Contemporary state of the zooplankton in Lake Peipsi

L. Peipsi is one of the richest fish lakes in Europe. Planktivorous smelt dominates in the fish fauna. The abundance of zooplankton fluctuates between 43 600–2241 500 ind m^–3, with the average 974 000 ind m^–3, biomass ranges from 0,09–3,69 g m^–3, with the average 1,86 g m^–3. Since the 1960s the abundance of rotifers has risen considerably while the mean zooplankter weight (B/N) has decreased from 0.005 mg to 0.004 mg. Zooplankton production (herbivores 20.6, predators 1.8, whole zooplankton community 22.4 g C m^–2 per period between May and October) can be considered high. Predatory zooplankton eats on an average 50% of the production of herbivorous zooplankton; about 50% of the whole zooplankton production (P_{Filt + Pred}) reaches fishes. The production of herbivorous zooplankton constitutes 10.1% of primary production. This ratio indicates a direct relationship between zoo- and phytoplankton in the food chain; the detrital food chain seems of little importance. About 6% of phytoplankton energy reaches fishes. The transformation of energy in the food web is efficient. On the basis of zooplankton L. Peipsi can be considered a moderately eutrophic or meso-eutrophic lake.     

A comparison of shallow Danish and Canadian lakes and implications of climate change

1. Winter temperatures differ markedly on the Canadian prairies compared with Denmark. Between 1 January 1998 and 31 December 2002, average weekly and monthly temperatures did not drop below 0 °C in the vicinity of Silkeborg, Denmark. Over this same time, weekly average temperatures near Calgary, Alberta, Canada, often dropped below −10 °C for 3–5 weeks and the average monthly temperature was below 0 °C for 2–4 months. Accordingly, winter ice conditions in shallow lakes in Canada and Denmark differed considerably. 2. To assess the implications of winter climate for lake biotic structure and function we compared a number of variables that describe the chemistry and biology of shallow Canadian and Danish lakes that had been chosen to have similar morphometries. 3. The Danish lakes had a fourfold higher ratio of chlorophyll‐a: total phosphorus (TP). Zooplankton : phytoplankton carbon was related to TP and fish abundance in Danish lakes but not in Canadian lakes. There was no significant difference in the ratio log total zooplankton biomass : log TP and the Canadian lakes had a significantly higher proportion of cladocerans that were Daphnia. These differences correspond well with the fact that the Danish lakes have more abundant and diverse fish communities than the Canadian lakes. 4. Our results suggest that severe Canadian winters lead to anoxia under ice and more depauperate fish communities, and stronger zooplankton control on phytoplankton in shallow prairie lakes compared with shallow Danish lakes. If climate change leads to warmer winters and a shorter duration of ice cover, we predict that shallow Canadian prairie lakes will experience increased survivorship of planktivores and stronger control of zooplankton. This, in turn, might decrease zooplankton control on phytoplankton, leading to ‘greener’ lakes on the Canadian prairies.     

Diversity of communities and quantitative parameters of hydrobionts in lakes of the Onon-Toreisk Plain

The diversity of communities and the quantitative parameters of hydrobionts of 17 lakes of the Onon-Toreisk Plain in Northeastern Mongolia during the low-water period are analyzed. The organisms revealed in the lakes include 35 species of phytoplankton, 35 species of zoobenthos, 31 species of zooplankton, and 14 species of hydrophytes. Chara virgata, a representative of charophytes, was revealed in Mongolia for the first time. The greatest species diversity of the communities of hydrobionts is seen in oligohaline lakes. The highest quantitative parameters of communities of phytoplankton and zoobenthos are typical for Gurmiin Lake.     

The role of zoobenthos in energy flow in two shallow lakes

Net production of zoobenthos in two shallow and eutrophic lakes, i.e. the S-basin of Mývatn, Iceland (maximum depth 4.2 m, mean depth 2.3 m) and Hjarbæk Fjord, Denmark (maximum depth 6.5 m, mean depth 1.9 m) were calculated as 878 and 1093 kJ m^-2 yr^-1, respectively. The zoobenthos in both lakes was dominated by Chironomidae (Diptera) living partly as filtrators feeding on suspended particles (phytoplankton) and partly as surface feeders foraging on benthic algae and/or seston. Respiration and consumption were estimated from the literature. Net production efficiency averaged 0.41 and 0.48 in Hjarbæk Fjord and Mývatn, respectively. Ingestion was dominated by herbivorous chironomids, while detritivorous tubificids were insignificant. Zoobenthic production made up 86% of total secondary production (zooplankton plus zoobenthos) in both lakes. The trophic efficiency between net primary production and benthic net secondary production was 8% and 11% in Hjarbæk Fjord and Lake Mývatn S-basin, respectively.     

Larger zooplankton in Danish lakes after cold winters: are winter fish kills of importance?

Winter fish kills can be intense under ice in shallow lakes, and have cascading effects on the food web and ultimately on lake water clarity. In maritime Western Europe, winters are usually mild, but occasional colder periods may also have strong effects on lake fish communities. Global warming may have disproportionate effects by delaying freezing and shortening the period of ice coverage. We studied differences in zooplankton (cladocerans, copepods, and rotifers): phytoplankton biomass, zooplankton community structure, and individual body size among 37 Danish lakes of various depths, chemical characteristics, and trophy, by comparing four winters of different severity (mean winter temperatures ranging from −1.19°C in 1996 to +2.9°C in 1995). We found that crustacean mean body sizes were significantly larger in the summer following a severely cold winter. The zooplankton communities in the summer after a cold winter had a significantly larger proportion of larger-bodied species and taxa. Phytoplankton biomass, expressed as chlorophyll-a (chl-a), was lower and zooplankton herbivory (chl-a:TP index), higher, in the summer after the severely cold winter of 1995/1996. All these effects were stronger in shallow lakes than in deep lakes. Changes in zooplankton during summer 1996, compared with other years, were likely caused by fish kills under ice during the preceding severe winter of 1995–1996. Fish kills due to under ice oxygen depletion would be expected to occur earlier and be more complete in the shorter water columns of shallow lakes. With climate change, severe winters are predicted to become less frequent and the winters to be milder and shorter. In general, this is likely to lead to higher winter survival of fish, lower zooplankton grazing of phytoplankton the following summer and more turbid waters, particularly in shallow eutrophic lakes.     

A LIMNOLOGICAL SYNOPSIS OF BHANGAZI SOUTH,A DYSTROPHIC COASTAL LAKE IN THE GREATER ST LUCIA WETLAND PARK (KWAZULU/NATAL), WITH COMMENTS ON ITS CONSERVATION VALUE.

Summary

Base-line limnological and biological data are given for Lake Bhangazi South, a small lake on the coastal plain of Kwazulu-Natal, South Africa. This shallow (Z_max < 6 m), subtropical lake is a warm and seemingly continuously polymictic system, but experiences severe (continuous?) deoxygenation of its deeper muddy sediments. Nutrient status (of N in particular), light attenuation, phytoplankton productivity and zooplankton biomass are high relative to other regionally comparable coastal lakes investigated. While the ichthyofauna is quite rich, zooplankton and especially zoobenthos communities are species-poor, and lack the relict estuarine components which often dominate these latter assemblages in comparable coastal lakes. The zooplankton is typically freshwater in composition, and contains a new species of copepod (Tropodiaptomus bhangazii Rayner); the lake may be older, or seen longer evolutionary divergence than its proximate counterparts. Benthic species diversity in the lake is especially low amongst the Crustacea and Mollusca, possibly reflecting the effect of relatively acid waters (pH ca 6.5) on the calcium budgets of these groups. Scanty observational data indicative of ecosystem changes over the past 35 years are reported and briefly evaluated from a conservation/management perspective. Along with the endemic copepod, several rare fish species give the lake added conservation status.     

Modelling production and biomasses of zoobenthos in lakes

This work presents a dynamic model to predict zoobenthos in lakes. The model has been developed within the framework of a more comprehensive lake ecosystem model, LakeWeb, which also accounts for the following functional groups of organisms, phytoplankton, bacterioplankton, two types of zooplankton (herbivorous and predatory), macrophytes, prey fish and predatory fish. This work also presents a new data-base for zoobenthos in lakes. Many of the lakes included in this study are situated in the former Soviet Union. They were investigated during the Soviet period and those results have been largely unknown in the West. Using this data-base, this work also presents new empirical models for zoobenthos. The new dynamic model gives seasonal variations (the calculation time, dt, is 1 week using Euler's method and enough iterations to get stable solutions). The basic aim of the dynamic model is that it should capture general functional and structural patterns in lakes. We have demonstrated by several model tests along limnological gradients (total phosphorus concentrations, pH, lake colour, latitude and lake size) that the dynamic model gives predictions that agree well with the values given by the empirical regressions, and also expected and requested divergences from these regressions when they do not provide sufficient resolution. It would have been very difficult indeed to carry out such tests regarding ecosystem responses using traditional methods with extensive field studies in a few lakes. We have given algorithms for (1) production of zoobenthos from eating macrophytes, benthic algae and sediments, (2) elimination (related to the turnover time of zooplankton), and (3) zoobenthos consumption by prey fish, and the factors influencing these processes/rates. The model is driven by data easily accessed from standard monitoring programs or maps a prerequisite for practical utility in contexts of lake management.     

Trophic structure, species richness and biodiversity in Danish lakes: changes along a phosphorus gradient

1. Using data from 71, mainly shallow (an average mean depth of 3 m), Danish lakes with contrasting total phosphorus concentrations (summer mean 0.02–1.0 mg P L?l), we describe how species richness, biodiversity and trophic structure change along a total phosphorus (TP) gradient divided into five TP classes (class 1–5: <0.05, 0.05–0.1, 0.1–0.2, 0.2–0.4,> 0.4 mg P L?1).
2. With increasing TP, a significant decline was observed in the species richness of zooplankton and submerged macrophytes, while for fish, phytoplankton and floating‐leaved macrophytes, species richness was unimodally related to TP, all peaking at 0.1–0.4 mg P L?1. The Shannon–Wiener and the Hurlbert probability of inter‐specific encounter (PIE) diversity indices showed significant unimodal relationships to TP for zooplankton, phytoplankton and fish. Mean depth also contributed positively to the relationship for rotifers, phytoplankton and fish.
3. At low nutrient concentrations, piscivorous fish (particularly perch, Perca fluviatilis) were abundant and the biomass ratio of piscivores to plankti‐benthivorous cyprinids was high and the density of cyprinids low. Concurrently, the zooplankton was dominated by large‐bodied forms and the biomass ratio of zooplankton to phytoplankton and the calculated grazing pressure on phytoplankton were high. Phytoplankton biomass was low and submerged macrophyte abundance high.
4. With increasing TP, a major shift occurred in trophic structure. Catches of cyprinids in multiple mesh size gill nets increased 10‐fold from class 1 to class 5 and the weight ratio of piscivores to planktivores decreased from 0.6 in class 1 to 0.10–0.15 in classes 3–5. In addition, the mean body weight of dominant cyprinids (roach, Rutilus rutilus, and bream, Abramis brama) decreased two–threefold. Simultaneously, small cladocerans gradually became more important, and among copepods, a shift occurred from calanoid to cyclopoids. Mean body weight of cladocerans decreased from 5.1 μg in class 1 to 1.5 μg in class 5, and the biomass ratio of zooplankton to phytoplankton from 0.46 in class 1 to 0.08–0.15 in classes 3–5. Conversely, phytoplankton biomass and chlorophyll a increased 15‐fold from class 1 to 5 and submerged macrophytes disappeared from most lakes.
5. The suggestion that fish have a significant structuring role in eutrophic lakes is supported by data from three lakes in which major changes in the abundance of planktivorous fish occurred following fish kill or fish manipulation. In these lakes, studied for 8 years, a reduction in planktivores resulted in a major increase in cladoceran mean size and in the biomass ratio of zooplankton to phytoplankton, while chlorophyll a declined substantially. In comparison, no significant changes were observed in 33 ‘control’ lakes studied during the same period.     

Differences in nutrient limitation and grazer suppression of phytoplankton in seepage and drainage lakes of the Adirondack region,NY, U.S.A

1. For seepage and drainage lakes of the Adirondack mountain region (NY, U.S.A) hydrologic regime is correlated with physical and chemical differences that can affect phytoplankton and planktonic food webs (e.g. presence and influence of wetlands, dissolved organic carbon concentration, anoxia, nutrient cycling). We conducted short‐term (48 h), in situ enclosure experiments to evaluate the relative importance of macrozooplankton grazing and nutrient limitation of phytoplankton biomass in small Adirondack seepage and drainage lakes (N = 18, 1–137 ha). Epilimnetic dissolved organic carbon (DOC) concentrations and pH values represented the diversity of the region. We measured chlorophyll a changes in response to grazer removal (> 120 μm) and nutrient addition (～ 10× ambient N, P, or N + P), and evaluated changes with respect to in situ light, temperature, NO₃, NH₄, SRP, and crustacean assemblage characters. 2. Nutrient addition stimulated significant increase in chlorophyll a concentration at 11 of 18 sites (GLM, Tukey–Kramer). Phytoplankton of clearwater drainage lakes were P‐limited, whereas clearwater and brownwater seepage lakes responded to additions of N and/or N + P. Relative light availability explained half the variance in response to nutrient addition in drainage (r2 = 0.48), but not seepage lake experiments (P > 0.05). 3. We observed responses to grazer removal at eight of 18 sites, usually clearwater drainage lakes. Crustacean grazing may be as significant as nutrient limitation of [chl a] for many drainage lake phytoplankton assemblages. Responses were related to in situ density of zooplankton only in drainage lakes. Light explained some variability in response to grazer removal for drainage (r2 = 0.35) and seepage lake experiments (r2 = 0.35). 4. These experiments provide evidence that hydrology may ultimately play an important role in determining nutrient and grazer regulation of phytoplankton. Proximate mechanisms affecting our results may be associated with differences in wetland vegetation, [DOC], and nutrient cycling.     

Secondary production of crustacean zooplankton and biomass of major rotifer species in Lake Bosumtwi/Bosomtwe,Ghana, West Africa

Studies have shown a strong linkage between zooplankton and fisheries' potential in tropical lakes. High zooplankton production provides the basis for fish production, but knowledge of zooplankton production dynamics in African lakes is extremely limited. Crustacean zooplankton production and the biomass of dominant rotifers in Lake Bosumtwi were assessed over a 2‐year period. The crustaceans comprised an endemic and extremely abundant cyclopoid copepod, Mesocyclops bosumtwii and the cladoceran Moina micrura. Mean standing stock of the crustaceans was 429 mg dw m^?3, whilst annual production averaged 2.1 g dw m^?3 y^?1. Production doubled from 1.4 g dw m^?3 y^?1 in 2005 to 2.8 g dw m^?3 y^?1 in 2006. Copepods accounted for 98.5% of crustacean production. The biomass of the dominant rotifers Brachionus calyciflorus and Hexarthra intermedia was less than 1% of total zooplankton biomass. Daily turnover rate and turnover time of the crustaceans was 0.19 day^?1 and 6.2 days respectively. Crustacean production yielded no statistical relationship with phytoplankton biomass. Production was well within the range of tropical lakes. Peak crustacean production synchronized maximum rainfall, lake mixing and phytoplankton production. Most importantly, no one year's set of dynamics can be used to characterize zooplankton production in the lake.     

Predictability and possible mechanisms of plankton response to reduction of planktivorous fish

The predictability of plankton response to reductions of planktivorous fish was investigated by comparing the plankton community in three biomanipulated lakes and ten unmanipulated lakes differing in intensity of fish predation. Data collected on total phosphorus, phytoplankton and zooplankton biomass and share of cyanobacteria and large grazers, as well as specific growth rate of phytoplankton, were further used to test some of the proposed underlying response-mechanisms. In the biomanipulated lakes the algal biomass and share of cyanobacteria decreased, specific growth rate of phytoplankton increased, and zooplankton biomass and share of large grazers increased or remained unchanged. This pattern was largely reflected in the differences in food-chain structure between the unmanipulated lakes with highversus those with low fish predation. The qualitative response to planktivorous fish reduction thus seems largely predictable. The biomanipulated lakes differed, however, in magnitude of response: the smallest hypertrophic, rotenone-treated lake (Helgetjern) showed the most dramatic response, whereas the large, deep mesotrophic lake (Gjersjøen), which was stocked with piscivorous fish, showed more moderate response, probably approaching a new steady state. These differences in response magnitude may be related to different perturbation intensity (rotenone-treatmentversus stocking with piscivores), food-chain complexity and trophic state. Both decreased phosphorus concentration and increased zooplankton grazing are probably important mechanisms underlying plankton response to biomanipulation in many lakes. The results provide tentative support to the hypothesis that under conditions of phosphorus limitation, increased zooplankton grazing can decrease algal biomassvia two separate mechanisms: reduction of the phosphorus pool in the phytoplankton, and reduction of the internal C:P-ratio in the phytoplankton cells.

     

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.

     

Restoration of shallow lakes by nutrient control and biomanipulation—the successful strategy varies with lake size and climate

Major efforts have been made world-wide to improve the ecological quality of shallow lakes by reducing external nutrient loading. These have often resulted in lower in-lake total phosphorus (TP) and decreased chlorophyll a levels in surface water, reduced phytoplankton biomass and higher Secchi depth. Internal loading delays recovery, but in north temperate lakes a new equilibrium with respect to TP often is reached after <10–15 years. In comparison, the response time to reduced nitrogen (N) loading is typically <5 years. Also increased top-down control may be important. Fish biomass often declines, and the percentage of piscivores, the zooplankton:phytoplankton biomass ratio, the contribution of Daphnia to zooplankton biomass and the cladoceran size all tend to increase. This holds for both small and relatively large lakes, for example, the largest lake in Denmark (40 km²), shallow Lake Arresø, has responded relatively rapidly to a ca. 76% loading reduction arising from nutrient reduction and top-down control. Some lakes, however, have proven resistant to loading reductions. To accelerate recovery several physico-chemical and biological restoration methods have been developed for north temperate lakes and used with varying degrees of success. Biological measures, such as selective removal of planktivorous fish, stocking of piscivorous fish and implantation or protection of submerged plants, often are cheap versus traditional physico-chemical methods and are therefore attractive. However, their long-term effectiveness is uncertain. It is argued that additional measures beyond loading reduction are less cost-efficient and often not needed in very large lakes. Although fewer data are available on tropical lakes these seem to respond to external loading reductions, an example being Lake Paranoá, Brazil (38 km²). However, differences in biological interactions between cold temperate versus warm temperate-subtropical-tropical lakes make transfer of existing biological restoration methods to warm lakes difficult. Warm lakes often have prolonged growth seasons with a higher risk of long-lasting algal blooms and dense floating plant communities, smaller fish, higher aggregation of fish in vegetation (leading to loss of zooplankton refuge), more annual fish cohorts, more omnivorous feeding by fish and less specialist piscivory. The trophic structures of warm lakes vary markedly, depending on precipitation, continental or coastal regions locations, lake age and temperature. Unfortunately, little is known about trophic dynamics and the role of fish in warm lakes. Since many warm lakes suffer from eutrophication, new insights are needed into trophic interactions and potential lake restoration methods, especially since eutrophication is expected to increase in the future owing to economic development and global warming.     

Winter ecology of shallow lakes: strongest effect of fish on water clarity at high nutrient levels

While the structuring role of fish in lakes is well studied for the summer season in North temperate lakes, little is known about their role in winter when fish activity and light irradiance potentially are lower. This is unfortunate as the progressing climate change may have strong effects on lake winter temperature and possibly on trophic dynamics too. We conducted an enclosure experiment with and without the presence of fish throughout winter in two shallow lakes with contrasting phosphorus concentrations. In hypertrophic Lake Søbygård, absence of fish led to higher biomass of zooplankton, higher grazing potential (zooplankton:phytoplankton ratio) and, accordingly, lower biomass of phytoplankton and chlorophyll a (Chl a), while the concentrations of total nitrogen (TN), total phosphorus (TP), oxygen and pH decreased. The average size of egg-bearing Daphnia and Bosmina and the minimum size of egg-bearing specimens of the two genera rose. In the less eutrophic Lake Stigsholm, zooplankton and their grazing potential were also markedly affected by fish. However, the decrease in Chl a was slight, and phytoplankton biovolume, pH and the oxygen concentration were not affected. TN was higher when fish were absent. Our results indicate that: (i) there is a notable effect of fish on zooplankton community structure and size during winter in both eutrophic and hypertrophic North temperate lakes, (ii) Chl a can be high in winter in such lakes, despite low light irradiance, if fish are abundant, and (iii) the cascading effects on phytoplankton and nutrients in winter may be more pronounced in hypertrophic lakes. Climate warming supposedly leading to reduced winter mortality and dominance of small fish may enhance the risk of turbid state conditions in nutrient-enriched shallow lakes, not only during the summer season, but also during winter.     

Plankton community and the relationship with the environment in saline lakes of Onon-Torey plain,Northeastern Mongolia

The plankton community of sixteen saline lakes located on Onon-Torey plain (Northeastern Mongolia) during the filling phase and the raising of the water level was investigated in July 2011. Thirty-five taxa of phytoplankton and thirty-one species of zooplankton were found. For phytoplankton, blue-green algae (Merismopedia elegans, Anabaenopsis elenkinii, Arthrospora fusiformis, Spirulina major, Lyngbya sp., Oscillatoria sp.) and green algae (Monoraphidium minutum, Tetrastrum komarekii, Ankyra ocellata, Oocystis sp.) were dominant. For zooplankton, Filinia longiseta, Brachionus plicatilis, B. variabilis, Hexarthra mira (Rotifera), Daphnia magna, Moina brachiata, M. mongolica (Cladocera), Arctodiaptomus bacillifer, Mixodiaptomus incrassatus, Metadiaptomus asiaticus (Copepoda) dominated. Mineralization, active hydrogen ratio, dissolved oxygen and water temperature were the main factors influencing the diversity, structure and distribution of plankton organisms in the steppe lakes during low water level. The RDA analysis for phytoplankton and zooplankton from different lakes was carried out for selected two groups which included lakes and a subset related species. The first group is of oligohaline and mesohaline lakes in which mostly green algae, rotifers and copepods inhabit. The second group is of mesohaline and polyhaline lakes with mainly blue-green algae, some crustaceans and rotifers inhabiting. High abundance and biomass of Spirulina major, Oscillatoria sp. and Brachionus variabilis were observed in lakes with high mineralization, pH and temperature.     

Changes in the biota of Chany Lake along a salinity gradient

Relationships among salinity and diversity, abundance, biomass of major biological components of Chany Lake (western Siberia, Russia) are examined across a salinity gradient. As salinity increased from 0.8 to 6.4 g l⁻¹, the species richness of aquatic vascular plants decreased from 16 to 2 species, of phytoplankton from 98 to 52 species, and of zooplankton from 61 to 16 species, but changes in species diversity of zoobenthos were negligible. Guest Editor: John M. Melack Saline Waters and their Biota     

Stable isotope analysis of the origins of zooplankton carbon in lakes of differing trophic state

Carbon stable isotope analysis was carried out on zooplankton from 24 United Kingdom lakes to examine the hypothesis that zooplankton dependence on allochthonous sources of organic carbon declines with increasing lake trophy. Stable isotope analysis was also carried out on particulate and dissolved organic matter (POM and DOM) and, in 11 of the lakes, of phytoplankton isolates. In 21 of the 24 lakes, the zooplankton were depleted in ¹³C relative to bulk POM, consistent with previous reports. δ¹³C for POM showed relatively little variation between lakes compared to high variation in values for DOM and phytoplankton. δ¹³C values for phytoplankton and POM converged with increasing lake trophy, consistent with the expected greater contribution of autochthonous production to the total organic matter pool in eutrophic lakes. The difference between δ¹³C for zooplankton and that for POM was also greatest in oligotrophic lakes and reduced in mesotrophic lakes, in accordance with the hypothesis that increasing lake trophic state leads to greater dependence of zooplankton on phytoplankton production. However, the difference increased again in hypertrophic lakes, where higher δ¹³C values for POM may have been due to greater inputs of ¹³C-enriched organic matter from the littoral zone. The very wide variation in phytoplankton δ¹³C between lakes of all trophic categories made it difficult to detect robust patterns in the variation in δ¹³C for zooplankton. Received: 2 November 1998 / Accepted: 3 December 1999     

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.     

Mesocosm experiments on nutrient and fish effects on shallow lake food webs in a Mediterranean climate

1. Nutrient and fish manipulations in mesocosms were carried out on food‐web interactions in a Mediterranean shallow lake in south‐east Spain. Nutrients controlled biomass of phytoplankton and periphyton, while zooplankton, regulated by planktivorous fish, influenced the relative percentages of the dominant phytoplankton species. 2. Phytoplankton species diversity decreased with increasing nutrient concentration and planktivorous fish density. Cyanobacteria grew well in both turbid and clear‐water states. 3. Planktivorous fish increased concentrations of soluble reactive phosphorus (SRP). Larger zooplankters (mostly Ceriodaphnia and copepods) were significantly reduced when fish were present, whereas rotifers increased, after fish removal of cyclopoid predators and other filter feeders (cladocerans, nauplii). The greatest biomass and diversity of zooplankton was found at intermediate nutrient levels, in mesocosms without fish and in the presence of macrophytes. 4. Water level decrease improved underwater light conditions and favoured macrophyte persistence. Submerged macrophytes (Chara spp.) outcompeted algae up to an experimental nutrient loading equivalent to added concentrations of 0.06 mg L^?1 PO₄‐P and 0.6 mg L^?1 NO₃‐N, above which an exponential increase in periphyton biomass and algal turbidity caused characean biomass to decline. 5. Declining water levels during summer favoured plant‐associated rotifer species and chroococcal cyanobacteria. High densities of chroococcal cyanobacteria were related to intermediate nutrient enrichment and the presence of small zooplankton taxa, while filamentous cyanobacteria were relatively more abundant in fishless mesocosms, in which Crustacea were more abundant, and favoured by dim underwater light. 6. Benthic macroinvertebrates increased significantly at intermediate nutrient levels but there was no relationship with planktivorous fish density. 7. The thresholds of nutrient loading and in‐lake P required to avoid a turbid state and maintain submerged macrophytes were lower than those reported from temperate shallow lakes. Mediterranean shallow lakes may remain turbid with little control of zooplankton on algal biomass, as observed in tropical and subtropical lakes. Nutrient loading control and macrophyte conservation appear to be especially important in these systems to maintain high water quality.     

Macrozooplankton and the persistence of the deep chlorophyll maximum in a stratified lake

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