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.     

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.     

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.     

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 hatching success in the calanoid copepods Calanus helgolandicus and Calanus finmarchicus in the North Sea from March to September 2001

Spatial and seasonal egg production rates (E_r) and egg hatchingsuccess in the copepods Calanus finmarchicus and Calanus helgolandicuswere measured in the North Sea from March to September. Foodavailability was monitored by chlorophyll and protist concentrationsand three size fractions of seston fatty acids. Seasonal andspatial distribution and production differed between the species.Calanus finmarchicus was found only offshore of the 50-m isobath,with decreasing E_r (37–28 eggs female^–1 day^–1)from March to July. Calanus helgolandicus had two abundancepeaks, in spring and autumn, with a low in May during whichtime the highest E_r were observed (38 eggs female^–1 day^–1).At other times, E_r in C. helgolandicus remained lower than inC. finmarchicus (     

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.     

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.     

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.     

Determining the mass density of marine copepods and their eggs with a critical focus on some of the previously used methods

We collected Calanus finmarchicus copepodites CIV, CV and CVIfemales in a deep fjord on the west coast of Norway during April1996, May 1997 and November 1998. Eggs of C. finmarchicus andCalanus glacialis were collected during May 1989 in the BarentsSea. The sinking speeds of animals and eggs were measured ina homogeneous column with seawater of known density, and Stokeslaw was applied to estimate their mass density. Also the densitycontrast between the organisms and seawater was calculated.The mean mass density of C. finmarchicus ranged from 1.0274to 1.0452 g cm^–3. During spring copepodite stage CV hada significantly lower mean mass density (1.0345 g cm^–3)compared to CIV (1.0381 g cm^–3) and CVI females (1.0408g cm^–3). Copepods collected during winter had a distinctlylower mass density. The sinking speed of C. glacialis eggs followeda unimodal distribution, with a mean of 25.9 m day^–1,while sinking speeds of C. finmarchicus eggs were bimodal, thetwo groups of eggs having a mean sinking speed of 23.3 m day^–1and 35.4 m day^–1 respectively. Correspondingly the meanmass density was 1.0556 g cm^–3 for C. glacialis eggs andfor the two groups of C. finmarchicus eggs 1.0639 g cm^–3and 1.0812 g cm^–3. Results of earlier work, particularlyusing density gradient methods to determine mass density ofzooplankton, are critically reviewed, and it is suggested thatthis method should not be used to determine the mass densityof small organisms the size of C. finmarchicus.     

Growth and development of Metridia pacifica (Copepoda: Calanoida) in the northern Gulf of Alaska

Juvenile growth and development rates for Metridia pacifica,one of the dominant larger copepods in the subarctic Pacific,were investigated from March through October of 2001–2004in the northern Gulf of Alaska. The relationship between prosomelength (PL, µm) and dry weight (DW, µg) was determined:log₁₀ DW = 3.29 x log₁₀ PL – 8.75. The stage durationsof copepodites ranged from 3 to 52.5 days but were 8–15days under optimal condition. Seasonally, growth rates increasedfrom March to October and typically ranged between 0.004 and0.285 day^–1, averaging 0.114 ± 0.007 day^–1(mean ± SE). After standardization to 5°C (Q₁₀ of2.7), growth rates averaged 0.083 ± 0.005 day^–1and were significantly correlated to chlorophyll a, with saturatedgrowth rates of 0.149 day^–1 for C1–C3, 0.102 day^–1for C4–C5 and 0.136 day^–1 for all stages combined.Measured juvenile growth rates were comparable with specificegg production rates in this species. The comparisons of ourrates in this study with those predicted by the global modelsof copepod growth rates suggested that further refinement ofthese models is required.     

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 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.     

Microzooplankton grazing in a coastal embayment off Concepcion, Chile, (~36{degrees} S) during non-upwelling conditions

The impact of grazing by natural assemblages of microzooplanktonwas estimated in an upwelling area (Concepción, Chile)during the non-upwelling season in 2003 and 2004. Seawater dilutionexperiments using chlorophyll a (Chl a) as a tracer were usedto estimate daily rates of phytoplankton growth and microzooplanktongrazing. Initial Chl a concentrations ranged from 0.4 to 1.4mg Chl a m^–3 and phytoplankton prey biomass and abundancewere numerically dominated by components <20 µm. Phytoplanktongrowth and microzooplankton grazing rates were 0.19–0.25day^–1 and 0.26–0.52 day ^–1, respectively.These results suggest that microzooplankton exert a significantremoval of primary production (>100%) during the non-upwellingperiod.     

Size-fractionated primary production studies in the vicinity of the Subtropical Front and an adjacent warm-core eddy south of Africa in austral winter

Results are presented of size-fractionated primary productionstudies conducted in the vicinity of the Subtropical Front (STF),an adjacent warm-core eddy, and in Sub-antarctic waters duringthe third South African Antarctic Marine Ecosystem Study (SAAMESIII) in austral winter (June/July) 1993. Throughout the investigation,total chlorophyll (Chl a) biomass and production were dominatedby small nano- and picophytoplankton. No distinct patterns intotal Chl a were evident. At stations (n = 7) occupied in thevicinity of the STF, total integrated biomass values rangedfrom 31 to 53 mg Chl a m^–2. In the vicinity of the eddy,integrated biomass at the eddy edge (n = 3) ranged from 24 to54 mg Chl a m^–2 and from 32 to 43 mg Chl a m^–2 inthe eddy (n = 2). At the station occupied in the Sub-antarcticwaters, total integrated biomass was 43 mg Chl a m^–2.Total daily integrated production was highest at stations occupiedin the vicinity of the STF and at the eddy edge. Here, totalintegrated production ranged from 150 to 423 mg C m^–2day^–1 and from 244 to 326mg C m^–2 day^–1, respectively.In the eddy centre, total integrated production varied between134 and 156 mg C m^–2 day^–1. At the station occupiedin the Sub-antarctic waters, the lowest integrated production(141 mg C m^–2 day^–1) during the entire survey wasrecorded. Availability of macronutrients did not appear to limittotal production. However, the low silicate concentrations duringthe survey may account for the predominance of small nano- andpicophytoplankton. Differences in production rates between theeddy edge and eddy core were related to water column stability.In contrast, at stations occupied in the vicinity of the STF,the control of phytoplankton production appears to be relatedto several processes, including water column stability and,possibly, iron availability.     

Description of the developmental stages in Acartia bifilosa Giesbrecht (Copepoda: Calanoida)

The developmental stages and increase in length of Acartia bifilosa,a dominant planktonic copepod in Kyeonggi Bay, Yellow Sea, Korea,were investigated in a laboratory. Adults laid eggs at a rateof 2.5 eggs day^–1 female^–1 and up to 97% of eggshatched. The eggs were 92 µm in diameter and larger thanthe eggs of other Acartia species. The body segmentation wasrecognizable at the fifth nauplius stage and sex distinctionwas possible at the fourth copepodite stage. Thesix naupliusstages took -7.5 days and the five copepodite stages -8.5 days.Sexually mature males and females appeared at the 16th day,and by the 20th day the adults laid eggs that hatched and grewsuccessfully into the nauplius stages. The detailed morphologicalcharacteristics of each developmental stage are presented anddiscussed in relation to A.dausi.     

Fluxes of diatoms in the Dona Paula Bay, west coast of India

Sediment traps were deployed at a station in the Dona PaulaBay to collect sedimenting particles at weekly intervals fromNovember to May during 1995–1997. Sedimented particleswere analysed for total diatom flux, chlorophyll a (Chl a) andparticulate organic carbon (POC). The highest diatom flux wasrecorded in April–May for both the years. Fluxes of diatomsvaried from0.6 x 10⁴ cells m^–2 day^–1 (November 1995)to 121.47 x 10⁴ cells m^–2 day^–1 (December 1996).In all, 19 diatom genera were identified in the sedimented material.Navicula, Nitzschia, Pleurosigma, Licmophora, Coscinodiscus,Rhizosolenia and Surirella were the most abundant genera inthe sedimented material throughout the sampling period. Meanflux of POC and diatom carbon was 251 and 0.39 mg C m^–2day^–1, respectively. The diatom carbon accounted for 0.15%of the POC flux. Mass flux of diatoms showed significant negativecorrelation with the concentration of nitrate and phosphate.This suggests that the nutrient concentration played an importantrole in influencing the sedimentation of diatoms at this coastalstation.     

In situ filtration responses of Daphnia galeata to changes in food quality

In the stoichiometric study of phyto–zooplankton interactions,a controversy exists about how Daphnia species regulate theirfeeding rate when submitted to low-quality food (i.e. high dietarycarbon:phosphorus [C:P] ratio). In this study, we gathered dataover 3 years on in situ clearance rates of a Daphnia galeatapopulation, by conducting grazing experiments from April 1998to October 2000 in the Esch-sur-Sûre reservoir (Grand-Duchyof Luxembourg). Observed clearance rates (2.5–13.5 mLindividual^–1 day^–1, mean 7.0 mL individual^–1day^–1) were correlated with population and environmentalvariables. Mean body size of Daphnia individuals was the bestpredictor of clearance rate (r² = 0.639), followed by watertemperature (r² = 0.262) and P concentration in the seston (r²     

Seasonal reproductive biology of Centropages tenuiremis (copepoda) in Xiamen waters, People's Republic of China

The egg production rate (EPR) and the effects of environmentalvariations on diapause induction in the copepod Centropagestenuiremis in Xiamen waters were studied in 2002 and 2003. TheEPR ranged from 7.33 ± 8.14 (on 18 February 2003) to91.08 ± 20.61 (on 6 June 2003) eggs female^–1day^–1(mean ± 95% confidence limits), and displayed a significantseasonal pattern. Before the period of maximum egg production,EPR increased with seawater temperature and chlorophyll a concentration;but after the maximum, no significant relationship was foundbetween EPR and chlorophyll a concentration, and there was anegative linear relationship between EPR and temperature. Seawatertemperature and photoperiod, respectively, played a primaryrole in the switch to diapause egg production by the femalesas shown by single-factor probit analysis. However, the differencein effect between the two factors was demonstrated by binarylogistic multivariate analysis. These results suggest that thiscopepod rapidly increases subitaneous EPR by taking full advantageof conditions in favorable seasons (winter and spring). SubitaneousEPR decreases and there is a switch to the production of diapauseeggs that survive in the bottom sediments during unfavorableseasons (summer and autumn) in Xiamen waters.     

On temperature-independent growth of phytoplankton

Growth rates of two freshwater diatoms and three chlorophyteswere compared under 3 h days at 10 µmol m^–2 s^–1.Specific growth rates ranged between µ=0.03 day^–1and µ=0.055 day^–1 for the different species andwere in every species independent of temperature between 8 and16°C.     

Community structure, biomass and productivity of size-fractionated summer phytoplankton populations in lower Narragansett Bay, Rhode Island

Seventeen size-fractionation experiments were carried out duringthe summer of 1979 to compare biomass and productivity in the< 10, <8 and <5 µm size fractions with that ofthe total phytoplankton community in surface waters of NarragansettBay. Flagellates and non-motile ultra-plankton passing 8 µmpolycarbonate filters dominated early summer phytoplankton populations,while diatoms and dinoflagellates retained by 10 µm nylonnetting dominated during the late summer. A significant numberof small diatoms and dinoflagellates were found in the 10–8µm size fraction. The > 10 µm size fraction accountedfor 50% of the chlorophyll a standing crop and 38% of surfaceproduction. The <8 µm fraction accounted for 39 and18% of the surface biomass and production. Production by the< 8 µm fraction exceeded half of the total communityproduction only during a mid-summer bloom of microflagellates.Mean assimilation numbers and calculated carbon doubling ratesin the <8 µm (2.8 g C g Chl a^–1 h^–1; 0.9day^–1)and<5 µm(1.7 g C g Chl a^–1h^–1; 0.5day^–1)size fractions were consistently lower than those of the totalpopulation (4.8 g C g Chl a^–1 h^–1; 1.3 day^–1)and the <10 µm size fraction (5.8 g C g Chl a^–1h^–1; 1.4 day ^–1). The results indicate that smalldiatoms and dinoflagellates in fractionated phytoplankton populationscan influence productivity out of proportion to their numbersor biomass. ¹Present address: Australian Institute of Marine Science, P.M.B.No. 3, Townsville M.S.O., Qld. 4810, Australia.