Relative impacts of Daphnia grazing and direct stimulation by fish on phytoplankton abundance in mesocosm communities

• 1 Planktivorous fish were hypothesised to influence the abundance of algal biomass in lakes by changing zooplankton grazing, affecting zooplankton nutrient recycling and by direct recycling of nutrients to phytoplankton. The relative roles of direct fish effects vs. zooplankton grazing were tested in mesocosm experiments by adding to natural communities large grazing zooplankton (Daphnia carinata) and small planktivorous fish (mosquitofish or juveniles of Australian golden perch).
• 2 The addition of Daphnia to natural communities reduced the numbers of all phytoplankton less than 30 µm in size, but did not affect total biomass of phytoplankton as large Volvox colonies predominated.
• 3 The addition of Daphnia also reduced the abundance of some small (Moina, Bosmina, Keratella) and large (adult Boeckella) zooplankton, suggesting competitive interactions within zooplankton.
• 4 The addition of mosquitofish to communities containing Daphnia further reduced the abundance of some small zooplankton (Moina, Keratella), but increased the numbers of Daphnia and adult Boeckella. In spite of the likely increase in grazing due to Daphnia, the abundance of total phytoplankton and dominant alga Volvox did not decline in the presence of mosquitofish but was maintained at a significantly higher level than in control.
• 5 The addition of juveniles of golden perch to communities containing Daphnia reduced the abundance of small zooplankton (Moina), increased the abundance of large zooplankton (adult Boeckella) but had no significant effect on Daphnia and total phytoplankton abundance.
• 6 The results of the present study suggest that some planktivorous fish can promote the growth of phytoplankton in a direct way, probably by recycling nutrients, and even in the presence of large grazers. However, the manifestation of the direct effect of fish can vary with fish species.

     

Effects of zooplankton on nutrient availability and seston C : N : P stoichiometry in inshore waters of Lake Biwa, Japan

Forty-eight-hour experimental manipulations of zooplankton biomass were performed to examine the potential effects of zooplankton on nutrient availability and phytoplankton biomass (as measured by seston concentration) and C : N : P stoichiometry in eutrophic nearshore waters of Lake Biwa, Japan. Increasing zooplankton, both mixed-species communities and Daphnia alone, consistently reduced seston concentration, indicating that nearshore phytoplankton were generally edible. The zooplankton clearance rates of inshore phytoplankton were similar to rates measured previously for offshore phytoplankton. Increased zooplankton biomass led to increased concentrations of nutrients (NH₄-N, soluble reactive phosphorus [SRP]). Net release rates were higher than those found in previous measurements made offshore, reflecting the nutrient-rich nature of inshore seston. Zooplankton nutrient recycling consistently decreased TIN : SRP ratios (TIN = NH₄ + NO₃ + NO₂). This effect probably resulted from the low N : P ratios of nearshore seston, which were lower than those commonly found in crustacean zooplankton and thus resulted in low retention efficiency of P (relative to N) by the zooplankton. Thus, zooplankton grazing inshore may ameliorate algal blooms due to direct consumption but tends to create nutrient supply conditions with low N : P, potentially favoring cyanobacteria. In comparison with previous findings for offshore, it appears that potential zooplankton effects on phytoplankton and nutrient dynamics differ qualitatively in inshore and offshore regions of Lake Biwa. Received: September 4, 2000 / Accepted: January 23, 2001     

Response of zooplankton to nutrient enrichment and fish in shallow lakes: a pan-European mesocosm experiment

1. Responses of zooplankton to nutrient enrichment and fish predation were studied in 1998 and 1999 by carrying out parallel mesocosm experiments in six lakes across Europe. 2. Zooplankton community structure, biomass and responses to nutrient and fish manipulation showed geographical and year‐to‐year differences. Fish had a greater influence than nutrients in regulating zooplankton biomass and especially the relative abundances of different functional groups of zooplankton. When fish reduced the biomass of large crustaceans, there was a complementary increase in the biomasses of smaller crustacean species and rotifers. 3. High abundance of submerged macrophytes provided refuge for zooplankton against fish predation but this refuge effect differed notably in magnitude among sites. 4. Large crustacean grazers (Daphnia, Diaphanosoma, Sida and Simocephalus) were crucial in controlling algal biomass, while smaller crustacean grazers and rotifers were of minor importance. Large grazers were able to control phytoplankton biomass even under hypereutrophic conditions (up to 1600 μg TP L^?1) when grazer biomass was high (>80–90 μg dry mass L^?1) or accounted for >30% of the grazer community. 5. The littoral zooplankton community was less resistant to change following nutrient enrichment in southern Spain, at high temperatures (close to 30 °C), than at lower temperatures (17–23 °C) characterising the other sites. This lower resistance was because of a greater importance of nutrients than zooplankton in controlling algal biomass. 6. Apart from the reduced role of large crustacean grazers at the lowest latitude, no consistent geographical patterns were observed in the responses of zooplankton communities to nutrient and fish manipulation.     

Trophic structure in the pelagial of 25 shallow New Zealand lakes: changes along nutrient and fish gradients

To gain better insight into the importance of predator and resourcecontrol in New Zealand lakes we surveyed the late summer trophicstructure of 25 shallow South Island lakes with contrastingnutrient levels (6–603 µg TP l^–1) and fishdensities. Total catch of fish per net (CPUE) in multi-meshgillnets placed in the open water and the littoral zones waspositively related with the nutrient level. Trout CPUE was negativelycorrelated with total phosphorus (TP) and total nitrogen (TN).Zooplankton seemed largely influenced by fish, as high fishCPUE coincided with low zooplankton and Daphnia biomass, lowaverage weight of cladocerans, low contribution of Daphnia tototal cladoceran biomass, low ratio of calanoids to total copepodbiomass and low ratio of zooplankton biomass to phytoplanktonbiomass. However, chlorophyll a was only slightly negativelyrelated to Daphnia biomass and not to zooplankton biomass ina multiple regression that included TN and TP. Ciliate abundancewas positively related to chlorophyll a and negatively to Daphniabiomass, but not to total zooplankton biomass, while no relationshipswere found between heterotrophic nanoflagellates and zooplankton.The relationships between fish abundance and nutrients and fishabundance and zooplankton:phytoplankton ratio and between chlorophylla and TP largely followed the pattern obtained for 42 northtemperate Danish lakes. We conclude that fish, including trout,have a major effect on the zooplankton community structure andbiomass in the pelagial of the shallow oligotrophic to slightlyeutrophic New Zealand lakes, but that the cascading effectson phytoplankton and protist are apparently modest.     

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.     

A 2-year experimental study on nutrient and predator influences on food web constituents in a shallow lake of north-west Spain

1. A 2‐year study was carried out on the roles of nutrients and fish in determining the plankton communities of a shallow lake in north‐west Spain. Outcomes were different each year depending on the initial conditions, especially of macrophyte biomass. In 1998 estimated initial ‘per cent water volume inhabited’ (PVI) by submerged macrophytes was about 35%. Phytoplankton biomass estimated as chlorophyll a was strongly controlled by fish, whereas effects of nutrient enrichment were not significant. In 1999 estimated PVI was 80%, no fish effect was observed on phytoplankton biomass, but nutrients had significant effects. Water temperatures were higher in 1998 than in 1999. 2. In the 1998 experiment, cladoceran populations were controlled by fish and cyanobacteria were the dominant phytoplankton group. There were no differences between effects of low (4 g fresh mass m^?2) and high (20 g fresh mass m^?2) fish density on total zooplankton biomass, but zooplankton biomass was higher in the absence of fish. With the high plant density in 1999, fish failed to control any group of the zooplankton community. 3. Total biovolume of phytoplankton strongly decreased with increased nutrient concentrations in 1998, although chlorophyll a concentrations did not significantly change. At higher nutrient concentrations, flagellate algae became more abundant with likely growth rates that could have overcompensated cladoceran feeding rates. This change in phytoplankton community composition may have been because of increases in the DIN : SRP ratio. Both chlorophyll a concentration and total phytoplankton biovolume increased significantly with nutrients in the 1999 experiment. 4. A strong decline of submerged macrophytes was observed in both years as nutrients increased, resulting in shading by periphyton. This shading effect could account for the plant decline despite lower water turbidity at the very high nutrient levels in 1998.     

Effect of larval fish and nutrient enrichment on plankton dynamics in experimental ponds

The hypotheses that larval fish density may potentially affect phytoplankton abundance through regulating zooplankton community structure, and that fish effect may also depend on nutrient levels were tested experimentally in ponds with three densities of larval walleye, Stizostedion vitreum (0, 25, and 50 fish m^–3), and two fertilizer types (inorganic vs organic fertilizer). A significant negative relationship between larval fish density and large zooplankton abundance was observed despite fertilizer types. Larval walleye significantly reduced the abundances of Daphnia, Bosmina, and Diaptomus but enhanced the abundance of various rotifer species (Brachionus, Polyarthra, and Keratella). When fish predation was excluded, Daphnia became dominant, but Daphnia grazing did not significantly suppress blue-green algae. Clearly, larval fish can be an important regulator for zooplankton community. Algal composition and abundance were affected more by fertilizer type than by fish density. Inorganic fertilizer with a high N:P ratio (20:1) enhanced blue-green algal blooms, while organic fertilizer with a lower N:P ratio (10:1) suppressed the abundance of blue-green algae. This result may be attributed to the high density of blue-green algae at the beginning of the experiment and the fertilizer type. Our data suggest that continuous release of nutrients from suspended organic fertilizer at a low rate may discourage the development of blue-green algae. Nutrient inputs at a low N:P ratio do not necessarily result in the dominance of blue-green algae.     

Biomanipulation and food-web dynamics — the importance of seasonal stability

Responses of phytoplankton biomass were monitored in pelagic enclosures subjected to manipulations with nutrients (+N/P), planktivore roach (Rutilus rutilus) and large grazers (Daphnia) in 18 bags during spring, summer and autumn in mesotrophic Lake Gjersjøen. In general, the seasonal effects on phytoplankton biomass were more marked than the effects of biomanipulation. Primary top-down effects of fish on zooplankton were conspicuous in all bags, whereas control of phytoplankton growth by grazing was observed only in the nutrient-limited summer situation. The effect of nutrient additions was pronounced in summer, less in spring and autumn; additions of fish gave the most pronounced effect in spring. The phytoplankton/zooplankton biomass ratio remained high (10–100) in bags with fish, with the highest ratios in combination with fertilization. The ratio decreased in bags without fish to<2 in most bags, but a real grazing control was only observed in bags with addition ofDaphnia. No direct grazing effects could be observed on the absolute or relative biomass of cyanobacteria (mainlyOscillatoria agardhii). The share of cyanobacteria in total phytoplankton biomass was lowest in summer (7–26%), higher in spring (39–63%) and more than 90% in the autumn experiment. The development of the cyanobacterial biomass was rather synchronous in all bags in all the three experiments. A high biomass ofDaphnia gave no increase in the pool of dissolved nutrients in spring, a slight increase in summer and a pronounced increase in autumn. While a strong decrease in the P/C-cell quota of the phytoplankton was observed from spring to autumn, no effect of grazing or nutrient release could be related to this P/C-status. The experiments indicate that such systems, with high and stable densities of inedible cyanobacteria, are rather insensitive to short-term (3–4 weeks) biomanipulation efforts. This is supported by observations on the long-term development of the lake.

     

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.     

Nutrient Enrichment and Planktonic Biomass Ratios in Lakes

Phosphorus (P) to chlorophyll ratios and zooplankton–phytoplankton (Z:P) biomass ratios were assessed in 400 temperate lakes over a gradient of phosphorus (P) and with different fish communities. Most of the lakes in this survey were oligotrophic, with a median total P of 7.3 μg P L⁻¹. Thus, the survey provided information on food web effects during the early phase of eutrophication. There was no tendency toward a reduced yield of autotrophs per unit of P over the gradient covered in this survey. The zooplankton yield per unit of P or chlorophyll a decreased slightly with increased nutrient concentrations, and Z:P biomass ratios decreased with fish community classes, reflecting increased fish predation pressure. However, the variability in biomass ratios within a given range of P and fish class was some 100 times higher than the difference over the gradients. This finding suggests that lake-specific properties, community composition, and food quality are by far the most important determinants of biomass ratios and probably also trophic efficiency in lakes; it further suggests that these factors are superimposed on the general effect of eutrophication, at least up to 30 μg P L⁻¹.     

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

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

The impact of year-to-year changes in the weather on the dynamics of Daphnia in a thermally stratified lake

The factors influencing the seasonal dynamics of Daphnia in a thermally stratified lake (Esthwaite Water) are described and related to long-term changes in the weather. The Daphnia produced three cohorts in the year and the strength of the cohorts was determined by year-to-year variations in the physical characteristics of the lake and the abundance of edible algae. Food was most abundant in early summer when small, fast-growing flagellates were particularly common. In late summer, the phytoplankton community was dominated by large, inedible species but edible forms re-appeared when nutrients were entrained by wind mixing. Examples are presented to demonstrate the effect that year-to-year variations in the weather have on the growth of the phytoplankton and the dynamics of the Daphnia. In ‘good’ years, when the lake stratifies early and there are periods of episodic mixing in summer, there are two ‘pulses’ of edible algae and two strong cohorts of Daphnia. In ‘bad’ years when stratification is delayed and there is little episodic mixing, the growth of the edible algae is suppressed and the Daphnia produce two weak cohorts. The results are discussed in relation to the impact of intermediate disturbances on growth of phytoplankton and current theories of population regulation in Daphnia. The evidence suggests that the dynamics of the Daphnia in the lake are strongly influenced by seasonal variations in the mixing regime, the recycling of nutrients and the episodic growth of edible algae.     

Changes in the fish and zooplankton communities of Ringsjön,a Swedish lake undergoing man-made eutrophication

During the 20th century Lake Ringsjön has developed from a mesotrophic to a eutrophic lake, and the phytoplankton community has changed from a rather diverse community to a monoculture of blue-green algae. The eutrophication process has accelerated during the last decade. The most important external nutrient loading of today comes from agriculture.Although phosphorus has been shown to be the primary nutrient leading to excessive algal growth in fresh water, several biotic factors — such as interactions between nutrients, phytoplankton, zooplankton and planktivorous fish — may play a decisive role in the occurrence and maintenance of large algal blooms.The aim of this investigation was to study the changes in the fish community of Lake Ringsjön, especially the most dominant planktivores, and the state of the zooplankton community during the seventies. The fish fauna is dominated by cyprinids, especially roach, and there are relatively few predatory fish. During the seventies the mean size of roach decreased, and measurements of the zooplankton community indicated that the predation pressure on zooplankton had increased. The mean sizes of cladocerans such as Daphnia and Bosmina, which were selected for by the planktivorous fish, decreased; the size of the calanoid Diaptomus, which was not preyed upon by the dominating fish species, did not change. The growth of zooplankton-feeding stages of several fish species was retarded, which meant that the growth of young perch decreased, while older roach were mainly affected. In the prevailing situation, planktivorous roach can maintain a numerous population of small individuals, whereas the predatory perch is at a disadvantage, and predation on zooplankton is intense.     

Does stocking of filter-feeding fish for production have a cascading effect on zooplankton and ecological state? A study of fourteen (sub)tropical Chinese reservoirs with contrasting nutrient concentrations

Stocking of filter-feeding fish is a common tool used in Chinese reservoirs to increase fish production because of low natural recruitment. Whether such stocking has important negative effects on zooplankton with cascading effects on phytoplankton is debated. We compared the zooplankton communities in fourteen reservoirs with different nutrient concentrations and fish densities. Both chlorophyll a (Chla) and fish catch were positively related with total phosphorus (TP), whereas zooplankton biomass did not show a similar relationship with TP. Zooplankton seemed to be influenced by fish as high fish catches coincided with a low proportion of calanoids of the total copepod biomass, a high proportion of rotifers of the total zooplankton biomass, a low zooplankton:phytoplankton biomass ratio, and the absence of Daphnia irrespective of TP concentration. Both zooplankton biomass and most of the zooplankton:phytoplankton biomass ratios were among the lowest reported in the literature for the nutrient range studied. Furthermore, the Chla:TP ratio was higher than what is typically observed in temperate lakes. We conclude that top-down control of zooplankton is of key importance in reservoirs in South China where frequent stocking of filter-feeding fish seems to contribute to poor water quality in the form of higher algal biomass and reduced clarity.     

Goose-mediated nutrient enrichment and planktonic grazer control in arctic freshwater ponds

A dramatic increase in the breeding population of geese has occurred over the past few decades at Svalbard. This may strongly impact the fragile ecosystems of the Arctic tundra because many of the ultra-oligotrophic freshwater systems experience enrichment from goose feces. We surveyed 21 shallow tundra ponds along a gradient of nutrient enrichment based on exposure to geese. Concentrations of total phosphorus (P) and dissolved inorganic nitrogen (DIN) in the tundra ponds ranged from 2–76 to 2–23 μg l⁻¹ respectively, yet there was no significant increase in phytoplankton biomass (measured as chlorophyll a; range: 0.6–7.3 μg l⁻¹) along the nutrient gradient. This lack of response may be the result of the trophic structure of these ecosystems, which consists of only a two-trophic level food chain with high biomasses of the efficient zooplankton grazer Daphnia in the absence of fish and scarcity of invertebrate predators. Our results indicate that this may cause a highly efficient grazing control of phytoplankton in all ponds, supported by the fact that large fractions of the nutrient pools were bound in zooplankton biomass. The median percentage of Daphnia–N and Daphnia–P content to particulate (sestonic) N and P was 338 and 3009%, respectively, which is extremely high compared to temperate lakes. Our data suggest that Daphnia in shallow arctic ponds is heavily subsidized by major inputs of energy from other food sources (bacteria, benthic biofilm), which may be crucial to the persistence of strong top–down control of pelagic algae by Daphnia.     

Zooplankton community structure and environmental conditions in a set of interconnected ponds

We studied the zooplankton community structure in a set of 33 interconnected shallow ponds that are restricted to a relatively small area (`De Maten', Genk, Belgium, 200 ha). As the ponds share the same water source, geology and history, and as the ponds are interconnected (reducing chance effects of dispersal with colonisation), differences in zooplankton community structure can be attributed to local biotic and abiotic interactions. We studied zooplankton community, biotic (phytoplankton, macrophyte cover, fish densities, macroinvertebrate densities), abiotic (turbidity, nutrient concentrations, pH, conductivity, iron concentration) and morphometric (depth, area, perimeter) characteristics of the different ponds. Our results indicate that the ponds differ substantially in their zooplankton community structure, and that these differences are strongly related to differences in trophic structure and biotic interactions, in concordance with the theory of alternative equilibria. Ponds in the clear-water state are characterised by large Daphnia species and species associated with the littoral zone, low chlorophyll-a concentrations, low fish densities and high macroinvertebrate densities. Ponds in the turbid-water state are characterised by high abundances of rotifers, cyclopoid copepods and the opposite environmental conditions. Some ponds show an intermediate pattern, with a dominance of small Daphnia species. Our results show that interconnected ponds may differ strongly in zooplankton community composition, and that these differences are related to differences in predation intensity (top-down) and habitat diversity (macrophyte cover).     

The influence of plant-associated filter feeders on phytoplankton biomass: a mesocosm study

Low phytoplankton biomass usually occurs in the presence of submerged macrophytes, possibly because submerged macrophytes enhance top-down control of phytoplankton by offering a refuge for efficient grazers like Daphnia against fish predation. However, other field studies also suggest that submerged macrophytes suppress phytoplankton in the absence of Daphnia. In order to investigate these mechanisms further, we conducted an outdoor mesocosm experiment to study the effect of submerged macrophytes (Elodea nuttallii) on phytoplankton and zooplankton biomass. The experiment combined four nutrient addition levels (0, 10, 100, and 1000 μg P l⁻¹; N/P ratio: 16) with three macrophyte levels (no macrophytes, artificial macrophytes, and real macrophytes). We inoculated the tanks with species-rich inocula of phytoplankton and zooplankton but excluded fish or macro-invertebrates. Probably due to the lack of predators in the mesocosms, potential grazing rates of pelagic zooplankton (estimated from zooplankton biomass) did not differ between the macrophyte treatment combinations. Compared to the treatment combinations without macrophytes, lower phytoplankton biomass occurred in the treatment combinations with real macrophytes at all the nutrient addition levels and in those with artificial macrophytes at all the nutrient levels except the highest. Significantly, higher abundances of plant-associated filter feeders (Simocephalus vetulus and Ceriodaphnia spp.) occurred in the treatment combinations with real and artificial macrophytes. The estimated potential grazing rate of these plant-associated filter feeders indicated that these filter feeders could be responsible for the lower phytoplankton biomass in the presence of real and artificial macrophytes. Our results suggest that the plant-associated filter feeders may be significant grazers in vegetated shallow lakes.     

Impact of the fish Garra on the ecology of reservoirs and the occurrence of Microcystis blooms in semi-arid tropical highlands: an experimental assessment using enclosures

1. Many man‐made reservoirs in the semi‐arid highlands of Northern Ethiopia (Tigray) are characterised by the occurrence of intensive blooms of cyanobacteria and a dominance of small riverine fishes belonging to the genus Garra. 2. We carried out enclosure experiments to test for the effect of these small fish on abiotic characteristics, phytoplankton biomass and zooplankton community structure in the pelagic of two reservoirs (Gereb Awso and Tsinkanet). Two experiments were carried out in each of the reservoirs, one at the end of the rainy season (highest water level) and one at the end of the dry season (lowest water level). 3. The presence of Garra in general increased the amount of suspended matter, nutrient concentrations (total nitrogen and total phosphorus), phytoplankton and Microcystis biomass (including the proportion of Microcystis in the phytoplankton community), and reduced water transparency. The positive effect of the presence of Garra on nutrient concentrations and phytoplankton productivity indicate that Garra has the potential to affect food web functioning indirectly through bottom‐up effects, by enhancing nutrient concentrations through sediment resuspension and excretion of nutrients. Indeed, population densities of the cladoceran zooplankton taxa Ceriodaphnia and Diaphanosoma also showed an overall increase in enclosures with Garra. 4. However, our data also provide some evidence for a potential of Garra to exert top‐down control on large bodied daphnids (Daphnia carinata, D. barbata), although such effect varied among experiments. The limited capability of Garra to control zooplankton communities mainly reflects the low efficiency of these small, riverine and benthos‐oriented fish in foraging on zooplankton and suggests the existence of an unoccupied niche for zooplanktivorous fish in the majority of the reservoirs. 5. Although the main effects of Garra on the pelagic food web seemed to be mediated by bottom‐up mechanisms, our results also indicate that one of the key variables, the relative abundance of Microcystis, was impacted by Daphnia‐mediated trophic cascade effects.     

Studies on the Effects of Silver Carp (Hypophthalmichthys molitrix) on the Phytoplankton in a Shallow Hypereutrophic Lake through an Enclosure Experiment

The responses of nutrients, water transparency, zooplankton and phytoplankton to a gradient of silver carp biomass were assessed using enclosure methods. The gradient of four silver carp biomass levels was set as follows: 0, 116, 176 and 316 g m^—2. Nutrients did not show any statistically significant differences among the treatments. An outburst of Daphnia only occurred in fishless enclosures where phytoplankton biomass was the lowest and water clarity significantly increased. While among fish enclosures, the small‐sized Moina micrura dominated throughout the experiment and both zooplankton and phytoplankton biomasses decreased with increased fish biomass. No large colonial cyanobacterial blooms occurred in the fishless enclosures as predicted. This might be due to low water temperature, short experiment time and the occurrence of large bodied Daphnia in our experiment. Cryptophyta was the most dominant group in most of the enclosures and the lake water throughout the experiment. The fishless enclosure had much lower proportion of Cyanophyta but higher proportion of Trachelomonas sp.     

Elemental stoichiometry of lower food web components in arctic and temperate lakes

Lakes were surveyed to assess the potential patterns of latitudinalvariation in carbon:nitrogen:phosphorus (C:N:P) stoichiometryof lower food web components. Thirty-four lakes were surveyedat an arctic latitude (68°38'N, 149°38'W) and 10 lakesat a temperate latitude (46°13'N, 89°32'W) during 1997.The temperate data set was augmented with earlier survey resultsemploying similar methods. It was hypothesized that differencesin environmental variables across latitude would cause differencesin community C:N:P ratios, leading to differences in trophicinteractions. Physical measurements (light, temperature), sestonand zooplankton were collected from each lake. Seston and zooplanktonwere analyzed for C, N and P content, and zooplankton were countedand measured for biomass estimates. The degree of trophic interactionbetween seston and zooplankton was assessed by (i) measuringelemental imbalances between seston and zooplankton and (ii)calculating the potential recycling ratio by the zooplanktoncommunity available for seston. Seston C:nutrient, but not N:P,ratios were higher in temperate than arctic lakes. Conversely,arctic zooplankton had higher C:nutrient, but not N:P, ratiosthan zooplankton in temperate lakes. Elemental imbalances weregreater in temperate than in arctic lakes, but N:P stoichiometryof potential zooplankton recycling was nearly identical betweenthe two latitudes. Zooplankton community C:N:P ratios were notrelated to either latitude or seston C:N:P. In accordance withstoichiometric theory, relative abundances of calanoid copepodswere positively correlated with seston C:N in temperate lakes.Additionally, relative abundances of Daphnia were negativelycorrelated with seston C:N ratios in temperate and arctic lakes,and positively correlated with N:P ratios in the arctic. Ingeneral, these results suggest that seston and zooplankton communitystoichiometry differ across latitude, and these differenceshave the potential to affect trophic interactions.