The ecological history of a mediaeval man-made lake. Hickling Broad,Norfolk, United Kingdom

Changes in Hickling Broad, since its creation in the 14th or 15th centuries by the flooding of peat diggings, have been deduced from dating and analysis of a sediment core, historical information and current limnological studies. Until the 1930's there was little major change. Increased agricultural land fertilization led to markedly increased organic sedimentation from the 1930's onwards, due to increased growth of submerged macrophytes. Inorganic sedimentation increased concurrently as more powerful pumps were installed to help drain the adjacent fens and marshes.There was no evidence of increased plankton populations during this phase, but epiphytic diatom populations increased. In the mid 1960's the current period of hypereutrophication began. Epiphytic diatom numbers increased markedly and in the early 1970's the previous luxuriant macrophytes became sparse and the water became turbid with phytoplankton. These changes are attributable mainly to increases in the size of a roost of migratory black headed gulls (Larus ridibundus L.) on the lake in autumn and winter.     

Can macrophytes be useful in biomanipulation of lakes? The Lake Zwemlust example

Lake Zwemlust (area 1.5 ha, Z_m 1.5 m) has been the object of an extensive limnological study since its biomanipulation involving removal of planktivorous fish (bream) in March 1987 and emptying of the lake. In the subsequent summer period of 1987 the Secchi depth increased to the lake bottom (2.5 m), compared withca 30 cm in the earlier summers. The reaction of submerged macrophytes to improving under-water light climate was rapid. In summer 1987, besides the introducedChara globularis, 5 species of submerged macrophytes occurred and colonized 10% of the lake area. In 1988 and 1989 only quantitative changes were observed; new species did not appear, but the area colonized by macrophytes increased by 7 and 10 times, respectively.Elodea nuttallii was dominant among the macrophytes andMougeotia sp. among the filamentous green algae. Their abundance, contributed to transient N-limination of phytoplankton causing a persistent clear water phase in 1988 and 1989, unlike in 1987 when zooplankton grazing contributed chiefly to the water clarity. Laboratory bioassays on macrophytes confirmed nitrogen limitation.     

Morphological and Morphometrical Variations of Selected Rotifer Species in Response to Predation: a Seasonal Study of Selected Brachionid Species from Lake Xochimilco (Mexico)

Allelopathy of filamentous green algae (FGA) has been less studied than that of macrophytes. Little Budworth Pool, Cheshire, UK is a small, shallow, clear-water lake with high TP concentrations, very high NO_3-N concentrations, only moderate phytoplankton density, high FGA growth (mainly Spirogyra sp.) and no submerged plants. Experiments were carried out to test the possible allelopathic effects of Spirogyra on the phytoplankton of this lake and on a submerged plant Elodea nuttallii. Changes in phytoplankton growth, phytoplankton species dynamics and species composition were apparently not influenced by allelopathy of live or decaying Spirogyra. A shift from diatom (Cyclotella sp) – cryptomonad (Chroomonas acuta and Cryptomonas erosa) dominance to Chlorococcales (Micractinium pusillum, Monoraphidium contortum and Scenedesmus opoliensis) – Volvocales (Chlorogonium elongatum and Pandorina morum) dominance was recorded in both control and FGA treatments, suggesting an effect of nutrient enrichment. Nutrient concentrations and differences in competitiveness among phytoplankton species can also explain differences in their growth rates in Spirogyra filtrate. Spirogyra also did not influence apex number per plant, shoot length or growth rate of E. nuttallii. This FGA species probably cannot control phytoplankton or E. nuttallii growth in nutrient rich conditions through allelopathy.     

A study of Lough Corrib,western Ireland and its phytoplankton

During 1980/1981, a twelve month study was carried out on the phytoplankton of Lough Corrib, a large freshwater lake in western Ireland. Cell counts indicated low phytoplankton productivity and this was reflected in low chlorophyll a values and high Secchi readings. Diatoms predominated in spring and were succeeded by blue-green algae in late summer. The spring diatom crop was apparently limited by either silicate or nitrogen. Mineral uptake by rooting macrophytes and benthic algae and phosphate recycling as a result of bottom sediment disturbance may be important elements in the nutrient cycling regime of the lake. Differences in morphology between the upper and lower basins of the lake are reflected, to some extent, in species composition and areal productivity.     

Movement of plankton through lake-stream systems

1. River plankton are often assumed to come from upstream lakes, but the factors controlling the movement of plankton between lakes and rivers into outflow streams are unclear. We tested the possibility that the physical structure of the littoral zone near the lake outlet (depth, presence of macrophytes) and diurnal differences in plankton composition at the lake surface influence the movement of plankton from the lake into the stream and determine their persistence downstream. 2. Zooplankton and phytoplankton biomass, community composition and mean body size were compared between two deep lakes without macrophytes at the lake edge and two shallow lakes with macrophytes at the lake edge. Samples were collected day and night on three dates, in the lake centre, in the littoral zone adjacent to the lake outlet, at the outlet and at two sites downstream in Algonquin Park, Ontario, Canada. 3. The morphology of lake edges clearly affects the movement of lake zooplankton into outlet streams. Outlets draining deeper littoral zones had higher zooplankton biomass than shallow littoral outlets (P < 0.0001), but these differences disappeared within 50 m downstream of the lake. There was no difference in mean zooplankton body size among lake outlets or between littoral and outlet samples. However, shallow littoral zones were dominated by cyclopoid copepods and deeper littoral zones were dominated by Bosmina longirostris. In contrast, phytoplankton biomass entering the outlet was similar to that found within the lake and did not vary with lake outlet morphology. These effects were consistent across several sampling weeks and were not affected by surface zooplankton biomass changes associated with diurnal vertical migration in the lake centre. 4. A comparison with published river zooplankton data suggests that zooplankton are rapidly eliminated from shallow outlet streams (≤1 m deep) but persist in most deeper outlet rivers (≥2 m deep). Because the depth of an outlet river determines downstream zooplankton community development, the contribution of lakes to river plankton communities may be influenced by the location of each lake within the drainage basin. These findings suggest that lake and outflow physical structure influences connection strength between spatially successive habitats.     

Trophic interactions in a shallow lake following a reduction in nutrient loading: a long-term study

After the diversion of a nutrient-rich inflow, the eutrophic lake, Alderfen Broad, initially showed reduced total phosphorus concentrations and phytoplankton populations, clear water and the establishment of submerged macrophytes. Internal P loading then increased, perhaps stimulated by the senescence of submerged macrophytes and exacerbated by the lack of flushing. Cyanophytes appeared in the summer of two years. As a consequence of poor recruitment of roach (Rutilus rutilus (L.)), the chief zooplanktivore, and a summerkill of the fish population, populations of large-bodied Cladocera (Daphnia hyalina/ longispina and ultimately D. magna) developed. In the long-term, these may have limited the further development of phytoplankton populations and clear water and submerged macrophytes returned. During this latter period, internal P release has remained high (> 380 µg l^-1), thereby indicating the scope for biomanipulation even in eutrophic conditions. However, isolation of the lake has led to a decrease in water level (which through increased temperatures and lowered dissolved oxygen levels was probably responsible for the fish deaths) and further concentration of internal P load. Sediment is now being removed to reestablish greater water depth.     

Long term study on the influence of eutrophication, restoration and biomanipulation on the structure and development of phytoplankton communities in Feldberger Haussee (Baltic Lake District, Germany)

Feldberger Haussee provides a classic example of eutrophication history of hardwater lakes in the Baltic Lake District (Germany) and of changes in their algal flora during the 20th century. The lake originally was regarded as slightly eutrophic. A process of drastic eutrophication from the 1950s until the end of the 1970s caused mass developments of blue-green and green algae. A restoration program was started in the 1980s to improve the water quality of the lake using both diversion of sewage outside the catchment area, and biomanipulation by altering the fish community. This restoration program led to positive changes in the lake ecosystem. Direct effects of biomanipulation resulted in an increase of herbivorous zooplankton, a decrease of phytoplankton biomass, and an increase of water transparency. The recovery of Feldberger Haussee also may have been indirectly enhanced by an increase in nutrient sedimentation as a consequence of intensified calcite precipitation, decrease in phosphorus remobilization due to a pH-decrease, increased NIP-ratio, and recolonization of the littoral zone by macrophytes. This paper concentrates on the long term development of the phytoplankton community as a response to changes in the food web structure as well as to alterations in the chemical environment of the algae. Both are reflected in four major stages passed by the algal assemblage between 1980 and 1994: (1) From 1980-summer 1985 dense green algal populations were found indicating similar conditions as in the 1970s during the period of maximum eutrophication. (2) A diverse phytoplankton community during summer 1985–1989 showed the first effects of a recovery. (3) From 1990–1992 the phytoplankton was characterized by ungrazeable filamentous blue-green algae first of all as a response to increased herbivory of zooplankton on edible species and to increasing N/P-ratios. (4) Finally, the algal species diversity increased in 1993 and 1994 whereas the phytoplankton biomass decreased showing the success of the combined restoration measures.     

Planktonic food chains of a highly humic lake

The development and metabolism of the plankton of a highly humic lake were followed over the vernal primary production maximum. The study was made in a mesocosm in which large filter feeders, typical of this lake in summer, were absent. During the rising phase of phytoplankton, the community was predominantly autotrophic. The most important constituents in the algal biomass were a dinoflagellate, Gymnodinium sp. (40–50%), and a prasinophycean, Scourfieldia cordiformis (7%). The biomasses of Chlamydomonas spp. and Chrysococcus spp. reached their maxima a few days later and Cryptomonas sp. became most abundant at the end of the experiment. After the phytoplankton maximum, about one week from the beginning ofthe experiment, grazing of algae by phagotrophic protozoans and phosphate depletion led to a rapid decrease of algal biomass and the community became predominantly heterotrophic. In spite of a large variation in algal biomass and primary production, the biomass of bacteria remained of the same order of magnitude as in algae both before and after the algal maximum. Bacteria were mostly responsible for the plankton respiration, which also showed no dependence on primary production. Since exudation by phytoplankton was also low, the nutrition of bacterioplankton was probably mainly based on allochthonous dissolved organic matter rather than or primary production. Thus the production of bacteria was an additional food source for higher trophic levels along with phytoplankton. Because filter feeding zooplankton was absent in the experiment, protozoans were the only grazers utilizing algae and bacteria. Essentially all growth of bacteria was used by bacterivores.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

Phosphorus cycling in a dense Potamogeton pectinatus L. Bed

Summary An analysis of phosphorus concentrations and standing stocks in the various components of a dense Potamogeton pectinatus L. community in a brackish lake showed that the amounts of P per unit area varied in the following order: Sediments > Above ground macrophytes > Detritus > Benthic Invertebrates > Below ground macrophyte tissue > Epiphytic algae > Water soluble P > Water borne particulate P. Seasonal changes in the variability of these stocks are described, and shown to be controlled by the annual growth and decomposition pattern of the Potamogeton. The sediments were suggested as the major source of P for the plant biomass. Studies using ³²P tracer showed that of a given input of P to the water, 32% went to large epiphytic algae, 17% to the Potamogeton, 16% to the benthic fauna (mostly filter feeding bivalves), 28% to the sediments (almost all incorporated in the top 1 cm), and the remaining 7% was adsorbed on to, or absorbed by microorganisms associated with detritus. Analysis of ³²P uptake curves indicated that of the P absorbed by the Potamogeton, a significant proportion went to the complex adnate periphyton on the leaf surface. We were unable to separate this fraction. Movement of P in the community was shown to be a closed cycle, and any release of P from decaying macrophytes would be rapidly reabsorbed by epiphytic algae. It is unlikely that phosphorus, once cycling in the macrophyte community, would become transferred to the circulation in the open lake.     

Algal Assemblages in Multiple Habitats of Restored and Extant Wetlands

The functional and structural attributes of algal assemblages were studied in 25 restored and 20 extant depressional wetlands in southern central Michigan. Environmental conditions and algal assemblages were compared between restored and extant wetlands and among habitats within wetlands. Restored marshes generally had lower shading by macrophytes, nutrient concentrations, and sediment organic matter. Relative biovolume of non-diatom algae was significantly different among plankton, macrophyte and sediment habitats in restored wetlands, but did not differ between macrophytes and sediments in extant wetlands. Species composition of diatom assemblages was not significantly different between plants and sediments in both restored and extant wetlands. The observed differences in non-diatom algae could not be attributed to any measured environmental variable; however, diatom assemblage differences between habitats increased with light irradiance. Differences in sediment diatom assemblages were observed between restored and extant wetlands and were related to differences in nutrients, pH, and canopy cover. Differences were also observed between epiphytic diatom assemblages in restored and extant wetlands and they were related to light and dissolved oxygen. In summary, differences in light and nutrient availability were the main environmental factors differentiating algal communities in wetlands.     

以浮游植物评价达赉湖水质污染及营养水平

1987—1988年在30个样点,四季采样,对达赉湖的浮游植物进行了种类组成,生物量、种群数量、优势种、污染指示种、硅藻指数、综合指数等群落生态学的初步研究,应用后五项参数对达赉湖水质污染及营养水平进行了评价。达赉湖浮游植物年均值达54.7×106个/L(细胞数,或个体数为2.3×106个/L),硅藻指数为149.3,综合指数为5.6。群落组成中污染指示种占65%,春季以绿藻的十字藻、卵囊藻为优势种,其它3个季节均以蓝藻中的微囊藻、鱼腥藻、腔球藻占优势,表明达赉湖已受到中等程度污染,属于蓝、绿藻型富营养湖。       

Carbon flow in the littoral food web of an oligotrophic lake

Benthic food web dynamics and carbon flow were examined in the littoral zone of Lake Coleridge, a large deep oligotrophic lake, using radioactive and stable isotope techniques in conjunction with analyses of stomach contents of the fauna. We specifically address two hypotheses: (1) that macrophytes only contribute to the carbon flow to higher trophic levels when they have decayed; and (2) that epiphytic algae is the major source of carbon for macroinvertebrates, and thus fish, with only minor contributions from phytoplankton or terrestrial sources. Epiphytic diatoms were a major component of the stomach contents of the gastropod snail Potamopyrgus antipodarum, and of chironomids. Animal remains were also common in the diet of some chironomids, while amorphous organic matter predominated in the stomachs of oligochaetes. A variety of epiphytic algal taxa was found in trichopteran larvae. Feeding rate of P. antipodarum measured with radioactive tracers increased by 10× on decayed macrophytes (Elodea) compared with live material, while feeding rates on characean algae increased by a factor of 3 when decayed material was presented. However, assimilation rates were less than 20% on decayed material compared with 48–52% on live material. Potential carbon sources were easily distinguished based on their ¹³C values, although isotopic ratios showed significant variation among sites. Epiphytic algae showed less variation among sites than macrophytes and were depleted by 4–5 compared with macrophytes. Detrital material, organic matter in the sediments and plankton were significantly depleted in ¹³C relative to macrophytes and slightly depleted relative to epiphytic algae. Most macroinvertebrate taxa showed a similar pattern among sites to macrophytes and epiphytic algae. P. antipodarum and chironomids were slightly enriched compared with epiphytic algae. Ratios for the common bully (Gobiomorphus cotidianus) were generally consistent with a diet dominated by chironomids, while there was some evidence for terrestrial inputs for koaro (Galaxias brevipinnis) and juvenile brown trout. Epiphytic algae appear to underpin much of the production in the littoral zone of this oligotrophic lake, with trichopteran and chironomid larvae mediating carbon flows from algae to fish. Macrophytes do not make a major contribution directly to carbon flow to higher trophic levels even when decayed. The lack of a direct link between macrophytes and higher trophic levels is due to the faunal composition, including a lack of large herbivores.     

THE INFLUENCE OF SUBMERGED MACROPHYTES ON SEDIMENTARY DIATOM ASSEMBLAGES1

Submerged macrophytes are a central component of lake ecosystems; however, little is known regarding their long‐term response to environmental change. We have examined the potential of diatoms as indicators of past macrophyte biomass. We first sampled periphyton to determine whether habitat was a predictor of diatom assemblage. We then sampled 41 lakes in Quebec, Canada, to evaluate whether whole‐lake submerged macrophyte biomass (BiomEpiV) influenced surface sediment diatom assemblages. A multivariate regression tree (MRT) was used to construct a semiquantitative model to reconstruct past macrophyte biomass. We determined that periphytic diatom assemblages on macrophytes were significantly different from those on wood and rocks (ANOSIM R = 0.63, P < 0.01). A redundancy analysis (RDA) of the 41‐lake data set identified BiomEpiV as a significant (P < 0.05) variable in structuring sedimentary diatom assemblages. The MRT analysis classified the lakes into three groups. These groups were (A) high‐macrophyte, nutrient‐limited lakes (BiomEpiV ≥525 μg · L^?1; total phosphorus [TP] <35 μg · L^?1; 23 lakes); (B) low‐macrophyte, nutrient‐limited lakes (BiomEpiV <525 μg · L^?1; TP <35 μg · L^?1; 12 lakes); and (C) eutrophic lakes (TP ≥35 μg · L^?1; six lakes). A semiquantitative model correctly predicted the MRT group of the lake 71% of the time (P < 0.001). These results suggest that submerged macrophytes have a significant influence on diatom community structure and that sedimentary diatom assemblages can be used to infer past macrophyte abundance.     

Periphyton in the Volta Lake. II. Seasonal changes on wooden blocks with depth

A seasonal study of the periphyton developed on wooden blocks suspended at five constant depths (0, 0.5, 1, 2, 5 m) in the Pawm-pawm arm of the Volta Lake is reported. The early colonizers at all depths were diatom species, they gradually declined in diversity and numbers and blue-green algae and green algae became more common. After one year the shallower (0–1 m) blocks were covered by a luxuriant growth of filamentous green algae though at lower levels diatoms predominated. Diversity and cell numbers of algae decreased with depth, this was most pronounced below 2 m. It appears that filamentous green algae eventually come to dominate wooden surfaces floating at relatively shallow depths in the Volta Lake.     

Establishing realistic restoration targets for nutrient-enriched shallow lakes: linking diatom ecology and palaeoecology at the Attenborough Ponds, U.K.

Eutrophication is a major problem for shallow lakes in the U.K. lowlands. Over the last few decades issues relating to the management and restoration of these lakes have come to the fore, including the need to develop methods for establishing realistic restoration targets. One group of lakes for which restoration is a key concern are the Attenborough Ponds in the English Midlands, an interconnected series of shallow, nutrient-enriched gravel-pit lakes. In November 1972, the highly polluted River Erewash was diverted into the Attenborough Ponds. To determine the ecological effects of this event and to establish restoration goals for this lake system, two complimentary studies were undertaken here; (i) monitoring of the ecology and sedimentary representation of diatom communities in a disturbed lake connected to the R. Erewash (Tween Pond: high nutrient concentrations, no submerged macrophytes) and a relatively undisturbed lake isolated from the R. Erewash (Clifton Pond: lower nutrient concentrations, abundant submerged macrophytes), and; (ii) examination of the sedimentary diatom record in a short sediment core collected from the chosen disturbed lake (Tween Pond). The species composition, seasonality, relative productivity and sedimentary representation of planktonic and periphytic diatom communities were very different in Tween and Clifton Ponds. In Tween Pond two major phases of planktonic diatom production/sedimentation were observed during March–May and July–September. By contrast, planktonic diatoms were restricted to a single spring peak in Clifton Pond and after May there was a switch to the dominance of periphytic diatoms associated with the development of submerged macrophytes. The 1972 diversion of the River Erewash was clearly reflected in the sedimentary diatom record from Tween Pond, by; (i) the abrupt shift to planktonic diatom dominance, and; (ii) increasing percentages of late summer–autumn associated planktonic diatom species. These changes suggest both significant nutrient enrichment and the switch from submerged macrophyte to phytoplankton dominance. Numerical matching of pre-1972 diatom assemblages with surface sediment assemblages in the gravel-pits using Principal Components Analysis (PCA) and a squared chord distance measure revealed no close analogues. Nevertheless, similarities between; (i) percentages of different diatom habitat and seasonality groups, and; (ii) pre-1972 and contemporary macrophyte survey data suggested that Clifton Pond is probably a good pre-diversion analogue for habitat structure (e.g. macrophyte biomass, composition and architecture) and phenology (e.g. diatom and macrophyte seasonality) in the Ponds. The practical value of combining space-for-time substitution and palaeoecological approaches in restoration ecology studies of shallow lakes is discussed.     

Abundance and size change of Hannaea baicalensis in Lake Baikal

Hannaea baicalensis is a benthic pennate diatom that predominantly grows at depths of less than a metre attached to rocks and filamentous algae in Lake Baikal, Russia. This narrow zone at the edge of the lake is subject to frequent wave action and lake level fluctuations, which combine with other factors to affect seasonal abundance. During ice cover from January to May in 2008, when lake levels decreased from 42 to 14 cm above datum, H. baicalensis cell abundance remained low (0.39 × 10⁶ cells cm^–2). The main period of net cell increase occurred in autumn, when there was a period of stable lake level (±10 cm changes in water depth) that coincided with the return of nutrients during autumn overturn. Cell abundance reached 1.52 × 10⁶ cells cm^–2 on 31 October. Alongside the changes in abundance, cyclic size changes in cell apical lengths were found (40 to 144 µm), which were associated with timing of the length of the life cycle. Size decline occurred in both spring and autumn, with an average decrease in apical length of 36 µm per year. It took two years for the mean apical length of a single cohort to decrease from 128 µm to 56 µm, which was then below the threshold (< 65 µm) for initiation of size regeneration.     

The responses of littoral invertebrates to eutrophication-linked changes in plant communities

The decay of submerged macrophytes in lakes of high trophic level drastically limits the extent of habitat available to littoral invertebrates. The loss can be partially compensated by growth of filamentous algae. Our results show that macroinvertebrates typically associated with submerged macrophytes as well as planktonic crustaceans and rotifers occurred within algal mats at high densities.Aggregations of filamentous algae are usually short-term, with frequent appearances and disappearances. The rate of colonization of algal mats by invertebrates is rapid. In locations with a high degree of water exchange, animals colonize both living and decomposing algal mats at a similar rate, but in sheltered habitats, decomposing filamentous algae are colonized by a smaller number of animals.Comparison was made between the occurrence of invertebrate macrofauna on Chara spp., Potamogeton perfoliatus, P. lucens and Myriophyllum spicatum in several lakes. Although these macrophytes differ visibly in morphology and phenology, the number and composition of macroinvertebrates during summer was associated more closely with trophic state of a lake than with plant species.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.     

A multivariate analysis of phytoplankton and food web changes in a shallow biomanipulated lake

1. Phytoplankton dynamics, food chain changes and resilience in Lake Zwemlust, a shallow lake in The Netherlands, are described for the period 1986–94.
2. After biomanipulation in 1987, the lake moved through two alternative states, while the external nutrient loadings were maintained. A clear-water phase, mostly dominated by macrophytes, persisted from 1987 to 1991, and a rather turbid state, dominated by algae, occurred in the summers of 1992–94, after several consecutive and sustained perturbations affecting different parts of the food web in the lake. These two periods were characterized by different community structures.
3. The phytoplankton assemblage gradually changed in a pattern that reverted in later years towards that of the pre-biomanipulation stage, although the same species composition was not regained. This agrees with some mathematical models. During the clear-water phase, nutrient shortage, light climate and zooplankton feeding selected in favour of small, high surface : volume ratio and rapidly reproducing algae. However, in mid-summer of 1992–94, nutrient availability and cladoceran grazing on edible algae favoured cyanophytes.
4. Nutrients were transferred to higher trophic levels or lost from the system at relatively high rates when the lake was in a piscivore–macrophyte-dominated state, while they tended to accumulate in the algae in a planktivore-dominated chain without macrophytes. The role of weed beds was central for nutrient competition (mostly nitrogen) with algae, as well as a refuge and a base for alternative food sources to grazers. Weed beds seemed to have a strong effect in increasing connectedness, resilience and stability of the lake community.
5. The complete return of Zwemlust to a turbid state dominated by phytoplankton seems to have depended upon turnover of the limiting nutrient, which was retarded by macrophytes and stimulated by planktivorous fish and waterfowl.