (prosomelength and dry weight), were investigated at a series of stationsin the Southern Yellow Sea by onboard incubation. Calanus sinicuswas spawning in all the 11 cruises investigated, and the annualvariation of EPR was bimodal. Monthly average EPR was highestfrom May to July, respectively, 5.97, 5.36 and 6.30 eggs female^–1d^–1, then decreased dramatically to only 1.37 eggs female^–1d^–1 in August and attained the lowest 1.07 eggs female^–1d^–1 in October. In November, average EPR increased againto 4.31 eggs female^–1 d^–1. Seasonal variation ofclutch size was similar to EPR, except that it decreased graduallyafter August rather than dramatically as did EPR. Prosome lengthof females was maximum in May and minimum in October, but dryweight was highest in November. Monthly average EPR correlatedbetter with prosome length than dry weight, while clutch sizewas rather determined by dry weight of females. It is suggestedthat egg production of C. sinicus was active during two discontinuousperiods when both surface and bottom temperature fell into itsfavorite range (i.e. 10–23°C), and different reproductivestrategies were adopted in these two reproductive peaks: otherthan the highest EPR, longer prosome length was also achievedby C. sinicus from May to July, while females in November developedshorter bodies but accumulated more energy for reproduction.     

Egg production and growth rates of Calanus euxinus (Copepoda) in the Black Sea

Egg production rates (EPRs) of Calanus euxinus were measuredin the Black Sea during October 2000 and May 2001. EPRs weregenerally low, on average 1.7 eggs female^–1 day^–1in October 2000 and 3.9 eggs female^–1 day^–1 in May2001. The relationships between EPRs and gonad maturity, depth-integratedchlorophyll a (Chl a) and mean surface layer temperature wereexamined. The EPRs were not related to depth-integrated Chla, but were negatively correlated with temperature. EPRs werestrongly related to the proportion of mature females. Growthrates of C. euxinus were derived from the EPRs. The mean growthrate was 0.011 day^–1 in October 2000 and 0.03 day^–1in May 2001. Growth rates were not significantly correlatedwith Chl a concentrations, but were negatively related to femaleweight and temperature.     

Feeding and reproduction of Calanus finmarchicus during non-bloom conditions in the Irminger Sea

Simultaneous ingestion and egg production experiments were conductedwith female Calanus finmarchicus in April/May and July/August2002 in the Irminger Sea. Experimental animals were providedwith natural microplankton food assemblages and incubated underin situ conditions for 24 h. The quantity of food consumed wassignificantly related to the concentration of prey cells, withtotal daily ingestion rates ranging from 0.6 to 8.1 µgof carbon female^–1 day^–1, corresponding to carbon-specificrates of 0.6–4.7% day^–1. Egg production rates (EPRs)remained relatively low (0.3–11 eggs female^–1 day^–1)during both periods of investigation and were not influencedby food availability. The data were used to construct energeticbudgets in which the microplankton carbon ingested, includingciliates, was compared with the carbon utilized for egg productionand respiration. These budgets showed that ingestion alone couldnot provide the necessary carbon to sustain the observed demandsfor growth and metabolism. Although ciliates constituted >80%of the total material ingested at times, they were not sufficientto provide the metabolic shortfall. Indeed, the females weretypically lacking 5 µg of carbon each day, 5% of theircarbon biomass. Our study results highlight the possible importanceof internal reserves in sustaining reproduction in C. finmarchicusduring periods of food scarcity.     

A comparison of seasonal growth and development of the copepods Calanus marshallae and C. pacificus in the northern Gulf of Alaska

The juvenile growth rates and development times of subarcticCalanus marshallae and temperate/sub-tropical C. pacificus wereinvestigated during nine cruises (May through October, 2001–04)in the northern Gulf of Alaska. The artificial cohort methodbased on a length-weight regression was used for growth estimatesand the reciprocal of the molting rate for developmental time.The copepodite stage duration ranged from 3 to 16 days for C.marshallae (C1–C4) and 3–23 days for C. pacificus(C1–C5). Seasonally, copepodid growth rates increasedfrom May to October, ranging between 0.055 and 0.291 day^–1(mean ± SE: 0.176 ± 0.008 day^–1) for C.marshallae, while growth rates increased from August to Octoberbetween 0.018 and 0.296 day^–1 (mean ± SE: 0.142± 0.016 day^–1) for C. pacificus. After standardizationto 5°C (Q₁₀ of 2.7), growth rate averaged 0.118 ±0.007 day^–1 and 0.075 ± 0.009 day^–1 for C.marshallae and C. pacificus, respectively. Calanus marshallaegrowth rate is satisfactorily described by a Michaelis–Mentenmodel using chlorophyll-a concentration (r² = 0.33) after temperaturecorrection, but the prediction improves with a composite nonlinearmodel combining body weight into the Michaelis–Mentenfunction (r² = 0.55). Considering the limited range of dataavailable for C. pacificus, the combination of the data forboth species suggests that C. pacificus has a similar functionalresponse to growth despite the differences in the geographicand temporal distributions with C. marshallae. Measured juvenilegrowth rates of the two Calanus species in this study were comparableto other calanoid species in the same area and showed reasonableagreement to Calanus growth models but less with global copepodgrowth models.     

Pigment ingestion and egg production rates of the calanoid copepod Temora longicornisr. implications for gut pigment loss and omnivorous feeding

We examined the relationship between egg production rate (E)and pigment ingestion rate (I, from gut content corrected for33% loss) for adult female Temora longicornis in Long IslandSound on 47 occasions. Linear regression of E on I [both variablesexpressed in mass of nitrogen (N) female^–1 day^–1]was: E_N = 0.0016 + 0.770 x I_N. The slope, 0.77, is the apparentgross efficiency of egg production, equivalent to the grossgrowth efficiency (GGE) assuming that females partition allnitrogen for growth into egg production. Published work suggeststhat a GGE of 0.37 would be expected for herbivorous copepods.The discrepancy between the expected value of 0.37 and observedvalue of 0.77 could result from unquantified losses of gut pigmentor because T.longicomis ingested a significant amount of nitrogenby feeding as a carnivore. We suggest that if T.longicomis femalesderive all of their nitrogen for growth by feeding on phytoplankton,and if no correction for pigment loss is employed, then thegut pigment method underestimates pigment ingestion by no morethan a factor of two.     

Distribution and species-specific egg production of Pseudocalanus in the Gulf of Alaska

Pseudocalanus species are important contributors to the secondaryproduction of the northern hemisphere mid- to high-latitudeoceans. In the coastal Gulf of Alaska, Pseudocalanus are presentyear round and are represented by three species. In 2001, Pseudocalanusmimus was the dominant Pseudocalanus species on the shelf duringspring and summer, comprising 30–100% of the total, whilePseudocalanus newmani dominated in Prince William Sound (10–90%).Pseudocalanus minutus were only abundant in Prince William Soundduring early spring. Egg production (by number and volume) wasa function of female prosome length and decreased from springto summer; however, significant variability was attributableto regional influences that were independent of size. For thesame sized female, P. newmani produced more eggs per clutchthan P. mimus. Pseudocalanus mimus, however, tended to havea larger mean egg size than P. newmani. Consequently, clutchvolumes of the two species were indistinguishable. Pseudocalanusegg production rates (EPRs) (eggs female^–1 day^–1)were lower in July and August (ca. 2–4) than April andMay (ca. 1–9), but total egg production by the population(eggs day^–1) was nearly equivalent for the two time periodsdue to higher female concentrations in summer.     

Growth and development of Neocalanus flemingeri/plumchrus in the northern Gulf of Alaska: validation of the artificial-cohort method in cold waters

In situ growth and development of Neocalanus flemingeri/plumchrusstage C1–C4 copepodites were estimated by both the artificial-cohortand the single-stage incubation methods in March, April andMay of 2001–2005 at 5–6°C. Results from thesetwo methods were comparable and consistent. In the field, C1–C4stage durations ranged from 7 to >100 days, dependent ontemperature and chlorophyll a (Chl a) concentration. Averagestage durations were 12.4–14.1 days, yielding an averageof 56 days to reach C5, but under optimal conditions stage durationswere closer to 10 days, shortening the time to reach C5 (fromC1) to 46 days. Generally, growth rates decreased with increasingstage, ranging from 0.28 day^–1 to close to zero but weretypically between 0.20 and 0.05 day^–1, averaging 0.110± 0.006 day^–1 (mean ± SE) for single-stageand 0.107 ± 0.005 day^–1 (mean ± SE) forartificial-cohort methods. Growth was well described by equationsof Michaelis–Menten form, with maximum growth rates (G_max)of 0.17–0.18 day^–1 and half saturation Chl a concentrations(K_chl) of 0.45–0.46 mg m^–3 for combined C1–3,while G_max dropped to 0.08–0.09 day^–1 but K_chl remainedat 0.38–0.93 mg m^–3 for C4. In this study, in situgrowth of N. flemingeri/plumchrus was frequently food limitedto some degree, particularly during March. A comparison withglobal models of copepod growth rates suggests that these modelsstill require considerable refinement. We suggest that the artificial-cohortmethod is the most practical approach to generating the multispeciesdata required to address these deficiencies.     

Reproductive response of the copepods Calanoides carinatus and Calanus agulhensis to varying periods of starvation in the southern Benguela upwelling region

Egg production by the calanoid copepods Calanoides carinatusand Calanus agulhensis fed excess Thalassiosira weissflogiiwas monitored in the laboratory following starvation periodsof 1, 3, 5, 7 and 9 days. Following short (1–3 day) periodsof starvation, egg production by C.agulhensis returned to thesatiated rate (51.1 eggs {female} day^–1) more rapidly(after 0.9–2.4 days of excess food) than that of Ca. carinatus(after 2.8–3.1 days). However, following longer (5–9day) periods of starvation, Ca. carinatus regained satiatedlevels of egg production (55.8 eggs {female}^–1 day^–1)more rapidly (after 3.1–4.0 days of excess food) thanC. agulhensis (after 3.8–5.2 days of feeding following5–7 days of starvation). Moreover, many C. agulhensisfemales did not regain normal rates of egg production after9 days of starvation. For both species, the time required foregg production to recover was proportional to the starvationperiod, although only up to 7 days for C. agulhensis, and wasthe same following 4.25 days of starvation. Previously fed Ca.carinatus terminated egg production more rapidly than C. agulhensiswhen starved. The ability of Ca. carinatus to tolerate, andrecover rapidly from, prolonged periods of starvation, combinedwith a comparatively fast development time and high rate ofegg production, provides this species with a strong competitiveadvantage over C. agulhensis in the highly pulsed food environmentof the southern Benguela upwelling region.     

Feeding rates and selectivity among nauplii, copepodites and adult females of Calanus finmarchicus and Calanus helgolandicus

This study focuses on selective feeding by developmental stages of two oceanic copepods, Calanus finmarchicus and Calanus helgolandicus from nauplii to adults. A mixture of four algal species of different biochemical composition, Prorocentrum nanum (dinoflagellate), Thalassiosira minima (diatom), Rhodomonas baltica (cryptophyte) and Dunaliella tertiolecta (chlorophyte), added in an equal biovolume, was used in three different experimental set-ups. In set-up 1 the algal species were present as single cells of similar size (14 μm). In set-up 2 the diatom T. minima was present in chains of two or three cells and was therefore larger than the other algae, while the biovolume of all species remained the same. In set-up 3, the diatom T. minima was excluded from the mixture. Feeding selectivity of the copepods was assessed in relation to the quality of the algal species expressed in terms of carbon and nitrogen content, fatty acid composition, and chain length of the diatom. The results show that younger stages and adult females of C. finmarchicus and C. helgolandicus did not show a preference for an algal species when the algae were of similar size. In the feeding experiments where the diatoms were offered as chains, both copepod species showed a selective behaviour only on the basis of algal size. Individual ingestion rates increased from 0.4 to 0.7 μg C day^–1 for nauplii of both species to 5 μg C day^–1 for adult females of C. helgolandicus to 12 μg C day^–1 for C. finmarchicus. Individual filtration rates ranged from 5 ml day^–1 for C. finmarchicus nauplii to 70–98 ml day^–1 for adult females, and from 3 ml day^–1 for C. helgolandicus nauplii to 35–46 ml day^–1 for adult females. Ingestion and filtration rates per unit body carbon decreased gradually in both copepod species with increasing body carbon. The daily ingested amount of food decreased for C. finmarchicus from 124–134% of the body carbon for nauplii to 19% of the body carbon for adult females, and for C. helgolandicus from 117–137% of the body carbon for nauplii to 13–26% of the body carbon of adult females. Electronic Publication     

Copepod egg production, moulting and growth rates, and secondary production, in the Skagerrak in August 1988

Measurements of hydrography, chlorophyll, moulting rates ofjuvenile copepods and egg production rates of adult female copepodswere made at eight stations along a transect across the Skagerrak.The goals of the study were to determine (i) if there were correlationsbetween spatial variations in hydrography, phytoplankton andcopepod production rates, (ii) if copepod egg production rateswere correlated with juvenile growth rates, and (iii) if therewas evidence of food-niche separation among co-occumng femalecopepods The 200 km wide Skagerrak had a stratified water columnin the center and a mixed water column along the margins. Suchspatial variations should lead to a dominance of small phytoplanktoncells in the center and large cells along the margins; however,during our study blooms of Gyrodinium aureolum and Ceratium(three species) masked any locally driven differences in cellsize: 50% of chla was >11 µm, 5% in the 11–50µm fraction and 45% <50 µm. averaged for allstations. Chlorophyll ranged from 0.2 to 2.5 µg l^–1at most depths and stations. Specific growth rates of copepodsaveraged 0.10 day^–1 for adult females and 0.27 day^–1for juveniles The latter is similar to maximum rates known fromlaboratory studies, thus were probably not food-limited. Eggproduction rates were food-limited with the degree of limitationvarying among species: 75% of maximum for Centropages typicus, 50% for Calanus finmarchicus, 30% for Paracalanus parvus and 15% for Acartia longiremis and Temora longicornis. Thedegree of limitation was unrelated to female body size suggestingfood-niche separation among adults. Copepod production, summedover all species, ranged from 3 to 8 mg carbon m^–3day^–1and averaged 4.6 mg carbon m^–1 day^–1. Egg productionaccounted for 25% of the total.     

Plankton community respiration in Bransfield Strait (Antarctic Ocean) during austral spring

Community respiration (R) was determined in Bransfield Straitfrom oxygen changes in water samples incubated in borosilicatebottles maintained at in situ temperature. The respiratory electrontransport system (ETS) activity of seawater communities wasalso measured from the same samples. Both data sets were relatedby the regression equation: log R (mg O₂ m^–3 day^–1)=0.462+0.730xlogETS activity mg O₂ m^–3 day^–1) (r=0.80, n=23). Fromthis equation and 37 ETS activity depth profiles, we calculatedthe integrated (0–100 m) community respiration as beingin the range 1.2–4.5 g O₂ m^–2 day^–1 (mean=2.2).These values do not differ significantly from other publishedresults for the Arctic and Antarctic Oceans. Assuming a respiratoryquotient of unity, the areal respiration ranges between 0.45and 1.69 g C m^–2 day^–1 (mean=0.8). This would representan important sink for the primary production reported for BransStrait. The spatial distribution of community respiration showedhigher values associated with the warmer and phytoplankton-richwaters outflowing from Gerlache Strait into Bransfield Strait,and with the front that separates Bellingshausen Sea watersfrom Weddell Sea waters. We suggest that this pattern of distributionmay be related to the transport of organic matter by the BransfieldCurrent along the front.     

Seasonal Cycles of Egg Production of Two Planktonic Copepods, Centropages typicus and Temora stylifera, in the North-western Mediterranean Sea

Reproduction of the dominant copepods Centropages typicus andTemora stylifera was studied at a permanent station in the LigurianSea (north-western Mediterranean). Seasonal patterns of eggproduction, clutch size, egg size and female prosome lengthwere followed from January 1998 to December 1999. Female carboncontent and weight-specific egg production were compared inautumn 1998 and spring 1999. Reproductive patterns of C. typicusand T. stylifera were very similar, indicating that reproductionwas affected by the same environmental factors. Reproductiveactivity was highest in autumn in both species and years. Asecond peak of egg production was observed in early summer,which was less intense in 1999 after a bloom of salps. Egg productionrates reached maximal values of 33.5 and 33.3 eggs female^–1day^–1 and annual means of 10.8 and 11.7 eggs female^–1day^–1 in Centropages and Temora, respectively. Maximalweight-specific egg production was 0.21 day^–1 in bothspecies in November 1998, when female carbon contents were 6.7(C. typicus) and 12.0 µg (T. stylifera). No statisticalrelationship between egg production and food availability ortemperature was detected. Reproductive activity did not reflectthe seasonal abundance patterns, with C. typicus dominatingin spring and T. stylifera in autumn.     

Seasonal and fine-scale spatial variations in egg production and triacylglycerol content of the copepod Acartia tonsa in a river-dominated estuary and its coastal plume

We measured egg production rates of the estuarine calanoid copepodAcartia tonsa in Mobile Bay, an estuary in the northern Gulfof Mexico. Two stations were sampled approximately monthly,one at the mouth of the bay and the other just beyond the mouthin the salinity front between bay and coastal waters. Over thewhole year, temperature was the most important environmentalvariable controlling egg production. Rates increased with temperatureup to 30°C and 140 eggs female^–1 day^–1. We foundno evidence of food limitation. There was no correlation betweenegg production and phytoplankton abundance, nor increased eggproduction in response to supplements of phytoplankton addedto natural food, suggesting that non-phytoplankton food wasimportant in the diet. At the highest egg production rates,the amount of the storage lipid triacylglycerol (TAG) in adultfemales was greatly diminished, to <50 ng female^–1.This suggests that lipids in the diet can be very tightly coupledto egg production. Both egg production and TAG content of femalesshowed significant variability on spatial scales of 3–15km, especially in relation to the salinity front separat ingwater outwelling from the bay from open coastal water. For organismsthat are using copepods and their eggs as food, this variabilitywould result in a heterogeneous food environment, both in termsof the abundance of food and its nutritional content.     

Growth and production of Euphausia lucens in the southern Benguela Current

The growth rate of a population of Euphausia lucens from thewest coast of South Africa was estimated from laboratory studiesand from monthly size-frequency distributions of samples collectedover a 1-year period. Laboratory studies indicated that growthrates ranged from 0.131 (larvae) to 0.047 mm day^–1 (juveniles),while size-frequency distributions suggested a growth rate of{small tilde}0.026 mm day^–1 for the adults. The mean annualbiomass from the inshore, intermediate and offshore regionsranged from 9.75 to 47.29 mg dry wt m^–3 with the highestbiomass being found in the inshore region. Calyptopis larvaewere present for most months of the year, indicating continuousrecrwtment. The relative contribution of flesh, moults and eggsto the total annual production was estimated separately forall three regions. Production due to growth (P_g) was estimatedto be 92.71–185.60 mg dry wt m^–3 year^–1, whileexuviai production (P_e) varied between 60.01 and 281.38 mg drywt m year Production of eggs (P_r) was estimated to range from5.07 to 12.39 mg dry wt m year the lowest value being obtainedin the inshore region. Moult production represented {small tilde}6times the mean biomass in each region, while the P/B ratio forflesh production varied from 3.92 to 8.91, the highest ratiobeing obtained in the offshore region. Total P/B ratios rangedfrom 10.14 to 16.01.     

The ecology of the Calanus ponticus population in the deeper layer of its concentration in the Black Sea

The layer of daytime concentration of Calanus ponticus (VC andVI C) performing daily vertical migrations and the layer of‘winter stock’ aggregation are confined to the depthof maximal gradient of the main pycnocline under an unusuallysharp oxycline. The concentration layer thickness ranges from2 to 20–30 m and the Calanus concentration in it is >250ind. m^–3, sometimes being 3500 ind. m^–3 and evenmore. The population in the concentration layer is divided intotwo ecological groups: I, feeding and migrating specimens ofcopepodite stages V and VI, their body lipid contents being25–60 µg min.^–1; and II, non-feeding and non-migratingspecimens of copepodite stage V, their body lipid contents being100–150 µg ind.^–1. The relationship with oxygenconcentration was studied in both ecogroups. The experimentsshow that specimens of ecogroup II can exit at an oxygen concentrationof 0.06 ml 1^–1, but at such concentration falling intoanabiosis. They die at 0.04 ml O₂ 1^–1. Estimates of respirationof the group II specimens (‘winter stock’) showthat lipids they store are sufficient for 7 months' survival.Depth of Calanus concentration is determined by water densityrather than concentration of oxygen.     

Planktonic primary production in the German Wadden Sea

By combining weekly data of irradiance, attenuation and chlorophylla concentrations with photosynthesis (P) versus light intensity(E) curve characteristics, the annual cycle of planktonic primaryproduction in the estuarine part of the Northfrisian WaddenSea was computed for a 2 year period. Daily water column particulategross production ranged from 5 to 2200 mg C m^–2 day^–1and showed a seasonal pattern similar to chlorophyll a. Budgetcalculation yielded annual gross particulate primary productionsof 124 and 176 g C m^–2 year^–1 in 1995 and 1996,respectively. Annual amounts of phytoplankton respiration, calculatedaccording to a two-compartment model of Langdon [in Li,W.K.W.and Maestrini,S.Y. (eds), Measurement of Primary Productionfrom the Molecular to the Global Scale. International Councilfor the Exploration of the Sea, Copenhagen, 1993, pp. 20–36],and dissolved production in 1996, were both in the range of24–39 g C m^–2 year^–1. Annual total net productionwas thus very similar to particulate gross production (127 and177 g C m^–2 year^–1 in 1995 and 1996, respectively).Phytoplankton growth was low or even negative in winter. Inspring and summer, production/biomass (Pr/B) ratios varied from0.2 up to 1.7. Phytoplankton growth during the growth seasonalways surpassed average flushing time in the area, thus underliningthe potential of local phytoplankton bloom development in thispart of the Wadden Sea. The chlorophyll-specific maximum photosyntheticrate (P^B_max) ranged from 0.8 to 9.9 mg C mg^–1 Chl h^–1and was strongly correlated with water temperature (r² = 0.67).By contrast, there was no clear seasonal cycle in ^B, which rangedfrom 0.007 to 0.039 mg C mg^–1 Chl h^–1 (µmolphotons m^–2 s^–1)^–1. Its variability was muchless than P^B_max and independent of temperature. The magnitudeand part of the variability of P^B_max and ^B are presumably causedby changes in species composition, as evidenced from the rangeof these parameters found among 10 predominant diatom speciesisolated from the Wadden Sea. The ratio of average light conditionsin the water column (E_av) to the light saturation parameterE_k indicates that primary production in the Wadden Sea regionunder study is predominantly controlled by light limitationand that nutrient limitation was likely to occur for a few hoursper day only during 5 (dissolved inorganic nitrogen) to 10 (PO₄,Si) weeks in the 2 year period investigated.     

Production of Oikopleura longicauda (Tunicata: Appendicularia) in Toyama Bay, southern Japan Sea

Oikopleura longicauda occurred throughout the year in ToyamaBay, southern Japan Sea, and analysis of its size compositionand maturity revealed that reproduction was continuous overtheyear. Somatic growth production (P_g) varied with season from0.03 to 103 mg carbon (C) m^–2day^–1 (annual P_g 4.5g C m^–2), and house production (P_e) from 0.11 to 266 mgC m^–2 day^–1 (annualP_e 11.3 g C m^–2). The annualP_g/B ratio was 176. Compared with production data of some predominantzooplankton species in Toyama Bay, it is suggested that despitetheir smaller biomass, appendicularians are an important secondaryproducer.     

Seasonal changes in zooplankton composition in the Barents Sea, with special attention to Calanus spp. (Copepoda)

The Barents Sea is an important area with respect to fisheriesresources (i.e. capelin and cod). In May, June and August 1981zooplankton biomass was measured along a transect at 30°E,from the ice border southwards. A maximum was recorded in Atlanticwater by the end of June (>100 g wet weight m^–2 InAugust the biomass values were relatively low south of the Polarfront and increased northwards into Arctic water (–50g m^–2 The species composition was influenced by the distributionof cold Arctic water and warmer Atlantic water. The zooplanktonwas dominated by the copepods Calanus finmarchicus and C. glacialis;the former is regarded as an Atlantic species and C. glacialisas an Arctic species.