Contribution of nanoprotists to metazooplankton diet in a mesocosm experiment in the coastal northern Baltic

Grazing by a metazooplankton community on nanoprotists <10µm was studied four times during a 21 day enclosure experimentcarried out off the SW coast of Finland in late summer. Duringthe study, the pelagic community was manipulated through nutrientenrichment (N + P) and through predation by stickleback fry.Grazing experiments were conducted in the laboratory using 5µm prefiltered, ³H-labelled nanoplankton as tracer food. Grazing by mesozooplankton (Meso) and metazaan microplankton(Micro), screened through 140 and 100 µm mesh, was studied.Owing to enrichment effects and weak predation control by fish,the biomass of Meso and Micro increased during the study peridMeso biomass consisted mostly of the copepod Eurytemora affinisand the cladoceran Bosmina longispims maritima, and Micro biomassof copepod nauplii NIII–NVI. The community clearance rateof Meso usually exceeded that of Micro when feeding on nanoprotistsThe opposite was found for the biomass-smc clearance rate, revealingnanoprotists to be a more important f d source for Micro thanfor Meso. Metazmplankton were not able to control nanoprotists,because Meso and Micro were estimated to remove on average 4and 2% of nanoprotowan biomass daily. When integrated throughthe study period, grazing on nanoprotists could meet 5 and 17%of the carbon need for Meso and Micro, 3 and 12% of their productionbeing estimated to consist of bacterial carbon transferred bynanoprotists Micro were estimated to be more closely Linkedto the microbial food web than Meso, suggesting that the trophicpusition of copepods changes slightly during their maturationfrom nauphi to larger copepodites. ¹ Present address: Department of Ecology and Systematics, Divkionof Hydrobiology, University of Helsinki, FIN-00014, Helsinki,Finland     

Omnivorousness in marine planktonic copepods

Adult females of Centropages furcatus and Temora stylifera hadsignificantly different feeding behaviours. At equal averagefood concentrations, C. furcatus ingested more nauplii of thecalanoid copepod Pseudodiaptomus coronatus and fewer diatomsthan T. stylifera. At 20°C and 2.0 µg of nitrogen.1.1.^–1of each food source, C. furcatus ingested 17.2% and 4.4% ofits body nitrogen daily as nauplii and diatoms, respectively.Ingestion rates of the same food sources by T. stylifera were5.1% and 14.5%, respectively. At the environmental concentrationssimulated in these experiments, the presence of diatoms didnot affect the ingestion of nauplii nor did the presence ofnauplii affect diatom ingestion. The behaviour of predator andanimal prey largely dictates the types and quantities of preyingested.     

The effect of container size on the feeding rate of Heterocope septentrionalis, a freshwater predaceous copepod

The feeding rates of freshwater predaceous copepods have beenmeasured many times using enclosure experiments. Typically theseexperiments involve enclosing a known number of zooplanklonprey with a known number of predaceous copepods and determiningthe number of prey remaining after a set length of time. Variousfactors such as prey species and size have been shown to influencethe feeding rate of copepods. However, little attention hasbeen paid to the influence that container size may have on feedingrates. Using previously published data and some new data itwas found that container size has a major impact on the feedingrate of Heterocope septentrionalis, a predaceous freshwatercopepod common in North American arctic waters. In experimentswith Daphnia puiex, where container size varied from 0.3 to54 l (180 times), the measured feeding rate coefficient variedfrom –0.04 to –1.7 (42.5 times). Other prey speciesshowed similar changes, but the vulnerability of particularprey species to Heterocope predation remained consistent acrossthis range of expenmental containers. The reason for the containereffect is not known, but it is postulated that edge avoidanceby Heterocope may account for the observed change in feedingrate.     

Diel patterns of zooplankton grazing in a shallow lake

A diel survey of in situ species-specific zooplankton clearancerates (with radioactively labelled Chlorella cells) was conductedin the shallow lake of Cr?teil characterized by small-sizedplanktonic forms (algae<10µm and zooplankton <1.3mm). Experiments were performed every 4h at two depths (1 and4m). Power functions relating individual filtering rates tobody length were established for the three most abundant cladoceransand for calanoids (nauplii being included in this feeding group),for each depth and time. No filtering penodicity was observedin Ceriodaphnia spp., adults and copepodites of Eudiaptomusgracilis and Eurytemora velox, and nauplii. On the contrary,clear nocturnal filtering peaks were obtained for Daphnia spp.and for Diaphanosoma brachyurum, these being more pronouncedfor the larger individuals at 1 m deep. The observed diel periodicitycannot be explained by variations in physico-chemical parametersor food concentration alone. The ecological significance ofthis phenomenon in the polymictic lake of Cr?teil is discussedin the light of previously published data and the hypothesesaccounting for it.     

Feeding efficiency of the larval ctenophore Mnemiopsis leidyi A. Agassiz (Ctenophora, Lobata)

Larval stages of the ctenophore Mnemiopsis leidyi rely on metazoanprey, such as Acartia tonsa nauplii and copepodites, to supporthigh growth rates. However, M. leidyi larvae <0.5 mm (totallength) had low retention efficiencies (REs) (proportion ofencountered prey actually ingested), 5.78 ± 2.6% (mean± SE), of nauplii and were often damaged by their encounters.REs of nauplii rapidly increased, 38.94 ± 3.73%, as larvaegrew to a size of     

Vulnerability of Daphnia middendorffiana to Parabroteas sarsi predation: the role of the tail spine

The predaceous calanoid copepod Parabroteas sarsi and Daphniamiddendorffiana co-exist in South Andes ponds. Daphnia middendorffianajuveniles have a tail spine with negative allometric development.A series ofexperiments was carried out with tailed and tail-removedjuveniles of three different instars. In all cases tested, feedingrates were significantly higher on tail-removed prey. Directobservations showed a higher frequency of unsuccessful attackson spined juveniles when compared with tail-removed juveniles.The proportion of dorsal attacks also increased in spined juveniles.Prey total length was a much better predictor of feeding ratethan prey body size. Three groups of juveniles with equal totallength, but differentage, body size and biomass, showed no significantdifferences in their vulnerability to predation.     

Effect of prey size on vulnerability of copepods to predation by the scyphomedusae Aurelia aurita and Cyanea sp.

A growing number of studies correlate changes in zooplanktonpopulations with abundance of medusae, but we cannot yet explainor predict the specific factors driving these interactions.This study demonstrates that the size of copepods has a significantinfluence on their vulnerability to predation by scyphomedusae.This finding is important because prey size, independent ofbehavior, has been neglected in theoretical models of predationby medusae. In experiments in a planktonkreisel, we used liveand heat-killed prey (Acartia hudsonica adults and copepodites)to separate the effects of copepod size and behavior on feedingrates by two medusae (Aurelia aurita and Cyanea sp.). Resultsrevealed that: differences in copepod size had a significantimpact on feeding rates, and thus small size can provide a refugefrom predation; behavior of adults diminished the liabilityassociated with larger size; and medusae with different morphologiesingested A.hudsonica at similar rates. Other experiments demonstratedthat medusae digested copepods at different rates based on preysize and predator species, findings that have implications forall future laboratory and field studies that assess feedingby scyphomedusae. Finally, this study illustrates how laboratorystudies serve as critical supplements to field observations.The effect of prey size on feeding rates can be confounded bydifferences in prey behavior, yet explains why small copepodswere typically ingested at relatively low rates by medusae.Size was clearly a dominant factor influencing copepod vulnerabilityacross scyphomedusan species, even those with very differentmorphologies. Future work should focus on the mechanisms ofsize selection, or the factors influencing contact and retentionrates.     

Predation and respiration by the small cyclopoid copepod Oithona similisr: How important is feeding on ciliates and heterotrophic flagellates?

Feeding on natural plankton populations and respiration of thesmall cyclopoid copepod Oithona similis were measured duringthe warm season in Buzzards Bay, Massachusetts, USA. AlthoughO.similis did not significantly ingest small autotrophic andheterotrophic flagellates (2–8 µn), this copepodactively fed on >10 µm particles, including autotrophic/heterotrophic(dino)flagel-lates and ciliates, with clearance rates of 0.03–0.38ml animal^–1 h^–1. The clearance rates increased withthe prey size. O.similis also fed on copepod nauplii (mainlycomposed of the N1 stage of Acartia tonsa with a clearance rateof 0.16 ml animal^–1 h^–1. Daily carbon ration fromthe combination of these food items averaged 148 ng C animal^–1day^–1 (41% of body C day^–1), with ciliates and heterotrophicdino-flagellates being the main food source ({small tilde}69%of total carbon ration). Respiration rates were 020–0.23µl O₂ animal^–1 day^–1. Assuming a respiratoryquotient of 0.8 and digestion efficiency of 0.7, the carbonrequirement for respiration was calculated to be 125–143ng C animal^–1 day^–1, close to the daily carbon rationestimated above. We conclude that predation on ciliates andheterotrophic dinoflagellates was important for O.similis tosustain its population in our study area during the warm season.     

A method for in situ estimation of prey selectivity and predation rate in large plankton, exemplified with the jellyfish Aurelia aurita (L.)

A method to estimate predation rates of large predatory zooplankton, such as jellyfish and ctenophores, is outlined. Large plankton size allows direct visual tracking of the predator during the process of foraging. The presented method is novel in the sense that it measures predation rate of a specific individual plankton predator in situ.After prey has been evacuated from the gut of an individual predator, the predator is incubated in situ, and observed by SCUBA-divers who recapture the individual after a defined time. Given that this incubation time is shorter than prey digestion time, predation rate can be calculated as increase in gut content over time. Clearance rates for different prey can be calculated from predation rates and prey concentrations in the water, allowing accurate estimates of prey selectivity. Thus, the problem of unknown feeding history and feeding environment, which can otherwise be a problem in prey selectivity studies of in situ-captured predators, is circumvented. Benefits and limitations of the method are discussed.The method was applied to adult medusae of the common jellyfish Aurelia aurita. A large variation in number of captured prey was detected both among individual jellyfish and among the various oral arms and gastric pouches within individuals. Clearance rates varied strongly with prey type. The medusae selected large crustacean prey (cladocerans and copepods/copepodites) over echinoderm larvae and copepod nauplii. Prey distribution within the medusae indicates that both tentacles and oral arms were used as prey capturing sites. Food passage time from prey capturing organs to gastric pouches was estimated.     

Implications of the feeding current structure of Euchaeta rimana, a carnivorous pelagic copepod, on the spatial orientation of their prey

Many marine planktonic organisms create water currents to entrainand capture food items. Rheotactic prey entrained within thesefeeding currents often exibit escape reactions. If the directionof escape is away from the feeding current, the prey may successfullydeter predation. If the escape is towards the center of thefeeding current, the prey will be re-entrained towards its predatorand remain at risk of predation. The direction of escape isdependent on (i) the ability of the prey to escape in a directiondifferent than its pre-escape orientation and (ii) the orientationcaused by the interaction of the prey's body with the movingfluid. In this study, the change in orientation of Acartia hudsonicanauplii as a result of entrainment within the feeding currentof Euchaeta rimana, a planktonic predatory copepod, was examined,When escaping in still water, A.hudsonica nauplii were ableto vary their pre-escape direction by only 10. This allowsonly a limited ability to escape in a direction different thantheir pre-escape orientation. Analyses of the feeding currentof E.rimana show the flow speed to be most rapid in the centralregion with an exponential decrease in speed distally. In contrast,flow vorticity is minimal in the center of the feeding currentand maximal at 1.75 mm along the antennae. As a result, thedegree of rotation of the prey towards the center of the feedingcurrent shows a strong dependency on the prey's location withinthe feeding current. The feeding current of E.rimana rotatedthe prey 14 when near the center of the flow field and up to160 when located more distal in the feeding current Since theprey's escape abilities cannot compensate for the rotation dueto the flow, this mechanism will maintain the escaping preywithin the feeding current of their predator. Therefore, thefeeding current facilitates predatory copepods in capturingprey by (i) increasing the amount of water which passes overtheir sensors and through their feeding appendages and (ii)controlling the spatial orientation of their prey prior to escape.     

In situ predatory behavior ofMysis relicta in Lake Michigan

Selectivity coefficients (W) and predation rates on Lake Michigan zooplankton were determined forMysis relicta during spring through fall using anin situ method. W values indicated the following ranked order of prey preference: Cladocera > copepod copepodites and copepod nauplii > adult diaptomids and cyclopoids. With few exceptions, W values for different prey categories remained fairly constant despite greatly changing relative abundances of prey. Predation rates and prey selectivity were similar in most cases to those determined in laboratory studies. Ingestion rates (percent dry body weight · day^–1) were correlated to total prey biomass (r = 0.38) and to effective prey biomass (r = 0.85), where the weighting factors were overall mean selectivity coefficients for the different prey categories. This result suggested that seasonally varying composition of prey caused much of the variation in ingestion rates among experiments. Feeding trials performed at the same depth with daytime and nighttime assemblages of zooplankton indicated that Cladocera may escape heavyMysis predation at night by migrating from the metalimnetic-hypolimnetic interface into the epilimnion.Contribution 333 from the Great Lakes Research Division, The University of Michigan and contribution 287 from the Great Lakes Environmental Research Laboratory, National Oceanic and Atmospheric Administration.Contribution 333 from the Great Lakes Research Division, The University of Michigan and contribution 287 from the Great Lakes Environmental Research Laboratory, National Oceanic and Atmospheric Administration.     

Predation impact of Cyclops vicinus on the rotifer community in Lake Constance in spring

The predation impact of Cyclops vicinus on rotifers was studiedunder near-natural conditions in small enclosures to evaluatewhether copepod predation is responsible for the decline ofrotifers in Lake Constance in spring. Cyclops vicinus fed selectivelyon Synchaeta spp.; Keratella and Polyarthra spp. were not selectedfor. Predation rates increased with prey density up to a maximumof 37 Synchueta day^–1 at a density of 1.6 x 10⁶ Synchaetam^–2, i.e. at -1200 Synchaeta l^–1. Calculation ofcropping rates suggests that Cyclops alone can control the abundanceof Synchaeta in spring, i.e. that mainly Cyclops is responsiblefor the decline of Synchaeta species in Lake Constance in May.     

Prey Selection by the Water Bug Aphelocheirus aestivalis Fabr. (Heteroptera: Aphelocheiridae)

Feeding experiments were conducted in situ in flow through boxes to determine the vulnerability of different prey types (the mayfly larvae Baetis, Ephemerella and Ecdyonurus spp., the caddisfly larva Hydropsyche spp. and the crustacean Gammarus spp.) to the predaceous water bug Aphelocheirus aestivalis and to estimate the predation impact of Aphelocheirus on prey populations. Experiments with equal densities of mixed prey and experiments where each prey was tested individually revealed that Baetis and Ephemerella were most vulnerable to Aphelocheirus predation: Hydropsyche, Ecdyonurus and Gammarus were little/not preyed upon. The present results suggest that vulnerability of prey depends mainly upon prey mobility and microhabitat overlap between predator and prey and that feeding behaviour of Aphelocheirus resembles more that of megalopterans than of stoneflies. Experiments with different prey densities (120–720 prey m⁻²) showed that the predation rate of Aphelocheirus increased with increasing prey density. Experiments with different substrates documented that mortality rates of prey decreased with increasing substrate complexity. When substrate conditions were complex mortality rates in the control and in the experimental boxes were the same which suggest little importance of Aphelocheirus predation on mayflies in the study site.     

The influence of individual variation on length-biomass regressions in three crustacean zooplankton species

Length-biomass regressions were established for the cladoceranBosmina longispina maritima and for the copepods Euryremoraaffinis hirundoides and Limnocalanus macrurus, all common inthe Baltic Sea. Biomass was analysed as individual organic carboncontent. For Bosmina, length was a weak estimator of biomass;only 51% of the variation in carbon content was explained bylength. For pre-adult stages of copepods length was a rathergood estimator of biomass. For Euryremora nauplii. the length-carbonrelationship was best described by linear equations (r² wasbetween 0.49 and 0.69), and for copepodite stages (CI-V) byexponential functions (r² = 0.81). The regressions for two differentsampling areas (Baltic proper and Bothnian Bay) did not showstatistically significant differences. Among the copepodites(CI-V) of Limnocalanus there was a linear relationship betweenlength and carbon content (r² = 0.76). In both copepod speciesthere was no or a weak correlation between length and biomassamong adults.     

Differences in predation among morphotypes of the rotifer Asplanchna silvestrii

1. Interactions were observed between three morphotypes of the predatory rotifer Asplanchna silvestrii and six different prey (Brachionus plicatilis, B. rotundiformis, B. pterodinoides, B. satanicus, Hexarthra jenkinae and copepod nauplii) to understand the differences in feeding abilities among morphotypes that may have led to the evolution of this predator polymorphism. The outcome of predation events was affected significantly, both by predator morphotype and prey type. Predator morphotypes also interacted differently with different prey types. 2. The two smaller morphotypes, the saccate and the cruciform, responded similarly to prey overall, except that the smallest morphotype (saccate) was unable to ingest the most mobile prey (nauplii) and less able to ingest relatively large prey (B. plicatilis). The largest morphotype, the campanulate, had the highest encounter rate with prey, but the lowest probability of attack after encounter, so that it consumed far fewer prey per feeding bout than did the smaller morphotypes. This may have been because campanulates prefer larger prey than used in this study. 3. Highly mobile prey (H. jenkinae and copepod nauplii) were much less susceptible to predation than the less mobile Brachionus species. While evasiveness reduced attacks by saccates and cruciforms, campanulates did not have a significantly lower attack rate on H. jenkinae and copepod nauplii than on less evasive prey. Large body size moderately defended B. plicatilis against ingestion by saccates only. The short-spined B. satanicus was the only prey that was rejected after capture, resulting in lower ingestion probabilities for B. satanicus than other brachionid prey.     

The role of pelagic water mites in the control of cladoceran population in a temperate lake of the southern Andes

In a small temperate lake of the southern Andes, Bosmina longirostrisand Ceriodaphnia dubia coexist with the predaceous water miteLimnesia patagonica. Sampling of natural populations and laboratoryexperiments were carried out. The field population of Limnesiadid not show a numerical response to the density or biomassof its prey. Laboratory experiments showed that the water miterejected C.dubia adults and juveniles as prey, but consumedBosmina. The maximum predation rate was 40 prey predator^–1day^–1 and a linear relationship between predation rateand prey density was obtained (R² = 66%). The contribution ofmortality due to predation and the predation risk are too lowto provoke a prey suppression. By means of computer modelling,the densities of predator necessary to explain half of the totalprey mortality were calculated. These densities were one ortwo orders of magnitude higher than those in the field. It isconcluded that Limnesia could really be a suppressor, but thepotential depends greatly on its density.     

Copepod grazing on a phytoplankton community containing the toxic dinoflagellate Dinophysis acuminata

Copepod grazing on the toxic dinofiagellate Dinophysis acuminatafrom the west coast of France (La Rochelle) was studied witha concentrated (40–70 µm) phytoplankton assemblagedominated by Leptocylindrus danicus, D.acuminata, Ceratium fususand Ceratium furca. Copepod nauplii were also present. Threeto five copepods/copepodites (Acartia clausi, Isias clavipesand Centropages typicus) were incubated together with the phytoplankton.Dinophysis acuminata was grazed upon by all copepod species.However, to some extent, I.clavipes and C.typicus avoided itas food. Dinophysis acuminata cells represented for them only5–10% of total ingested carbon during the first 24 h,and almost all individuals survived and thrived well. In contrast,A.clausi did not avoid D.acuminata. which represented 30% ofingested carbon in 1 day. Acartia clausi then had a lower survivalthan the two other copepod species. However, the survival ofA.clausi was high in control incubations, where a plankton communitywithout D.acuminata was used as food. It is concluded that theokadaic acid of D acuminata is potentially toxic to some grazers,and/or might function as an allelopathic grazer repellent.     

Differential grazing on protozoan microplankton by developmental stages of the calanoid copepod Eurytemora affinis Poppe

Nauplii and adults of the copepod Eurytemora affinis Poppe collectedfrom the Multiscale Experimental Ecosystem Research Center (MEERC)mesocosms and from the Choptank River (a subestuary of ChesapeakeBay) reduced ciliate and dinoflagellate microplankton densitiessignificantly during grazing experiments. Protozoan microplanktongenerally were consumed in proportion to their availability,although both adult copepod and naupliar clearance rates werehigher for larger prey ({small tilde}40µm). Ingestionof ciliate microplankton was confirmed by examining copepodgut fluorescence after brief (1 h) incubations with ciliateslabeled with a fluorescent vital stain (5-chloromethylfluoresceindiacetate). In addition, both adults and nauplii of E.affiniscleared protozoan microplankton at considerably higher ratesthan chlorophyll a. Naupliar clearance rates were generallyan order of magnitude lower than adult rates, but given naupliarabundances in copepod assemblages, they should contribute substantiallyto total grazing on protozoan microplankton. Grazing by thetotal copepod assemblage in mesocosms and in the field may beunderestimated if juvenile stages are ignored.     

Seasonal variations in copepod size: effects of temperature, food abundance, and vertebrate predation

Seasonal dry weights of female and male Cyclops bicuspidatusthomasi, Diaptomus ashlandi and Diaptomus minutus were studiedin southeastern Lake Michigan during 1975–1981. Smallestanimals occurred during summer and early fall, and largest animalsin winter and spring, a pattern reported for other copepods.The range of weights for the species and sexes decreased fromwinter to summer, and converged in summer to a value of approximately2 µg/animal. Surface water temperature and abundancesof young-of-the-year (YOY) fish were inversely correlated withweights of males and females of the three copepod species. Standingstocks of important phytoplankton groups, especially pennatediatoms and flagellates, were positively correlated with copepodweight. Seasonal change of copepod body size appears to be morethan simply a function of temperature; seasonal predation byYOY fish may be a factor influencing sizes of adult copepods.     

Capelin larvae (Mallotus villosus) and community structure of zooplankton off the coast of northern Norway

Capelin larvae (Mallotus villosus) and mesozooplankton werecollected in May 2001 in the southern part of the Barents Sea.During the period of hatching and first feeding, capelin larvaeexperience high mortality rates. In order to understand theunderlying mechanisms influencing larval survival, we monitoreda selected site for 10 days using a variety of different samplingapproaches. Hydrological parameters conductivity–temperature–depth–fluorescence(CTDF) were recorded and zooplankton samples obtained simultaneouslyusing MOCNESS at three time periods. The mesozooplankton consistedessentially of copepods (85), dominated by Calanus finmarchicus.The zooplankton community consisted of different assemblagesderived from different areas (i.e. different water masses; Atlanticand coastal). Capelin larvae were found in surficial coastalwaters. Copepod nauplii and C. finmarchicus were less abundantin coastal waters where small copepodites (such as Acartia spp.and Temora longicornis) were found together with early stagesof coastal invertebrates that are possible prey for capelinlarvae. We suggest that capelin larvae base their foraging onsmall prey organisms frequently found among the coastal zooplanktonand that these larvae are less dependent on the propagationof the recruiting generation of C. finmarchicus present fromApril to June.