Diel and vertical variability of seston food quality and quantity in a small subalpine oligomesotrophic lake

Vertical profiles of seston food quality and quantity were measuredin subalpine Castle Lake for particulate carbon, chlorophylla, fatty acids and phosphorus in addition to abiotic parametersincluding water temperature and dissolved oxygen levels. Fieldand laboratory incubation experiments were employed to manipulateDaphnia rosea growth environments. Sestonic eicosapentaenoicacid (EPA) content was much lower, whereas carbon to phosphorus(C:P) ratios were much higher in the epilimnion compared tothe deep-water chlorophyll maximum (16–20 m). In a temperature-controlledincubation, Daphnia grew faster when fed seston from the deep-waterchlorophyll maximum. In in situ cage incubations, D. rosea grewfastest in the epilimnion. Daphnia rosea in a migrating treatmentexhibited intermediate growth rates between the ones for epilimnionand hypolimnion. A projection of D. rosea growth rates by growthmodels without regard to water temperature showed much highergrowth potentials in the hypolimnion. However, with inclusionof water temperature, epilimnetic water always gave higher potentialgrowth rates. In this lake, warmer temperatures of epilimneticwater override the higher food quality and quantity providedby the deep-water chlorophyll maximum.     

Seston food quality and Daphnia production efficiencies in an oligo-mesotrophic Subalpine Lake

Because of major biochemical imbalances between plants and animals, ecological efficiency at this interface may have a major impact on overall energy flow in ecosystems. In order to study relationships between seston food quality and energy transfer between primary producers and herbivores, we conducted five microcosm experiments in Castle Lake, California, USA during the summer of 1996. We simultaneously performed life table experiments to determine the effects of highly unsaturated fatty acids (HUFA) on Daphnia rosea growth, reproduction and survival. The results of these experiments suggest strong energy limitation of D. rosea growth in Castle lake during the study. D. rosea production was coupled with primary production in Castle Lake and in the microcosm experiments. D. rosea production efficiencies, i.e., the ratios of D. rosea productivity to primary productivity, decreased towards the end of the summer. A food quality index based on phytoplankton species composition and seston carbon to phosphorus (C:P) ratio were good predictors of D. rosea production efficiencies. The predicted D. rosea production pattern based on phytoplankton composition and primary productivity matched the zooplankton biomass dynamics in Castle Lake during 1991. Life table experiments showed HUFA effects on D. rosea population growth rates, reproduction and survival in support of the HUFA limitation hypothesis.     

Limiting effect of abioseston on food ingestion, postembryonic development time and fecundity of daphnids in Lake Balaton (Hungary)

During ice-free periods, seston ranges from 10 to 600 and averages29.2 mg dry wt 1^–1 in Lake Balaton, the largest shallowlake in Central Europe. Most (80–99%) of the seston consistsof 1–10 µm sized mineral particles. The abiosestoncauses permanent food limitation in the ingestion of the ediblephytoplankton (1–22 µm fraction) by Daphnia cucullataand D.galeata. The postembryonic development time of D.galeataincreases and its fecundity decreases with increasing abioseston.The fecundity and the mortality of D.galeata are balanced atan abioseston concentration of 20.0 mg dry wt 1^–1.     

In situ filtration responses of Daphnia galeata to changes in food quality

In the stoichiometric study of phyto–zooplankton interactions,a controversy exists about how Daphnia species regulate theirfeeding rate when submitted to low-quality food (i.e. high dietarycarbon:phosphorus [C:P] ratio). In this study, we gathered dataover 3 years on in situ clearance rates of a Daphnia galeatapopulation, by conducting grazing experiments from April 1998to October 2000 in the Esch-sur-Sûre reservoir (Grand-Duchyof Luxembourg). Observed clearance rates (2.5–13.5 mLindividual^–1 day^–1, mean 7.0 mL individual^–1day^–1) were correlated with population and environmentalvariables. Mean body size of Daphnia individuals was the bestpredictor of clearance rate (r² = 0.639), followed by watertemperature (r² = 0.262) and P concentration in the seston (r²     

Is competition important to arctic zooplankton community structure?

1. Daphnia pulex and Daphnia middendorffiana are commonly found in the Toolik Lake region of arctic Alaska. These two species are very similar morphologically, although their natural distributions differ markedly: D. pulex is restricted to shallow ponds, while D. middendorffiana is widely distributed and found in a variety of ponds and lakes. We compared the reproductive capabilities of D. pulex and D. middendorffiana grown under similar resource conditions and in the absence of the invertebrate predator Heterocope septentrionalis. In situ life table and mesocosm experiments were conducted in Toolik Lake and Dam Pond, habitats that have historically contained natural populations of D. middendorffiana, but never D. pulex. 2. Daphnia pulex exhibited a significantly higher net growth rate than D. middendorffiana in both life table and mesocosm experiments although D. pulex has never been found in either Toolik Lake or Dam Pond. Daphnia middendorffiana exhibited a negative net growth rate in Dam Pond, which had lower resource levels then Toolik Lake. Therefore, the smaller D. pulex appears to have a lower food threshold concentration than the larger D. middendorffiana. 3. Our results indicate that D. pulex is a superior resource competitor in the Toolik Lake region. These results combined with distributional patterns suggest that the restricted distribution of D. pulex in these arctic lakes and ponds cannot be explained by resource competition alone. We suggest that in the presence of H. septentrionalis, predation is an important factor structuring arctic zooplankton communities in the Toolik Lake region.     

Seasonal patterns of food limitation in Daphnia galeata: separating food quantity and food quality effects

Food limitation for the cladoceran Daphnia galeta was measuredgrowing them with natural lake seston for one growing seasonunder standardized conditions in the laboratory. Growth rateswere related to several measures of food quantity. Particulateorganic carbon (POC) <30 µm was the best predictorof total food availability measured. It was better than chlorophyll-a(Chl-a) of the same size fraction, algal volume calculated fromcell counts or total particle volume from Coulter Counter measurements.Daphnids were also grown with Scenedesmus acutus under the sameconditions and their growth rates were compared. Sestonic carbonconcentrations were related to Scenedesmus carbon concentrations,that provided equal growth rates. The ratio of the carbon concentrationsof both food types (seston, Scenedesmus) represented the foodquality of seston. Thus, the nutritional quality of naturallake seston relative to a standard alga was determined overone growing season. Growth rates of seston fed animals revealedthat they were food limited during long periods of the year.This was also ascertained by standard clutch sizes of fieldanimals. However, the causes of food limitation varied withinthe duration of the studied period. Whereas during the clear-waterphase the food quantity was limiting (while the quality washigh: 100% Scenedesmus), shortly after on 10 June and 17 Junefood conditions were close to threshold concentrations due tolow food quality. For the remainder of the summer, a POC of0.3–0.6 mg C I^–1 with a quality of     

Microcrustacean communities of high-elevation lakes in the Sierra Nevada, California

Seventy-five high-elevation lakes in the Sierra Nevada mountainsof California were sampled for microcrustacean species and majorion compositions. Cluster analysis was used to delineate fiverecurrent community types. Distributions of both individualspecies and of community types were related to chemical, morphometric,topographic and geologic variables, as well as fish presenceor absence, by stepwise logistic multiple regression. Fish distributionwas an important predictor of the distributions of all individualspecies for which significant regression models could be built;other common predictors included nitrate concentration, elevation,basin area and lake depth. One common community type (n = 22)consisted of Dapbnia rosea, Diaptomus signicauda, Bosmina longirostrisand Holopedium gibberum. A second, more species-rich, community(n = 27) included many of these same species, in addition toCyclopsvemalis, Diaphanasoma brachyurum, Polyphemus pediculis and/orCeriodaphnia affinis. Two further communities (n = 9 and n =6 respectively) contained Daphnia middendorffiana with eitherDiaptomus shoshone or Diaptomus eiseni. A fifth community type(n = 11) either lacked microcrustaceans or contained only Chydorusand/or Alona spp. The distributions of the first four of thesecommunity types could be well predicted on the basis of fishpresence and elevation (community types I and II), or fish absenceand lake depth (community types III and IV). The distributionof the fifth community type was independent of fish presenceor absence; phosphate concentration was the only significantpredictor of this community. ¹Present address: New York City Department of EnvironmentalProtection, PO Box 184, Valhalla, NY 10595, USA     

Primary and bacterial production compared to growth and food requirements of Daphnia longispina in Lake Kvernavatnet, west Norway

Primary production, and bacterial production as measured byincorporation of [³H-methyl]thymidineinto ice cold TCA insolublematerial were investigated during 1984 in Lake Kvernavatnet,west Norway. Primary production averaged 222 mg C m^–2day^–1 and bacterial production averaged 163 mg C m^–2day^–1. The bacterial production in the euphotic pelagiczonecontributed -60% of the total pelagic bacterial production.The zooplankton was dominated byDaphnia longispina. From growthexperiments with animals fed only natural food in coarse filteredlake water, the population daily growth increments were calculated.The average production of D.longispina was 151 mg C m^–2day^–1 during the period investigated. The estimated primaryproduction was too low to sustain both the bacterial productionand the zooplankton food requirements. These results imply thatthe carbon cycle of the lake is dependent on the supply of allochtonousmaterial, or that the current methods for measuring productionrates in aquatic environments are systematical erratic.     

Relationships between fecal pellet production and feeding in the calanoid copepod Boeckella

The number, size (mm³ pellet ^–1) and total volume of fecalpellets produced (mm³ h^–1) by Boeckella titicacae increasedwith concentration of Chlorella and natural seston in threedifferent lakewaters from Bahia de Puno and Lago Grande of LakeTiticaca and nearby Laguna Arapa. The three lakewaters differedin the size and number of fecal pellets produced at seston concentrations>0.5 mg dry wt l^–1. However, the total volume of fecalpellets egested varied little. Large-sized food (>10 µg)resulted in larger pellets but smaller total fecal production.Food concentration had parallel effects on the production offeces and on feeding rates. Both fecal production and feedingincreased linearly with food concentration with saturation between5 and 7.5 x 10⁵ Chlorella ml^–1 (3.7–5.6 mg dry wtl^–1). Varied responses in fecal production when fed differentsize fractions of lake seston suggest Boeckella has a complexand flexible feeding behavior. Feeding preferences of B.titicacaeand B.occidentalis (Ivlev index) for two seston size fractionsare generally reflected by fecal production.     

The influence of experimental scale on estimating the predation rate of Gammarus lacustris (Crustacea: Amphipoda) on Daphnia in an alpine lake

We examined the effect of experimental scale on the predationrate of Gammarus lacustris preying on Daphnia middendorffianain 20 l cubitainers, 2800 l mesocosms and diurnal whole-lakesurveys. The predation rate differed significantly among experimentalscales. It was highest in the 20 l cubitainers, intermediatein the 2800 l mesocosms and lowest in the lake. At all experimentalscales, the predation of G.lacustris on D.middendorffiana increasedas a linear function of prey density (Type I functional response).Very different predation rates were calculated for the wholelake depending on whether D.middendorffiana densities were estimatedfrom whole-water column hauls or stratified depth hauls. Thishighlights the importance of accurately determining the spatialdistribution of prey for estimating reliable in situ predationrates. We also determined prey selection by G.lacustris. Daphniamiddendorffiana was strongly selected over the calanoid copepodHesperodiaptomus arcticus. When presented with different sizedDaphnia, G.lacustris preyed size selectively on individuals2 mm. Our results show that the predation rate of small invertebratepredators is sensitive to a wide range of experimental conditionsand needs to be corroborated at a variety of scales. We concludethat the impact of G.lacustris on zooplankton communities wouldbe strongest in small fishless lakes or ponds.     

Feeding rate of Diaptomus sicilis and its relation to selectivity and effective food concentration in algal mixtures and in Lake Michigan

The concept of effective food concentration (EFC), a means ofpredicting food consumption from selectivity and food concentrationdata, is explained, tested, and applied to understanding foodconsumption by the freshwater copepod Diaptomus sicilis on mixturesof algae of different sizes and on Lake Michigan seston. Experimentson mixtures of different sized Chlamydomonas spp. showed thatselection (W') was an invariant function of particle size whenthe algae were counted microscopically. When the Coulter counterwas used, a more variable pattern of selectivity — similarto the peak tracking response reported by some investigators- was obtained. This was due to bias of zooplankton-producedparticles. Size-selective selectivity coefficients (W') wereused to weight the food concentration in each size categoryand the weighted values summed to give EFC. Food consumptionin experiments with seston and with cultured algae was betterdescribed by EFC than by total food concentration (TFC), theunweighted sum. Moreover, use of EFC diminished the magnitudeof the apparent threshold concentration required for feedingto commence. Although selectivity in algal mixtures and lakeseston was approximately the same, the food consumption versusEFC curve saturated more quickly for the algal mixtures thanfor the lake seston. Since expression of food concentrationas EFC allowed direct comparison of experiments having differentparticle-size spectra of food, we concluded the difference resultedfrom the lower food quality of lake seston, that is, its lowerdigestibility and sensory quality for zooplankton capture. ¹GLERL Contribution No. 368 ²Also Division of Biological Sciences, University of Michigan,Michigan, USA     

Impact of Daphnia pulex on a metalimnetic microbial community

A well-structured metalimnetic community, composed mainly ofphototrophic purple bacteria, Cryptomonas, and a few speciesof ciliates and rotifers, was regularly found between 1984 and1991 in Lake Cis. These populations appeared during spring,reached high concentrations and remained in the metalimnionuntil winter mixing. We had previously postulated that thesemetalimnetic populations could persist in such high numbersbecause they were free of efficient predation, since the oxygen-sulfideinterface provided a refuge from predation. In September 1992,an abundant Daphnia pulex population developed in the lake whichresulted in drastic changes in the formerly stable community.The presence of D.pulex induced an extreme clear-water phasein the epilimnion and the metalimnetic populations were severelyreduced. Feeding experiments with Cryptomonas and heterotrophicand phototrophic bacteria as food for D.pulex resulted in clearancerates around 0.7 ml individual^–1 h^–1. The impactwas very high on the microaerophilic populations, whereas theanaerobic populations were least affected, since sulfide limitedthe vertical distribution of ^D.pulex. We conclude from thisevidence that the metalimnetic populations in previous yeanwere mostly food limited (bottom-up control) and that theirhigh abundances were in great part due to a lack of efficientpredation.     

Chaoborus americanus predation on various zooplankters; Functional response and behavioral observations

Summary Predation rates for Chaoborus americanus on different types of noncyclomorphic zooplankton prey were determined in the laboratory as a function of both prey species and density. The sequence of events leading to consumption of prey was also observed, and probabilities determined for the various components.Predation rates generally reached an upper limit as the density of prey increased to high levels. Larger prey were always less vulnerable to Chaoborus predation. Among the species of zooplankton prey examined Holopedium gibberum suffered the lowest rates of predation, while those of the other plankters (Daphnia pulex, D. rosea, Moina hutchinsoni, Diaptomus birgei, D. leptopus, Cyclops vernalis) were approximately equivalent for any given size category.Observation of the predatory sequence indicates that for all the plankters except H. gibberum there were relatively high rates of evasion of attack by C. americanus and low rates of escape after capture. For H. gibberum the situation was reversed, and escape after capture was particularly important in determining the predation rate.The gelatinous envelope of Holopedium gibberum seems to operate as an effective anti-predator device which reduces the likelyhood of retention by Chaoborus after contact has been made.     

Mycosporine-like amino acids in planktonic organisms living under different UV exposure conditions in Patagonian lakes

Mycosporine-like amino acids (MAAs) were studied in zooplanktonfrom 13 Argentinian lakes covering a broad range in altitude,maximum depth and physico-chemical properties of the water.Four to nine different MAAs (predominantly porphyra-334 andshinorine) were found in the copepods Boeckella gibbosa, B.gracilipes, B. meteoris and Parabroteas sarsi, and in the ciliateStentor amethystinus, while MAAs were undetectable in the cladoceranDaphnia middendorffiana. Among the different copepods, maximumMAA concentrations accounted for 0.25–1.31% of the dryweight, and contents were generally about three to seven times(up to 43 times) higher in the animals living in the clearestlakes compared to those occurring in low-UV systems. This variabilityin the content of MAAs was related to the lake altitude (r²= 0.71), and the fraction of the water column to which 1% ofthe surface UV radiation at 320 nm penetrated (r² = 0.57). Ourdata therefore underscore the role of MAAs as sunscreens todecrease the potential negative effects of solar radiation,but they also indicate that other environmental factors besidesUV transparency play a role in determining MAA concentrations.One lake was selected to obtain additional information on thequalitative composition of MAAs in seston of <100 µmbetween two sampling sites and over a 2 month study period (australsummer). Six different MAAs were detected in the samples, withporphyra-334 and palythine being predominant. In the copepodscollected simultaneously, there was low variation in MAA concentrationsbetween the two sites and over time. Thus, our results suggestthat under similar UV exposure conditions MAA contents of planktonicorganisms show low temporal variation.     

Positive effects of UV radiation on a calanoid copepod in a transparent lake: do competition, predation or food availability play a role?

Zooplankton tolerant to ultraviolet radiation (UVR) could beindirectly affected by UVR through interactions with UV-sensitivespecies in the same ecosystem. In Lake Giles, Pennsylvania,USA, the calanoid copepod Leptodiaptomus minutus is more UVRtolerant than the cohabiting species Daphnia catawba and Cyclopsscutifer. We asked whether L. minutus is affected by UV-inducedmortality of a food competitor (D. catawba) or a predator ofits nauplii (C. scutifer). We conducted two in situ enclosureexperiments with six treatments: L. minutus alone, L . minutus+ Daphnia and L. minutus + Cyclops in the presence and absenceof UVR. There were few differences in survival among treatmentsin Experiment 1, which had enhanced food and a cumulative UVR(320 nm) dose of 9.3 kJ m^–2. In Experiment 2, which hadambient food and a UVR (320 nm) dose of 20.0 kJ m^–2, L.minutus survival and reproduction were higher in the +UVR comparedto –UVR, regardless of competitors or predators. Chlorophylla (Chl a) in Experiment 2 was higher in the +UVR than –UVR.While interactions between zooplankton species of differingUVR tolerances are potentially important, these results insteaddemonstrate that the beneficial UVR effect on L. minutus isindependent of concurrent detrimental UVR effects on competitorsand predators. Further research on the phytoplankton communityis necessary to determine whether UVR alleviates bacterial competition,increases nutrient availability or affects phytoplankton byother mechanisms.     

Feeding of freshwater filter-feeders at very low food concentrations: Poor evidence for "threshold feeding" and "optimal foraging" in Daphnia longispina and Eudiaptomus gracilis

The feeding behaviour of two freshwater zooplankton species,the calanoid copepod Eudiaptomus gracilis and the cladoceranDaphnia longispina, at extremely low food concentrations hasbeen studied in order to test whether the animals react as predictedby the models of optimal foraging. Three different sized algae were offered in concentrations downto 1µg C/litre at 7°C during winter and 19°C duringsummer. Ingestion rates were measured by use of a flow-through¹⁴C-technique In contrast to the findings of several authors working withmarine copepods a threshold concentration where the animalsceased filtering could be detected only in one experiment byregression analysis (Daphnia, 19°C, food: Stichococcus).A depression of the filtering rate at low concentrations wasobserved only in Daphnia at 19°C with Stichococcus and Scenedesmusas food. It is supposed that in this case the reduction of thefiltering rate is not a direct response of the filtrator, butis caused by exhaustion due to the experimental conditions. As freshwater zooplankton is very rarely exposed to food abundancesas low as in the marine environment, there seems to be no selectionpressure favouring the evolution of an "optimal forager" behaviour. ⁽¹⁾Supported by Deutsche Forschungsgemeinschaft     

Effects of invading vendace (Coregonus albula L.) on species composition and body size in two zooplankton communities of the Pasvik River System, northern Norway

Species composition and body-size distribution were studiedin the crustacean zooplankton communities of two limnologicallysimilar lake localities situated 50 km apart in the Pasvik RiverSystem, northern Norway. A recent invasion and successive downstreamexpansion of vendace (Coregonus albula), a specialized zooplanktivorousfish, allowed comparisons between sites with different predationpressures. Vendace had established a high population densityand was the dominant fish species in the pelagic of the upperlocality, but had just invaded the lower locality with a smallnumber of individuals. Whitefish (Coregonus lavaretus), a closelyrelated but less specialized zooplanktivore species, dominatedthe native fish community of both lakes. The zooplankton communityof the upper locality was dominated in June and August by Bosminalongirostris, the smallest zooplankton species represented inthe water course, and in September by Daphnia cristata. Thelower locality was dominated by the larger Holopedium gibberumand Eudiaptomus graciloides in June, by D.cristata in August,and by D.cristata and B.longirostris in September. The meanbody size of the three most abundant cladoceran species wassignificantly smaller in the upper locality, compared to thelower locality. It was concluded that the invasion and establishmentof a dense vendace population in the upper locality had increasedthe predation pressure in the pelagic, resulting in a reductionof body size and a shift towards smaller species in the zooplanktoncommunity.     

The recurrence of Daphnia rosea in Lake Tahoe: analysis of a population pulse

Following the disappearance of all Lake Tahoe cladocera in 1970,Daphnia rosea reappeared in the lake during the summer and fallof 1983. Since the introduction and population increase of theomnivorous opossum shrimp, Mysis relicta, was heavily implicatedin the initial disappearance, we investigated the D. rosea recurrencewith respect to fluctuating M. relicta abundance and steadilyincreasing primary productivity. An analysis of D. rosea populationdynamics, fecundity, feeding characteristics and M. relictafeeding selectivity, suggested that D. rosea was able to coexistwith M. relicta in 1983 only because D. rosea birth rates weresubstantially higher than historical levels due to the increasein algal productivity since 1970. The variation of cladoceransuccess in lakes of M. relicta introductions may be due, inlarge part, to variations in primary productivity.     

Are zooplankton food resources poor in the vegetated littoral zone of shallow lakes?

1. The distribution of zooplankton in shallow lakes is negatively related to macrophyte density. However, the abundance of their food along density gradients of macrophytes is unknown. A common but untested assumption is that food quantity and quality for pelagic zooplankton is poor in the littoral zone owing to the deleterious influence of macrophytes on phytoplankton. 2. We tested this assumption with a combination of a field survey and laboratory experiments. We collected seston samples from the littoral and pelagic zones of four shallow temperate lakes and related food quantity (phytoplankton biovolume) and quality to macrophyte abundance (per cent volume infested). Seston food quality was assessed in three ways: N/C and P/C ratios, polyunsaturated fatty acid content and phytoplankton community composition. In the laboratory, we measured the growth and reproduction of Daphnia pulicaria on diets consisting of seston from the littoral and pelagic zones in one lake. 3. In our four study lakes, food quantity was not significantly influenced by macrophyte abundance, and food quality was generally high. Laboratory experiments showed increased juvenile growth, but no significant change in D. pulicaria reproduction, when feeding on littoral resources compared to pelagic resources. 4. Our results suggest that there is no nutritional cost to pelagic zooplankton inhabiting the littoral zone. Therefore, it is likely that other factors (e.g. predation, abiotic factors) are involved in determining zooplankton habitat use.     

Demography of a natural population of Daphnia retrocurva in a lake with low food quality

The hypothesis was tested that Daphnia exhibit low populationrecruitment in a lake with phosphorus (P)-deficient food, whereDaphnia populations failed to increase in response to greatlylowered minnow density. Egg ratio analysis was performed onDaphnia retrocurva from L110 of the Experimental Lakes Area,Ontario, in 1994. L110 has high food abundance (seston carbon(C) 100 µM C), but the seston is P deficient (C:P atomic- 600). Chrysophytes dominated the algal community. The fishcommunity in this lake was manipulated in 1993 and 1994 by theaddition of northern pike, Esox lucius. Zooplanktivorous minnowswere greatly reduced after the piscivore addition, but the expectedincrease in abundance of the Daphnia to this altered predationregime was not observed, If food quality constrains the responseof the Daphnia to changed predation, there should be low recruitment in the relatively sparse Daphnia population in this lake.Birth rates estimated by standard egg ratio analysis were <0.1day^–1 for most of the sample period. In comparison toannual averages from the literature, recruitment in L110 wasmoderately, but not dramatically, low. Daphnia exhibited pronounced‘normal’ vertical migration (up at night, down duringthe day) all year, and a strongly declining size at first reproductionthrough the season. Population density and birth rates seemedto show much less short-term change than previously observedin other lakes. Although the demo graphic predictions for foodquality constraints seemed clear, in situ demography unfortunatelyneither supported nor strongly rejected the overall hypothesisof food quality limitation in this lake with very high C:P ratios.Possibly, direct demographic effects of low food quality willbe difficult to observe in nature.