Long- and short-term dynamics of submerged macrophytes in two shallow eutrophk lakes

1. Periods with clear water and abundant submerged vegetation have alternated with periods of turbid water and sparse vegetation during recent decades in Lake Tåkern and Lake Krankesjön, two shallow, calcium-rich, moderately eutrophic lakes in southern Sweden, Between 1983 and 1991, submerged vegetation (predominant species: Chara tomentosa, Nitellopsis obtusa, Myriophyllum spicatum) covered about 50% of the open lake area in Lake Tåkern. In Lake Krankesjön, submerged vegetation was sparse during 1983–84, but increased continuously in the following years and covered about 50% of the open lake area by 1990 and 1991. Potamogeton pectinatus was the first species to expand in Lake Krankesjön, but was later replaced by C. tomentosa. 2. During 1983–84, turbidity was high in Lake Krankesjön, which indicated that submerged macrophytes were light-limited. During 1986–91, there was a negative correlation between the areal coverage of charophytes and angiosperms, indicating that competition for space had become an important limiting factor. The same negative correlation was found in Lake Tåkern for 1983–91. 3. Charophytes had much higher biomass per unit area than angiosperms in both lakes and reduced water movement considerably. This was probably one reason for the increase of water transparency in Lake Krankesjön during the spatial expansion of these plants. Charophytes also stored large amounts of phosphorus and nitrogen, Charophytes are probably superior competitors for both space and nutrients and thus have competitive advantage over angiosperms in this lake type. 4. In Lake Krankesjön, both P. pectinatus and C. tomentosa were negatively affected by high water level during the growing period. Total disappearance of submerged vegetation occurred in both lakes after catastrophic events (dry-out during summer or mechanical damage by ice) caused by extremely low water level. Changes in water level are thus one of the most important reasons for among-year fluctuations in areal coverage of submerged macrophytes in these lakes.     

The species composition,occurrence and temporal stability of submerged aquatic macrophyte patches along the main channel border of pool 5A,Upper Mississippi River

Species composition, relative abundance, distribution and physical habitat associations of submerged aquatic macrophytes in the main channel border (MCB) habitat of Pool 5A, Upper Mississippi River (UMR) were investigated during the summers of 1980 and 1983. The submerged aquatic macrophytes in Pool .5A MCB were a small and stable component of the river ecosystem. Submerged plants occurred primarily in small, monospecific clumps. Clumps in close proximity to each other formed plant patches. Plant patches were stable in location and number between 1980 and 1983; 82.5% of the patches first observed in 1980 were present in 1983. Submerged macrophytes covered about 10–12 ha of the 201 ha MCB in Pool 5A. Submerged plants were most common in the lower two-thirds of the pool. Ten species of aquatic macrophytes occurred on rock channel-training structures and eleven occurred on non-rock substrates in the MCB. The most common submerged plants, in order of abundance, were Vallisneria americana Michx., Heteranthra dubia Jacq., Potamogeton pectinatus L., Ceratophyllum demersum L. and Potamogeton americanus C. & S.     

Loss of Submerged Macrophytes in Shallow Lakes in North‐Eastern Germany

During the 1950s, the submerged vegetation of shallow lakes in north‐eastern Germany was dominated by nutrient tolerant species, with Ceratophyllum demersum and Myriophyllum sp. being most common. Almost one third of 300 investigated lakes had already lost their submerged macrophytes at that time. Very shallow lakes showed either high or low macrophyte abundance. Increasing depth resulted in medium macrophyte abundances, which may contribute to the stabilisation of local or temporary clearwater states. Forty years later, the percentage of lakes without macrophytes had dramatically increased. Between 55 and 85% of the investigated lakes showed a low abundance. The decline was most pronounced in very shallow lakes. The majority of the investigated lakes showed summer TP concentrations below 100 μg L^–1, but no colonisation by submerged macrophytes, which indicates a resilience against re‐colonisation.     

Stabilizing the clear-water state in eutrophic ponds after biomanipulation: submerged vegetation versus fish recolonization

Biomanipulation through fish removal is a tool commonly used to restore a clear-water state in lakes. Biomanipulation of ponds is, however, less well documented, although their importance for biodiversity conservation and public amenities is undisputed. In ponds, a more complete fish removal can be carried out as compared to lakes and therefore a stronger response is expected. Fish recolonization can, however, potentially compromise the longer term success of biomanipulation. Therefore, we investigated the impact of fish recolonization on zooplankton, phytoplankton, and nutrients for several years after complete drawdown and fish removal in function of submerged vegetation cover in 12 peri-urban eutrophic ponds situated in Brussels (Belgium). Fish recolonization after biomanipulation had a considerable impact on zooplankton grazers, reducing their size and density substantially, independent of the extent of submerged vegetation cover. Only ponds with <30% cover of submerged vegetation shifted back to a turbid state after fish recolonization, coinciding with an increase in density of small cladocerans, rotifers, and cyclopoid copepods. In ponds with >30% submerged vegetation cover, macrophytes prevented an increase in phytoplankton growth despite the disappearance of large zooplankton grazers. Our results suggest that macrophytes, rather than by providing a refuge for zooplankton grazers, control phytoplankton through other associated mechanisms and confirm that the recovery of submerged macrophytes is essential for biomanipulation success. Although the longer term effect of biomanipulation is disputable, increased ecological quality could be maintained for several years, which is particularly interesting in an urban area where nutrient loading reduction is often not feasible.     

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     

Wave exposure related growth of epiphyton: implications for the distribution of submerged macrophytes in eutrophic lakes

The distribution of submerged macrophytes in eutrophic lakes has been found to be skewed towards sites with intermediate exposure to waves. Low submerged macrophyte biomass at exposed sites has been explained by, for instance, physical damage from waves. The aim of this study was to investigate if lower biomass at sheltered sites compared to sites with intermediate exposure to waves can be caused by competition from epiphyton.Investigations were performed in eutrophic lakes in southern Sweden. Samples of submerged macrophytes and epiphytic algae on the macrophytes were taken along a wave exposure gradient. The amount of epiphyton (AFDW) per macrophyte biomass decreased with increased exposure. Biomass of submerged macrophytes, on the other hand, increased with increased exposure until a relatively abrupt disappearance of submerged vegetation occurred at high exposures. Production of epiphytic algae was monitored on artificial substrates from June to September at a sheltered and an exposed site in three lakes. It was higher at sheltered sites compared with exposed sites.We suggest that epiphytic algae may be an important factor in limiting the distribution of submerged macrophytes at sheltered sites in eutrophic lakes.     

Echosounding observations of coverage,height, PVI,and biomass of submerged macrophytes in the southern basin of Lake Biwa,Japan

We have developed a procedure to process echosounding data to map the distribution of submerged aquatic macrophytes in the southern basin of Lake Biwa, a water body that has a surface area of 52 km² and a mean depth of 4 m. Echosounding observations were made along 27 transect lines spaced at 500-m intervals on August 4 and September 2 and 30, 2003. Quantitative vegetation data including percent coverage, mean vegetation height, and percent vegetation infestation were directly determined using image data from the echosounder recorded digitally on videotape. Based on the image data from an echosounder, a regression model was developed for estimating biomass of submerged macrophytes. The regression model using the total echo strength as the explanatory variable could reliably estimate macrophyte biomass up to 300 g m⁻². Distribution maps of macrophyte height and biomass suggest that the recent summer decline of submerged macrophytes started earlier in shallow areas (<3 m of depth) than deep areas (>4 m) in the southern basin of Lake Biwa.     

Effects of aquatic vegetation type on denitrification

In a microcosm ¹⁵N enrichment experiment we tested the effect of floating vegetation (Lemna sp.) and submerged vegetation (Elodea nuttallii) on denitrification rates, and compared it to systems without macrophytes. Oxygen concentration, and thus photosynthesis, plays an important role in regulating denitrification rates and therefore the experiments were performed under dark as well as under light conditions. Denitrification rates differed widely between treatments, ranging from 2.8 to 20.9 ??mol N m^?2 h^?1, and were strongly affected by the type of macrophytes present. These differences may be explained by the effects of macrophytes on oxygen conditions. Highest denitrification rates were observed under a closed mat of floating macrophytes where oxygen concentrations were low. In the light, denitrification was inhibited by oxygen from photosynthesis by submerged macrophytes, and by benthic algae in the systems without macrophytes. However, in microcosms with floating vegetation there was no effect of light, as the closed mat of floating plants caused permanently dark conditions in the water column. Nitrate removal was dominated by plant uptake rather than denitrification, and did not differ between systems with submerged or floating plants.     

六种沉水植物的克隆生长特征

为构建种群动态模型以指导沉水植被修复工程实践, 研究采用同质园实验方法对6种常见沉水植物(竹叶眼子菜(Potamogeton wrightii)、眼子菜(P. distinctus)、光叶眼子菜(P. lucens)、穿叶眼子菜(P. perfoliatus)、扭叶眼子菜(P. intortifolius)和苦草(Vallisneria natans)的克隆生长模式进行了连续观测研究, 获取了分株形成速率、空间扩张速率、株高增加速率等种群扩张动态参数,及分株数、间隔子长度、分株高度等克隆构件特征参数。结果表明, 6种沉水植物的分株数从28d开始增长, 其中苦草的分株形成速率最高, 平均为1.09株/d, 分株形成最大速率出现在55d之后; 穿叶眼子菜和扭叶眼子菜的分株形成速率低于苦草, 但是高于竹叶眼子菜、眼子菜和光叶眼子菜, 最大速率出现在41d之后。虽然苦草的分株最多, 但是分株的株高最低, 其株高增长速率均值为0.2 cm/d。眼子菜属物种中竹叶眼子菜和眼子菜株高增长速率最高, 光叶眼子菜的株高增长速率和分株形成速率都最低。克隆系占据面积随时间的扩张速率为穿叶眼子菜(113.22 cm2/d)>扭叶眼子菜(71.70 cm2/d)>苦草(35.48 cm2/d)>竹叶眼子菜(12.09 cm2/d)>眼子菜(3.07 cm2/d)>光叶眼子菜(0.53 cm2/d)。此外, 研究还发现眼子菜属植物普遍表现出匍匐茎上“节”的形成, 而苦草则不具备这种特性, 匍匐茎“节”的形成及随之形成的不定根在眼子菜属植物空间扩张过程中具有重要的生态功能, 并在种群构建方面与苦草等其他物种发生分异。基于眼子菜属植物匍匐茎上的“节”可以形成跳跃性的分株, 在种群面积扩张方面更具优势; 而苦草形成分株的数量更多、速度更快, 在提高种群密度保障种群稳定方面更有优势。     

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.     

The role of solid waste materials as habitats for macroinvertebrates in a lowland dam reservoir

Eight hypereutrophic phytoplankton dominated ponds from the Brussels Capital Region (Belgium) were biomanipulated (emptied with fish removal) to restore their ecological quality and reduce the risk of cyanobacterial bloom formation. Continuous monitoring of the ponds before and after the biomanipulation allowed the effects of the management intervention on different compartments of pond ecosystems (phytoplankton, zooplankton, submerged vegetation and nutrients) to be assessed. Fish removal resulted in a drastic reduction in phytoplankton biomass and a shift to the clear-water state in seven out of eight biomanipulated ponds. The reduction in phytoplankton biomass was associated with a marked increase in density and size of large cladocerans in six ponds and a restoration of submerged macrophytes in five ponds. The phytoplankton biomass in the ponds with extensive stands of submerged macrophytes was less affected by planktivorous fish recolonisation of some of the ponds later in the summer. The two non-vegetated ponds as well as one pond with sparse submerged vegetation showed a marked increase in phytoplankton biomass associated with the appearance of fish. Phytoplankton biomass increase coincided with the decrease in large Cladocera density and size. One pond lacking submerged macrophytes could maintain very low phytoplankton biomass owing to large Cladocera grazing alone. The results of this study confirmed the importance of large zooplankton grazing and revegetation with submerged macrophytes for the maintenance of the clear-water state and restoration success in hypereutrophic ponds. They also showed that large Cladocera size is more important than their number for efficient phytoplankton control and when cladocerans are large enough, they can considerably restrain phytoplankton growth, including bloom-forming cyanobacteria, even when submerged vegetation is not restored. The positive result of fish removal in seven out of eight biomanipulated ponds clearly indicated that such management intervention can be used, at least, for the short-term restoration of ecological water quality and prevention of noxious cyanobacterial bloom formation. The negative result of biomanipulation in one pond seems to be related to the pollution by sewage water. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

A new sampler for collecting invertebrates in submerged vegetation

A new sampler is described that can rapidly enclose a specific volume of water at a given depth, including submerged macrophytes and associated invertebrates. The sampler is a tong-shaped instrument consisting of two metal rods connected by a flexible joint. At the end of each arm is a metal cylinder (diameter 10 cm); one end of the cylinder has a sharpened edge and the other is equipped with a net. Powerful metal springs force the cylinders together at high speed, and macrophytes are enclosed in the nets. The sampler is suitable for studying vertical and horizontal distribution of invertebrates, even fast-moving taxa, within submerged vegetation. The device causes little disturbance of the vegetation, hence it is also suitable for repeated sampling within mesocosms.     

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.     

Amphipoda (Crustacea) in the Littoral Zone of Lake Balaton (Hungary). Qualitative and Quantitative Studies

Species composition, size distribution, egg number and biomass of different amphipod species on littoral submerged aquatic vegetation (Myriophyllum, spicatum and Potamogeton perfoliatus stands) and on the stony shores of Lake Balaton at Tihany were estimated in August 1983 and 1985. Three species occurred in the material: Corophium curvispinum, Dikerogammarus haemobaphes and Dikerogammarus villosus. The bulk was formed by C. curvispinum. The mean number of eggs per female of the three investigated species showed differences at the different sampling stations and ranged between the following values: C. curvispinum from 3.3 to 8.8, D. haemobaphes from 16.4 to 22.8 and D. villosus from 8.0 to 17.5. The total biomass of amphipods was 1.8 and 3.5 mg dry weight/g fresh weight of macrophytes in 1983 and 1985 respectively, and 4.7 mg animal per dm² stone surface in the stony littoral zone in 1983.     

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.     

The aquatic macrophytes of Lake Vechten. Species composition,spatial distribution and production

The macrophytic species compositions in Lake Vechten of 1963 and 1979–80 were compared and showed a considerable change. The main vegetation types were mapped in 1973, 1978, 1979 and 1980. All macrophytic communities, i.e. submerged, floating-leaved and emergent vegetation types, declined mainly due to increasing water turbidity, increasing tree-shading and, from 1978 onwards, grazing and trampling by cattle.Production rates, derived from different combinations of measurement and calculation, were compared. The macrophytes contributed only about 7% to the total lake production in 1980.     

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．     

Macrophyte loss drives decadal change in benthic invertebrates in peatland drainage ditches

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