Natural filtration rates of zooplankton in a closed system: the derivation of a community grazing index

Measurements of individual filtration rates by natural suspensionsof zooplankton are presented. The products filtration rate xanimal concentration, calculated for each of several species-sizecategories, are summated to estimate the total volume of waterfiltered by the zooplankton per unit time, here defined as the"Community Grazing Index" (CGI). Seasonally, CGI varies overnearly three orders of magnitude. The Daphnia hyalina individualspresent frequently contributed >90% of CGI. Individual filtrationrates were highest when the concentrations of suitable foodswere limiting (equivalent to 0.1–0.2 µg C ml^–1),but were depressed after long (3-week) periods of low food availabilityand when large Microcystis colonies dominated the phytoplankton.Approximately one order of magnitude separated these extremes.Fluctuations in CGI owe relatively more to change in the concentration,size and species distribution of the animal population thanto changes in individual filtration rate. ¹Present address: Anglian Water Authority, Oundle, PeterboroughPE8 4AS, UK     

Experimental studies of food and suspended sediment effects on growth and reproduction of six planktonic cladocerans

Life-long somatic growth and egg production responses of Daphniabarbata, Daphnia gibba, Daphnia laevis, Daphnia pulex, Moinamicrura and Diaphanosoma excisum were examined in relation todifferent food and suspended sediment levels, in factorial experimentsat 20°C. Food treatments involved additions of 0, 100, 500and 2500 µg C l^–1 of Selenastrum to 20 µmfiltered water, generally from source lakes. These food levelswere tested at two suspended sediment levels in all species,viz. (i) natural source-lake nephelometric turbidities of 7–65nephelometric turbidity units (NTU) ({small tilde}10–70mg l^–1 TSS) and (ii) {small tilde}3- to 4-fold enrichmentof natural levels (8-fold in D.excisum). A third sediment-freetreatment was tested in all species of Daphnia other than D.gibba.Growth in body length and reproductive output were assesseddaily for individual animals. Main and interactive effects offood and sediment on growth were assessed at selected ages fromday 3 through subsequent life using analysis of variance (ANOVA).Strong and highly significant positive food effects were apparentat all ages in all species examined, with the curious exceptionof D.excisum. The onset of maturity and fecundity were alsopositively related to food level. Sediment effects on growth,age of maturity and fecundity were variable, but tended to bemuch weaker than food effects. Both stimulatory (positive, especiallyat low food levels) and inhibitory (negative) effects of sedimenton growth were apparent, both between and within species. Performancein sediment-free water was very poor, except in D.pulex. Food-sedimentinteraction effects were variable. Generally, turbidwater specieswere influenced less adversely by sediment than clear-waterspecies, indicating the existence of environmentally appropriateadaptive responses. Overall, the findings indicate that lightlimitation of algal resources by suspended sediments, ratherthan the condition of high turbidity per se, is responsiblefor the apparently negative impact of suspended sediments frequentlyobserved on natural populations of cladoceran zooplankters.     

Experimental studies into the feeding biology of rotifers in brackish water

Mass developments of rotifers of the genus Brachionus, and especiallyof B.quadridentatus, occur regularly in the largely hypertrophicchain of shallow waters (‘boddens’) south of theDarss-Zingst peninsula (Southern Baltic). Interest in the autecologyof the species is, therefore, considerable. Various food sourceswere used in laboratory experiments to ascertain the food requirementsof B.quadridentatus, determine its filtration and ingestionrates, and assess its food particle size-selection ability.Growth experiments showed that the chlorophyceans Nannochlorissp. and Chlorella vulgaris possess considerable nutritionalvalue for the species, whereas abundances declined when Microcystisfirma, detritus from Enteromorpha sp. and only bacteria (Pseudomonas),respectively, were provided as food sources. Filtration ratesvaried between 0.02 and 1.73 µl ind.^–1 h^–1,and ingestion rates between 121 and 5560 cells ind.^–1h^–1, depending on the filtration rate and algal concentration.Investigations into food particle size selection using fluorescentlatex particles revealed that particle size influences foodparticle intake. When particles of different sizes were mixed,the animals showed a preference for the larger particles andingested the smaller ones with a diameter of 1–2 µmless efficiently. The brackish water species Brachionus plicatiliswas studied besides B.quadridentatus in all experiments. Theformer species proved to be superior both in its range of utilizableparticle sizes and its growth rate. The experiments with laboratorycultures were backed up by studies performed with various rotiferspecies taken from natural populations.     

Carbon dynamics in the 'grazing food chain' of a subtropical lake

Studies were conducted over a 13 month period at four pelagicsites in eutrophic Lake Okeechobee, Florida (USA), in orderto quantify carbon (C) uptake rates by size-fractionated phytoplankton,and subsequent transfers of C to zooplankton. This was accomplishedusing laboratory ¹⁴C tracer methods and natural plankton assemblages.The annual biomass of picoplankton (<2 µm), nanoplankton(2–20 µm) and microplankton (<20 µm averaged60, 389 and 100 µg C 1^–1 respectively, while correspondingrates of C uptake averaged 7, 51 and 13 µg C1^–1h^–1. The biomass of microzooplankton (40–200 µm)and macrozooplankton (<200 µm averaged 18 and 60 µgC 1^–1, respectively, while C uptake rates by these herbivoregroups averaged 2 and 3 µg C 1^–1 h^–1. Therewere no strong seasonal patterns in any of the plankton metrics.The ratio of zooplankton to phytoplankton C uptake averaged7% over the course of the study. This low value is typical ofthat observed in eutrophic temperate lakes with small zooplanktonand large inedible phytoplankton, and indicates ineffectiveC transfer in the grazing food chain. On a single occasion,there was a high density (<40 1^–1) of Daphnia lumholrzii,a large-bodied exotic cladoceran. At that time, zooplanktoncommunity C uptake was <20 µg C 1^–1 h^–1and the ratio of zooplankton to phytoplankton C uptake was near30%. If D.lumholrzii proliferates in Lake Okeechobee and theother Florida lakes where it has recently been observed, itmay substantially alter planktonic C dynamics.     

Sedimentation of phytoplankton populations dominated by Microcystis in a shallow lake

The sedimentary flux of phytoplankton was measured using sedimenttraps in a shallow hypertrophic lake (Lake Kasumigaura), whereMicrocystis bloomed, from June to November 1983 The sedimenttraps were set at 0.5, 1.5 and 3.0 m depth in Takahamairi Bay(3.5 m depth). Microcystis spp. (including M.aerugmosa and M.viridis)in the traps were rare until early August, but increased thereafter.Sinking rates of Microcystis were 0.0045, 0.020 and 0.24 m day^–1in June–August, September and October respectively, whichwere far lower than those of Melosira (0.2–1.7 m day^–1)and Syncdra (0.2–1.0 m day^–1). The total sedimentaryfluxes of POC and that of algal carbon during the study periodwere 283.2 and 96.7 gC m^–2 which were 59.5% and 20.3%of the gross primary production (475.8 gC m^–2) respectively.The sedimentary flux of living algae measured by algal countswas large in June but small in August and September. On theother hand, the flux of detritus obtained by subtracting totalalgal carbon from POC was small in June and July but large inAugust and September. Therefore diatoms, which appeared mostlyin June, tended to sink as live algae, while Microcystis sankas detritus after being decomposed or consumed in the waterIt was concluded from the results of carbon budget calculationsand the respiration rate of the 1- to 20-µm fraction thatthe activity of decomposers or consumers increased greatly inthe short period at the end of the bloom of Microcystis.     

Respiratory rates in the cladoceran Ceriodaphnia dubia in Lake Rotongaio, a monomictic lake

The experimentally measured oxygen consumption rate by the cladoceran,Ceriodaphnia dubia, showed a linear increase between 5 and 20°C.Oxygen consumption rates of C. dubia were estimated in situfrom respiratory electron transport system (ETS) activity inLake Rotongaio during summer stratification and winter mixing.Oxygen consumption was 0.002 µl O₂ animal^–1 h^–1in the hypolimnion and 0.076 µl O₂ animal^–1 h^–1in the epilimnion during stratification. Implications of respiredoxygen for metabolic carbon requirements are discussed.     

Relationships between fecal pellet production and feeding in the calanoid copepod Boeckella

The number, size (mm³ pellet ^–1) and total volume of fecalpellets produced (mm³ h^–1) by Boeckella titicacae increasedwith concentration of Chlorella and natural seston in threedifferent lakewaters from Bahia de Puno and Lago Grande of LakeTiticaca and nearby Laguna Arapa. The three lakewaters differedin the size and number of fecal pellets produced at seston concentrations>0.5 mg dry wt l^–1. However, the total volume of fecalpellets egested varied little. Large-sized food (>10 µg)resulted in larger pellets but smaller total fecal production.Food concentration had parallel effects on the production offeces and on feeding rates. Both fecal production and feedingincreased linearly with food concentration with saturation between5 and 7.5 x 10⁵ Chlorella ml^–1 (3.7–5.6 mg dry wtl^–1). Varied responses in fecal production when fed differentsize fractions of lake seston suggest Boeckella has a complexand flexible feeding behavior. Feeding preferences of B.titicacaeand B.occidentalis (Ivlev index) for two seston size fractionsare generally reflected by fecal production.     

Diel variations in gut pigment content, diel vertical migration and estimates of grazing impact for copepods in the southern Benguela upwelling region in October 1987

The gut fluorescence technique was used to estimate ingestionand filtration rates of the adult female copepods Paracalanusparvus, Cenlropages brachiatus and Calanus austrails, and copepoditestages 3, 4 and 5 of C.australis in the southern Benguela upwellingregion. During the study period chlorophyll concentrations withinthe upper 20 m of the water column were high, 5 µg I^–1in mid-shelf waters and 15–30 µg I^–1 in innershelf waters. Copepod gut pigment content was low and constantduring the day then increased sharply during the first 2 h aftersunset. Gut pigment content was 2–6 times higher duringthe night compared with daytime values. Small non-migratingcopepods (Paracalanus parvus) showed the smallest diel differencein gut pigment content and large migrating copepods (Centropagesbrachiatus and Calanus australis) the largest difference. Eggproduction rates were 20 and 50% of maximum at the mid-shelfand inner shelf stations respectively, suggesting food-limitation.Comparison of ingestion rates calculated from egg productiondata with ingestion rates calculated from gut pigment data suggestedthat the copepods were feeding omnivorously at the inner shelfstations but herbivorously at the mid-shelf stations. Assumingthat all of the phytoplankton was available as food, the nearshorecopepod assemblage grazed {small tilde}1% of the standing cropeach day, and the mid-shelf assemblage grazed 5% day^–1.Because of errors and uncertainties associated with the gutfluorescence technique, the feeding impact could be underestimatedby 2–4-fold. We discuss several approaches which couldlead to more precise estimates of feeding rates. ³Present address: Marine Sciences, SUNY, Stony Brook, NY, 11794-5000,USA     

Feeding and growth rates of the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

The goal of this research is to enhance our knowledge of thecontributions of doliolids to the planktonic community as consumersand secondary producers. The objectives are to quantify feedingand growth rates of Dolioletta gegenbauri gonozooids at fourfood concentrations and four temperatures in order to determinetheir impact as grazers throughout the water column. Althoughdoliolids are abundant in numerous regions of the coastal ocean,and are considered to be major planktonic grazers, data on ratesof feeding and growth are scarce. Laboratory experiments wereconducted at 16.5, 20, 23.5 and 26.5°C to quantify removalof a 50:50 volumetric concentration of Thalassiosira weissflogiiand Rhodomonas sp. at four different food concentrations of20, 60, 160 and 390 µg C l^–1. Results from theseexperiments suggest that clearance rates are similar at concentrationsfrom 20 to 60 µg C l ^–1, and decrease as the foodconcentrations increase to 160 and 390 µg C l ^–1.The ingestion rates increase over a range of phytoplankton concentrationsfrom 20 to 160 µg C l ^–1, then decrease when abnormallyhigh concentrations of 390 µg C l ^–1 are offered.Clearance and ingestion rates increase as temperature increasesfrom 16.5 to 26.5°C. The exponential growth rates rangefrom k = 0.2–0.7, with the lowest rates occurring at thehighest food concentration. Growth rates increase with increasingtemperature from K = 0.1–0.3 at 16.5°C to 0.45–0.7at 26.5°C. In each case, the small- and medium-sized zooidshad higher growth rates than the larger gonozooids. These resultssuggest that doliolid feeding and growth rates are a functionof environmental food concentrations and temperatures, and implythat they can be important consumers in a changing neritic environment.     

On the causes of interspecific differences in the growth-irradiance relationship for phytoplankton. Part I. A comparative study of the growth-irradiance relationship of three marine phytoplankton species: Skeletonema costatum, Olisthodiscus luteus and Gonyaulax tamarensis

Three marine phytoplankton species (Skeletonema costatum, Olisthodiscusluteus andGonyaulax tamarensis) were grown in batch culturesat 15°C and a 14:10 L:D cycle at irradiance levels rangingfrom 5 to 450 µEinst m^–2 s^–1. At each irradiance,during exponential growth, concurrent measurements were madeof cell division, carbon-specific growth rate, photosyntheticperformance (both O₂ and POC production), dark respiration,and cellular composition in terms of C, N and chlorophyll a.The results indicate that the three species were similar withrespect to chemical composition, C:N (atomic) = 6.9 ±0.4, photo-synthetic quotient, 1.43 ± 0.09, and photosyntheticefficiency, 2.3 ±0.1 x 10^–3 µmol O₂ (µgChl a)^–1 h^–1 (µEinst m^–2 s^–1)^–1.Differences in maximum growth rate varied as the –0.24power of cell carbon. Differences in growth efficiency, werebest explained by a power function of Chl a:C at µ = 0.Compensation intensities, ranged from 1.1 µEinst m^–2s^–1 for S. costatum to 35 forG. tamarensis and were foundto be a linear function of the maintenance respiration rate.The results indicate that interspecific differences in the µ–Irelationship can be adequately explained in terms of just threeparameters: cell carbon at maximum growth rate, the C:Chl aratio (at the limit as growth approaches zero) and the respirationrate at zero growth rate. A light-limited algal growth modelbased on these results gave an excellent fit to the experimentalµ–I curves and explained 97% of the observed interspecificvariability. ¹Present address: Lamont-Doherty Geological Observatory Columbiaof University, Palisades, NY 10964, USA     

Seasonal patterns of food limitation in Daphnia galeata: separating food quantity and food quality effects

Food limitation for the cladoceran Daphnia galeta was measuredgrowing them with natural lake seston for one growing seasonunder standardized conditions in the laboratory. Growth rateswere related to several measures of food quantity. Particulateorganic carbon (POC) <30 µm was the best predictorof total food availability measured. It was better than chlorophyll-a(Chl-a) of the same size fraction, algal volume calculated fromcell counts or total particle volume from Coulter Counter measurements.Daphnids were also grown with Scenedesmus acutus under the sameconditions and their growth rates were compared. Sestonic carbonconcentrations were related to Scenedesmus carbon concentrations,that provided equal growth rates. The ratio of the carbon concentrationsof both food types (seston, Scenedesmus) represented the foodquality of seston. Thus, the nutritional quality of naturallake seston relative to a standard alga was determined overone growing season. Growth rates of seston fed animals revealedthat they were food limited during long periods of the year.This was also ascertained by standard clutch sizes of fieldanimals. However, the causes of food limitation varied withinthe duration of the studied period. Whereas during the clear-waterphase the food quantity was limiting (while the quality washigh: 100% Scenedesmus), shortly after on 10 June and 17 Junefood conditions were close to threshold concentrations due tolow food quality. For the remainder of the summer, a POC of0.3–0.6 mg C I^–1 with a quality of     

Diet selection by copepods in the presence of cyanobacteria

Boeckella, the dominant calanoid in many Southern Hemispherelakes, can survive, grow and reproduce to varying extents onmonocultures of cyanobacteria. In this study, we determinedthe effects of algal and cyanobacterial foods of different nutritionalvalue and concentration on food preferences of adult femaleBoeckella trianiculata and Boeckella hamata. Four species ofcyanobacteria (Anabaena flos-aquae, Nostoc sp. 2, OscillatoriatenuisandMicrocystis aeruginosa) were offered alone and mixedwith equal biomasses of Cryptomonas sp., Choricystis or a cyanobacterium.Food preferences were calculated as ratios of the rates at whichthe copepods removed each food at high and low food concentrations.In high-concentration mixtures with cyanobacteria, Cryptomonaswas consistently preferred by both Boeckella spp. In low-concentrationmixtures, both Boeckella spp. preferred Anabaena and Nostoc,which they removed at high rates(81–142 ml mg^–1h^–1), although Cryptomonas was selected in preferenceto Oscillatoria and Microcystis. When fed mixtures of filamentouscyanobacteria, both species of Boeckella showed invariant discriminationagainst Nostoc, andshifts in preference between Anabaena andOscillatoria that were related to food concentration. Microcystis,the least favouredfood, appeared to have a toxic effect on B.triarticulata. ¹Present address: Nursing and Midwifery Department, Otago Polytechnic,Forth Street, Dunedin, New Zealand     

Feeding selectivities and food niche separation of Acartia clausi, Penilia avirostris (Crustacea) and Doliolum denticulatum (Thaliacea) in Blanes Bay (Catalan Sea, NW Mediterranean)

Selectivity-size spectra, clearance and ingestion rates andassimilation efficiencies of Acartia clausi (Copepoda), Peniliaavirostris (Cladocera) and Doliolum denticulatum (Doliolida)from Blanes Bay (Catalan Sea, NW Mediterranean) were evaluatedin grazing experiments over a wide range of food concentrations(0.02–8.8 mm³ L^–1 plankton assemblages from BlanesBay, grown in mesocosms at different nutrient levels). Acartiaclausi reached the highest grazing coefficients for large algae>70 µm (longest linear extension), P. avirostris forintermediate food sizes between 15 and 70 µm, and D. denticulatumfor small sizes from 2.5 to 15 µm. Penilia avirostrisand D. denticulatum acted as passive filter-feeders. Acartiaclausi gave some evidence for a supplementary raptorial feedingmode. Effective food concentration (EFC) decreased linearlywith increasing nutrient enrichment for D. denticulatum andfollowed domed curves for A. clausi and for P. avirostris withmaximum values at intermediate and high enrichment levels, respectively.Clearance rates of crustacean species showed curvilinear responseswith narrow modal ranges to increasing food concentration. Clearancerates of D. denticulatum increased abruptly and levelled intoa plateau at low food concentrations. Mean clearance rates were13.9, 25.5 and 64.1 mL ind.^–1 day^–1, respectively.No clearance could be detected for A. clausi at food concentrations<0.1 mm³ L^–1 and for P. avirostris at food concentrations     

Impacts of nutrients and zooplankton on the microbial food web of an ultra-oligotrophic lake

In ultra-oligotrophic lakes and the sea, calanoid copepods arethe dominant mesozoo-plankton and cladocerans are generallysparse or absent. To determine the effects of predation andnutrient enrichment on the pelagic microbial food web of anultra-oligotrophic lake, we added copepods and cladocerans atlow biomasses (<60 µg l^–1 to in situ enclosuresin Lake Wakatipu, New Zealand, in the presence and absence ofadded nutrients (nitrogen and phosphorus). In response to nutrientfertilization, the concentrations of phototrophs >3 µmand heterotrophic bacteria increased by 50 and 15%, respectively,over 4 days, but those of cyanobacterial picoplankton decreasedby 68%. The presence of calanoid copepods (Boeckella dilatata)at ambient densities (1 and 4 l^–1) rapidly and severelysuppressed ciliate population growth over 4 days and also loweredthat of flagellates >3 µm, even when microbial growthwas enhanced by added nutrients. The presence of a small cladoceran,Ceriodaphnia dubia, at double the densities, but similar biomasses,to those of copepods, depressed the net growth rates of ciliatesand flagellates to a lesser degree. The net growth rate of heterotrophicbacteria after 4 days declined with flagellate abundance, consistentwith the possibility of regulation by flagellates. Althoughbacteria and algae increased in response to nutrient fertilization(bottom-up control), predation (top-down control) appeared toplay an important role in structuring the microbial food webof this ultra-oligotrophic lake in summer.     

Growth and production of heterotrophic nanoflagellates in a meso-eutrophic lake

The growth of heterotrophic nanoflagellates (HNF) in mesotrophicLake Constance was measured in situ during a 13 month period.Experiments were conducted with 10 µm pre-filtered lakewater incubated in diffusion chambers at 3 m water depth atthe sampling location for 24 h. Growth rates were calculatedfrom changes in cell numbers occurring during the period ofincubation. Growth rates of all dominant taxa showed pronouncedseasonal variation (–0.13 to 1.76 day^–1 and weregenerally highest in summer at high water temperatures. In situgrowth rates were well below maximum growth rates known forthe respective and similar species from laboratory experiments.While water temperature was a key parameter positively relatedto the growth of all HNF species, the effect of various potentialfood items was taxon specific and less clear. Bacterial abundancewas equally important as temperature for growth in the smallbactenvorous Spumella sp., but was insignificant for growthrates of the larger omnivorous Kathablepharis sp. In Spuniellasp., 84% of the observed seasonal variation of its growth ratecould be explained by temperature and bacterial food supply.Based on these results, a multiple linear regression equationwith temperature and bacterial concentration as dependent variableswas calculated for the growth rate of Spumella. Taxon-specificproduction rates were derived from growth rates and averagebiomass of these two species, and compared to total HNF productionestimated from previously measured community growth rates andbiomass in Lake Constance. Production peaks of Spumella sp.and Kathablepharis sp. alternated seasonally. Total HINF productionranged from –0.01 to 10 mg C m^–3 day^–1. Theaverage seasonal production varied between 1.4 and 33 mg C m^–3day^–1 over 6 consecutive years. These small protozoa thuscontribute a substantial amount to total zooplankton productionin Lake Constance.     

Photosynthesis and primary production of Microcystis aeruginosa Kutz. in Lake Kasumigaura

Seasonal changes in the photosynthesis and primary productionof Microcystis aeruginosa Kütz. were investigated in LakeKasumigaura during 1981–1982. Microcystis always showeda light-saturated photosynthesis-light curve. Both P_max andthe initial slope of the photosynthesis-light curve of Microcystisin early summer were very high, so it was concluded that Microcystisutilized both low and high light intensities efficiently. TheP_max of Microcystis was found to be a function of the watertemperature except in August and September. The linear regressionon the temperature-P_max relationship discontinued at 11°C,where the P_max value dropped; Microcystis did not photosynthesizebelow 4°C. The initial slope of the curve was also descendingbelow 11°C. It is suggested that Microcystis changes itsphysiological properties below 11°C. The highest value ofgross production calculated for M. aeruginosa was 5.4 gC m^–2d^–1 in July; the annual gross production was estimatedto be 300 gC m^–2year^–1 (i.e., 40% of the total primaryproduction in this lake).     

Feeding, growth and food conversion of the marine cladoceran Penilia avirostris

Ingestion and clearance rates, feeding behaviors and life historyvariables of the marine cladoceran Penilia avirostris were evaluatedover a range of food concentrations encountered in nature (0.01–3.0mm³ 1^–1 of Isochrysis galbana). Ingestion rates increasedand clearance rates decreased with increasing food concentration.No maximum feeding thresholds were observed over the range ofalgal concentrations offered. Weight-specific ingestion ratesdecreased with increasing body weight. Feeding behaviors suchas mandibular activity, abdominal claw rejections of cloggedfeeding structures and feeding appendage activity decreasedat a food level of 0.3 mm³ l^–1 of l. galbana. Peniliaavirostris had very poor survivorship at extremely low (0.01mm³ l^–1 and high (3.00 mm³ l^–1) food levels. Mortalitywas hardly affected at food levels of 0.03–1.0 mm³ l^–1Reproduction did not occur at food levels of     

Colony growth and evidence for colony multiplication in Phaeocystis pouchetii (Prymnesiophyceae) isolated from mesocosm blooms

Using well plates of Phaeocystis pouchetii colonies isolatedfrom experimental mesocosms in western Norway, increases incolony size and division were documented. Median longest lineardimensions increased 0–7 µm h^–1; literaturePhaeocystis globosa values are 0.9–4.7 µm h^–1.Ten to twelve percent of colonies divided at rates of 0.21–0.28divisions day^–1. Daughter colonies were 100 µm smallerthan mother colonies. Colonies delayed 3.5–4.9 days tofirst division, compared with literature values of 4–5days for P. globosa. This study provides the first experimentalevidence for colony division of wild P. pouchetii.     

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.     

Laboratory-measured grazing and ingestion rates of the salp, Thalia democratica Forskal, and the doliolid, Dolioletta gegenbauri Uljanin (Tunicata, Thaliacea)

Grazing and ingestion rates of laboratory-born Thalia democraticaaggregates and Dolioletta gegenbauri gonozooids, phorozooidsand oozooids were determined while fed Isochrysis galbana (4–5µm diameter) alone or in combination with Peridinium trochoideum(16–18 µm diameter) at concentrations of 0.15–0.70mm³ x 1^–1. Grazing rates (ml x zooid^–1 x 24 h ^–1)ranged from 10 to 355, and at zooid weights greater than 5 µgcarbon were in order oozooid > gonozooid > aggregate.Grazing rates increased exponentially with increasing zooidweight. Weight-specific grazing rates (ml x µgC^–1x 24 h^–1) were independent of the four-fold initial foodconcentration. Mean weight-specific grazing rates increasedlinearly with increasing zooid weight for the aggregates andoozooids, but gonozooid mean rates were independent of zooidweight. Aggregate and gonozooid ingestion rates (10⁶ µm³x zooid^–1 x 24 h^–1) ranged from 4 to 134 while oozooidrates ranged from 3 to 67. All ingestion rates were independentof the initial food concentration but increased linearly withincreasing zooid weight at similar rates. All mean weight-specificingestion rates (ml x µgC^–1 x 24 h^–1) wereindependent of zooid weight. The mean aggregate daily ration(µgC ingested x µg body C^–1) was 59% and themean doliolid ration was 132%. Field studies indicate that normalconcentrations of D. gegenbauri in the Georgia Bight clear theirresident water volume (1 m³) in about 4 months, but that highlyconcentrated, swarm populations which occur along thermohalinefronts clear their resident water volume in less than 1 day. ¹Current address: MacLaren Plansearch Ltd., P.O.Box 13250, sta.A.,St.John's, Nfld. A1B 4A5