Food-web manipulations influence grazer control of phytoplankton growth rates in Lake Michigan

Stocking piscivorous salmonids in Lake Michigan produced dramaticalterations in food-web structure, including higher numbersof large-bodied zooplankton (especially Daphnia pulicaria),lower summer chlorophyll concentrations and increased watertransparency. Experimental determinations of epilimnetic phytoplanktongrowth rates and of zooplankton grazing rates indicate thatherbivorous zooplankton controlled algal dynamics during thesummer of 1983 because grazers occupied the surface waters throughoutthe day. In 1985, however, both large- and small-bodied Daphniamade approximately equal contributions to total grazer biomass,and all grazers displayed pronounced diel vertical migrations,visiting epilimnetic waters only at night. This prohibited zooplanktonfrom controlling algal dynamics because grazing losses did notexceed phytoplankton growth rates. The changes in zooplanktoncommunity composition and behavior observed in summer 1985 probablyresulted from increased predation by visually orienting planktivorousfish, especially bloater chub (Coregonus hoyi). Effects of food-webmanipulations on phytoplankton dynamics were evident only duringJuly and August. During spring and early summer copepods dominateLake Michigan's zooplankton community. Owing to their smallbody size, copepods are less susceptible to fish predation andexhibit much lower filtering rates than Daphnia. Variabilityin zooplanktivorous fish abundance probably has little effecton phytoplankton dynamics during spring and early summer.     

Zooplankton responses to predation by larval bluegill: an enclosure experiment

SUMMARY 1. Larval fish are gape-limited predators that forage on prey of specific sizes, and thus may be expected to differentially affect members of a zooplankton community, possibly altering the size-structure or species composition.
2. I used an enclosure experiment to look at the effect of predation by larval bluegill on the dynamics of two zooplankton communities, one dominated by large-bodied individuals and the other by small-bodied individuals. Enclosures containing these zooplankton received a zero, low, medium, or high density of larval bluegill predators.
3. Increasing larval density had a negative effect on zooplankton abundance and abundance declined similarly in the large-bodied and small-bodied communities.
4. Zooplankton size-structure, as estimated by the length of the average zooplankton, increased and then decreased during the experiment, decreasing faster at higher larval fish densities. When zooplankton size-structure was estimated as the length of the average cladoceran, size-structure declined in the large-bodied but not in the small-bodied community and the greatest decline in size-structure was seen in the medium and high larval density treatments.
5. Ordination of each community using multidimensional scaling (MDS) indicated that the trajectory of change in species composition differed between the presence and absence of larval fish. In both communities, the degree of response by individual taxa depended on the density of bluegill larvae. This effect on zooplankton abundance, size-structure and community composition suggests that larval fish may make an important contribution to zooplankton dynamics in many lakes and ponds.     

Effects of the blue-green alga Microcystis aeruginosa on zooplankton competitive relations

Summary Field distribution patterns and laboratory feeding experiments have suggested that blooms of colonial blue-green algae strongly inhibit relatively large-bodied daphnid cladocerans. We conducted laboratory experiments to test the hypothesis that blooms of the colonial blue-green alga Microcystis aeruginosa would shift competitive dominance away from large-bodied daphnid cladocerans toward smaller-bodied cladocerans, copepods, and rotifers. In laboratory competition experiments, increasing the proportion of M. aeruginosa in the algal food supply resulted in a shift from dominance by the relatively largebodied cladoceran Daphnia ambigua to dominace by the copepod Diaptomus reighardi. The small-bodied cladoceran Bosmina longirostris was always numerically heavily dominant over D. ambigua, but its estimated population biomasses were only slightly higher than those of D. ambigua. Daphnia ambigua consistently outcompeted the rotifer Brachionus calyciflorus. Our results demonstrate that blooms of M. aeruginosa can alter zooplankton competitive relations in laboratory experiments, favoring small-bodied cladocerans and copepods at the expense of large-bodied cladocerans. However, contrary to predictions, blooms of M. aeruginosa did not improve the competitive ability of rotifers.     

Trophic interactions in rockpool food webs: regulation of zooplankton and phytoplankton by Notonecta and Daphnia

We studied trophic interactions in experimental rockpools with three different food web structures: phytoplankton and small-bodied zooplankton; phytoplankton, small-bodied zooplankton and Daphnia ; and phytoplankton, small-bodied zooplankton, Daphnia and Notonecta . Nutrients, primary productivity, chlorophyll a and zooplankton species composition and biomass were measured over eight weeks.
2. Daphnia had a negative impact on other zooplankton and reduced the phytoplankton biomass and primary productivity. In the absence of Daphnia , small-bodied zooplankton species were abundant, in particular cyclopoid copepods. Concentrations of dissolved nutrients were lower and the standing crop of primary producers was higher when Daphnia was absent.
3. The presence of the invertebrate predator Notonecta produced a top-down effect which was similar to that reported for planktivorous fish, i.e. a selective reduction of daphnids followed by an increase of small-bodied zooplankton species and phytoplankton biomass.
4. The study showed that consumer regulation of Daphnia by Notonecta and of algae by Daphnia are important, but also demonstrated that trophic level biomasses were controlled by a combination of predation and resource limitation.     

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.     

An artificial neural network approach for studying phytoplankton succession

Artificial neural networks are used to model phytoplankton succession and gain insight into the relative strengths of bottom-up and top-down forces shaping seasonal patterns in phytoplankton biomass and community composition. Model comparisons indicate that patterns in chlorophyll aconcentrations response instantaneously to patterns in nutrient concentrations (phosphorous (P), nitrite and nitrate (NO₂/NO₃–N) and ammonium (NH₄–H) concentrations) and zooplankton biomass (daphnid cladocera and copepoda biomass); whereas lagged responses in an index of algal community composition are evident. A randomization approach to neural networks is employed to reveal individual and interacting contributions of nutrient concentrations and zooplankton biomass to predictions of phytoplankton biomass and community composition. The results show that patterns in chlorophyll aconcentrations are directly associated with P, NO₂/NO₃–N and daphnid cladocera biomass, as well as related to interactions between daphnid cladocera biomass, and NO₂/NO₃–N and P. Similarly, patterns in phytoplankton community composition are associated with NO₂/NO₃–N and daphnid cladocera biomass; however show contrasting patterns in nutrient– zooplankton and zooplankton–zooplankton interactions. Together, the results provide correlative evidence for the importance of nutrient limitation, zooplankton grazing and nutrient regeneration in shaping phytoplankton community dynamics. This study shows that artificial neural networks can provide a powerful tool for studying phytoplankton succession by aiding in the quantification and interpretation of the individual and interacting contributions of nutrient limitation and zooplankton herbivory on phytoplankton biomass and community composition under natural conditions.     

Low fish predation pressure in London reservoirs: II. Consequences to zooplankton community structure

The 1992 survey of zooplankton structure in fourteen London supply reservoirs showed the overall dominance of large-bodied zooplankton, mainly species of Daphnia. These reservoirs can be considered as ‘anti-fish’ by virtue of their steeply sloping concrete or brick sides. The average biomass of large Daphnia spp (retained on a 710 μm sieve) in the total zooplankton biomass was higher than 20% for twelve out of fourteen reservoirs. The cladoceran-copepod ratio was inversely correlated with both dominance of large-bodied Daphnia magna and cladoceran body-size structure. Parallelly, there were tendency of more efficient utilization of lowered algal crops in reservoirs dominated by large-bodied Daphnia spp. A graphical model is presented which relates daphnid species composition and zooplankton size structure to a presumed gradient of fish biomass in these reservoirs.     

The potential of artificial refugia for maintaining a community of large-bodied cladocera against fish predation in a shallow eutrophic lake

The Norfolk Broads are a series of shallow lakes which are highly eutrophic and typified by dense populations of phytoplankton and an absence of submerged aquatic plants. The zooplankton community is subject to intense predation pressure by young fish and is dominated by small-bodied organisms which have a low potential for reducing phytoplankton populations through grazing. Various designs and densities of artificial refugia for zooplankton against fish predation were established in Hoveton Great Broad in order to enhance populations of large-bodied Cladocera. Initially some of the refuges contained higher densities and larger individuals ofDaphnia andCeriodaphnia than the surrounding open water. However, towards the end of the first season and throughout the subsequent two years, population densities and size-structure were similar both within and outside the refuges, although there was still evidence of enhanced body-size ofDaphnia within the refuges compared with the open water. The provision of habitat structures designed as refugia from fish predation did not enhance large-bodied cladoceran populations enough to promote this restoration strategy as feasible for eutrophic and shallow lakes.     

Interactions between Nile tilapia (Oreochromis niloticus) and cladocerans in ponds (Khartoum,Sudan)

Juvenile Nile tilapia (Oreochromis niloticus) are omnivorous, and the question asked in this study is how they affect on their environment? Do they mainly act as predators on the cladoceran zooplankton or do they compete with the cladocerans for phytoplankton? This problem was studied in three ponds with and three ponds without small tilapia (3–5 cm). The fish growth rate, the succession of plankton species and the changes in abiotic conditions, were monitored over a period of 67 days. The fish biomass was kept low and the mean was approximately constant (12.6 g m^?2) during the experiment. Phosphate was added to avoid phytoplankton nutrient limitation. Although the diet of Nile tilapia contained both phytoplankton and zooplankton, the fish affected the ecosystem in a similar way as zooplanktivorous fish. The fish ponds got more phytoplankton due to increase of Chlorophyta. Effects on the other phytoplankton groups Euglenophyta, Bacillariophyta, Cryptophyta and Cyanophyta could not be registered. The ponds without fish had higher densities of Daphnia lumholtzi and D. barbata. The other Cladocerans seemed less influenced by fish presence. The relative fish growth rate was most positively correlated with the density of Daphnia lumholtzi, Diaphanosmoa excisum and Bosmina longirostris. Tilapia seemes to have two feeding modes: (1) preying on large zooplankton and (2) unselective filtration of small planktonic organisms such as phytoplankton. In our experiment the first feeding mode affected the ecosystem more than the second.     

The loss of submerged plants with eutrophication II. Relationships between fish and zooplankton in a set of experimental ponds, and conclusions

SUMMARY. 1. In 1982 and 1983 sets of experimental ponds were left with their submerged plant communities intact (plant ponds) or were cleared manually of them (cleared ponds). The ponds were all fertilized with ammonium nitrate and with variable amounts of phosphate. In 1982 fish were removed from the ponds. Zooplankton communities were dominated by large Cladocera with Daphnia prominent in the cleared ponds and Simocephalus in the plant ponds. There was no detectable effect of differential phosphorus additions on zooplankton communities or populations.
2. In 1983 zooplanktivorous fish (mainly roach) were stocked in the ponds. In the plant ponds the fish did not survive, probably through severe deoxygenation and the zooplankton community again included large-bodied Simocephalus. Fish survival was variable in the cleared ponds. Where fish stocks were absent or low (0.5–1 g m⁻²) a Daphnia- dominated community persisted; at intermediate fish stocks (18.1 g m⁻²) Eudiaptomus gracilis was predominant and where fish stock was high (22.8–29.1 g m⁻²) Bosmina longirostris , and cyclopoid copepods dominated the communities. Mean biomass of the zooplankton community declined with increase in fish stock to between 5.1 and 18.1 g m⁻² then increased.
3. On the basis of results from the experimental ponds and elsewhere, a new hypothesis is put forward to account for the switch from aquatic plant to phytoplankton dominance in eutrophicated shallow lakes. It envisages dominance by either group to be possible as alternative states over a wide range of high nutrient loadings. It suggests that each state is buffered against increased loading by mechanisms involving plant and algal physiology and zooplankton grazer populations. The nature of the buffers and the reasons by which one state may be switched to the other are, discussed.     

A review: limnological management and biomanipulation in the London reservoirs

Low algal biomasses and high water transparencies are a feature of the storage reservoirs that supply most of London's treated water. This is a result of knowledgeable limnological management and biomanipulation and despite the eutrophic nature of the River Thames with its high nutrients (7 gN m⁻³; 1 gP m⁻³) and particulate organic carbon (2 gC m⁻³). Built-in possibilities of jetting input water are managed to prevent stratification, to ensure isothermy, to mix chemicals and plankton vertically and horizontally and to manipulate the mixed-depth of the algal populations such that their potential for biomass growth is reduced by light-energy limitation. Spring algal growth is delayed and the spring peak is reduced and curtailed by the grazing impact of considerable biomasses of large-bodied daphnid populations (Daphnia magna, pulicaria & hyalina) whose development is also supported by the continuous input of high riverine algal crops. The existence of a large-bodied daphnid zooplankton in the reservoirs is associated with low levels of fish predation since the late 1960s. Variations in the intensity and nature of this vertebrate predation during the subsequent twenty years (1968–88) are illustrated by the changes that have occurred in the relationship between the phytoplankton and zooplankton biomasses of the April-May-June quarter of the year. This example of the London reservoirs serves to illustrate biomanipulation in deep water bodies by bottom-up as well as top-down effects.     

Predation-driven dynamics of zooplankton and phytoplankton communities in a whole-lake experiment

Summary 1. Species compositions of zooplankton and phytoplankton were followed in Tuesday Lake before and after experimental manipulation of its fish populations (addition of piscivorous largemouth bass, removal of planktivorous minnows). Plankton dynamics were compared to those of adjacent, unmanipulated Paul Lake, where piscivorous fish have been dominant historically. 2. Indices of similarity for the zooplankton communities in the two lakes in 1984 prior to the manipulation were low; however, following the manipulation in spring, 1985, similarity of the zooplankton in the two lakes rose considerably and remained high throughout 1986. This was the result of an increase in Tuesday Lake of previously rare large-bodied cladocerans (Daphnia pulex, Holopedium gibberum) which were the dominants in Paul Lake, and the disappearance in Tuesday Lake of the dominant small-bodied copepod Tropocyclops prasinus, a minor component of the Paul Lake zooplankton. These observations are consistent with prior observations of the effects of size-selective predation on zooplankton communities. 3. Phytoplankton communities also responded strongly to the manipulation, with similarity indices for the two lakes rising from low levels in 1984 to high levels of similarity in 1985 and 1986, reflecting the decrease of formerly dominant Tuesday Lake taxa which were unimportant in Paul Lake and the appearance or increase in Tuesday Lake of several taxa characteristic of the Paul Lake phytoplankton assemblage. these results clearly show that food web structure can have pronounced effects on community composition at all levels of the food web, and that, just as zooplankton communities are structured by sizeselective predation, phytoplankton communities are structured by herbivory. These observations may provide some insight into factors governing the complex distributions of phytoplankton species among various lakes.A contribution from the University of Notre Dame Environmental Research Center, funded by NSF grants BSR-83-08918 and BSR-86-06271     

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.

     

Algal picoplankton production and contribution to food-webs in oligotrophic British Columbia lakes

The pelagic communities of two contrasting oligotrophic lakes in British Columbia were studied to determine why an interior, dimictic lake (Quesnel) supports a greater biomass of zooplankton and produces larger planktivorous sockeye salmon (Oncorhynchus nerka) than a coastal warm-monomictic lake (Sproat). The ultra-oligotrophic status and differing planktivore densities in Sproat Lake increased the relative importance of algal picoplankton, diminished the abundance of large zooplankton, and increased the significance of rotifers and other small-bodied zooplankton. These picoplankton based food webs result in longer, indirect and less efficient pathways of carbon flow from phytoplankton to fish. In contrast, Quesnel Lake is a more productive oligotrophic lake and its pelagic food webs are based more on nanoplankton and small microphytoplankton that support larger-bodied zooplankton (Daphnia, Diaptomus), and a more direct and efficient two-step transfer to fish. The greater variability of the annual recruitment of sockeye fry in interior lakes may keep zooplankton communities in a non-steady state, this in turn may perpetuate the occurrence of quadrennial cyclic dominance in adult salmon returning to these systems.     

Food web complexity affects stoichiometric and trophic interactions

The stoichiometry of trophic interactions has mainly been studied in simple consumer–prey systems, whereas natural systems often harbour complex food webs with abundant indirect effects. We manipulated the complexity of trophic interactions by using simple laboratory food webs and complex field food webs in enclosures in Lake Erken. In the simple food web, one producer assemblage (periphyton) and its consumers (benthic snails) were amended by perch, which was externally fed by fish food. In the complex food web, two producer assemblages (periphyton and phytoplankton), their consumers (benthic invertebrates and zooplankton) and perch feeding on zooplankton were included. In the simple food web perch affected the stoichiometry of periphyton and increased periphyton biomass and the concentration of dissolved inorganic nitrogen. Grazers reduced periphyton biomass but increased its nutrient content. In the complex food web, in contrast to the simple food web, perch affected periphyton biomass negatively but increased phytoplankton abundance. Perch had no influence on benthic invertebrate density, zooplankton biomass or periphyton stoichiometry. Benthic grazers reduced periphyton biomass and nutrient content. The difference between the simple and the complex food web was presumably due to the increase of pelagic cyanobacteria ( Gloeotrichia sp.) with fish presence in the complex food web, thus fish had indirect negative effects on periphyton biomass through nutrient competition and shading by cyanobacteria. We conclude that the higher food web complexity through the presence of pelagic primary producers (in this case Gloeotrichia sp.) influences the direction and strength of trophic and stoichiometric interactions.     

Zooplankton body size and community structure: Effects of thermal and toxicant stress

Episodic heat waves and an increase in pesticide use are widely cited as consequences of climatic warming. Recent studies show that these stressors often cause declines in the mean body size of zooplankton. Results from laboratory and field studies, as well as observations from both thermal- or toxicant-stressed natural systems, show (1) reductions in mean body size within stressed populations, or (2) changes in community composition that favor small-bodied over large-bodied species. During the past decade, it has become widely accepted that a shift in zooplankton body size can dramatically affect water clarity, rates of nutrient regeneration and fish abundances. Thus, climatic warming and associated change in pesticide use has the potential to cause striking change in the structure and functioning of temperate-zone lakes.     

Grazing effects of a freshwater bivalve (Corbicula leana Prime) and large zooplankton on phytoplankton communities in two Korean lakes

This study examined the effects of a freshwater filter feeding bivalve (Corbicula leana Prime) and large zooplankton (>200 μm, mostly cladocerans and copepods) on the phytoplankton communities in two lakes with contrasting trophic conditions. A controlled experiment was conducted with four treatments (control, zooplankton addition, mussel addition, and both zooplankton and mussel addition), and each established in duplicate 10-l chambers. In both lakes there were significant effects of mussel grazing on phytoplankton density and biomass. The effects were greater in mesotrophic Lake Soyang than in hypertrophic Lake Ilgam. Effects of zooplankton grazing did not differ between these lakes, and zooplankton effects on phytoplankton were much less than the effects of mussels. Although mussels exerted a varying effect on phytoplankton according to their size, mussels reduced densities of almost all phytoplankton taxa. Total mean filtering rate (FR) of mussels in Lake Soyang was significantly greater than that in Lake Ilgam (p=0.002, n=5). Carbon fluxes from phytoplankton to mussels (977–2,379 μgC l^?1d^?1) and to zooplankton (76–264 μgC l^?1 d^?1) were always greater in Lake Ilgam due to the greater phytoplankton biomass (p<0.01, n=6). Based on the C-flux to biomass ratios, the mussels consumed 170–754% (avg. 412%) of phytoplankton standing stock in Lake Soyang, and 38–164% (avg. 106%) in Lake Ilgam per day. The C-flux to biomass ratio for mussels within each lake was much greater than for large zooplankton. Mussels reduced total phosphorus concentration by 5–34%, while increasing phosphate by 30–55% relative to the control. Total nitrogen also was reduced (by 9–25%), but there was no noticeable change in nitrate among treatments. The high consumption rate of phytoplankton by Corbicula leana even in a very eutrophic lake suggests that this mussel could affect planktonic and benthic food web structure and function by preferential feeding on small seston and by nutrient recycling. Control of mussel biomass therefore might be an effective tool for management of water quality in shallow eutrophic lakes and reservoirs in Korea.     

Cascading effects of the flathead grey mullet Mugil cephalus in freshwater eutrophic microcosmos

Flathead grey mullet (Mugil cephalus L.) stocked in fish ponds were long considered to feed primarily on detritus. However, recent research has found that they obtain much of their food from plankton and that they have a detrimental effect on pond zooplankton and large phytoplankton, whilst enhancing small phytoplankton. It has been suggested that flathead grey mullet may also increase the internal phosphorus loading of the ecosystem, which would also increase phytoplankton density. To test whether zooplankton removal or nutrient release from the sediment is the better explanation for phytoplankton enhancement in the presence of flathead grey mullet, the ecosystems of fish-less tanks, tanks with a 60 m mesh filter and tanks stocked at a fish density of 243 g m^-3 were compared. In the presence of flathead grey mullets, cladocerans, ostracods and chironomid larvae became scarcer than in the control tanks, while there were more small phytoplankton and mud snails. The green algae Cladophora sp. did not occur at all. The presence of a mechanical filter also reduced cladoceran, ostracod and chironomid densities and increased phtyoplankton and mud snail density. However, the situation observed in filter tanks was intermediate between that observed in the fish tanks and the control tanks, due to the lower filtering efficiency of the mechanical filter. The organic matter content of the sediment decreased throughout the experiment in the control and filter tanks, but remained stable in fish tanks. Phosphorus and nitrogen concentrations were not affected by any treatment. These results showed that flathead grey mullet enhanced phytoplankton development due to the removal of large cladocerans and not as a consequence of nutrient release. Furthermore, the flathead grey mullet strongly modified the benthic community, probably due to direct predation.     

Relationship between nutrient concentration,phytoplankton density,and zooplankton density in nutrient enriched experimental ponds

The effects of different levels of nutrient input on the plankton community was investigated in a two-year controlled fertilization study of eight experimental ponds. There were four treatments, each replicated: a control, to which no fertilizer was added, and three levels of nutrient addition. Limnological parameters including phytoplankton and zooplankton densities were measured frequently during both summers and less frequently during the rest of the year. Inorganic nitrogen and phosphorus concentrations in the treated ponds increased. Phytoplankton and zooplankton density increased with treatment level but was variable. There was a limited relationship between the average chlorophyll a concentration per summer and the average cladoceran dry weight per summer. Above chlorophyll a concentrations greater than 60–70 mg/m³ other factors such as a pH zooplankton mortality effect, prevailed.     

Long-term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Müggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Müggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N₂‐fixing species Aphanizomenon flos‐aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re‐colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Müggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.