Subsurface chlorophyll maxima and vertical distribution of zooplankton in the Gulf of Maine

The vertical disthbution of chlorophyll, zooplankton and physicalstructure were measured using a pumping system and CTD on twocruises in the Gulf of Maine during June and September 1982.The vertical distribution of chlorophyll was closely relatedto the density structure of the water column. In waters witha pronounced pycnocline subsurface chiorphyll maxima (SCM) werelocated at or just above the pycnocline. Chlorophyll concentrationswere maximal in the surface waters at those stations sampledin June where the pycnocline was not well defined. The relationshipbetween the zooplankton and chlorophyll distribution differedbetween cruises. In June, the zooplankton, particularly post-naupliarcopepods, were associated with the depth of the chlorophyllmaxirnum, while in September the post-naupliar copepods weremost abundant in the surface waters above the SCM at the stratifiedstations. During the September cruise we observed that the copepodnauplii were most abundant at the depth of the SCM, and thatthe larger protozoans (>35 µm) were most abundant atdepths of 55–85 m, which were well below the SCM and pycnocline. ^*Bigelow Laboratory for Ocean Sciences Contribution No. 83025     

Influence of the pycnocline on the vertical migration of chaetognaths in the northern Benguela

The vertical distribution and migration patterns of chaetognathswere studied at a sampling station off northern Namibia (18°00'S,10°30'E during a 48 h sampling cycle. The sampling areawas characterized by mixing of the Angola current with the northernmostwaters of the Benguela current in the surface region. The continuousflow of the Angola current during the study period gave riseto a thermocline and a halocline at a depth of 40 m. The samplingintervals used to study the behaviour of epipelagic chaetognathspecies under these conditions were 200–100, 100–60,60–40, 40–20 and 20–0 m. A total of 10 differentchaetognath species were captured. Sagitia enflata was the predominantspecies, with a mean density of 4400 individuals per 1000 m³in daytime hauls, accounting for 54.1% of all individuals collectedin the samples. Sagitia minima was the next most predominantspecies, with a mean density of 2400 individuals per 1000 m³in daytime hauls, accounting for 16.6% of the chaetognath communitysampled. Three maturity stages were considered in analysingpossible ontogenetic migrations. Nearly all the species wereaggregated above the pycnocline, and stages I and II of manyspecies carried out short migrations in the surface layers.The limited migration pattern, together with a staggered distributionof the different species and stages above the pycnocline, hasbeen interpreted as a space partitioning mechanism to preventintra- and interspecific competition.     

Remote detection of algae by copepods: responses to algal size, odors and motility

Selective feeding on large algae by copepods involves remotedetection of individual particles and subsequent active captureresponses In this study we use radiotracer experiments to quantifythe clearance rates of five coexisting freshwater copepods andto investigate the relative merits of the chemoreception andmechanoreception hypotheses of remote detection Tropocyclopsand three diaptomid copepods exhibited relatively high clearancerates when feeding on low concentrations of large algae, suggestingthat most previous studies with freshwater copepods have underestimatedmaximal clearance rates and the degree of size selectivity.All five species of copepods exihibited strong selection foran intermediate-sized flagellate (25 µm Cartena) or alarge-sized nonmotile alga (80 µ.m Pediastrum) over asmall-sized flagellate (6 µ.m Chlamydomonas). The weight-specificclearance rate for Tropocylops prasmus feeding on motile Cartena(271 ml mg^–1 h^–1) was about twice that of threediaptomid copepods and more than an order-of-magnitude higherthan the estimate for Epischura lacustris feeding on its preferredalga, Pediastrum Assuming that distance chemoreception (‘smell’)is important in remote detection, we predicted that the additionof high concentrations of ‘algal odors’ would obscureany chemical gradients emanating from individual algal cellsand would thereby hinder the remote detection and active captureof large algae Contrary to this hypothesis, the addition ofamino acids, sucrose, and algal extracts had no effect on theclearance rates and selectivity of Diaptomus birgei. These results,together with recent cinematographic studies (Vanderploeg etal.,1990), suggest that mechanoreception is the primary mechanismfor the remote detection of large particles by diaptomid copepods.A raptorial cyclopoid, Tropocyclops prasinus, exhibited strongpreferences for motile algae, whereas a suspension-feeding calanoid,D birgei, did not select between motile and nonmotile cells.Motility appears to be an important factor in algal detectionfor small cyclopoid copepods but not for suspension-feedingdiaptomids     

Environmental factors influencing diurnal distribution of zooplankton and ichthyoplankton

The diurnal vertical distribution of a large number of speciesof zooplankton, icbthyoplankton and micronekton were determinedin the top 150 m in three locations in the Shelf Water, on theNova Scotia Shelf, and Slope and on Georges Bank during springand fall periods. Species were categorized as to their trophiclevel and their type of diurnal migration behaviour. The influenceof temperature, salinity, and water density on the diurnal verticaldistribution of the species was examined. Temperature was foundto have the greatest influence on the distribution of the largestnumber of species. Diurnal migration behavior of the same speciesin Shelf and Slope water and at different times of the yearwas examined. Results showed that species changed their behaviorin the two water masses, while some species changed their migrationbehavior at different times of the year. During the night inApril the most abundant copepod species, Calanus finmarchicus,making up about 80% of the biomass, was found concentrated abovethe thermocline and the main chlorophyll layer. The majorityof the less abundant species of copepods were found below thethermocline and the chlorophyll layer. At night in August thetwo most abundant copepod species, Centropoger typicus and Paracalanusparvus, making up at least 80% of the zooplankton biomass, werealso concentrated above the thermocline and the main chlorophyllLayer. Three species of copepods were concentrated at the depthof the main chlorophyll layer and two species were concentratedbelow the chlorophyll layer and thermocline. The vertical distributionof other zooplankton and ichthyoplankton species was examinedin relation to the thermocline and chlorophyll layer. Relationshipsbetween concentrations of six species of fish larvae and allspecies of copepods in the same samples showed a general increasein the numbers of larvae m^–3 as the numbers of copepodsm^–3 increased in a range of 500–4000 m^–3.However, the concentration of Merluccius bilinearis decreasedas the concentration of copepods exceeded 4000 m^–3 suggestingthat high concentrations of copepods may not be a favourableenvironment for the larvae.     

Selective predation by herring and mysids, and zooplankton community structure in a Baltic Sea coastal area

In lakes, fish and invertebrate predation are recognized asstrong structuring forces on zooplankton communities. The objectof this investigation was to study whether predation has a similarpotential in a coastal area of the brackish Baltic Sea and ifit could explain observed differences in zooplankton communitystructure between a reference area and an eutrophied area. Speciescomposition and daily vertical migration of zooplankton andzooplanktivores, and the diets of the latter, were studied inJuly and August at two 30 m deep stations differing in primaryproductivity. The biomass of zooplankton >35 µm wasdominated by copepods, but cladocerans and rotifers also occurredin significant numbers. The dominating zooplanktivores wereherring (Clupea harengus) and the mysid shrimp Mysis mixta.They fed almost exclusively on zooplankton, mainly copepods,and their estimated food consumption equalled or exceeded thesummer copepodite production. The structure of the zooplanktoncommunity cannot be explained by effects of predation or byfeeding conditions alone. Increases in rotifer and cladoceranabundances with increased primary production suggests effectsof food supply. However, a generally rapid decline in the annualsummer peak of cladocerans may be caused by predation. The totalabundance of copepods did not increase with improved feedingconditions, but there was a shift in species dominance. Thecopepod Ewytemora affinis hirundoides, which was intensivelypreyed upon, increased with increased phytoplankton production,while Acartia bifilosa and/or A.longiremis, which was less eaten,decreased. Predation may explain a pronounced daily verticalmigration of the most predated copepods. They occurred in deeperwater during the day, when the visually feeding herring wereactive, and moved closer to the surface at night when M.mixtaleft the bottom, to forage in the water column.     

Vertical migrations of herring,Clupea harengus,larvae in relation to light and prey distribution

Synopsis The influence of light and prey abundance on the vertical distribution of herring larvae was evaluated by three investigations made under calm weather conditions in the North Sea off the Scottish coast. The investigations took place at different time after hatching and the vertical distributions of three size groups of larvae (mean sizes 8,15 and 19 mm) were related to time of day and the vertical distribution of copepods. No migratory behaviour of copepods was observed but their vertical distribution differed between investigations. In the investigation on intermediate sized larvae, copepod density peaked at the pycnocline (40 m). Larvae concentrated at this depth at noon. At dawn and dusk larvae migrated towards the surface and the vertical distributions fluctuated semidielly. In the two other investigations, copepods were homogeneously distributed in the water column and after migration towards the surface at dawn larvae stayed in the upper water column during the day. The observations suggest that the daytime vertical distribution of larvae in calm weather is mainly determined by feeding conditions: the larvae move to depths were light is sufficient for feeding, and refinement within that zone is made according to a compromise between optimal light conditions for feeding and optimal prey densities.     

Short-term changes in the mesozooplanktonic community in theSeine ROFI (Region of Freshwater Influence)(eastern English Channel)

Discrete-depth, hourly mesozooplankton samples were collectedover a 92 h period in May 1992 at an anchor station within theSeine Region of Freshwater Influence (ROFI) (English Channel).The mesozooplankton community defined as a euryhaline marineassemblage was dominated by the calanoid copepods Acartia spp.,Temora longicornis and Centropages hamatus, the cladoceran Evadnenordmanni and the appendicularian Oikopleura dioica. The semi-diurnaltidal current was the dominant factor in determining the short-termtemporal changes in the community in terms of density and speciescomposition so that zooplankton patches displayed oscillatingmotion in relation to tidal advection. Although a few species(e.g. Pleurobrachia pileus) exhibited higher densities aroundlow tide, maximum densities were observed for most species (e.g.T.longicornis and E.nordmanni) around high tide, according tosalinity variations. Diurnal changes were only reported forcyclopoid copepods (i.e. Halicyclops sp. and Cyclopina sp.)which wer$$$ mainly endobenthic during the day and moved intothe water column at night. Besides temporal changes in depth-averageddensities, most species exhibited vertical migrations at dieland/or tidal periods. Tidal vertical migrations were reportedonly for a few taxa and could be the result of passive mechanisms(e.g. vertical mixing) rather than of active behaviour. Dielvertical migrations were observed in most of the abundant taxa.While this migration pattern did not appear to be an adaptationto predator avoidance within the Seine ROFI, it could regulatehorizontal transport of organisms and promote their retention.The consequences of the short-term mesozooplankton fluctuationsfor sampling designs are discussed.     

Vertical structure of small metazoan plankton, especially noncalanoid copepods. I. Deep Arabian Sea

The abundance and vertical distribution of micro-metazoans sampledwith fine nets of 0.05 mm mesh size were studied at three stationsin the Arabian Sea during the intermonsoon period (April/May1987) and down to 1850 m depth. In the epipelagic zone (0–100m). values of biomass and metazoan abundance tended to be higherthan those reported for other tropical oceanic areas. In themesopelagic zone, which is characterized by an extreme oxygendeficiency between 100 and 1000 m depth, the abundance of metazoantaxa and species numbers of non-calanoid copepods were largelyreduced. However, intermediate abundance maxima occurred withinthis zone, which were dominated by specific metazoan taxa (copepods.appendicularians) and species of non-calanoids (Oncaea sp. C).In the bathypelagic zone below 1050 m, the species diversityof the dominant copepod family Oncaeidae increased substantially.Two-thirds of a total of 69 oncaeid species recovered were confinedto this layer. As most of them were small in size and occurredin low abundance only, the increase in total oncaeid densityand/or plankton biomass was less conspicuous. Dominant Oncaeaspecies in the bathypelagic zone were O.longipes and O.brodskii.The results are compared with published data from the ArabianSea and other tropical oceanic areas with and without an extrememesopelagic oxygen minimum zone Possible causes of the intermediateabundance maxima within the oxygen deficiency zone are discussed.     

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.     

Neocalanus scattering layers near the western Aleutian Islands

Dense copepod scattering layers were detected in the upper 50m of the water column in the Bering Sea north of the westernAleutian Islands during June 1992. Acoustic data were collectedat frequencies of 200 and 420 kHz, and compared with simultaneousestimates of wet weight biomass from net tows. Regression ofnet and acousticdata produced a scaling factor of –69.6dB g^–1 to convert volume scattering to biomass. Applicationof the fluid sphere scattering model produced an average equivalentspherical radius for the targets of 0.54 mm. The major sound-scatteringorganism, Neocalanus cristatus copepodid stage V, dominatedthe biomass in the pycnocline between the upper mixed layerand the underlying cold intermediate layer. Populations weredominated by Neocalanus plumchrus and Neocalanus flemingeri,withgreatest abundance above the pycnocline.     

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.     

Response of Gonyaulax tamarensis to the presence of a pycnocline in an artificial water column

In nature, large concentrations of the toxic bloom-forming dinoflagellate,Gonyaulax tamarensis, are frequently observed in the vicinityof the pycnocline. In the absence of a pycnocline the organismis usually recorded near the surface, where light levels aremore advantageous for photosynthesis. In this paper we examinethe swimming behaviour of G.tamarensis when exposed to varyingdegrees of stratification and investigate whether the maintenanceof a subsurface (pycnocline) population is the result of retentionof the algae by a physical barrier or active accumulation ofthe organisms at a density interface. The study indicates thatG.tamarensis cells presented with a halocline of S<{smalltilde}6–7 (occurring over a few centimeters) cross thissalinity barrier and accumulate at the highest available photonflux density ({small tilde}100 µmol m^–2 s^–1).Cells exposed to a gradient of S>{small tilde}7remain atthe halocline (pfd={small tilde}40 µmol m^–2 s^–1).However, when light above the pycnocline is attenuated by theaddition of food colour to the medium, the cells cross a haloclineof S=10 and accumulate at the highest available photon fluxdensity. In the absence of added nutrients (inorganic N andP) the organism fails to exhibit a phototactic response. Thus,the presence of a strong halocline does not represent an inpenetrablephysical barrier for G.tamarensis and the development of pycnoclinepopulations of this organism is a function of density, lightand nutrient climate.     

Weak 'top-down' control of dinoflagellate growth in the coastal Skagerrak

Effects of planktivorous fish (Sprattus sprartus), copepods(mainly Oithona sp.) and benthic filter feeders (Mytilus edulis)on the biomass and species composition of a natural marine planktoncommunity, dominated by dinoflagellates, was studied experimentallyin September 1988. Plastic 300 I cylinders were filled withSkagerrak surface water, filtered through a 90 µm nylonnet in order to remove mesozooplankton. Copepods were addedin three different concentrations (0, 1x and 6x the ambientdensity) with and without nutrient additions (nitrate, silicateand phosphate). In two nutrient-enriched cylinders with addedcopepods (1x and 6x), 5 cm sprats were also added (10 per cylinder).In two nutrient-enriched cylinders without added copepods. bluemussels (20 per cylinder. 3 cm length) were introduced. Chlorophylla. inorganic nutrients and densities of phytoplankton, ciliatesand copepods were measured throughout the experimental periodof 10 days. Gyrodinium aureolum increased in cell numbers inall but the mussel cylinders, while the other dominating dinoflagellate,Ceratium furca, decreased. This decrease was not due to grazing,but probably a container artefact. Diatoms, Rhizosolenia fragilissinia,Leptocylindrus danicus and Chaetoceros spp., increased slowlyin treatments without fish. In the fish treatments, diatomshad appreciably higher growth rates and cell numbers, probablydue to predation by fish on the copepods. Gyrodiniun aureolunalso had a somewhat higher growth rate in the presence of fish.although this was probably not due to reduction of copepod grazing.The results show that planktivorous fish can affect the biomassand species composition of natural phytoplankton communitiesin marine waters, but that the effect on the late summer-autumndinoflagellate dominated community is weak. Blue mussels areefficient grazers of the whole phytoplankton community and arenot particularly selective with respect to either size or species.     

Chemical, mechanical and visual cues in the vertical migration behavior of the marine planktonic copepod Acartia hudsonica

Recent experimental evidence in both marine and freshwater systemsindicates that predators can induce vertical migration behaviorin individual zooplankters, yet the specific cues by which zooplankterssense their predators appear to vary. In situ manipulation experimentswere carried out with enclosed populations of the marine planktoniccopepod Acartia hudsonica to re-examine the potential role ofchemical cues in the behavior of A.hudsonica, and to test explicitlyfor the role of mechanical or visual stimuli in triggering verticalmigration behavior in this species. Adult female copepods wereinduced to vertically migrate (descend) when exposed to fishmimics during the day, but no such response occurred when thecopepods were exposed to Fish mimics during the night. Moreover,copepods exhibited no changes in vertical distribution whenexposed to water which, having recently held a natural predator(the threespine stickleback, Gasterosteus aculeatus), was presumedto be laden with predator-produced chemical exudates. Predator-mediatedmechanical or visual cues, or a hierarchy of both, are responsiblefor eliciting vertical migration behavior in adult female A.hudsonica.These results, together with those of other investigations demonstratingthe inducing role of chemical exudates, indicate that the stimulieliciting vertical migration in zooplankton can be expectedto vary between species.     

Experimental development of a model for the feeding of neritic copepods on phytoplankton

Grazing of female Paracalanus parvus and Acartia lonsa on mixturesof phytoplankton was explainable by a model involving only ageneral filtration rate and an efficiency of retention whichis itself a function of cellular size but is not easily relatedto the setular spacing on the maxillae. Acartia was less efficientat capturing very small cells than was Paracalanus. A sensitivityanalysis was used to determine the relative importance of thesize distribution of food, the age distribution of the copepods,and the vertical distributions of food and copepods in estimatingthe feeding of a natural population.     

Predator-induced diet vertical migration in a planktonic copepod

Predator evasion is the most commonly hypothesized reason fordiel vertical migrations undertaken by a wide variety of planktonicorganisms in lakes and seas, yet direct evidence remains elusive.We tested the predation hypothesis by exposing enclosed populationsof a marine copepod Acartia hudsonica to caged or free-rangingindividuals of their natural predator, the planktivorous fishGasterosteus aculeatus. After little more than a week, adultcopepods changed their vertical distribution and migration behaviordepending on the presence or absence of predation. Only free-rangingfish induced vertical migration in the copepod population. Cagedfish had no effect, indicating that vertical migration was nota simple chemically mediated response of copepods to the predator.Rather, copepods seemed to react to the presence of predatorsby other means, perhaps visual or mechanical stimuli, and toexhibit a downward escape response which, because encounterswith visually orienting fish occur chiefly in the daytime, effectivelylimited the copepods' occurrence in the upper water column tothe night-time hours. Alternatively, because fish imposed heavymortality on copepods, it is possible that selective predationaltered the proportions of individuals with fixed, geneticallydetermined migration behaviors. We suggest experiments to distinguishthese alternatives.     

Near-bottom depletion of zooplankton over a coral reef II: relationships with zooplankton swimming ability

Intense predation by corals and associated fauna can generate vertical gradients of their zooplankton prey. The goal of this study was to characterize the small-scale distribution of zooplankton above the coral reef of Eilat, Israel. Four vertical arrays, each consisting five underwater pumps attached 0.5–3 m apart on a taut mooring, were deployed for 10 days at two sites over the reef slope. A distinct layer of depleted zooplankton was repeatedly found in the 1.5 m high benthic boundary layer. The gradient was sharpest for strong swimmers (copepods and polychaeta), intermediate for weak swimmers (nauplii, mollusks and appendicularia), and lacking for passive taxa (eggs and foraminifera). Spatio-temporal changes in the abundance of copepods and polychaetes were highly correlated, more so in the water aloft (Pearson r>0.9) than near the bottom (r>0.8). The spatio-temporal correlations between weak swimmers and between passive taxa were much weaker (r<0.8 and r<0.4, respectively). Flow and shear stress did not affect the distribution patterns. The correspondence between the zooplankton distribution and their swimming ability indicates that bottom avoidance contribute to the formation of depleted layer over the reef.     

Spatial and temporal patterns of zooplankton on baleen whale feeding grounds in the southern Gulf of Maine

This study addresses the regional variation of zooplankton inthe Great South Channel area in the southern Gulf of Maine betweenCape Cod, Massachusetts and Georges Bank. This is a region ofparticular interest because of the intense concentrations ofthe copepod Calanus finmarchicus in the spring, along with theco-occurrence of right whales that feed upon these copepod aggregations.Zooplankton in the Great South Channel were sampled with theMOCNESS plankton sampler during spring 1988 and 1989 as partof the SCOPEX (South Channel Ocean Productivity Experiment)project. Zooplankton variation was addressed through comparisonsof taxonomic composition and water column abundances among towswithin and between years, and between locations with and withoutright whales. Results showed that zooplankton community compositionwas highly similar between tows within each year average percentsimilarity index (PSI) for pairwise comparisons of tows = 82.2and 88.8% in 1988 and 1989, respectively] as well as betweenyears (mean PSI = 84.4). The copepods C.finmarchicus and Pseudocalanusspp. dominated the zooplankton in terms of total water columnabundance (>94% of all zooplankton), with C.finmarchicuscomprising an average of 84% of the copepods. Highest abundancesof these copepods (particularly the younger life stages) coincidedwith a region of low-salinity transport from the north. In addition,these copepods had higher abundances in 1989, which may be relatedto the fact that low-salinity transport was approximately twiceas large in 1989 as in 1988. Given the physical dynamics ofthe region, it is possible that developing copepod populationswere adverted into the Great South Channel from the northwesternGulf of Maine via the low-salinity plume. Each year, whaleswere located in areas of both high and low copepod abundance,and tended to concentrate near the leading edge of the low-salinityplume. In 1988, there were no significant differences in zooplanktonabundance between right whale areas and non-whale areas forany taxon. In 1989, whale areas had a greater proportion ofC.finmarchicus relative to other zooplankton in 1989, whichsuggests that whales preferred regions enriched in C.finmarchicus.     

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     

The effect of ultraviolet radiation on the vertical distribution and mortality of estuarine zooplankton

The effect of UV-B radiation on the vertical distribution ofthree calanoid copepod species (Tortanus dextrilobatus, Acartiuraspp. and Acanthacartia spp.) and three larval stages of Pacificherring, Clupea pallasi (1-, 7-, and 14-day-old larvae) wasinvestigated. A series of 2 m high columnar tanks equipped withinfra-red light and video-microscopy was used to investigatethe vertical distribution of zooplankton in the presence andabsence of UV-B radiation. In the presence of UV-B radiation,T.dextrilobatus and 1-day-old C.pallasi resided about 50 cmdeeper than in the absence of UV-B, while Acartiura spp. andAcanthacartia spp. showed no (or only minimal) change in verticaldistribution. Mortality experiments were also conducted outdoorsin which each copepod species was exposed to full or reducednatural radiation levels. Only T.dextrilobatus showed an increasein mortality when exposed to full radiation. Our results showedthat T.dextrilobatus and 1-day-old C.pallasi larvae were sensitiveto UV radiation (UVR), and to reduce or eliminate UV-inducedstress, they avoided the surface of the water column when UV-Bradiation was present. Copepod species were chosen to span arange of pigmentation: T.dextrilobatus (heavily pigmented),Acartiura spp. (moderately pigmented) and Acanthacartia spp.(not pigmented). The pigmentation did not appear to play a rolein UVR tolerance of the copepods, but may be a factor determiningUV tolerance of C.pallasi.