Planktonic predators and copepod abundance near the Dutch coast

Hypotheses that planktonic predators are responsible for thespring-summer decrease in copepod abundance and that the dominantpredator, Pleurobrachia pileus, is associated with high concentrationsof copepods were investigated at a station near the Dutch coast.Neither hypothesis was supported. Predators and copepods weresampled together with a 156 L ‘water box’ from lateApril through early July, including the season of P.pileus abundance.Using predators and copepods from the same box samples, hencefrom the same water parcel, feeding rates on copepod naupliiand copepodites + adults were measured onboard ship. Less than6% of the copepods, the sensitivity of the method, were removedper day. In additional shipboard feeding experiments net-caughtP.pileus were added to ambient copepod densities. By combiningwater volume cleared of copepods with ambient P.pileus densityin the sea, the predicted impact was 0–1.6% of copepodseaten per day from late April to early July. The hypothesisthat P.pileus associates with copepod concentrations was testedby comparing abundances of both groups from the same box samples.The correlations were not significant for any sample series.The ways measurement methods have restricted progress in understandingthe predatory impact of Pleurobrachia sp. are also considered.     

Mass density contrast in relation to the feeding currents in calanoid copepods

Theoretical analyses show that positively buoyant copepods areable to generate feeding currents by adopting upside-down bodypositions and pushing water upward. Thus, the excess buoyancyacting on the copepods will be balanced and cone-shaped feedingcurrents generated to transport water to the capture areas.The intensities of the feeding currents, which can be measuredin the present modeling study by calculating the volumetricflux going through the capture areas, are proportional to themass density contrasts between the copepods and the ambientseawater. The mass density contrasts may vary spatially andtemporally depending on copepod body contents and on the propertiesof the seawater immediately surrounding them. We focus on thecase where the mass density contrast between a wax ester-richcopepod and its ambient seawater can vary strongly with depthbecause wax esters are more compressible and 6–10 timesmore thermally expansible than seawater. These theoretical analysesshow that the intensities of the feeding currents generatedby wax ester-rich copepods vary strongly with depth. Our conclusionsfrom these theoretical analyses need to be tested by directobservations. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

Effects of prey motility and concentration on feeding in Acartia tonsa and Temora longicornis: the importance of feeding modes

Feeding experiments were conducted with the ambush-feeding copepodAcartia tonsa and the feeding-current-generating copepod Temoralongicornis. The copepods were offered a mixed diet of the dinoflagellateHeterocapsa triquetra and the ciliate Balanion comatum of similarcell size. The dinoflagellate was offered at a constant concentrationof 10–15 cells mL^–1, whereas the ciliate was offeredat a variety of concentrations, ranging from 7 to 57 cells mL^–1.Copepods with different feeding modes possess different mechanismsfor prey detection, suggesting that the two copepods would responddifferently to the two prey types. Both copepods had significantlyhigher clearance rates on the highly motile ciliate than onthe less motile dinoflagellate. In encounters between A. tonsaand its prey, we argue that this is due to the higher hydromechanicalsignal generated by the ciliate. The advection feeding copepodT. longicornis fed on the two prey according to their relativeconcentrations; in this case, we suggest that although B. comatumis capable of detecting feeding-current-generating predators,the feeding current velocity generated by T. longicornis isgreater than the escape velocity of this ciliate.     

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.     

Diel periodicity and selectivity in the feeding rate of the predatory copepod Mesocyclops edax

The diel periodicity and selectivity in the feeding behaviorof the predatory cyclopoid copepod Mesocyclops edax was investigatedat 3 h intervals over two 24 h sampling periods in nature. Gutcontent analyses revealed an increase in gut fullness at 11.30and 20.30 in July, and at 20.30 in August. The increase in gutfullness at these times could not be explained by an increasein prey density, changes in predator-prey overlap or differencesin prey vulnerability. We suggest that M.edax exhibits a truediel periodicity in the intensity of its predatory activities,although the alternative hypothesis of a diel periodicity ingut passage time cannot be ruled out. Vanderploeg and Scavia'sE^* selectivity index indicated a preference for rotifer andcladoceran prey over copepods.     

Perceptive performance and feeding behavior of calanoid copepods

The goal of this study was to determine variables associatedwith calanoid feeding behavior, and thus, to improve our understandingof the basics of calanoid feeding rates. These variables includedperiods and frequency of appendage motion, rates of cell clearance,distance at which a copepod first reacts to a cell which iseventually captured, and rate of water flow through the areacovered by the motions of a copepod's feeding appendages. Theeffects of these variables on feeding rates were determinedfor copepodids and adult females of the calanoid copepod Eucalanuspileatus at phytoplankton concentrations covering the rangeencountered by this species on the south-eastern shelf of theUSA. Our results indicate that the distance at which E.pileatusperceives phytoplankton cells increases {small tilde}2-foldas food concentrations decrease from 1.0 to 0.1 mm³ l^–1.These results lead us to hypothesize that this is due to increasedsensitivity of chemosensors on the copepods' feeding appendages.This 2-fold increase in perceptive distance amounts to a near4-fold increase in perceived volume which is close to the 6-foldincrease in volume swept clear (VSC) from 1.0 to 0.1 mm³ l^–1of Thalassiosira weissflogii. We assume that the increases inVSC by planktonic copepods, when food levels are below satiation,are largely a function of the sensory performance of the individualcopepod.     

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.     

The feeding strategies of two large marine copepods

We compared the feeding behaviour of the two copepods Paraeuchaetanorvegica and Chiridius armatus, allowing them to prey on othercalanoids in small-scale laboratory experiments. Several differenceswere found. When fed either live, free-swimming or dead, non-movingprey, P.norvegica seemed unable to locate the dead prey itemswhile C.armatus foraged heavily on them. When starved, P.norvegicaincreased its feeding rate while C.armatus reduced its feedingafter an initial increase. None of the predators changed theirfeeding rates when exposed to light. Nighttime versus daytimefeeding was tested only with P.norvegcia, which seemed to possessan endogenous feeding rhythm with increased rates at night.Chiridius armatus infected with epizooic ciliates appeared tohave increased feeding rates.     

Fitness consequences of selfing and outcrossing in the cestode Schistocephalus solidus

Mixed-mating, that is reproduction by both self-fertilizationand cross-fertilization is common in hermaphroditic parasites.Its maintenance poses, however, a problem for evolutionary biology.The tapeworm Schistocephalus solidus Müller 1776, servedas a model to study experimentally the consequences of selfingand outcrossing in its 2 consecutive intermediate hosts, a copepod(Macrocyclops albidus Jurine) and the three-spined sticklebackfish (Gasterosteus aculeatus). Size-matched tapeworms were allowedto reproduce either alone or in pairs in an in vitro systemthat replaced the definitive bird host's gut. Selfed eggs fromsingletons had a 4 times lower hatching success than outcrossedeggs from pairs. Outcrossed offspring achieved both a higherinfection success and a higher weight in the copepod, and ahigher number of parasites per host in both intermediate hosts,but only under competition. Outcrossed offspring were generallymore successful. If a S. solidus plerocercoid has a partnerin the bird's gut, they should outcross unless they differ insize and thus cannot solve the Hermaphrodite's Dilemma cooperatively.Using microsatellite markers, the proportion of selfed offspringand the total reproductive output of each worm within pairsvarying in mean weight and in weight difference was measured.Worms produced more selfed offspring not only with increasingweight difference as expected but also with decreasing totalweight of the pair. If small worms were selfed, they have alreadypurged deleterious mutations and would thus be better selfersin a year with low parasite density when worms cannot find partners.To maintain this advantage they should self a higher proportionof their eggs even with a partner. Here I review recent exprimentalevidence.     

On assessment of prey ingestion by copepods

The consumption of photosynthetic and heterotrophic cells byan abundant calanoid copepod species feeding on natural planktoncommunities was quantified with a state-of-the-art image-analysissystem. Late copepodid stages of Eucalanus pileatus did notingest bactena, small photosynthetic and heterotrophic nanoplankton,or the abundant Ceratium spp. in quantifiable amounts Althoughdiatoms were by far the most abundant cells (in terms of POC1^–1), the copepods ingested a higher percentage of ciliatesin relation to their abundance than of diatoms and small heterotrophicdinoflagellates in the first experiment, and ingested a higherpercentage of dinoflagel lates and ciliates compared to diatomsin the second experiment Heterotrophic cells sufficiently largeto be captured were repeatedly preferred by E.pileatus overautotrophs of similar or larger size. More over, among the cellswhich could be individually perceived by this calanoid, largerones were not pre ferred over smaller cells, implying that someaspect of food quality can be as significant as prey size. Theseresults support the notion (e.g. Kleppel, Mar. Ecol Prog. Ser.,99, s183–195, 1993) that feeding by copepods will be underestimatedif ingestion of heterotrophic food organisms is not quantified.While the proposed microscope-based method is comparativelyslow (     

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.     

Across-shelf predatory effect of Pleurobrachia bachei (Ctenophora) on the small-copepod community in the coastal upwelling zone off northern Chile (23{degrees} S)

To estimate the predation effect of the predominant ctenophorePleurobrachia bachei on the small-copepod community in the upwellingarea off Mejillones (23°S), northern Chile, a series ofoceanographic cruises and predation experiments were conductedin the austral springs 2000, 2001 and 2002. The daily consumptionrates and predatory effect of P. bachei on the small copepods(in terms of % of standing stock and biomass removed daily)were determined at three stations located in relation to theshelf-break (coastal, shelf-break and oceanic) reaching valuesup to 4.5% per day of the <1500 µm copepod standingstock. Our results indicate that the ctenophores were most abundantat the coastal station, that small copepods dominated the copepodcommunity (being more abundant nearshore), and that the relativefrequency of ctenophores with copepods in their guts was alsohigher near the coast. The predatory effect of P. bachei onthe small-copepod community was also higher in the coastal zone.However, the effect of this predation on the copepod biomassin terms of carbon did not decrease steadily seawards, whichmay be due to the larger sized copepods consumed at the offshorestations. Determinations of predatory effect on the secondaryproduction of the more abundant small-copepod populations (i.e26% daily in 2000) suggest that this single species of Pleurobrachiais modulating the population growth rate of the small copepods,the copepod community size structure, and maybe even the alternanceof key species in the Mejillones coastal upwelling zone.     

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.     

Competitive Growth Strategies in Intermediate Hosts: Experimental Tests of a Parasite life-History Model Using the Cestode, Schistocephalus solidus

In parasites with a complex life cycle, the fitness of an individual depends on its probability of reaching the final host and on its fecundity. Because larval growth in intermediate hosts may affect both transmission and adult size, selection should optimize growth patterns that are conditional on the presence and number of conspecific competitors. A recent model predicts that the total parasite volume per host should increase with intensity if larvae are able to vary growth depending on the number of conspecifics in the host (Life History Strategy hypothesis, i.e. LHS). Further, we would here expect growth rates to increase with intensity. By contrast, under the simplest alternative hypothesis of Resource Constraints (i.e. RC), the total parasite volume should remain constant. We experimentally infected copepods Macrocyclops albidus with the cestode Schistocephalus solidus to achieve 1, 2 or 3 parasites per host taking care that hosts had similar quality status at each infection level, and compared larval growth trajectories at the three intensity levels. The asymptotic total parasite volume was larger in double and triple infections than in single infections. Furthermore, the asymptotic total parasite volume was significantly larger in triple than in double infections but only in larger copepods that were less constrained by a host-size ceiling effect. These results, together with the fact that growth rates increased with intensity, support the LHS hypothesis: procercoids of a tapeworm may “count” their conspecific competitors in their first intermediate host to harvest its resources strategically until the next step in their complex life cycle. Co-ordinating editor: A. Biere     

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.     

Inter- and intraspecific conflicts between parasites over host manipulation

Host manipulation is a common strategy by which parasites alter the behaviour of their host to enhance their own fitness. In nature, hosts are usually infected by multiple parasites. This can result in a conflict over host manipulation. Studies of such a conflict in experimentally infected hosts are rare. The cestode Schistocephalus solidus (S) and the nematode Camallanus lacustris (C) use copepods as their first intermediate host. They need to grow for some time inside this host before they are infective and ready to be trophically transmitted to their subsequent fish host. Accordingly, not yet infective parasites manipulate to suppress predation. Infective ones manipulate to enhance predation. We experimentally infected laboratory-bred copepods in a manner that resulted in copepods harbouring (i) an infective C plus a not yet infective C or S, or (ii) an infective S plus a not yet infective C. An infective C completely sabotaged host manipulation by any not yet infective parasite. An infective S partially reduced host manipulation by a not yet infective C. We hence show experimentally that a parasite can reduce or even sabotage host manipulation exerted by a parasite from a different species.     

Intersetule distances are a poor predictor of particle-retention efficiency in Diaptomus sicilis

In contrast to published results for the copepod Acartia, thecumulative frequency distribution of intersetule distances onthe second maxillae of Diaptomus sicilis is a poor predictorof the experimentally determined particle-retention efficienciesof feeding. Moreover, a simple model that includes intersetalretention also does not work. This may be because D. sicilisraptorially seizes particles, as well as filtering them. Also,certain assumptions about the hydrodynamics of the filteringprocess that are implicit in the intersetule-distance modelsmay be false for Diaptomus and other calanoid copepods whosesecond maxillae form a stationary filter chamber. ¹GLERL Contribution No. 246 ²Present address: Department of Biological Sciences, CaliforniaState University, Chico, CA 95929, USA     

Feeding interactions of the copepods Eurytemora affinis and Acartia bifilosa with the cyanobacteria Nodularia sp.

We measured ingestion and clearance rates of two Baltic Seacalanoid copepods, Eurytemora affinis and Acartia bifilosa,on toxic and non-toxic cyanobacteria Nodularia sp. using theisotope technique. Eurytemora affinis fed actively on the non-toxicstrain and moderately actively on the toxic strain, whereasA.bifilosa totally avoided feeding on both strains. This suggeststhat A.bifilosa rejected cyanobacterial filaments due to theirnutritional inadequacy or difficult manageability. The differentresponse of E.affinis to the non-toxic and toxic strains, inturn, shows that this copepod species was able to sense thepresence of the toxin in cyanobacterial filaments and thereforefed less on the toxic strain. The interaction between A.bifilosaand Nodularia sp. was further examined (with the particle countingmethod) by measuring the clearance rates of A.bifilosa on ediblegreen flagellates in the presence of cyanobacteria. The presenceor concentration of toxic Nodularia sp. did not affect grazingrates of A.bifilosa on Brachiomonas submarina. Since earlierstudies have shown that ingestion of Nodularia sp. decreasesegg production and increases mortality in E.affinis, we suggestthat the occurrence of Nodularia sp. blooms in the Baltic Seamay favour individuals of copepod species capable of selectivefeeding, such as A.bifilosa.     

A new species of Stephos T.Scott, 1892 (Copepoda: Calanoida) from coastal waters of Sicily, Italy

Stephos cryptospinosus sp. nov., a new species of calanoid copepod,is described from Western Mediterranean coastal waters. It differsfrom other species of the genus in the structure of the fifthlegs in both sexes, and in having a small spinous process onthe posterolateral margins of the last prosomite in both sexes.     

Predation, production and the organization of an estuarine copepod community

The copepod community of the estuaries near Beaufort, NC underwenta consistent seasonal succession from a spring assemblage dominatedby the medium-sized copepod Acartia tonsa (1 mm) to a summer—fallassemblage dominated by the small-bodied copepods Parvocalanuscrizsrirostris and Oithona colcarva (both 0.5 mm). However,in enclosure experiments during this period, A. tonsa dominatedthe community, due to higher growth rates and its predationon the nauplii of other species. Nutrient additions enhancedthe dominance by A. tonsa. The decline in abundance of A. tonsain the estuary was associated with increased abundance of planktivorousanchovies and silversides. In other enclosure experiments, planktivorousfish eliminated A. tonsa and other large copepods, althoughthey persisted in enclosures lacking fish. I conclude that predationby size-selective planktivorous fish prevents dominance by A.tonsa during summer—fall. ¹Present address: Institute of Marine Sciences, University ofNorth Carolina, 3407 Arendell Street, Morehead City, NC 28557,USA