Long-term Patterns of Shifts between Clear and Turbid States in Lake Krankesjön and Lake Tåkern

During the past century, Lake Tåkern and Lake Krankesjön, southern Sweden, have shifted repeatedly between a state of clear water and abundant submerged vegetation, and a state of turbid water and sparse vegetation. Long-term empirical data on such apparently alternative stable state dynamics are valuable as complements to modeling and experiments, although the causal mechanisms behind shifts are often difficult to identify in hindsight. Here, we summarize previous studies and discuss possible mechanisms behind the shifts. The most detailed information comes from monitoring of two recent shifts, one in each lake. In the 1980s, L. Krankesjön shifted to clear water following an expansion of sago pondweed, Potamogeton pectinatus. Water clarity increased when the pondweed was replaced by characeans. Zooplankton biomass in summer declined and the concentration of total phosphorus (TP) was reduced to half the previous level. The fish community changed over several years, including an increasing recruitment of piscivorous perch (Perca fluviatilis). An opposite directed shift to turbid water occurred in Lake Tåkern in 1995, when biomass of phytoplankton increased in spring, at the expense of submerged vegetation. Consistent with the findings in L. Krankesjön, phyto- and zooplankton biomass increased and the average concentration of TP doubled. After the shift to clear water in L. Krankesjön, TP concentration has increased during the latest decade, supporting the idea that accumulation of nutrients may lead to a long-term destabilization of the clear water state. In L. Tåkern, data on TP are inconclusive, but organic nitrogen concentrations oscillated during a 25-year period of clear water. These observations indicate that intrinsic processes cause gradual or periodic changes in system stability, although we cannot exclude the possibility that external forces are also involved. During such phases of destabilization of the clear water state, even small disturbances could possibly trigger a shift, which may explain why causes behind shifts are hard to identify even when they occur during periods of extensive monitoring.     

Long-term pattern of alternative stable states in two shallow eutrophic lakes

• 1 Lake Tåkern and Lake Krankesjön, two moderately eutrophic, shallow lakes in southern Sweden, have during the past few decades shifted several times between a clear-water state with abundant submerged vegetation and a turbid state with high phytoplankton densities.
• 2 Between 1985 and 1991, Lake Takern was in a clear state, whereas Lake Krankesjon shifted from a turbid to a clear state. During this shift, the area covered by submerged macrophytes expanded, followed by an increase in water transparency, plant-associated macroinvertebrates, and piscivorous fish. Nutrient concentrations, phytoplankton biomass and abundance of planktonic cladocerans decreased.
• 3 In both lakes, water level fluctuations were the most common factor causing shifts, affecting submerged macrophytes either through changes in light availability or through catastrophic events such as dry-out or mechanical damage by ice movement.
• 4 Our data give further support for the existence of two alternative stable states in shallow lakes maintained by self-stabilizing feedback mechanisms.

     

Waterfowl,macrophytes, and the clear water state of shallow lakes

The importance of lake ecosystems for waterfowl remains a topic of debate. In order to assess how temporal variations in lake features, specifically shifts between alternative stable states, may interact with the waterfowl fauna, we performed a long-term (22 years) study of the shallow Lake Krankesjön, southern Sweden. Lower total numbers of waterfowl occurred during periods with low macrophyte cover and turbid water, than when submersed macrophytes flourished and the water was clear. Some specific functional groups of waterfowl, such as herbivores, invertebrate, and fish feeders, showed a positive relation to clear water and high macrophyte cover. Hence, our data suggest that some migratory waterfowl may select lakes based on water quality, thereby adjusting their large-scale migratory routes. On the other hand, omnivorous waterfowl exhibited their highest abundances during turbid conditions. Furthermore, waterfowl not primarily relying on food from the lake showed no response to fluctuations in turbidity or macrophyte cover, but followed regional trends in population dynamics. In our study lake, L. Krankesjön, we estimated that waterfowl remove less than 3% of the macrophyte biomass during a stable clear-water state with lush macrophyte beds. However, during transition periods between alternative stable states, when macrophyte biomass is lower and the plants already stressed, the consumption rate of waterfowl may have a stronger effect on lake ecosystem functioning.     

Facilitation of clear-water conditions in shallow lakes by macrophytes: differences between charophyte and angiosperm dominance

A number of mechanisms result in a feedback between water clarity and macrophytes and, consequently, the occurrence of alternative stable states in shallow lakes. We hypothesize that bottom-up mechanisms and interactions within the benthic food web are more important in a charophyte-dominated clear-water state, while top-down mechanism and interactions in the planktonic food web prevail at angiosperm dominance. Charophytes, which dominate at lower nutrient concentrations and develop higher densities than most angiosperms, can have a higher influence on sedimentation, resuspension, and water column nutrients. During dominance of dense submerged vegetation like charophytes, zooplankton can be hampered by low food quality and quantity and by high predation pressure from juvenile fish, which in turn are favoured by the high refuge potential of this vegetation. Grazing pressure from zooplankton on phytoplankton can therefore be low in charophytes, but the main feedback in angiosperm-dominated ecosystems. Charophytes offer a higher surface than most angiosperms to periphyton, which favors benthic invertebrates. These support macrophytes by grazing periphyton and constitute a central link in a trophic cascade from fish to periphyton and macrophytes. To test these hypotheses, more experiments and field measurements comparing the effect of charophytes and angiosperms on water clarity are needed.     

Recovery of Potamogeton pectinatus L. stands in a shallow eutrophic lake under extreme grazing pressure

In shallow lakes, submerged macrophytes contribute to the stabilization of the clear water state. If lost, a number of mechanisms prevent re-colonization. Lake Müggelsee (730 ha) lost its submerged vegetation due to increasing eutrophication and switched to phytoplankton dominance in 1970. After the reduction of nutrient loading in 1990, Potamogeton pectinatus L. started re-colonizing the lake. During the following years, it spread at a mean rate of 2.5 ha per year to all available areas <80 cm depth. Between 1993 and 1999, decreasing maximum biomass indicated hampered growth. Exclosure experiments revealed that herbivory reduced the aboveground biomass by more than 90%. Both waterfowl and fish were found to contribute to the grazing pressure despite a low abundance of the known herbivorous fish species and waterfowl in spring and summer. Protection of stands against grazing resulted in higher biomass of shoots, whereas shoot and tuber density did not change. Both shading by phytoplankton and periphyton, as well as grazing pressure, prevented the submerged vegetation of Lake Müggelsee from developing back to a dense zone that contributed to the reduction of turbidity.     

Trophic web structure in a shallow eutrophic lake during a dominance shift from phytoplankton to submerged macrophytes

In Lake Krankesjön, southern Sweden, sago pondweed (Potamogeton pectinatus L.) and a stonewort (Chara tomentosa L.) expanded spatially during the second half of the 1980's after more than a decade of phytoplankton blooms and sparse submerged vegetation. During the expansion of submerged plants the number of resting and breeding waterfowl increased. The increase was significant for herbivorous birds such as coot (Fulica atra L.) and mute swan (Cygnus olor (Gmelin)), but also for omnivorous dabbling ducks. The shift from phytoplankton to submerged macrophytes caused structural changes on higher trophic levels, and an altered trophic web developed. The density of planktonic Cladocera decreased, which is suggested to be a result of decreased phytoplankton productivity and biomass as nutrient levels dropped. The benthic macroinvertebrate assemblage changed from low diversity and biomass dominated by Chironomidae and Oligochaeta on bare sediment, to high diversity and biomass characterized by plant-associated forms like snails and isopods in areas covered by macrovegetation. The mean size of perch (Perca fluviatilis L.) increased, probably as a result of higher availability of macroinvertebrates in the vegetation. The perch reached a mean size where the species is known to shift to a fish diet, permitting an increased top down effect on the ecosystem. The results support the idea that shallow eutrophic lakes can shift between two states, each one stabilized by feed-back mechanisms including both biotic and abiotic factors. Shifts between these states are suggested to be a possible explanation for observed drastic changes in abundance of waterfowl in shallow eutrophic lakes.     

The submerged macrophytes in Lake Maarsseveen I: Changes in species composition and biomass over a six year period

The submerged macrophytes of Lake Maarsseveen I were surveyed in 1983 using SCUBA diving techniques. Only 40% of the characeans and 75% of the angiosperms detected in 1977 remained. The area colonized by submerged macrophytes was 0.45% in 1983, compared with 25.10% in 1977. The observed decreases were largely attributable to a shift of the plantcolonized areas to shallower depths. By 1983, most of the earlier predominant vegetation types had disappeared and the biomass had decreased. The decline in submerged vegetation may be attributed to increasing eutrophication, fish populations and recreational activities.     

Dynamics of submerged macrophyte populations in response to biomanipulation

1. A 6‐year study (1992–97) of changes in submerged vegetation after biomanipulation was carried out in the eutrophicated Lake Finjasjön, Southern Sweden. Ten sites around the lake were revisited each year. At each site five samples of above‐ground biomass were taken at 10 cm water depth intervals. An investigation of the seed bank at the 10 sites, and a grazing experiment where birds and large fish were excluded was also conducted. 2. Between 1992 and 1996, in shallow areas (water depth < 3 m), vegetation cover increased from < 3 to 75% and above‐ground biomass from < 1 to 100 g DW m^–2. Mean outer water depth increased from 0.3 to 2.5 m. Elodea canadensis and Myriophyllum spicatum accounted for > 95% of the increase in biomass and plant cover. The following year (1997), however, cover and above‐ground biomass decreased, mainly attributable to the total disappearance of E. canadensis. Secchi depth increased after biomanipulation until 1996, but decreased again in 1997. 3. Total and mean number of submerged species increased after biomanipulation, probably as a result of the improved light climate. However, after the initial increase in species number there was a decrease during the following years, possibly attributed to competition from the rapidly expanding E. canadensis and M. spicatum. The lack of increase in species number after the disappearance of E. canadensis in 1997 implies that other factors also affected species richness. 4. A viable seed bank was not necessary for a rapid recolonization of submerged macrophytes, nor did grazing by waterfowl or fish delay the re‐colonization of submerged macrophytes. 5. Submerged macrophytes are capable of rapid recolonization if conditions improve, even in large lakes such as Finjasjön (11 km²). Species that spread by fragments will increase rapidly and probably outcompete other species. 6. The results indicate that after the initial Secchi depth increase, probably caused by high zooplankton densities, submerged vegetation further improved the light climate. The decrease in macrophyte biomass in 1997 may have caused the observed increase in phosphorus and chlorophyll a, and the decrease in Secchi depth. We suggest that nutrient competition from periphyton, attached to the macrophytes, may be an important factor in limiting phytoplankton production, although other factors (e.g. zooplankton grazing) are also of importance, especially as triggers for the shift to a clear‐water state.     

Abrupt shift from clear to turbid state in a shallow eutrophic, biomanipulated lake

Monitoring data were used to assess causes behind a recent shift from a clear-water to a turbid-water state in Lake Major, a 10 ha shallow lake in Hungary. In 1999–2000, fish manipulation was conducted in this hypertrophic lake. Reduced fish stock resulted in clearing water and the development of a dense (>80% coverage) submerged vegetation in 2005. During the recent abrupt shift, which occurred in 2007, submerged vegetation subsequently declined after a two-year period of clear water and abundant vegetation. An intense decay of macrophytes within the lake produced a rapid transition between the clear- and turbid-water states. During the clear-water state in 2005–2006, the most important variables predominantly correlating with macrophyte cover were Secchi transparency, temperature and TN, while TN, temperature, Secchi depth and chlorophyll-a were the most significant variables during the turbid-water state in 2007. Nitrogen may play a significant role in the cover of submerged macrophytes when TP is moderate. We argue that several factors in concert are necessary to initiate a shift. Water temperature likely has contributed to triggering shift through inter-year-dependent changes in cover of macrophytes, with fish recruitment having key roles in the dynamics of shallow lakes. Handling editor: Luigi Naselli-Flores     

Depth limits and minimum light requirements of freshwater macrophytes

1. Data for maximum colonization depth (Z_c) of five groups of submerged macrophytes and light attenuation were collected for forty-five Danish lakes and 108 non-Danish lakes. The macrophyte groups were bryophytes, charophytes, caulescent angiosperms, rosette-type angiosperms and Isoetes spp. 2. The data showed systematic differences among the groups in the relationship of Z_c to water transparency. In lakes with low transparency (Secchi disc transparency (Z_s) less than 7 m) caulescent angiosperms and charophytes penetrated deepest followed by bryophytes and Isoetes spp. In more transparent lakes bryophytes grew deepest, followed by charophytes, caulescent angiosperms and Isoetes spp. Rosette-type angiosperms had the lowest Z_c in all types of lakes. Charophytes and caulescent angiosperms had similar depth limits in lakes with Z_s < 4 m but charophytes grew deeper in more transparent lakes. The depth limits of both groups were independent of light penetration in lakes with very low transparency (Z_s < 1 m). The annual light exposure for the deepest growing macrophytes (bryophytes) was 20–95 mol photons m^–2. 3. The relationship between Z_c, macrophyte type and lake transparency could be explained by three distinct processes regulating Z_c. In lakes with low transparency (Z_s < 1 m), tall macrophytes (caulescent angiosperms and charophytes) compensate for light limitation by shoot growth towards the water surface and Z_c is therefore independent of transparency. In lakes with medium transparency (1 m < Z_s < 4 m) Z_c for angiosperms, charophytes and Isoetes spp. is constrained by light attenuation in the water column, corresponding to a linear relationship between Z_c and Z_s. This pattern also applies to bryophytes, despite lake transparency. In transparent lakes, the minimum light requirement at Z_c increased with increasing transparency for angiosperms, charophytes and Isoetes spp. 4. The minimum light requirements among submersed macrophytes (including marine macroalgae) depend on their plant-specific carbon value (plant biomass per unit of light-absorbing surface area) for the species/group, indicating that the light requirements of submersed plants are tightly coupled to the plants’ possibility to harvest light and hence to the growth form. 5. The light requirements increased on average 0.04% surface irradiance per degree increase in latitude corresponding to an average decrease in Z_c of 0.12 m per degree latitude.     

Submerged vegetation development in two shallow, eutrophic lakes

Submerged macrophyte vegetation in two shallow lakes in the Netherlands, Lake Veluwemeer and Lake Wolderwijd, has been affected by eutrophication in the late 1960's and 1970's. Recent changes in the vegetation occurred in the period following lake restoration measures. Between 1987 and 1993, the dominance of Potamogeton pectinatus decreased, while Charophyte meadows expanded over the same time interval. The pattern of change of the dominant macrophyte species might result from changes in the underwater light climate. Seasonally persistent clear water patches associated with the Chara meadows have been observed in the last few years. The interaction between submerged macrophyte vegetation succession and water transparency in the lakes is discussed.     

沉水植物重建对富营养水体氮磷营养水平的影响

利用富营养浅水湖泊(武汉东湖)中所建立的大型实验围隔系统,研究了沉水植物对水体N、P营养水平的影响．结果表明,沉水植物重建后N、P营养水平显著降低．在研究期间,水生植物围隔总N和总P水平均显著低于对照围隔和大湖水体,而且水生植物围隔的总P含量一般维持在0．1mg·L^-1左右。季节性波动远低于对照围隔和大湖水体．水生植物围隔水体中氨态氮和亚硝态氮含量较低．而硝态氮含量与对照围隔和和大湖水体差别不大．由此可见。恢复以沉水植物为主的水生植被,可以有效地降低N、P营养循环速度,控制浮游植物过度增长,是重建富营养湖泊生态系统的重要措施．     

西太湖水生植物时空变化

水生植物在浅水湖泊生态系统中具有十分重要的作用。根据中国科学院太湖湖泊生态系统研究站1989年以来的常规监测资料,将西太湖（除东太湖以外的湖区）划分为9个区,采用点截法（point intercept method）,于2002～2005年对各区水生植物的种类、生物量和空间分布情况进行了6次调查。结果表明：西太湖现有水生植物16种,分属于11科12属;水生植物总面积约10220hm^2,其中沉水植物分布面积约占64．58％;挺水植物约占0．29％;漂浮植物约占38．16％。各个种之间生物量差异显著,马来眼子菜、荇菜、芦苇的生物量在所有水生植物中居前3位。多样性分析表明,水生植物种类4a来未发生明显变化,但种类和生物量季节性差异较大。水生植物呈环状分布在距湖岸5km以内的水域和部分岛屿周围,东岸和南岸为水生植物的主要集中分布区域,分布区连续性好,且水草种类齐全。挺水植物种类单一,仅有芦苇（Phragmites communis）一种,分布区域多限于水深小于1．6m的湖岸;沉水植物共有8种,为水生植物的主要组成部分,马来眼子菜（Potamogeton malaianus）的分布频度最高,在西山岛周围水域逐年扩张,成为该区域的先锋种;漂浮植物3种,主要以荇菜（Nymphoides peltata）为主,在七都水域有逐渐扩张的趋势。马来眼子菜、芦苇、荇菜表现出对水环境较强的适应能力,目前为西太湖的3个优势种。20世纪50年代以来,西太湖水生植物种类减少了50种,其中水质下降是导致水生植物种类不断减少甚至消失的一个重要原因。围网养殖和不合理的捕捞方式也对局部水域的植物造成极大的破坏。水生植物生存环境日益严峻,种群单一化趋势日益明显。     

Influence of submerged vegetation and fish abundance on water clarity in peri-urban eutrophic ponds

Despite the presence of high nutrient concentrations, most ponds located around Brussels (Belgium) show a considerable variation in turbidity. The importance of submerged macrophytes in maintaining the clear-water state requires identification of the main factors determining macrophyte abundance and diversity in ponds and small lakes. In this study, the inter-relationships between submerged macrophyte cover, fish abundance and turbidity were investigated in 13 eutrophic peri-urban ponds. Along a turbidity gradient, vegetation switched from dominance by Stoneworts (Chara and Nitella spp.) in the clearest ponds, to dominance by Potamogeton pectinatus in ponds with a slightly lower water transparency. Despite the presence of both P. pectinatus and Stoneworts in each of the vegetated ponds, only one became dominant. Only a very low abundance (around 20%) of submerged vegetation was found in ponds of intermediate turbidity, while macrophytes were absent in turbid ponds. Multi- and univariate analysis showed a marked difference in chemical, physical and biological properties between ponds deliberately used for fish stocking and ponds that were not. Macrophyte cover was significantly negatively correlated with turbidity and plankti-benthivorous fish abundance. No such correlation was observed with piscivorous fish abundance, except for pike that were associated with a charophyte vegetation in the study ponds. The strong relationship found between fish abundance and turbidity, its negative effect on submerged vegetation cover, and the importance of submerged vegetation in controlling phytoplankton abundance, should be taken into account when selecting ponds for fish stocking. It also suggests that the study ponds have a good potential for ecological quality restoration by biomanipulation.     

Effects of elevated turbidity on shallow lake fish communities

Synopsis We compared the fish communities of two shallow lakes in the lower Waikato River basin, North Island, New Zealand, to determine the effects of elevated suspended solids (SS) and collapse of submerged macrophytes. Lake Waahi was turbid (20–40 g m^-3 SS) and devoid of submerged macrophytes whereas Lake Whangape was clearer (5 g m^-3 SS) and dominated by submerged macrophytes. The lakes had similar fish species richness and had nine major species in common; representing eight families including Anguillidae, Retropinnidae, Galaxiidae, Eleotridae, Mugilidae, Ictaluridae, Poeciliidae, and Cyprinidae (two species). The only major fish that was absent from Lake Waahi was a lacustrine form of the common smelt, Retropinna retropinna, which disappeared after the lake became turbid in the late 1970s. CPUE, condition, and size of most species in Lake Waahi were similar to, or greater than, those in Lake Whangape. Lake Whangape clearly exceeded Lake Waahi only for CPUE of two species. Within Lake Whangape two species displayed significantly greater condition, and one species greater size, in a turbid arm of the lake than in the main basin. Apart from lacustrine Retropinna retropinna, the fish in these lakes appear well adapted to cope with, or to avoid, the direct toxic effects of suspended and settleable solids on sensitive early developmental stages. In Lake Waahi loss of cover and food provided by submerged macrophytes appears to have been compensated for by increased turbidity and an associated increase in the biomass of the mysid, Tenagomysis chiltoni (a major prey item).     

Clear,crashing, turbid and back – long‐term changes in macrophyte assemblages in a shallow lake

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Colonization and succession of submerged macrophytes in shallow Lake Væng during the first five years following fish manipulation

Colonization of submerged macrophytes and changes in species composition were studied in shallow Lake Væng during the first five years (1987–91) following fish manipulation in 1986–1988 and a resultant significant improvement in lake water transparency. No submerged macrophytes were present in the lake from 1981–1986, during which time the summer mean Secchi depth ranged from 0.6 and 0.8 m. From 1987 to 1990, Secchi depth increased from 0.9 m to 1.8 m and macrophyte coverage consequently increased (1 % of the lake area in 1987, 2% in 1988, 50% in 1989, 80% in 1990 and 90% in 1991). At the same time, the macrophytes became taller, and the weedbeds more dense. The macrophytes colonized from the exposed and deeper part of the lake towards the sheltered and more shallow part of the lake, a colonization pattern that was confirmed by transplantation experiments. The delay in colonization of the shallow parts may be caused by waterfowl grazing. The vegetation was initially dominated by Potamogeton crispus L., but there was a gradual change during 1988–1989 and Elodea canadensis Michx became exclusively dominant in 1990–1991.     

Vegetation abundance in lowland flood plan lakes determined by surface area, age and connectivity

SUMMARY 1. We analysed the vegetation structure of 215 lakes in the flood plain of the river Lower Rhine in relation to environmental variables related to hydrological connectivity, lake morphometry, lake age and land use on adjacent land. 2. The frequency distribution of the cover of submerged macrophytes was not normal, implying that submerged macrophytes in any one lake were either scarce or abundant. 3. We observed clear water lakes with submerged macrophyte dominance over a wide range of total P concentration (0.020–0.40 mg total P L^?1). 4. Multiple logistic regression indicated that the probability of dominance by submerged macrophytes decreased markedly with the surface area, depth and age of the lakes. The surface area effect occurred independently of the depth. Further, there was a negative relationship between submerged macrophyte dominance and the long‐term annual duration of inundation by the river. 5. Nymphaeid cover showed a distinct optimum with respect to mean lake depth, being almost absent in lakes shallower than 0.5 m. In contrast to what was found for submerged plants, the probability of occurrence of nymphaeids increased with lake age. 6. The probability of helophyte occurrence increased with lake age, and decreased with the presence of trees, cattle grazing, surface area, use of manure and mean lake depth. 7. In all cases the critical level of one factor (e.g. mean lake depth) depended on other factors (e.g. surface area or age of lake). Thus, in the present study, small lakes tended to remain dominated by submerged macrophytes up to a greater depth than large lakes, and helophytes colonised smaller lakes in an earlier phase. 8. The effect of inundation by the river was modest. This could be because most of our lakes are rarely inundated during the growing season and experience only moderate current velocities while flooded. 9. The results have practical implications for future management of flood plains for conservation purposes. In new water bodies, macrophyte domination will be promoted if many small shallow lakes, rather than few large deep ones, are excavated.     

Seasonal changes of mechanisms maintaining clear water in a shallow lake with abundant Chara vegetation

The study is based on monitoring data on the seasonal variation during four (1996–1999) vegetation periods, as well as long-term summer data on submerged vegetation, nutrients, light, phytoplankton and zooplankton in Lake Krankesjön, a shallow, calcium-rich, moderately eutrophic lake in southern Sweden.The lake has been in the clear water state with abundant submerged vegetation since the end of the 1980s. Somewhat lower summer biomass of submerged macrophytes during 1997 and 1999 indicates a temporary instability of the clear water state. During these 2 years, summer transparency was about 1.2–2.1 m, while concentrations of total phosphorus and chlorophyll (Chl) a were about 26–40 and 8–18 μg l⁻¹, respectively.Summer biomass of submerged macrophytes was higher during 1996 and 1998. In both years, a distinct increase in light availability and decrease in concentrations of nutrients and chlorophyll occurred simultaneously with the development of dense Chara vegetation. Summer values for transparency were about 2.0–2.5 m, while concentrations of total phosphorus and Chl a were about 20–32 and 4–11 μg l⁻¹, respectively.Summer biomass of crustacean zooplankton was below 250 μg l⁻¹ during all 4 years. A peak abundance of Cladocera (mainly Bosmina longirostris) during May or June caused only a short-term reduction in chlorophyll concentrations that was more pronounced in 1997 than in 1996.Measured light attenuation during 1999 was closely correlated with light attenuation calculated from the amount of suspended solids, chlorophyll concentrations, and water colour. Detritus contributed most to the total amount of suspended solids, while chlorophyll was the main contributor to light attenuation.A long-term decrease of the ratios between chlorophyll and total phosphorus suggests that phytoplankton in the clear water state is limited by factors other than total phosphorus concentrations. Increased sedimentation rate, carbon limitation, allelopathy and a lower bioavailable fraction of the total amount of phosphorus are possible explanations, while nitrogen limitation and grazing from zooplankton probably are of minor importance.Possible reasons for the “instability” of the clear water state during 1997 and 1999 are discussed. Unusually high water level as well as cold and windy weather during the spring of 1996–1999 may have caused a slow and late growth of the plants and thus a temporary instability. However, a tendency for an increase in total phosphorus concentrations and sediment accumulation along the wind-protected shores during the clear water state indicate the possibility of a long-term destabilization which contradicts the alternative stable states model.     

Reduction of benthic macroinvertebrates due to waterfowl foraging on submerged vegetation during autumn migration

If present in large numbers, as during migration, herbivorous waterfowlmay reduce the amount of submerged vegetation. Because the vegetation is a keyfactor in shallow eutrophic lakes, removal of the green biomass can be expectedto affect also other biota that depend on the vegetation. We conducted anexperiment to determine how the abundance of chironomids andPisidium sp. were affected by intense foraging ofwaterfowlon the submerged plant Potamogeton pectinatus. This wasdone in Lake Ringsjön in southern Sweden, during the autumn migration ofthe birds. Three treatments, replicated six times, were used: (i) closed cagesthat excluded all waterfowl, (ii) semi-open cages that excluded only largewaterfowl (geese and swans), and (iii) open plots where all waterfowl couldfreely enter. Waterfowl densities were monitored during the experiment. Theresults suggest that the foraging of large waterfowl (swans) had a clearlynegative effect on macroinvertebrate abundance and aboveground biomass ofP. pectinatus. At the end of the experiment, the densityofchironomids was about 46% lower in the open than in the closed cages. Ingeneral, the density of Pisidium sp. tended to be lower inthe open plots. Small waterfowl alone did not seem to affect either thevegetation or macroinvertebrates. We suggest that thePisidium sp. was influenced at an early stage of grazing,when waterfowl foraged on aboveground biomass, whereas chironomids wereaffectedat a later stage, when swans were digging for below-ground tubers.