Seasonal succession of zooplankton in the north basin of Lake Biwa

To examine the seasonal succession of the entire zooplankton community in Lake Biwa, zooplankton biomass (on an areal basis) and its distribution patterns among crustaceans, rotifers and ciliates were studied in the north basin from April 1997 to June 1998. Seasonal changes in phytoplankton and population dynamics of Daphnia galeata were also examined to assess food condition and predation pressure by fish. From March to November, crustaceans dominated zooplankton biomass, but rotifers and ciliates were dominant from December to February. Among crustaceans, Eodiaptomus japonicus was the most abundant species, followed by D. galeata. Zooplankton biomass increased from January to a peak in early April, just before the spring bloom of phytoplankton, then decreased in mid-April when mortality rate of D. galeata increased. From mid-June, zooplankton increased and maintained a high level until the beginning of November. During this period, both birth and mortality rates of D. galeata were relatively high and a number of rotifer and crustacean species were observed. However, their abundances were very limited except for E. japonicus which likely preys on ciliates and rotifers. In Lake Biwa, food sources other than phytoplankton, such as resuspended organic matter from the sediments, seems to play a crucial role in zooplankton succession from winter to early spring, while zooplankton community seems to be regulated mainly by fish predation from summer to fall.     

Contrasting effects of a cladoceran (Daphnia galeata) and a calanoid copepod (Eodiaptomus japonicus) on algal and microbial plankton in a Japanese lake, Lake Biwa

Macrozooplankton may affect algal and microbial plankton directly through grazing or predation and indirectly through nutrient regeneration. They may also affect potential prey positively by removing alternative predators. Here, we examined the effects of a cladoceran (Daphnia) and a calanoid copepod (Eodiaptomus) on algal and microbial plankton in a Japanese lake using in situ experiments in which we manipulated the nutrient supply and biomass of these macrozooplankton. The response of algal and microbial plankton to macrozooplankton was diverse and varied depending on the level of nutrient supply. Eodiaptomus seemed to feed mainly on large algae (>20 µm) and microzooplankton, while direct grazing by Daphnia on algae, bacteria, heterotrophic nanoflagellates (HNF), and microzooplankton (ciliates, heliozoa, and rotifers) was pronounced. Trophic linkages within these microbial plankton was also suggested; bacteria were grazed by HNF and these in turn were grazed by microzooplankton. When the nutrient supply was high, both HNF and microzooplankton were exposed to higher amounts of algae and lower bacterial abundance. Moreover, nutrient regeneration by daphnids and Eodiaptomus copepods seemed to differentially stimulate the growth of algae and bacteria. The results suggest that the relationship between macrozooplankton and microbial plankton cannot be fully understood without taking into consideration not only the feeding characteristics of the macrozooplankton, but also the food web structure, the subsidized algal resource, and nutrient regeneration from the macrozooplankton.     

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.     

Effects of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community of a deep,tropical reservoir: an enclosure experiment

1. An in situ enclosure experiment was conducted in a deep reservoir of southern China to examine (i) the effects of a low biomass (4 g wet weight m^?3) of silver carp (Hypophthalmichthys molitrix) and nutrients on the plankton community and (ii) the response of Daphnia to eutrophication. 2. In the absence of fish, Daphnia galeata dominated the zooplankton community, whereas calanoids were dominant in the fish treatments, followed by D. galeata. Silver carp stocking significantly reduced total zooplankton biomass, and that of D. galeata and Leptodorarichardi, but markedly increased the biomass of smaller cladocerans, copepod nauplii and rotifers. In contrast, nutrient enrichment had no significant effect on the plankton community except for cyclopoids. 3. Chlorophyta, Cryptophyta and Bacillariophyta were dominant phytoplankton groups during the experiment. Chlorophyta with high growth rates (mainly Chlorella vulgaris in the fish enclosures and Ankyra sp. in the fishless enclosures) eventually dominated the phytoplankton community. Total phytoplankton biomass and the biomass of edible phytoplankton [greatest axial linear dimension (GALD) < 30 μm], Chlorophyta, Cryptophyta, Bacillariophyta and Cyanobacteria showed positive responses to fish stocking, while inedible phytoplankton (GALD ≥ 30 μm) was significantly reduced in the fish enclosures. However, there was no significant effect on the plankton community from the interaction of fish and nutrients. 4. Overall, the impact of fish on the plankton community was much greater than that of nutrients. High total phosphorus concentrations in the control treatment and relatively low temperatures may reduce the importance of nutrient enrichment. These results suggest it is not appropriate to use a low biomass of silver carp to control phytoplankton biomass in warmer, eutrophic fresh waters containing large herbivorous cladocerans.     

The importance of crustacean zooplankton in structuring rotifer and phytoplankton communities; an enclosure study

During the spring clear-water phase of 1993, an enclosure experimentwas performed in the mesotrophic Schöhsee (Plön, FRG)in order to assess the impact of crustacean zooplankton on therotifer and phytoplankton community. Among the crustacean plankton,calanoid and cyclopoid cope-pods were abundant, but Daphnia‘longispina’ reached the highest densities in thisexperiment. The colonial rotifer Conochilus unicomis was notaffected by crustacean plankton. The two most abundant species,Synchaeta peclinata and Keratella cochlearis, increased exponentiallywhen macrozooplankton had been excluded from the enclosures,but did not increase when crustaceans were present. Birth anddeath rates of K.cochlearis could be reliably determined inthis field experiment, suggesting that this rotifer specieswas mainly controlled by exploitative competition rather thanby mechanical interference or predation. Daphnia ‘longispina’generally grazed selectively on the smaller ciliates and algae,thus depriving the rotifers of their phytoplankton resources.The dominant alga, the chrysophycean Dinobryon, increased, whethercrustaceans were present or not, but appeared to be grazed uponto a certain extent despite its considerable cell size and colonialorganization.     

Zooplankton-phytoplankton interactions in a eutrophic lake

Enclosure experiments were made in a cyanobacteria dominatedlake (Lake Rotongaio) to assess the impact of zooplankton (>150µm) grazing on algal growth rates and determine the effectof diel and vertical changes in zooplankton grazing intensityand nutrient (NH₄-N) regeneration upon abundance of phytoplankton.The filamentous cyanobacterium Anabaena minutissima var. attenuataand diatom Cyclotella meneghiniana showed a negative linearchange in abundance with a gradient in zooplankton grazing intensity.Phytoflagellates were not grazed and showed a positive linearchange in abundance with increasing zooplankton biomass. Theseeffects, as well as shortening of filament length of Anabaena,were caused by raptorial feeding by the alanoid copepod Boeckellapropinqua which dominated the zooplankton. Phytoplankton growthwas not stimulated by addition of nutrients, suggesting nutrientregeneration was not important. Diel and vertical changes infeeding and NH₄-N regeneration rates were measured in Marchand June 1988. Diel differences were more pronounced in Marchwhen the water column was stratified. Specific feeding rateswere more important than vertical changes in zooplankton biomassin determining community grazing rates in March, but in Junewhen the water column was mixed, vertical distribution of zooplanktonbiomass was important. Zooplankton grazing was an importantloss process for phytoplankton in the lower part of the epilimnionin Lake Rotongaio.     

Erratum to: How do UV radiation, temperature, and zooplankton influence the dynamics of alpine phytoplankton communities?

Plankton in mountain lakes are confronted with generally higher levels of incident ultraviolet radiation (UVR), lower temperatures, and shorter growing seasons than their lower elevation counterparts. The direct inhibitory effects of high UVR and low temperatures on montane phytoplankton are widely recognized. Yet little is known about the indirect effects of these two abiotic factors on phytoplankton, and more specifically whether they alter zooplankton grazing rates which may in turn influence phytoplankton. Here, we report the results of field microcosm experiments that examine the impact of temperature and UVR on phytoplankton growth rates and zooplankton grazing rates (by adult female calanoid copepods). We also examine consequent changes in the absolute and relative abundance of the four dominant phytoplankton species present in the source lake (Asterionella formosa, Dinobryon sp., Discostella stelligera, and Fragilaria crotonensis). All four species exhibited higher growth rates at higher temperatures and three of the four species (all except Dinobryon) exhibited lower growth rates in the presence of UVR versus when shielded from UVR. The in situ grazing rates of zooplankton had significant effects on all species except Asterionella. Lower temperatures significantly reduced grazing rates on Fragilaria and Discostella, but not Dinobryon. While UVR had no effect on zooplankton grazing on any of the four species, there was a significant interaction effect of temperature and UVR on zooplankton grazing on Dinobryon. Discostella and Dinobryon increased in abundance relative to the other species in the presence of UVR. Colder temperatures, the presence of zooplankton, and UVR all had consistently negative effects on rates of increase in overall phytoplankton biomass. These results demonstrate the importance of indirect as well as direct effects of climate forcing by UVR and temperature on phytoplankton community composition in mountain lakes, and suggest that warmer climates and higher UVR levels may favor certain species over others.     

Interactions between Phytoplankton,Zooplankton, and Fish in the Nutrient Rich Shallow Lake Hjarbæk Fjord,Denmark

In the shallow, nutrient-rich dammed-up lake Hjarbæk Fjord the balance between phytoplankton, zooplankton, and fish was completely disturbed during the summer period. Extensive growth of small blue-green algae (Oscillatoria cfr. trichoides, Pseudanabaena galeata) and chlorococcal green algae (Scenedesmus spp., Monoraphidium contortum, Actinastrum hantzschii) induced a pH rise to 10.5 in July. Fish were driven into the rivers, and many that did not escape died. After the fish had disappeared daphniae multiplied explosively and grazed away the major part of the phytoplankton. Ammonia concentrations rose and oxygen concentrations dropped to noxious levels for highly sensitive fish in August–September. Larger phytoplankton species (Aphanizomenon flos-aquae, Pediastrum boryanum, Pandorina morum, a. o.) were not grazed by the daphniae and gave rise to a maximum of copepods in September. Fish returned to the lake in October, when pH, ammonia, and oxygen levels had normalized.     

Transpiration in response to variation in microclimate and soil moisture in southeastern deciduous forests

Responses of forests to changes in environmental conditions reflect the integrated behavior of their constituent species. We investigated sap flux-scaled transpiration responses of two species prevalent in upland eastern hardwood forests, Quercus alba in the upper canopy and Acer rubrum in the low to mid canopy, to changes in photosynthetically active radiation above the canopy (Q_o), vapor pressure deficit within the canopy (D), and soil moisture depletion during an entire growing season. Water loss before bud break (presumably through the bark) increased linearly with D, reaching 8% of daily stand transpiration (E_C) as measured when leaf area index was at maximum, and accounting for 5% of annual water loss. After leaves were completely expanded and when soil moisture was high, sap flux-scaled daily E_C increased linearly with the daily sum of Q_o. Species differences in this response were observed. Q. alba reached a maximum transpiration at low Q_o, while A. rubrum showed increasing transpiration with Q_o at all light levels. Daily E_C increased in response to daily average D, with an asymptotic response due to the behavior of Q. alba. Transpiration of A. rubrum showed a greater response to soil moisture depletion than did that of Q. alba. When evaluated at a half-hourly scale under high Q_o, mean canopy stomatal conductance (G_S) of individuals decreased with D. The sensitivity of G_S to D was greater in species with higher intrinsic G_S. Regardless of position in the canopy, diffuse-porous species in this and an additional, more mesic stand showed higher G_S and greater stomatal sensitivity to environmental variation than do ring-porous species.     

Analysis of phytoplankton-zooplankton relationships in an oligotrophic lake under natural and manipulated conditions

During August, 1987, we performed a series of Limnocorral experiments in lake La Caldera, a small winter-kill lake in which phytoplankton is strikingly nutrient-limited. The effects of biomanipulation on zooplankton-phytoplankton relationships were assessed by monitoring both individual species and whole-assemblage responses. Two sizes of enclosures were used (15 and 350 litres) and two treatments were assayed: 1) removal of zooplankton by 45 μm filter net and 2) doubling the natural grazing pressure by increasing the zooplankton concentration. Results show the two enclosure types to differ strikingly: flagellates disappeared from the small enclosures, resulting in four- to six-fold changes in chlorophylla concentration and three- to four-fold changes in number of individuals. Most species were grazed (a prey selectivity based on criteria other than size was observed) and their net growth rate increased with zooplankton concentration, causing a net increase in the phytoplankton growth, a stimulatory effect probably through nutrient regeneration that overrides the losses due to grazing.     

Facing multiple enemies: parasitised hosts respond to predator kairomones

During their lifetime most organisms are exposed to various enemies influencing their victims in multiple direct and indirect ways. Most studies concentrate on the effects of one enemy at a time, thereby not taking into account that in nature organisms are often simultaneously exposed to more than one enemy. We conducted a life-history experiment to investigate the simultaneous effects of predators (fish, Leuciscus idus) and parasites (microparasite, Caullerya mesnili) on their victim (Daphnia galeata). D. galeata were exposed to predator kairomones, parasites or both. D. galeata are able to sense the presence of fish predators via chemical cues (= kairomones). Both fish predator kairomones and microparasite infections influence the life history of Daphnia. Some of the effects of fish predator kairomones are directly opposed to microparasite effects; fecundity, for example, is increased in the presence of fish kairomones and decreased in Daphnia parasitised with C. mesnili. We investigated the influence of both threats on age at maturity, body size at different adult instars, fecundity and survival of one D. galeata clone. In the presence of fish kairomones, all D. galeata matured significantly earlier and increased the number of eggs in the second brood significantly. Parasitised D. galeata matured significantly earlier than non-parasitised ones in the absence and presence of fish kairomones. An infection with the microparasite C. mesnili led to significantly lower clutch sizes at the second adult instar, to significantly smaller body sizes from adult instar three onwards and to significantly reduced survival. No significant interaction effect between the responses to fish presence and to parasite infection was found for any of the investigated life-history traits. The lack of interaction effects between the exposure to predator kairomones and parasite infection was most likely due to the different timing of the effects. Fish kairomones affected D. galeata early in its life history whereas C. mesnili increased in its effects over time. Our results show that parasitised D. galeata are able to exhibit life-history responses to fish predator presence early in their lives. Thus, D. galeata parasitised with C. mesnili have a similar chance as non-parasitised D. galeata to escape from fish predation via life-history changes. Since older parasitised D. galeata are smaller, they may have an even better chance to escape visual predators under actual predation.     

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.     

Farthest north lake and fjord populations of calanoid copepods Limnocalanus macrurus and Drepanopus bungei in the Canadian high Arctic

The zooplankton assemblages of Lake A and Disraeli Fjord, northern Ellesmere Island (83°N, 75°W), were surveyed in early summer 1999. In permanently ice-covered Lake A, two glacial relict calanoid copepod species (Drepanopus bungei and Limnocalanus macrurus) were found in the top 30 m. All developmental stages of the more abundant D. bungei were present, whereas only adults of L. macrurus were found. Analysis of gut contents showed that L. macrurus preyed upon the smaller species. A net tow sample of zooplankton from Disraeli Fjord was mainly composed of D. bungei and L. macrurus, along with two marine cyclopoid copepods (Oncaea borealis and Oithona similis). These two zooplankton communities occur within unusual environments that are strongly influenced by perennial ice and snow. They will be subject to major habitat disruption should the current warming trends continue in the north polar region.     

<Emphasis Type="Italic">Daphnia</Emphasis> versus copepod impact on summer phytoplankton: functional compensation at both trophic levels

Here we report on a mesocom study performed to compare the top-down impact of microphagous and macrophagous zooplankton on phytoplankton. We exposed a species-rich, summer phytoplankton assemblage from the mesotrophic Lake Schöhsee (Germany) to logarithmically scaled abundance gradients of the microphagous cladoceran Daphnia hyalina×galeata and of a macrophagous copepod assemblage. Total phytoplankton biomass, chlorophyll a and primary production showed only a weak or even insignificant response to zooplankton density in both gradients. In contrast to the weak responses of bulk parameters, both zooplankton groups exerted a strong and contrasting influence on the phytoplankton species composition. The copepods suppressed large phytoplankton, while nanoplanktonic algae increased with increasing copepod density. Daphnia suppressed small algae, while larger species compensated in terms of biomass for the losses. Autotrophic picoplankton declined with zooplankton density in both gradients. Gelatinous, colonial algae were fostered by both zooplankton functional groups, while medium-sized (ca. 3,000 µm³), non-gelatinous algae were suppressed by both. The impact of a functionally mixed zooplankton assemblage became evident when Daphnia began to invade and grow in copepod mesocosms after ca. 10 days. Contrary to the impact of a single functional group, the combined impact of both zooplankton groups led to a substantial decline in total phytoplankton biomass.     

Interactions between metazoans,autotrophs, mixotrophs and bacterioplankton in nutrient‐depleted high DOC environments: a long‐term experiment

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Summer feeding activities of zooplankton in Prydz Bay, Antarctica

Grazing of dominant zooplankton copepods (Calanoides acutus, and Metridia gerlachei), salps (Salpa thompsoni) and microzooplankton was determined during the austral summer of 1998/1999 at the seasonal ice zone of the Prydz Bay region. The objective was to measure the ingestion rates of zooplankton at the seasonal ice zone, so as to evaluate the importance of different groups of zooplankton in their grazing impact on phytoplankton standing stock and primary production. Grazing by copepods was low, and accounted for <1% of phytoplankton standing stocks and 3.8-12.5% of primary production for both species during this study; even the ingestion rates of individuals were at a high level compared with previous reports. S. thompsoni exhibited a relatively high grazing impact on primary production (72%) in the north of our investigation area. The highest grazing impact on phytoplankton was exerted by microzooplankton during this investigation, and accounted for 10-65% of the standing stock of phytoplankton and 34-100% of potential daily primary production. We concluded that microzooplankton was the dominant phytoplankton consumer in this study area. Salps also played an important role in control of phytoplankton where swarming occurred. The grazing of copepods had a relatively small effect on phytoplankton biomass development.     

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.     

The structure and diurnal variations of the zooplankton of the Prince Edward Islands: Implications for the biomass build-up of higher trophic levels

Summary Short-term variations in the micro-, meso- and macrozooplankton communities at shelf and offshore stations in the vicinity of the Price Edward Islands were examined during April/May 1985 and 1986. Microzooplankton was dominated by copepod nauplii. Other holoplanktonic groups were represented in large numbers, while meroplanktonic larvae of benthic invertebrates were very scarce despite the large benthic population of the island shelf. Copepods dominated in terms of numerical abundance the meso- and macrozooplankton assemblage but cuphausiids were also very important in terms of biomass contribution. Dirunal vertical migrations were most pronounced for the euphausiids and the copepod Metridia gerlachei. The copepods Eucalanus longipes, Rhincalanus gigas and Microsetella sp. showed significant, but lesser, diurnal vertical migrations. A daylight decrease in zooplankton biomass was observed consistently on the island shelf, compared to the pattern found in deep-water. Data from daytime tows collected during April/May 1983 show that zooplankton biomass in the area increases with depth up to a stability level of about 50 mg/m³ (dry weight). It is suggested that an interaction between vertical migrations, surface Ekman drift and bottom topography results in zooplankton mass depletion by visual predators during daytime and replenishment during the night. The nocturnal advection of allochtonous zooplankton into the area may represent an input of food supply equivalent to as much as 2.2 times the local maximum phytoplankton production.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Nutrient enrichment and zooplankton effects on the phytoplankton community in microcosms from El Andino reservoir (Venezuela)

To quantify the effects of nutrient enrichment (N and P) and zooplankton grazing on the phytoplankton community structure of El Andino reservoir (Venezuela), in situ microcosms were installed for 6–7 days. Microcosms consisted of polyethylene bags (42 cm × 71 cm, non-cylindrical shaped) filled with 10 l of filtered epilimnetic water. Experiments were carried out on a monthly basis from January to December 1993. The lack/addition of nutrients was cross-classified with the absence/presence of zooplankton, resulting in an experimental design of four treatment levels: (1) no nutrient addition, zooplankton absent (C); (2) nutrient addition (150 NH₄Cl mol ml^–1 and 10 KH₂PO₄ mol ml^–1; 1 ml per l of sample), zooplankton absent (N); (3) no nutrient addition, zooplankton present (collected from the reservoir water column using a 6-m vertical tow with a 80-m plankton net) (Z); and (4) nutrient addition (as in [2]), zooplankton present (as in [3]) (NZ). Treatments were triplicated, and samples were collected at the start and end of each experiment. Significant differences between treatments were determined using a two-way ANOVA at p<0.05. Nutrient enrichment caused an increase in phytoplankton biomass, with the increase of all algal groups, except Pyrrhophyta. In spite of this, relative proportions of Cyanobacteria decreased in most cases. Chlorophyta and Bacillariophyta increased, probably due to their greater competitive abilities for phosphorus. After enrichment, Scenedesmus was the dominant species from January to June, while from July to December, Dactylococcopsis and Lyngbya dominated in the enriched microcosms. Zooplankton affected the phytoplankton community in microcoms through grazing and nutrient (mainly P) regeneration. Cladocerans (Ceriodaphnia cornuta, Moina micrura and Diaphanosoma sp.) mainly grazed on diatoms, although particulate material was present in almost all the gut contents analyzed. Particulate material probably consisted of micro-algae, detritus, bacteria, triturated algae and mineral particles. Ostracoda mainly fed on Peridinium and particulate material, whereas Thermocyclops sp. and rotifers (Brachionus spp. and Keratella spp.) mainly ingested particulate material. On the other hand, zooplankton excretion caused a slight increase in phytoplankton biomass and P concentrations in microcosms with the animals present. The effects of nutrient and zooplankton did not interact in most cases. Experimental results suggest that, at the initial stages of a eutrophication process, phytoplankton could increase their abundance and biomass, but might not change its community structure. Since there was a strong correlation between phosphorus and chlorophyll-a (bottom-up control), it is suggested that eutrophication could be avoided by controlling P input to the reservoir.