Food- and temperature-dependent function of moulting rate in copepods: An example of parameterization for population dynamics models

The effects of temperature and food concentration on moultingrates in copepods were studied. The distribution of transferrates in different stages of Euterpina acuhfrons in differenttemperature and food concentrations were simulated using a processmodel. These distributions were fitted by an appropriate asymmetricalprobability distribution: the gamma density function (GDF).A parameterization of moulting rates established a continuousrelationship between parameters of GDF, and the two externalfactors, temperature and food concentration. This function canalso represent the frequency distribution of successive developmentalstages in a cohort. Two examples of such representation aregiven, using literature data.     

The influence of food resources on the development, survival and reproduction of the two cyclopoid copepods: Cyclops vicinus and Mesocyclops leuckarti

Laboratory experiments were conducted to determine whether thetwo cyclopoid copepods. Cyclops vicious and Mesocyclops leuckaru.exploit the same food resources. The food requirements of juvenilesof the two cyclopoid copepods were investigated. Moreover, theimportance of algae for the predaceous adults was studied. Naupliiof both M leuckaru and C.vicinus successfully developed intocopepodites when fed the motile algae Chlamydomonas reinhardtii.Chlamydomonas sphaeroides and Cryptomonas sp. Threshold foodconcentrations for naupliar development varied between offeredalgae and between the two cyclopoid species. The food thresholdfor successful naupliar development, when reared on C.reinhardui,was lower for M.leuckarti (0.3 mg C 1^–1) than for C.vicinus(0.5 mg C l^–1) whereas a similar food threshold was foundusing Cryptornonas sp (0.3 mg C ^–1) and C.sphaeroides(<0.2 mg C 1^–1), Naupliar development time was inverselyrelated to food concentration. Food required for copepoditedevelopment differed for the two cyclopoid species. Cyclopsvicinus was able to develop to the adult stage on a pure dietof any one of the three algal species. whereas M.leuckarti requireda prey supply of the rotifer Brachionus rubens. Food composition.i.e. algal species, algal concentration and rotifer abundance,influenced copepodite survivorship of both cyclopoids and wasalways higher in the presence of B.rubens. Under similar foodconditions, mortality was higher for M.leuckarti than for Cvicinus. Mesocyclops leuckaru females were very dependent onanimal food. The predation rate of M.leuckaru was not lowerin the presence of algae. Egg production of M.leuckarti waslow on a pure algal diet and significantly higher when B rubenswas present. The results were used to discuss the life cyclestrategy and the possibility of exploitative competition ofthe two cyclopoid copepods.     

Top-down impact by copepods on ciliate numbers and persistence depends on copepod and ciliate species composition

Experiments with the copepods Acartia clausi, Centropages hamatusand Pseudocalanus sp. were performed to assess the species-specificeffect of these copepods on the development of monospecificalgae (Nephroselmis pyriformis) and ciliate communities (Strombidiumvestitum, Strombidium conicum, Strombidium sp. and Lohmanniellaoviformis). It was hypothesized that potentially switching copepodslike A. clausi will stabilize the algal community by switchingbetween ciliate and algal food, in contrast to copepods withstereotypic filter feeding behaviour (Pseudocalanus sp.). Intreatments with Pseudocalanus sp. and C. hamatus, all ciliatespecies were wiped out in 2 days, resulting in blooms of N.pyriformis. In treatments with A. clausi, two of the ciliatespecies were able to persist, but the combined ciliate and copepodcommunity was not able to control the algal bloom. Ciliatesbecame abundant in control treatments without copepods, butonly S. vestitum and S. conicum seemed able to establish grazingcontrol. Hence, when evaluating the role of ciliates in foodwebs, their actual numbers and species composition should betaken into account. Likewise, the species composition of copepodsmay be crucial; these experiments demonstrate that small filterfeeding copepods may have tremendous impact on ciliate numbers.     

Omnivory in cyclopoid copepods: comparisons of algae and invertebrates as food for three, differenfly sized species

We examined the importance of algal versus invertebrate preyfor three cyclopoid copepods by comparing reproductive successfor females fed on three diets: exclusively algal food, exclusivelyinvertebrate prey and a combination of algal plus invertebrateprey. The three cyclopoid species represent a distinct gradientin body size: Tropocyclops prasinus mexicanus (small body size),Diacyclops thomasi (medium body size) and Mesocyclops edax (largebody size). Our results provide strong evidence for the potentialimportance of algae in the diet of adult cyclopoid copepods.However, the role of herbivory is dependent on species bodysize. We found that algae are the key dietary factor for reproductivesuccess of the small T.p.mexicanus. In contrast, the availabilityof invertebrate prey had a greater influence on reproductionby the two larger species. Overall, cyclopoid copepods shouldbe generally considered as having the potential to derive theirenergy from either animal or algal sources. This has importantconsequences for interpretations of the forces influencing zooplanktoncommunity diversity as well as for general food web theory.     

The fate of the carotenoid pigment fucoxanthin during passage through the copepod gut: pigment recovery as a function of copepod species, season and food concentration

The amount of fucoxanthin, a taxonomically diagnostic carotenoid,recovered after passage through the guts of the copepods Acartiacalifomiensis and Calanus pacificus, was determined after thecopepods had fed on low (50 µg Cl^–1) and high (350µg C1^–1 for Acartia; 500 ug C H for Calanus) concentrationsof the diatom Thalassiosira weissftogii, during spring (May)and winter (December). Changes in pigment concentrations andcell abundances were assessed in experimental (with copepods)and control (without copepods) samples by standard incubationexperiments. Pigment recovery was assessed by (i) comparingthe amount of ingested pigment recovered in the experimentalgroups with that predicted to have been ingested from cell countdata and (ii) comparing fuco-xanthin/cell ratios in controland experimental samples. Both techniques suggested that pigmentloss is substantial (usually 60–100%), regardless of species,food availability or season. Patterns of pigment conservationdiffered between species, although pigment recovery was alwayshigher at high, than at low, food concentrations. Pigment recoveryin Acartia was higher (9.4–28.0%) in the spring than duringthe winter (0 recovery), regardless of food concentration. InCalanus, however, pigment recovery was always higher at high(34.9–67.8%) than at low (0 recovery) food concentrations,regardless of season.     

Vectorial transport of toxins from the dinoflagellate Gymnodinium breve through copepods to fish

Toxins from the dinoflagellate Gymnodinium breve were tracedthrough experimental food chains from dinoflagellates, throughcopepod grazers, to juvenile fish. The generality of this foodweb transfer was demonstrated using three different combinationsof copepods and juvenile fish during different seasons. Fishwere not exposed directly to the toxic dinoflagellates but werefed toxin-laden copepods in order to examine sublethal vectorialintoxication. Toxins were shown to move from fish viscera tomuscle tissue within periods of 2–6 h to 25 h. A new toxindetection method was used in this first stepwise demonstrationof multi-trophic-level intoxication of a planktonic food chainby G.breve. Micellar electrokinetic capillary electrophoresiswith laser-induced fluorescence allowed measurements of toxinsat trace levels and nanoliter-sized volumes critical for planktonicfood web transfer studies.     

Copepod foraging and predation risk within the surface layer during night-time feeding forays

Vertical distribution data seem to indicate that certain speciesof diel vertical migrating copepods avoid the surface high chlorophyll(Chl) region within coastal and estuarine environments, evenduring the night. Copepods may avoid this layer to reduce predationmortality, avoid advective loss or to avoid consuming too muchtoxic algae. We hypothesize that copepods make several intermittentfeeding ‘forays’ into shallow surface layers duringthe night, returning to intermediate depths between forays.Using an individual-based model (IBM) of Calanus pacificus,we examined the implications of this behavior on feeding successand mortality risk, and tested whether a practical field-samplingscheme would be able to detect foray-like behavior. In somecases, mortality of the foray-foraging copepods was up to 50%less than that of randomly behaving controls, for a given amountof food ingested. The trapping scheme devised should be ableto detect the occurrence of foray behavior (FB) in the fieldand should show differences in the gut contents of copepodsentering and leaving the uppermost food-rich layer. The presenceor absence of foray-like behavior significantly altered therelative concentration of copepods within various surface strataand thus could influence the temporal availability of copepodsas prey for the larvae and juveniles of several important managedfish species. This paper is one of six on the subject of the role of zooplanktonpredator–prey interactions in structuring plankton communities.     

Nutritional suitability of the dinoflagellate Ceratium furcoides for four copepod species

The nutritional suitability of Ceratium furcoides for Eudiaplomusgracilis, Mesocyclops leuckarti, Thermocyclops oithonoides andCyclops abyssorum was studied by observing ingestion and assimilation.The dinoflagellate was ingested by female adults of all species.The calanoid copepod E.gracilis could not utilize Ceratium;mortality was high and no egg production was observed when Ceratiumwas the only food source. Mortality was low for the adult cyclopoidson this food and reproduction indicated that Ceratium was assimilated.The first two copepodite instars of M.leuckarti were not ableto handle Ceratium, while older stages preyed on them. The dinoflagellatewas not ingested by female M.leuckarti when its densities werelow. Advanced copepodite stages of C.abyssorum developed intoadults on a diet of Ceratium only. Mesocyclops leuckarti femalesingested Ceratium when offered a mixed food source of Ceratiumand the rotifer Brachionus rubens, but the rotifer was positivelyselected, even if its density was low. The results show thatCeratium provides a suitable food source for advanced copepoditeinstars and adult cyclopoid copepods, although it is not a preferredfood source.     

Toxic and inhibitory effects of the blue-green alga Microcystis aeruginosa on herbivorous zooplankton

Blooms of blue-green algae are often associated with declinesin populations of large-bodied cladocerans and increased importanceof small cladocerans, copepods, and rotifers. We conducted toxicityand herbivory experiments, using a wide range of herbivore taxa,to test the hypothesis that the blue-green alga Microcystisaeruginosa most strongly inhibits large cladocerans. For a varietyof herbivore taxa, M. aeruginosa was toxic or non-nutritious,and inhibited feeding on co-occurring nutritious food. The rotiferBrachionus calyciflorus was unique in several respects: it wasunaffected by M. aeruginosa toxins, it showed some ability togrow and reproduce on a diet of M. aeruginosa, and it maintainedhigh feeding rates on co-occurring nutritious food in the presenceof bloom densities of M. aeruginosa. There was a strong relationbetween the toxicity of M. aeruginosa and its inhibitory effecton herbivore feeding rates. Copepods strongly avoided consumingM. aeruginosa, but all cladocerans and rotifers tested filteredunicellular M. aeruginosa at rates similar to or higher thannutritious Chlamydomonas reinhardi. Our results indicate thatthere are a variety of mechanisms whereby herbivorous zooplanktoncan coexist with blooms of M. aeruginosa, including resistanceto toxic chemicals (B. calyciflorus), and avoidance of consumptionof M. aeruginosa by chemosensory means (copepods), or by theinability to consume large colonies (some small cladocerans). ¹Present Address: Department of Biology, George Mason University,4400 University Drive, Fairfax, VA 22030, USA     

The swimming behavior of the freshwater calanoid copepods Limnocalanus macrurus Sars, Senecella calanoides Juday and Epischura lacustris Forbes

The swimming behavior of the freshwater calanoid copepods Epischuralacustris, Senecella calanoides, and Limnocalanus macrurus wasexamined using a videotape analysis technique. E. lacustrisand S. calanoides alternate between periods of slow glidingand passive sinking. L. macrurus does not exhibit sinking behavior;its swimming activity consists of slow and steady gliding interruptedoccasionally by jumps. All three copepods exhibit behavioralresponses when exposed to prey organisms, as expressed by anincrease in the duration of sinking for E. lacustris and S.calanoides, a decrease in sinking speed for S. calanoides, anda less circuitous swimming path for L. macrurus. These behavioralchanges may allow the copepods to allocate more time for preydetection, to be less detectable by potential prey and to reducethe frequency of path re-crossing. The results demonstrate therelationship between the swimming and feeding behavior of calanoidcopepods and are consistent with previous findings that copepodshave the ability to modify their foraging behavior in responseto changes in food conditions.     

Gut evacuation rates and ingestion rates of Eudiaptomus graciloides measured by means of the gut fluorescence method

Diel variation in feeding rates of Eudiaptomus graciloides wasmeasured in situ with the gut fluorescence method during Mayand June in the eutrophic Frederiksborg Slotss Pronounced dielperiodicity occurred in both periods with highest feeding ratesduring the night. The food concentration as well as the periodicitywas almost constant with depth, indicating that the feedingactivity represents a behavioral periodicity. The gut evacuationmeasured in the laboratory on field-collected copepods was exponentialand temperature-dependent. The exponential decline in gut contentwas found for both copepods feeding continuously and when feedinghad ceased. Considering spatial and temporal variation in feedingactivity, daily ratios were 61–68 body carbon per day,and higher than if calculated from existing laboratory dataof short-term feeding dates.     

The importance of plankton to Cassin's auklets during breeding

The breeding chronology, food habits, growth and reproductivesuccess of Cassin's Auklets, Ptychoramphus aleuticus, were investigatedon Triangle Island, British Columbia. The breeding of Cassin'sAuklets coincides with the plankton bloom in the eastern NorthPacific. Cassin's Auklets in British Columbia feed their youngmostly large copepods, Calanus cristatus, and euphausiids, chieflyThysanoessa spinifera, but fish was an important food item inone summer. Chicks which hatched early, grew and survived betterthan late-hatched chicks and this may relate to food availability.Adults and chicks on Triangle Island were heavier than thoseon South Farallon Island, California. The heavier weight andgreater numbers of Cassin's Auklets in British Columbia as comparedwith regions to the south may relate to Calanus cristatus availabilityand abundance in northern areas.     

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.     

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 (     

Trophic interactions of the freshwater jellyfish Craspedacusta sowerbii

The predatory impact and the trophic role of the freshwaterjellyfish, Craspedacusta sowerbii, was studied using microcosmand enclosure experiments as well as a 3-year pond survey. Theresults showed a significant decrease of small herbivorous crustaceans,i.e. Bosmina longirostris and juvenile cyclopoid copepods, inthe medusa treatments of the microcosms and the enclosure experiments.Chlorophyll concentrations in the enclosure experiment weresignificantly increased in the medusa treatment, suggestingthat C. sowerbii may cause cascading effects in the food chain.A comparison of daily zooplankton losses during the pond surveycaused by medusae and fish (roach, Rutilus rutilus), and theirfood selectivities suggest food separation of these two predatorsand reveal a strong negative impact of medusae on the copepodpond community. In the case of a jellyfish bloom, our resultsshow that both food chains can co-occur in lakes because ofa weak interaction between these top predators, fish and jellyfish,with simultaneous impacts on the zooplankton structure.     

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.     

Feeding and metabolism of the siphonophore Sphaeronectes gracilis

The in situ predation rate of the siphonophore Sphaeronectesgracilis was estimated from gut content analysis of hand-collectedsiphonophores and from laboratory data on digestion rates ofprey organisms. At daytime prey densities of 0.25 copepods 1^–1,S. gracilis was estimated to consume 8.1 – 15.4 prey day^–1siphonophore^–1. From data on abundances of siphonophoresand copepods, S. gracilis was estimated to consume 2–4%of the copepods daily. In laboratory experiments, ingestionrates averaged 13.8 prey day^–1 siphonophore^–1 atprey densities of 5 copepods 1^–1 and 36.9 at 20 copeods1^–1. This was equivalent to a specific ingestion rate(for both carbon and nitrogen) of –17% day^–1 and45% day^–1, respectively, while specific ingestion in situwas only 2% day^–1. Ammonium excretion averaged 0.095 µg-atsiphonophore^–1 day^–1 at 5 prey 1^–1, and 0.162at 20 prey 1^–1. The specific respiration (carbon) andspecific excretion (nitrogen as ammonium) were calculated tobe 3% day^–1 at the lower experimental food level, and5% day^–1 at the higher food level. ¹Contribution from the Catalina Marine Science Center No. 66. ²Present address: Dept. of Biology, University of Victoria,Victoria, B.C., Canada V8W 2Y2.     

Spatio-temporal distribution and production of calanoid copepods in the central Baltic Sea

The aim of our study was the exploration of species-specificdistribution and production patterns of dominant copepods inthe Central Baltic Sea (Bornholm Basin). Spatio-temporal distribution,egg and secondary production were studied by means of net-samplingand egg production experiments from April to August 1999. Verticaland horizontal distribution patterns appeared to be linked witheach other and were primarily species-dependent. Both Acartiaspp. and Temora longicornis preferentially inhabited the upper30 m of the water column and the shallower marginal regionsof the Bornholm Basin. In contrast, the C4–5 and C6 ofPseudocalanus spp. preferentially inhabited the halocline region(50–70 m) and the deep central part of the area. Observeddifferences in horizontal distribution among these three copepodsappear to result from different depth preferences and depth-dependentwater circulation patterns. Egg production rates (EPR) clearlyexhibited seasonal differences with maximum EPR in May and minimumEPR in July. Mean EPR was significantly different between species,being highest in Acartia spp. and lowest in Pseudocalanus spp.The variability in EPR was related to differences in hydrographyand modelled food environment.     

Copepod reaction to odor stimuli influenced by cestode infection

The cestode Schistocephalus solidus uses copepods as first andsticklebacks as second intermediate hosts. For transmission,an infected copepod has to be preyed upon by a stickleback.We used copepods of the species Macrocy albidus to test whetherinfected and uninfected copepods differ in their reaction totwo kind of simultaneously presented odors: odors of sticklebacksand odors of sticklebacks and conspecificz. By giving this choice,we attempted to force the copepods to make a trade-off betweenthe benefit of risk dilution and possible predator confusionand the costs of food competition and other disadvantages inducedby conspecifics. Within 1–8 h after last feeding, uninfectedcopepods clearly preferred the odors of conspeciflcs under thechemically simulated threat of predation. This was in contrastto the infected copepods, who tended to avoid the odor of conspecifics.When the time between experiment and last feeding varied, infectedcopepods showed an increas preference for fish water only (oravoided conspecthcs) with increasing hunger level This suggeststhat S. solidus benefits from hunger-induced behavioral changesof its copepod host by influencing its microhabitat selection.The same effect could be found in both sexes; however, it wassignificantly more pronounced in male than in female copepods.We propose several hypotheses that could explain the differencebetween the sexes in their infection-dependent microhabitatselection.     

Le zooplancton de la baie de Sh?diac (Nouveau-Brunswick)

Analyses of boreal zooplankton of Shediac Bay demonstrate theabundance of copepods (81%) and meroplankters (18%). Whetherexclusively pelagic or not, 67 species are mentioned for thefirst time in this Northumberland Strait area, out of 76 recordedwithin 23 higher taxa. The fluctuations were observed from Mayto November and pointed out the dominance of such copepods asAcartia tonsa, A. clausi, Oithona similis and Centropages hamatusin relation with temperature, salinity and food distributions. ¹Adresse actuelle: Station marine de Tul?ar, B.P. 141, Universit?de Madagascar, (R?p. Malgache)