Experimental studies of zooplankton–phytoplankton–nutrient interactions in a large subtropical lake (Lake Okeechobee, Florida, U.S.A.)

1. Over a 1-year period, twenty controlled experiments were performed using small mesocosms (20-l clear plastic carboys) and plankton communities collected from four sites in shallow, subtropical Lake Okeechobee, Florida. In replicated treatments, macrozooplankton grazers were excluded by size fractionation (115 μm), and/or nutrients (N and P) were added, and impacts on phytoplankton biomass and productivity were measured after 3-day incubations.
2. In most experiments (fifteen out of twenty), there was no significant effect of zooplankton exclusion on phytoplankton biomass or productivity, but there were significant increases in those attributes due to nutrient additions. The magnitude of the responses was a function of light availability at the collection sites.
3. In three experiments, zooplankton exclusion led to declines in phytoplankton biomass and productivity, suggesting that animals may sometimes have net positive effects on the phytoplankton, perhaps via nutrient recycling.
4. In only two experiments was there evidence of net negative impacts of grazers on the phytoplankton. In both instances, cladocerans ( Daphnia ambigua and Eubosmina tubicen ) were dominant in the zooplankton. However, the increases in chlorophyll a due to zooplankton exclusion were small (5–20%), probably because of the small size and relatively low grazing rates of the cladocerans.
5. The results support the hypothesis that phytoplankton biomass in Lake Okeechobee is little affected by herbivorous macrozooplankton. This may be a common feature of lowland tropical and subtropical lakes.     

Experimental study of the impacts of silver carp on plankton communities of eutrophic Villerest reservoir (France)

We examined the impact of five silver carp biomass levels (0, 8, 16, 20, and 32 g m⁻³) on plankton communities and water quality of Villerest eutrophic reservoir (France). We realized the experiments using outdoor mesocosms. The presence of silver carp led to changes in zooplankton and phytoplankton assemblages. High fish biomass strongly reduced cladoceran abundance (through predation). Silver carp inefficiently grazed down particles < 20 μm. More importantly, however, the suppression of herbivorous cladocerans resulted in the increase of small size algae which were relieved from grazing and benefit from high nutrient concentrations. In contrast, in mesocosms without fish, the dominance of cladocerans (mainly Daphnia) controlled small size algae and probably also larger size algae (colonial chlorophytes, cyanobacteria). Thus, the Secchi disc transparency increased markedly. Through cascade effects, the modification of grazers communities led to changes in the utilization patterns of the added nutrients by phytoplankton communities. In high fish biomass treatments, nutrients were more efficiently accumulated into particulate fractions compared with no-fish and low-fish biomass treatments that were characterized by higher dissolved nutrients concentrations. Zooplankton was an essential source of food for silver carp. The productivity of zooplankton sustained a moderate silver carp biomass (up to 16 g m⁻³). In the presence of the highest fish biomass, the productivity of zooplankton was not large enough and silver carps fed on additional phytoplankton. Although mesocosms with high fish biomass were characterized by a slight cyanobacteria development compared with other fish mesocosms, silver carp was not effective in reducing cyanobacteria dominance. This revised version was published online in August 2006 with corrections to the Cover Date.     

Drivers assessment of zooplankton grazing on phytoplankton under different scenarios of fish predation and turbidity in an in situ mesocosm experiment

Zooplankton play a key role in energy transfer within lake food webs, but we have a poor knowledge concerning their role as phytoplankton grazers in shallow subtropical lakes. In this study, we aimed to determine how zooplankton grazing upon phytoplankton is altered in different scenarios of fish predation and turbidity, and we explored the relevance of grazing compared to other environmental variables, to explain phytoplankton biomass changes. A mesocosm experiment was conducted by including the following treatments: fish, turbidity, fish + turbidity, and a control (without fish or varying turbidity). The experiment lasted 21 days, and samples were taken four times. Zooplankton grazing was only effective for the microphagous group upon Cryptophyceae, while large Chlorophyceae and small pennate Bacillariophyceae biomass were benefited in the presence of copepods and cladocerans, being negatively affected by depletions in nitrogen availability. In the turbidity treatment, a reduction in phytoplankton biomass was obtained, artificially increasing zooplankton grazing on phytoplankton, while fish presence inhibited grazing of adult copepods and cladocerans. The other groups of phytoplankton were only influenced by the environment. This experiment suggests that phytoplankton biomass variations would be more affected by the environment than by zooplankton grazing in shallow lakes from the Paraná River.

     

Impact of moderate silver carp biomass gradient on zooplankton communities in a eutrophic reservoir. Consequences for the use of silver carp in biomanipulation.

We examined the impacts of moderate gradient silver carp biomass (five levels from 0 to 36 g.m-3, i.e. about 0-792 kg.ha-1) on zooplankton communities of the eutrophic Villerest reservoir (France). During our mesocosm experiment changes in zooplankton assemblages were dependent on silver carp biomass. In the fishless and low fish biomass treatments, zooplankton abundance increased through time, owing to a peak in cladoceran density, but decreased (mainly cladocerans) at highest fish biomass. Copepods and rotifers were less affected at the highest fish biomass and dominated zooplankton communities. We highlighted that the presence of high silver carp biomass could lead to changes in phytoplankton assemblage via the impact on herbivorous zooplankton. Since silver carp efficiently graze on particles > 20 microns, the suppression of herbivorous cladocerans could result in an increase in small size algae (< 20 microns) abundance since these species would be released from grazers as well as competitors (large algae grazed by silver carp) and nutrients levels would be enhanced by fish internal loading. Our results showed that the use of low silver carp biomass (< 200 kg.ha-1) would allow us to minimize these negative effects.     

Structure et fonctionnement des ecosystemes du haut-rhone francais. 14-etude des preferences alimentaires de la loche franche (noemacheilus barbatulus L.) par une methode des points modifiee

The net effect of zooplankton on phytoplankton productivity was investigated through experiments using natural concentrations of plankton in the epilimnion of Castle Lake, California. Zooplankton enhanced algal productivity during July and August, and nocturnal grazers caused greater proportionate increases than did daytime densities. Consumers had neutral or negative effects in September. Animal biomass was associated significantly with enhanced productivity for the experiment in late July, and with decreased growth rates in the last trial of September. The direct relationship between the activities of zooplankton and primary productivity observed in other lakes is qualified by this seasonal change in net effect. The removal of algae by grazing, increases in the productivity to biomass ratio through nutrient regeneration and temporary photosynthetic inhibition from ammonia excretion appear to have shifted in relative impact during the 3 month experimental period.     

Top-down control of natural phyto- and bacterioplankton prey communities by Daphnia magna and by the natural zooplankton community of the hypertrophic Lake Blankaart

Biomanipulation, through the reduction of fish abundance resulting in an increase of large filter feeders and a stronger top-down control on algae, is commonly used as a lake restoration tool in eutrophic lakes. However, cyanobacteria, often found in eutrophic ponds, can influence the grazing capacity of filter feeding zooplankton. We performed grazing experiments in hypertrophic Lake Blankaart during two consecutive summers (1998, with and 1999, without cyanobacteria) to elucidate the influence of cyanobacteria on the grazing pressure of zooplankton communities. We compared the grazing pressure of the natural macrozooplankton community (mainly small to medium-sized cladocerans and copepods) with that of large Daphnia magna on the natural bacterioplankton and phytoplankton prey communities. Our results showed that in the absence of cyanobacteria, Daphnia magna grazing pressure on bacteria was higher compared to the grazing pressure of the natural zooplankton community. However, Daphnia grazing rates on phytoplankton were not significantly different compared to the grazing rates of the natural zooplankton community. When cyanobacteria were abundant, grazing pressure of Daphnia magnaseemed to be inhibited, and the grazing pressure on bacteria and phytoplankton was similar to that of the natural macrozooplankton community. Our results suggest that biomanipulation may not always result in a more effective top-down control of the algal biomass.     

Species-specific phytoplankton responses to nutrients and zooplankton manipulations in enclosure experiments

1. In situ enclosure experiments were performed in the mesotrophic Bermejales reservoir to evaluate the algal response to changes in the nutrient supply and in the zooplankton size structure and density in a 2 × 2 factorial design. The experiments were conducted during the spring bloom of nanoplanktonic diatoms in 1989. 2. Nutrient enrichment promoted a great increase of phytoplankton biomass indicating a strong nutrient limitation on phytoplankton growth. Total phytoplankton biomass was significantly lower in the Daphina-added enclosures at a given nutrient level and strong direct an indirect effect of zooplankton on phytoplankton community structure and nutrient availability were observed. 3. Most of the nanoplanktonic species were effectively grazed but species with protective coverings and large size colonies were favoured by grazers and small chlorococcales were unaffected probably because of their compensatory high growth rates. The decrease in total biomass imposed by grazers is attributable mainly to the decrease of Cyclotella ocellata, the most abundant species. This taxon suffers two net effects of zooplankton: direct grazing and the indirect decrease of Si availability caused by the growth of C. ocellata which was promoted by P excretion by zooplankton. Indirect effects of grazers on Si availability should, therefore, be taken into account in explaining phytoplankton succession and community structure. 4. In this experiment grazers affected considerably the nanoplanktonic community in Bermejales reservoir. The extent which they were affected, however, depended not only on the algal size as a determinant of edibility but also greatly on the specific nutrient requirements and taxonomic features of the algal species.     

Short-term productivity responses of algae and bacteria to zooplankton grazing in two freshwater lakes

SUMMARY. 1. The specific productivities of algae and bacteria were measured in short-term (4 day) experiments consisting of enclosures with natural or reduced zooplankton biomass. Experiments were repeated five times over a season in each of two lakes that differed in the background concentration of dissolved organic carbon (DOC).
2. Algal biomass as estimated by chlorophyll a was suppressed in enclosures with ambient grazer levels in six of ten experiments and enhanced in one experiment. Distribution of chlorophyll among net and nanoplankton was not significantly affected by grazing.
3. Relative to enclosures with reduced zooplankton, normal grazer biomass (97–466μg 1⁻¹ dry weight) enhanced specific algal productivity in only one of five experiments in the low DOC take and had no effect in all five experiments in the high DOC lake. The main effects of grazers on algae was through removal of biomass rather than through indirect changes in turnover rate.
4. Between experiments, bacterial density was either unaffected, or mildly enhanced (4–87%) in enclosures with ambient macrozooplankton compared to those with reduced levels. Bacterial productivity and turnover estimated by incorporation of [³H]thymidine into DNA showed different responses across experiments; increasing, declining or remaining the same with grazer minipulation. This variability was not related to differences in dissolved primary production or to background DOC between lakes or experiments. Comparison of bacterial productivities based on thymidine incorporation rates with changes in cell densities indicated that control of bacterial loss processes by macrozooplankton is more important than control of growth rates.     

Benthic and pelagic food resources for zooplankton in shallow high-latitude lakes and ponds

1. Shallow lakes and ponds are a major component of the northern landscape and often contain a high zooplankton biomass despite clear waters that are poor in phytoplankton. 2. In this study we quantified zooplankton food sources and feeding rates in the shallow waters of two contrasting high‐latitude biomes: subarctic forest tundra (Kuujjuarapik, Quebec) and high arctic polar desert (Resolute, Nunavut). Five substrate types were tested (beads, bacteria, picophytoplankton, filamentous plankton and microbial mats). Special attention was given to the role of benthos, a component that is usually poorly integrated into models of aquatic foodwebs. 3. Consistent with observations elsewhere in the circumpolar region, high concentrations of adult macrozooplankton occurred in all sites (up to 17 100 crustaceans m^?3) while phytoplankton concentrations and primary productivity were low. The communities were composed of multiple species, including Daphnia middendorfiana, Hesperodiaptomus arcticus, Leptodiaptomus minutus, Artemiopsis stefanssoni and Branchinecta paludosa. 4. Detritus made 89–98% of the planktonic resource pool and bacteria contributed the highest biomass (up to 29 mg C m^?3) of the planktonic food particles available to zooplankton. Benthic resources were dominated by microbial mats that grew in nutrient‐rich conditions at the base of the ponds and which dominated overall ecosystem biomass and productivity. 5. All species were flexible in their feeding but there were large, order of magnitude differences in clearance rates among taxa. These differences likely resulted from different grazing strategies among cladocerans, copepods and fairy shrimps, and possibly also from adaptation to specific food types and size ranges that occur locally in these waters. 6. The subarctic cladocerans Ceriodaphnia quadrangula and D. middendorfiana, and the arctic fairy shrimp B. paludosa were observed to graze directly on the microbial mats and the feeding experiments confirmed their assimilation of benthic substrates. The other zooplankton species showed a more pelagic feeding mode but were capable of using microbial mat filaments, thus may be indirectly linked to benthic processes via resuspension. 7. Our study indicates that the classical aquatic food web in which phytoplankton provide the sole production base for grazers does not apply to northern shallow lakes and ponds. Instead, microbial mats increase the physical complexity of these high latitude ecosystems and likely play a role in sustaining their high zooplankton biomass.     

Seasonal pattern in nutrient limitation and grazing control of the phytoplankton community in a non-stratified lake

1. The relative importance of zooplankton grazing and nutrient limitation in regulating the phytoplankton community in the non-stratified Lake Kvie, Denmark, were measured nine times during the growing season.
2. Natural phytoplankton assemblage bioassays showed increasing importance of nutrient limitation during summer. Growth rates at ambient nutrient concentrations were continually below 0.12 per day, while co-enrichment with nitrogen (N) and phosphorus (P) to above concentration-saturated conditions enhanced growth rates from May to the end of July.
3. Stoichiometric ratios of important elements in seston (C : N, C : P, N : P), in lake water (TN : TP), in external loading (TN : TP) and in internal loading (DIN : DIP) were measured to determine whether N or P could be the limiting nutrient. TN : TP molar ratio of both lake water, benthic fluxes and external loading suggested P limitation throughout the growing season. However, seston molar ratios suggested moderate P-deficiency only during mid-summer.
4. Abundance and community structure of the zooplankton varied considerably through the season and proved to be important in determining the responses of algal assemblages to grazing. High abundance of cladocerans and rotifers resulted in significant grazing impact, while cyclopoid copepods had no significant effect on the phytoplankton biomass.
5. Regeneration of ammonium and phosphate by zooplankton were periodically important for phytoplankton growth. A comparison of nutrient regeneration by zooplankton with nutrient inputs from sediment and external sources indicated that zooplankton may contribute significantly in supplying N and P for the growth of phytoplankton.     

How important is bacterial carbon to planktonic grazers in a turbid, subtropical lake?

This study examined the relative contributions of bacterialand phytoplankton production to the pelagic carbon flow of LakeOkeechobee, a large and shallow subtropical lake. Due to thepredominance of cyanobacteria in this lake, we hypothesizedthat bacterial carbon flow would be larger than phytoplanktoncarbon flow to grazers. Using epifluorescent and light microscopyand radiotracer techniques, we measured the carbon biomass ofplanktonic functional groups and carbon flow between these groups.The functional groups that we used in this study included: picophytoplankton,autotrophic nanoflagellates (ANAN), microphytoplankton, bacteria,heterotrophic nanoflagellates (HNAN), ciliates, microzooplankton(rotifers and copepod nauplii) and macrozooplankton (cladocerans,copepodites and adult copepods). Microphytoplankton dominatedthe carbon biomass of all plankton, whereas the calanoid copepod,Diaptomus, dominated the carbon biomass of the grazers. Phytoplanktoncarbon flow often was higher than bacterial carbon flow to grazers;however, bacterial carbon constituted a large percentage ofthe total carbon flow to grazers (33.7 ± 22.4%). Bacterialcarbon provided roughly one quarter of the carbon flow to macrozooplankton(27.1 ± 25.4%), whereas it provided half of the carbonflow to microzooplankton (57.4 ± 20.3%) and to protozoans(47.2 ± 25.8%). These results suggest that microbialpathways play an important role in the energetics of subtropicallake plankton communities. Although microbial loop pathwaysare important in many systems, direct bacterial carbon flowto macrozooplankton also may be important in copepod- and cyanobacteria-dominatedlakes.     

Field and experimental evidence of the effect of Jenynsia multidentata, a small omnivorous–planktivorous fish, on the size distribution of zooplankton in subtropical lakes

1. Small cladocerans, copepod nauplii and rotifers often dominate the zooplankton community in tropical and subtropical lakes. This is probably because of high predation pressure by small omnivorous–planktivorous fish, but experimental evidence is scarce.
2. This study used two approaches to test the effect of the small omnivorous–planktivorous fish species Jenynsia multidentata , which is frequently abundant in (sub)tropical eutrophic lakes in South America, on the size distribution of zooplankton. In Lake Blanca (Uruguay), which lacks any piscivores, we sampled seasonally for both fish and zooplankton. We also conducted an outdoor mesocosm experiment with treatments containing or lacking J. multidentata .
3. Together, the empirical and experimental data suggest that J. multidentata predation plays an important role in modulating the size structure of the zooplankton community in subtropical lakes. In the absence of J. multidentata , stocked large-sized zooplankters like Daphnia obtusa were abundant in the experiments, while small-sized zooplankton dominated in the presence of fish, as they did in the lake itself from spring to the end of the season.     

The zooplankton-phytoplankton interface in lakes of contrasting trophic status: an experimental comparison

We report here the results of an experimental study designed to compare algal responses to short-term manipulations of zooplankton in three California lakes which encompass a broad range of productivity (ultra-oligotrophic Lake Tahoe, mesotrophic Castle Lake, and strongly eutrophic Clear Lake). To assess the potential strength of grazing in each lake, we evaluated algal responses to a 16-fold range of zooplankton biomass. To better compare algal responses among lakes, we determined algal responses to grazing by a common grazer (Daphnia sp.) over a range ofDaphnia densities from 1 to 16 animals per liter. Effects of both ambient grazers andDaphnia were strong in Castle Lake. However, neither ambient zooplankton norDaphnia had much impact on phytoplankton in Clear Lake. In Lake Tahoe, no grazing impacts could be demonstrated for the ambient zooplankton butDaphnia grazing had dramatic effects. These results indicate weak coupling between phytoplankton and zooplankton in Clear Lake and Lake Tahoe, two lakes which lie near opposite extremes of lake trophic status for most lakes. These observations, along with work reported by other researchers, suggest that linkages between zooplankton and phytoplankton may be weak in lakes with either extremely low or high productivity. Biomanipulation approaches to recover hypereutrophic lakes which aim only to alter zooplankton size structure may be less effective if algal communities are dominated by large, inedible phytoplankton taxa.     

Grazer control and nutrient limitation of phytoplankton biomass in two Australian reservoirs

1. Grazer and nutrient controls of phytoplankton biomass were tested on two reservoirs of different productivity to assess the potential for zooplankton grazing to affect chlorophyll/phosphorus regression models under Australian conditions. Experiments with zooplankton and nutrients manipulated in enclosures, laboratory feeding trials, and the analysis of in-lake plankton time series were performed. 2. Enclosures with water from the more productive Lake Hume (chlorophyll a = 3–17.5 μg l^–1), revealed significant zooplankton effects on chlorophyll a in 3/6, phosphorus limitation in 4/6 and nitrogen limitation in 1/6 of experiments conducted throughout the year. Enclosures with water from the less productive Lake Dartmouth (chlorophyll a = 0.8–3.5 μg l^–1), revealed significant zooplankton effects in 5/6, phosphorus limitation in 5/6 and nitrogen limitation in 2/6 of experiments. 3. While Lake Hume enclosure manipulations of the biomass of cladocerans (Daphnia and Diaphanosoma) and large copepods (Boeckella) had negative effects, small copepods (Mesocyclops and Calamoecia) could have positive effects on chlorophyll a. 4. In Lake Hume, total phytoplankton biovolume was negatively correlated with cladoceran biomass, positively with copepod biomass and was uncorrelated with total crustacean biomass. In Lake Dartmouth, total phytoplankton biovolume was negatively correlated with cladoceran biomass, copepod biomass and total crustacean biomass. 5. In both reservoirs, temporal variation in the biomass of Daphnia carinata alone could explain more than 50% of the observed variance in total phytoplankton biovolume. 6. During a period of low phytoplankton biovolume in Lake Hume in spring–summer 1993–94, a conservative estimate of cladoceran community grazing reached a maximum of 0.80 day^–1, suggesting that Cladocera made an important contribution to the development of the observed clear-water phase. 7. Enclosure experiments predicted significant grazing when the Cladocera/Phytoplankton biomass ratio was greater than 0.1; this threshold was consistently exceeded during clear water phase in Lake Hume. 8. Crustacean length had a significant effect on individual grazing rates in bottle experiments, with large Daphnia having highest rates. In both reservoirs, mean crustacean length was negatively correlated with phytoplankton biovolume. The observed upper limit of its variation was nearly twice as high compared to other world lakes.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Complementary impact of copepods and cladocerans on phytoplankton

The differences in the impact of two major groups of herbivorous zooplankton (Cladocera and Copepoda) on summer phytoplankton in a mesotrophic lake were studied. Field experiments were performed in which phytoplankton were exposed to different densities of two major types of herbivorous zooplankton, cladocerans and copepods. Contrary to expectation, neither of the two zooplankton groups significantly reduced phytoplankton biomass. However, there were strong and contrasting impacts on phytoplankton size structure and on individual taxa. Cladocerans suppressed small phytoplankton, while copepods suppressed large phytoplankton. The unaffected size classes compensated for the loss of those affected by enhanced growth. After contamination of the copepod mesocosms with the cladoceran Daphnia , the combined impact of both zooplankton groups caused a decline in total phytoplankton biomass.     

Changes in the crustacean zooplankton community of Harp Lake, Canada, following invasion by Bythotrephes cederstrœmi

1. Detecting the impacts of invading Bythotrephes cederstrœmi (Crustacea, Onychopoda, Cercopagididae) on zooplankton in North American lakes has been hampered by the brevity of pre-invasion data, and by the difficulty of distinguishing the effects of the invader from other stresses. The data from Harp Lake in Ontario, Canada, circumvent these difficulties. Bythotrephes appeared in the lake in 1993. There is a 15-year pre-invasion data set, and no significant complicating concurrent stresses.
2. The species composition and the size structure of the crustacean zooplankton community of Harp Lake changed after the invasion. Several small species either declined dramatically in abundance (e.g. Bosmina longirostris , Tropocyclops extensus ) or disappeared ( Chydorus sphaericus , Diaphanosoma birgei , Bosmina ( Neobosmina ) tubicen ). In contrast the abundance of the larger cladocerans Holopedium gibberum and Daphnia galeata mendotae and the hypolimnetic copepod Leptodiaptomus sicilis increased. Several univariate and all multivariate summarizations of zooplankton abundance, biomass and size structure highlighted the uniqueness of the post-invasion community.
3. The alterations in the zooplankton community could not be attributed to changes in lake acidity, thermal regimes, penetration by ultraviolet light, nutrient status, fish stocking or the abundances of native invertebrate predators, but they were correlated with Bythotrephes abundance, both within and among years. Hence, we hypothesize that the invasion by Bythotrephes has significantly altered the crustacean zooplankton community of Harp Lake.     

Zooplankton response to extreme drought in a large subtropical lake

Plankton data from 1997 to 2005 were used to examine impacts of a managed draw-down, subsequent drought and resulting historic low water levels (during 2000 and 2001) on the zooplankton of Lake Okeechobee, Florida. Prior to the drought the lake supported less than 150 ha of submerged vegetation. Following the drought, over 15,000 ha of submerged vegetation developed around the lake shore and conditions favored greater survival of age 0 fish. The zooplankton changed significantly from the pre- to post-drought period, including: (a) a near-complete loss of all dominant species of cladocerans and rotifers; and (b) an abrupt transition to a community with over 80% of total biomass comprised of Arctodiaptomus dorsalis, a calanoid copepod previously described as being resistant to fish predation. These changes persisted over a 5 year post-drought sampling period. In contrast, there were no systematic changes in biomass of bacteria, phytoplankton, inedible cyanobacteria, algal cell size, suspended solids, or any other physical or chemical attributes known to affect zooplankton in shallow lakes. Evidence points towards increased predation by fish, and perhaps invertebrates, as factors responsible for loss of cladocerans and rotifers following the drought, and indicates a need for future research to link changes in water level to shifts in predation pressure in this and other shallow lakes. Handling editor: S.I. Dodson     

Response of phytoplankton and bacteria to nutrients and zooplankton: a mesocosm experiment

Although both nutrient inputs and zooplankton grazing are importantto phytoplankton and bacteria in lakes, controversy surroundsthe relative importance of grazing pressure for these two groupsof organisms. For phytoplankton, the controversy revolves aroundwhether zooplankton grazers, especially large cladocerans likeDaphnia, can effectively reduce phytoplankton populations regardlessof nutrient conditions. For bacteria, little is known aboutthe balance between possible direct and indirect effects ofboth nutrients and zooplankton grazing. However, there is evidencethat bacteria may affect phytoplankton responses to nutrientsor zooplankton grazing through direct or apparent competition.We performed a mesocosm experiment to evaluate the relativeimportance of the effects of nutrients and zooplankton grazingfor phytoplankton and bacteria, and to determine whether bacteriamediate phytoplankton responses to these factors. The factorialdesign crossed two zooplankton treatments (unsieved and sieved)with four nutrient treatments (0, 0.5, 1.0 and 2.0 µgphosphorus (P) l^–1 day^–1 together with nitrogen(N) at a N:P ratio of 20:1 by weight). Weekly sieving with 300µm mesh reduced the average size of crustacean zooplanktonin the mesocosms, decreased the numbers and biomass of Daphnia,and increased the biomass of adult copepods. Nutrient enrichmentcaused significant increases in phytoplankton chlorophyll a(4–5x), bacterial abundance and production (1.3x and 1.6x,respectively), Daphnia (3x) and total zooplankton biomass (2x).Although both total phytoplankton chlorophyll a and chlorophylla in the <35 µm size fraction were significantly lowerin unsieved mesocosms than in sieved mesocosms, sieving hadno significant effect on bacterial abundance or production.There was no statistical interaction between nutrient and zooplanktontreatments for total phytoplankton biomass or bacterial abundance,although there were marginally significant interactions forphytoplankton biomass <35 µm and bacterial production.Our results do not support the hypothesis that large cladoceransbecome less effective grazers with enrichment; rather, the differencebetween phytoplankton biomass in sieved versus unsieved zooplanktontreatments increased across the gradient of nutrient additions.Furthermore, there was no evidence that bacteria buffered phytoplanktonresponses to enrichment by either sequestering P or affectingthe growth of zooplankton.     

The effect of grazer size manipulation on periphyton communities

Summary We examined the effect of grazer size on periphyton biomass, size structure, and species compostion by removing the largest invertebrate grazers on artificial macrophytes planted in the littoral of Lake Memphremagog (Que-Vt). A series of exclosures with increasingly fine mesh prevented colonization by large invertebrates but allowed in smaller grazers. Oligochaetes, chironomids, and cladocerans effectively replaced snails so that total grazer biomass in the various treatments was not significantly different from the controls. With one exception, algal biomass, measured as chlorophyll a, did not differ significantly among the various treatments. However algal size and taxonomy were affected because the dominance of large blue-green colonies was apperantly related to the presence of large grazers. The results of the size manipulations were qualitatively similar to those induced in phytoplankton communities by size selective zooplankton grazing and are consistent with models based on general allometric equations.Contribution 181 of the Lake Memphremagog Project, Limnology Research Centre