Response of ciliates and Cryptomonas to the spring cohort of a cyclopoid copepod in a shallow hypereutrophic lake

The impact of a cyclopoid copepod population on the protozoacommunity (two ciliate categories and Cryptomonas) was assessedweekly during the spring cohort of Cyclops vicinus (one monthduration) in hypereutrophic Lake Søbygård by insitu gradient experiments with manipulation of ambient zooplanktonabundance. As C.vicinus always made up >92% of the zooplanktonbiomass, the response of protozoa is assumed to be a resultof predation by the copepod. Significant effects of copepodbiomass on protozoa net population growth rates were obtainedin the four experiments. Copepod clearance rates were significantlyhigher on oligotrichs than on prostomatids and Cryptomonas butdeclined for all three protozoa categories during the firstthree weeks of the copepod cohort, probably because of the changein developmental instar composition of the copepod population.Grazing impact on protozoa at ambient copepod abundance wasconsiderable (range, 0.05–0.87 day^–1) and could,together with the estimated reproductive potential of protozoans(range, –0.20–0.87 day^–1), account for thedecline in abundance and biomass of protozoa during the cohortdevelopment. Carbon flow from the protozoa to C.vicinus (range,2.8–23.5 µg C l^–1 day^–1) documents thepresence of a trophic link between protozoa and the spring cohortof C.vicinus in Lake Søbygård.     

Life cycle of Cyclops vicinus in relation to food availability, predation, diapause and temperature

The seasonal cycle of Cyclops vicinus was studied during a 5year period in the shallow and hypertrophic Lake Søbygård.The annual number of generations varied between three and sixand sometimes included midsummer generations. Naupliar and copepoditedensity was extremely high, the maximum recorded being 1313and 745 1^–1 respectively. The seasonal fluctuation inthe abundance of C.vicinus was related to the concentrationof edible phytoplankton (EDP): when diatoms and cryptomonadswere abundant, nauplii and copepodites were also abundant, whereaswhen greens such as Scenedesmus spp. were dominant, no cohortswere observed. The maximum density of copepodites in the separatecohorts and the amount of EDP, expressed as carbon content,were found to be significantly correlated. The development ofthe second cohort also seemed to depend on the occurrence ofa peak in rotifer density. High number of eggs per female generallycoincided with high biovolume of EDP, suggesting that algaeconstituted a major part of the diet of adult C. vicinus. Theduration of the naupliar stage was inversely related to temperatureand is one of the shortest times that have been recorded inthe literature. The short duration is probably attributableto the abundance of food. Fish composition and the calculatedsex ratio of C.vicinus indicate low predation pressure fromplanktivorous fish. The high density of C.vicinus during thesummer, when the whole population would be expected to haveentered diapause, is probably attributable to the concurrenceof low predation and the abundance of food.     

Metazooplankton dynamics and secondary production of Daphnia magna (Crustacea) in an aerated waste stabilization pond

The seasonal dynamics of metazooplankton biomass was monitoredin an aerated waste stabilization pond during three consecutiveyears (1994–1996). The pond showed a low diversity ofplanktonic metazoans because of elevated pH, relatively highconcentration of free dissolved ammonia and low oxygen concentration.The planktonic community was composed of the anomopod branchiopodDaphnia magna, and the cyclopoid copepods Cyclops vicinus andCyclops strenuus. Both predation by cyclopoids and competitionwith D.magna excluded rotifers from the pond, except duringa short period in spring 1996. Daphnia magna was the dominantorganism from a biomass point of view. In parallel with biomass,demographic parameters, secondary production and the spatialdistribution of D.magna were studied. A significant seasonaland interannual variation in the density, biomass and productionof D.magna was observed. The maximum density of daphnids variedfrom 264 x 10³ to 686 x 10³ individuals m^–2 and the maximumbiomass from 4 to 30 g dry weight (DW) m^–2. The annualnet production was high compared with the production of Daphniain natural environments, ranging from 288 to 593 g DW m^–2year^–1. The annual net production of exuviae accountedfor ~25% of the total annual net production. Harvesting of daphnidsfor commercial applications that took place during the productiveperiod did not have any discernible effect on the populationdynamics of D.magna. Sexual reproduction was not observed duringthe three studied years. Negative mortality rates, occurringduring early spring, however, indicated that recruitment fromephippia was effective in the pond of Differdange and that sexualreproduction took place before 1994. Swarming was regularlyobserved in relation to high densities.     

Fish-induced changes in zooplankton grazing on phytoplankton and bacterioplankton: a long-term study in shallow hypertrophic Lake Sobygaard

The impact of fish-mediated changes on the structure and grazingof zooplankton on phytoplankton and bacterioplankton was studiedin Lake Søbygaard during the period 1984–92 bymeans of in vitro grazing experiments (¹⁴C-labelled phytoplankton,³H-labelled bacterioplankton) and model predictions. Measuredzooplankton clearance rates ranged from 0–25 ml l^–1h^–1 on phytoplankton to 0–33 ml l^–1 h^–1on bacterioplankton.The highest rates were found during thesummer when Daphnia spp. were dominant. As the phytoplanktonbiomass was substantially greater than that of bacterioplanktonthroughout the study period, ingestion of phytoplankton was26-fold greater than that of bacterioplankton. Multiple regressionanalysis of the experimental data revealed that Daphnia spp.,Bosmina longirostris and Cyclops vicinus, which were the dominantzooplankton, all contributed significantly to the variationin ingestion of phytoplankton, while only Daphnia spp. contributedsignificantly to that of bacterioplankton. Using estimated meanvalues for clearance and ingestion rates for different zooplankters,we calculated zooplankton grazing on phytoplankton and bacterioplanktonon the basis of monitoring data of lake plankton obtained duringa 9 year study period. Summer mean grazing ranged from 2 to4% of phytoplankton production and 2% of bacterioplankton productionto maxima of 53 and 88%, respectively. The grazing percentagedecreased with increasing density of planktivorous fish caughtin August each year using gill nets and shore-line electrofishing.The changes along a gradient of planktivorous fish abundanceseemed highest for bacterioplankton. Accordingly, the percentagecontribution of bacterioplankton to the total ingestion of thetwo carbon sources decreased from a summer mean value of 8%in Daphnia-dominated communities at lower fish density to 0.7–1.1%at high fish density, when cyclopoid copepods or Bosmina androtifers dominated. Likewise, the percentage of phytoplanktonproduction channelled through the bacteria varied, it beinghighest (5–8%) at high fish densities. It is argued thatthe negative impact of zooplankton grazing on bacterioplanktonin shallow lakes is highest at intermediate phosphorus levels,under which conditions Daphnia dominate the zooplankton community.     

Regulation of Rotifer Community by Predation of Cyclops vicinus (Copepoda) in the Římov Reservoir in Spring

The structuring role and predation impact of Cyclops vicinus on the rotifer community was studied using in situ enclosure experiments in Římov Reservoir during the spring of 2000. Seasonal changes in abundance and birth rates of the rotifer species Synchaeta lakowitziana , Polyarthra spp., Keratella cochlearis and Kellicottia longispina were related to predation pressure, i.e. selectivity and cropping rates of C. vicinus . All four rotifer species were found to be consumed by C. vicinus whose predation rates showed a marked succession through the spring period and corresponded to changes in the relative abundance of the pelagic species. The highest predation rates exhibited by C. vicinus were on Synchaeta lakowitziana and Polyarthra spp., being 11.3 and 6.4 ind Cyclops ^–1 day^–1 respectively. The activity of the predator switched among three types of preferred food during the spring period. The most preferred, Synchaeta, was replaced by Polyarthra after Synchaeta became extinct and these species were replaced by Keratella when both soft‐bodied forms were not available in sufficient quantities. Loricated species (K. cochlearis and K. longispina) were consumed at a slower rate. We calculated the proportion of rotifer production cropped day^–1 in order to estimate the predation impact of C. vicinus , which ranged from 11.6 to more than 100% over time and rotifer species. This shows that relative predation impact can be high not only on species most strongly selected, but also on species which are not selected. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

The effects of food deprivation, prey density and volume on clearance rates and ingestion rates of Diacyclops thomasi

The ingestion rates of the copepod, Diacyclops thomasi, on thesoft-bodied rotifer, Synchaeta pectinata, increased 10-fold(0.07–0.77 Synchaeta h^–1) over the range 50–250prey l^–1. The saturating functional response curve appearedsigmoid but was statistically indistinguishable from a parabola.The response curve was more linear and 10 times lower over thesame range of density when Diacyclops was offered Kerarellacochleans, a species having a stiffened lorica. Diacyclops maximizedits ingestion rate on Synchaeta as a function of the availablegut space. Predation effort, measured as clearance rates, waslinked tentatively to changes in swimming speed of Diacyclopsand was a function of hunger level. Diacyclops, which were starvedfor varying periods of time, increased their ingestion rateson Synchaeta up to a maximum (-3.0 h^–1) after 7–10h of food deprivation. The gut passage time of Diacyclops wasestimated to be 7–8 h. Therefore, ingestion rates (andclearance rates) appeared to be strongly correlated to the volumeof food in the gut.     

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

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

Herbivorous nutrition of Cyclops vicinus: the effect of a pure algal diet on feeding, development, reproduction and life cycle

Nutritional requirements, functional response, development andreproduction of Cyclops vicinus were studied with exclusivelyalgal food. Phytoflagellates were found to be adequate foodresources for both juvenile development and egg production.Ingestion measurements were performed with Chlamydomonas reinhardii.The functional response data give evidence for low feeding efficiency,especially for the naupliar stages. A difference between naupliiand older instars was also found in their quantitative foodneeds. A higher threshold food concentration was observed fornaupliar development (0.4 mg C l^–1) than for copepoditedevelopment (0.2 mg C l^–1). The calculation of assimilationefficiencies suggests that the high food requirements are dueto low specific ingestion rates rather than poor assimilationefficiency. Development time decreased as algal density increasedand males developed more quickly than females at all food concentrations.Body size and carbon content increased with increasing foodconcentration. Continuous egg production was observed abovea food concentration of 0.5 mg C l^–1. The results haveimplications for the life cycle of C. vicinus. Summer diapauseis interpreted as a strategy to avoid starvation of the juvenilestages. The facultative herbivory of the adults might be anadvantage when competing against other more carnivorous cyclopoidcopepods. ¹Present address: Agricultural University of Wageningen, Departmentof Mathematics, Dreijenlaan 4, NL-6703 HA Wageningen, The Netherlands     

Laboratory and field observations on the scuticociliate Histiobalantium from the pelagic zone of Lake Constance, FRG

Histiobalantium sp. was found regularly in the pelagic zoneof Lake Constance, FRG, over five annual cycles. Maxima of upto 6400 cells l^–1 were recorded in late summer, with similarnumbers in the 0–8 and 8–20 m depth intervals. Onan annual average, the population accounted for 10–17%of the total biomass of planktonic ciliates. In the laboratory,Histiobalantium grew well on a diet of the cryptophyte Rhodomonassp. Maximum growth rates obtained in batch cultures were 0.21and 0.33 day^–11 at 9 and 18°C, respectively. In situexperiments using diffusion chambers yielded positive growthrates in autumn and winter. The highest values recorded at theambient temperatures 5, 14 and 17°C were 0.17, 0.32 and0.40 day^–1, respectively. Comparing these results withthe different seasonal distributions and higher measured growthrates of other ciliates from Lake Constance, we conclude thatHistiobalantium is a superior competitor at relatively low algalfood concentrations. ²Present address: Fisheries & Oceans Canada, 4160 MarineDrive, West Vancouver, BC, V7V 1N6, Canada     

Feeding of planktonic rotifers on ciliates: a method using natural ciliate assemblages labelled with fluorescent microparticles

A method was developed to allow direct measurements of predationexerted by metazooplankton on ciliates. The method relied onthe use of ciliates labelled with fluorescent microparticles(FMP). Optimal labelling conditions were determined with ciliatesfrom cultures (Tetrahymena pyriformis) and with natural ciliateassemblages sampled in a river. Labelled T. pyriformis wereused as tracer food to determine gut passage time (GPT) andingestion rates of the rotifer Brachionus calyciflorus in thelaboratory. Predation of metazooplankton from the lowland riverMeuse (Belgium) was determined by labelling natural assemblagesof ciliates and using them as tracer food for metazooplankterssampled in the river. Optimal labels of ciliates, i.e. sharpdistribution of FMP in cells, were obtained with short incubations(10 min) and low FMP concentrations (1 x 10⁵ mL^–1). GPTvaried between 30 and 45 min for B. calyciflorus and from 25up to >35 min for rotifers from the river. The ingestionrate of B. calyciflorus fed with T. pyriformis was 3.3 ±0.6 ciliate rot^–1 h^–1, i.e. 1.4 ± 0.3 ngCrot^–1 h^–1. Metazooplankton species for which theingestion of ciliates could be measured were the rotifers Keratellacochlearis, Euchlanis dilatata and Synchaeta spp. Ingestionrates measured ranged from 0.4 to 12.5 ngC rot^–1 h^–1.The method proposed proved to be useful in estimating the predationof microplankton on ciliates in semi- in situ conditions; infurther developments, labelled natural assemblages of ciliatescould be used for in situ incubations with the Haney chamber.     

Trophodynamics of the scyphomedusae Aurelia aurita. Predation rate in relation to abundance, size and type of prey organism

Ephyra larvae and small medusae (1.7–95 mm diameter, 0.01–350mg ash-free dry wt, AFDW) of the scyphozoan jellyfish Aureliaaurita were used in predation experiments with phytoplankton(the flagellate Isochrysis galbana, 4 µm diameter, {smalltilde}6 x 10^–6 µg AFDW cell^–1), ciliates (theoligotrich Strombidium sulcatum, 28 µm diameter, {smalltilde}2 x 10^–3 µg AFDW), rotifers (Synchaeta sp.,0.5 µg AFDW individual^–1) and mixed zooplankton(mainly copepods and cladocerans, 2.1–3.1 µg AFDWindividual^–1). Phytoplankton in natural concentrations(50–200 µg C I^–1) were not utilized by largemedusae (44–95 mm diameter). Ciliates in concentrationsfrom 0.5 to 50 individuals ml^"1 were consumed by ephyra larvaeand small medusae (3–14 mm diameter) at a maximum predationrate of 171 prey day^–1, corresponding to a daily rationof 0.42%. The rotifer Synchaeta sp., offered in concentrationsof 100–600 prey I^–1, resulted in daily rations ofephyra larvae (2–5 mm diameter) between 1 and 13%. Mixedzooplankton allowed the highest daily rations, usually in therange 5–40%. Large medusae (>45 mm diameter) consumedbetween 2000 and 3500 prey organisms day^"1 in prey concentrationsexceeding 100 I^–1. Predation rate and daily ration werepositively correlated with prey abundance. Seen over a broadsize spectrum, the daily ration decreased with increased medusasize. The daily rations observed in high abundance of mixedzooplankton suggest a potential ‘scope for growth’that exceeds the growth rate observed in field populations,and this, in turn, suggests that the natural populations areusually food limited. The predicted predation rate at averageprey concentrations that are characteristic of neritic environmentscannot explain the maximum growth rates observed in field populations.It is therefore suggested that exploitation of patches of preyin high abundance is an important component in the trophodynamicsof this species. ¹Present address: University of Bergen, Department of MarineBiology, N-5065 Blomsterdalen, Norway     

Predaceous feeding habits of Limnocalanus macrurus

Limnocalanus macrurus, a large, glacial-relict copepod, hasbeen assumed an omnivore or a herbivore; predaceous habits ofthe species are unknown. The predaceous feeding habits of Limnocalanusfrom Lake Michigan were studied in the laboratory using naturalprey. Predation rates were highest on copepod nauplii. Copepoditesof Diaptomus spp. and Cyclops spp. were preyed upon at lowerrates. Limnocalanus preyed selectively upon nauplii <300µm. Small cyclopoid copepodites (<–750 µm)were also selected over large copepodites. Experiments usingtwo prey types showed that nauplii were selected over all copepodites,and that no selectivity existed for either diaptomid or cyclopoidcopepodites. Predaceous feeding habits began in the fourth copepoditestage of Limnocalanus. Predaceous feeding rates of Limnocalanuschanged seasonally being highest in late spring and autumn andlowest in summer and early winter. Since Limnocalanus also feedson net-phytoplankton, predation rate changes may be relatedto changes in the relative abundance of large phytoplanktonand naupliar prey in nature. Limnocalanus predation may be animportant factor in structuring the zooplankton community. Present address: Great Lakes Research Division, University Michigan,Ann Arbor, MI 48109, USA     

Life cycles of two limnetic cyclopoid copepods, Cyclops vicinus and Thermocyclops crassus, in two different habitats

The life cycles of Cyclops vicinus and Thermocyclops crassusin two shallow eutrophic habitats, Junsainuma and Naganuma Ponds,Hokkaido, Japan, were investigated. Both ponds exhibited similarseasonal patterns of temperature, oxygen levels and pH duringice-free periods; however, oxygen levels were extremely lowerunder the ice in Naganuma Pond. Cyclops vicinus showed differentlife cycles in the two ponds; in Junsainuma Pond, it reproducedin winter and spring (January-May) and entered diapause duringsummer and autumn (June-October) as copepodite IV stage, whileit reproduced in autumn (October-November) and spring (April-May),and entered diapause in summer (June-September) and winter (Januaryand February) as copepodite V stage in Naganuma Pond. Thermocyclopscrassus entered diapause during winter (December-April) as copepoditeIV and V stages in both ponds, and egg-bearing females appearedonly during the warmseason, from early May to late October,when water temperatures were >10°C. Summer diapause inC.vicinus was suggested to be an adaptation against fish predation,whereas C.vicinus entered winter diapause in Naganuma Pond probablyto avoid low oxygen levels. Thermocyclops crassus entered diapausein both ponds to avoid low water temperature. These resultssuggest that biotic and abiotic factors are important for leadingto specific life cycles of cyclopoid copepods in small waterbodies.     

Post-embryonic development of cyclopoid copepods in various seasons at Lake Balaton (Hungary)

The development, generation number and diapausing state of threeCyclops species dominant in the open water of Lake Balaton werestudied by means of vials placed back into the lake, in naturalfeeding conditions. The males of all three species turned intoadults earlier than the females. The copepodite V stage wasof the longest duration. Three generations of Cyclops vicinusdeveloped yearly; in spring, autumn and winter. About 50% ofthe copepodite stage IV of the spring generation entered activediapause at the end of May, beginning of June, then entereddormancy between the middle of June and middle of July. Consideringthe literary data as well, it seems that the generation numberof the species is the function of temperature and of the water'strophic state. The Acanthocydops robustus f. limnetica is fondof warm water (> 14°C). Seven generations of this speciesdeveloped completely yearly, during an average of 2-week intervalsbetween May and September. The eightli generation died in thenaupliar stage. The generation number of A. robustus f. limneticadepends on the average temperature of the lake. Inactive diapausecould not be observed for this species. Only one generationof Mesocyclops latckarti developed in autumn. The copepoditeV of the winter generation entered active diapause below a watertemperature of 10°C, and inactive diapause from the endof January, which state ended in March.     

Pseudobalanion planctonicum (Ciliophora, Prostomatida): ecological significance of an algivorous nanociliate in a deep meso-eutrophic lake

Pseudobalanion plancioracum was the most abundant ciliate speciesin the pelagic zone of Lake Constance, FRG, over a 3 year period.Annual averages accounted for 30, 35 and 43% of total ciliatenumbers in the uppermost 20 m of the water column in 1987, 1988and 1989 respectively. Highest cell numbers were observed inearly spring, simultaneously with the first phytoplankton maximum.The small ciliate (mean length 15 µm, mean cell volume1300 µm³) is a raptorial feeder and predominantly consumesphytoplankton. In laboratory cultures, P planctonicum grew wellon a diet of Rhodomonas sp. (Cryptophyceae). Maximum growthrates increased from 0.46 day^–1 at 5 5°C to 1.52 day^–1at 18.5° C, while temperatures above 21°C were lethal.Depending on food concentration, 0.2–4.4 Rhodomonas cellswere ingested per ciliate and hour.     

Can phaeopigments be used as markers for Daphnia grazing in Lake Constance?

The formation of chlorophyll a degradation products was measuredwith natural phytoplankton from Lake Constance and Daphnia magnaand native Daphnia as grazers in grazing experiments duringspring bloom conditions using high-pressure liquid chromatography(HPLC). Chlorophyll a start concentrations were between 1.2and 16.3 µg l^–1; phaeopigment weights constituted5% of chlorophyll a weight. Only phaeophorbide a was a markerfor Daphnia grazing; concentrations of other phaeopigments (phaeophytina, chlorophyllide a and two unidentified phaeopigments) didnot increase during Daphnia grazing. Conversion efficiencies(chlorophyll a to phaeophorbide a) were between 0 and 43% ona weight basis, and between 0 and 65% on a molar basis. Conversionefficiencies were highest at high grazer density (40 Daphnial^–1) and after a 24 h exposure time. Grazing by microzooplanktonprobably led to the formation of the two unidentified phaeopigments.In Lake Constance, Daphnia density was significantly positivelycorrelated with the phaeophorbide a/chlorophyll a ratio whenit was <5000 Daphnia m^–3. However, when higher Daphniadensities were included in calculations, then Daphnia densitywas positively, but insignificantly, correlated with the phaeophorbidea/chlorophyll a ratio. This suggests that when the level offood per Daphnia is low, then grazing is more efficient withless production of phaeophorbide a and a higher production ofcolourless products.     

Microbial dynamics during the summer ice-loss phase in maritime Antarctic lakes

The dynamics of bacterioplankton and protozooplankton in twomaritime Antarctic lakes (Heywood Lake and Sombre Lake, SignyIsland, South Orkneys) were studied during the phase of icebreak-out (December and early January 1994/95). The lakes aresuffering animal-induced (fur seal) eutrophication, though HeywoodLake is most severely affected. Both lakes had morphologicallydiverse bacterial communities which increased during the studyperiod, reaching maxima of 80 x 10⁸ l^–1 in Heywood Lakeand 31.8 x 10⁸ l^–1 in Sombre Lake. Heterotrophic nanoflagellates(HNAN) reached a peak in late December with maxima of 40.6 x10⁸ l^–1 in Sombre Lake and 174 x 10⁵ l^–1 in HeywoodLake. Phototrophic nanoflagellates (PNAN) peaked in late Decemberafter ice loss in Heywood Lake (63 x 10⁵ l^–1), which coincidedwith a peak in a bloom of Chroomonas acuta which reached abundancesof 1.0 x 10⁸ l^–1. In Sombre Lake, ice persisted for alonger period and here PNAN reached their highest density atthe end of the study period (around 70.0 x 10⁵ l^–1). Ciliateabundance reached high levels in Heywood Lake (>60001^–1),while in Sombre Lake maximum abundance was 568l^–1. Protozooplanktondiversity was greater in the less-enriched Sombre Lake. Grazingrates of ciliates averaged 70.6 bacteria indiv.^–1 h^–1in Heywood Lake and 119.3 bacteria indiv.^–1 h^–1in Sombre Lake. The difference was a reflection of the differenttaxonomic make-up of the community in the lakes. HNAN grazingrates varied between 0.51 and 0.83 bacteria indiv.^–1 h^–1in Sombre and Heywood Lakes, respectively. Specific growth rates(r) h^–1 in Sombre Lake were 0.028 for ciliates and 0.013for HNAN, and in Heywood Lake 0.010 for ciliates and HNAN 0.012.These growth rates result in doubling times ranging between38 and 69 h for ciliates and around 55 h for HNAN.HNAN grazingon bacteria was curtailed in Heywood Lake in early January asa result of predation by microcrustacean larvae feeding on theplankton. Thus, for a short phase top-down control was apparentin the dynamics of Heywood Lake, a feature uncommon in Antarcticlake ecosystems. The impact of natural eutrophication on thesesystems is discussed in relation to other unaffected Antarcticlakes.     

The importance of macrophyte bed size for cladoceran composition and horizontal migration in a shallow lake

Cladoceran composition and diel horizontal migration were studiedin 2, 10 and 25 m diameter macrophyte exclosures establishedin the littoral zone of shallow Lake Stigsholm, Denmark. Theexclosures were protected from waterfowl grazing, but open tofish. The macrophyte community cornprized Potwnogeton pectinatus,Potamogeton pusillus and Callitriche hemaphroditica. Cladoceranswere sampled randomly every third hour inside and outside themacrophyte exclosures during a 24 h period. In the 2m exclosure,the pelagic species Ceriodaphnia spp. and Bosmina spp. dominatedduring the day, mean density being as high as 3430 indiv. l^–1During the night, density decreased to 10–20% of the daytimedensity thus indicating diel horizontal migration. In the 10and 25 m exciosures, the daytime mean density of Ceriodaphniaspp. was 865 and 202 indiv. l^–1, respectively, and didnot decrease at night. In contrast to the pelagic species, thedensity of macrophyte-associated species tended to be higherin the 10 and 25 m exclosure than in the 2 m exclosure. In thedaytime, Eurycercus lamellatus density in the 2, 10 and 25 rnmacrophyte exclosures was 7, 28 and 16 indiv. l^–1 respectively,while that of Simocephalus vetulus was 11, 171 and 92 mdiv.l^–1, respectively. There was no thy-night difference inthe density of macrophyte-associated species. We conclude thatcladoceran community composition varies with macrophyte bedsize, and that the edge zone between the bed and open wateris an important daytime refuge for potentially migrating pelagiccladocerans.     

An example of niche partitioning in three coexisting freshwater cyclopoid copepods

Vertical distributions at noon and midnight of copepodites,males and females of three coexisting species of freshwatercyclopoid copepods, Cyclops vicinus (a cold-water form), Thermocyclopscrassus and Mesocyclops leuckarti (warm-water forms), were studiedin Dreiangel Lake, a small eutrophic gravel pit. The largestspecies, C. vicinus, inhabits deep water layers, both at noonand midnight, while the smaller species, T.crassus and M.leuckarti,inhabit upper water layers. In all three species, copepoditesand males live closer to the water surface than females. Insummer, abundance of the smallest species (T.crassus) is highest,although finite birth rates are lowest. The distribution patternsand population characteristics suggest that the vertical distributionof cyclopoid copepods in Oreiangel Lake is mainly governed byfish predation; different temperature tolerances and preferencesof the three species may also be of significance.