Hydrobiologia 506–509: 29–35, 2003.

© 2003 Kluwer Academic Publishers. Printed in the Netherlands.

29

Eutrophication and restoration of shallow lakes – the concept of stable

equilibria revisited

Martin T. Dokulil & Katrin Teubner

Institute of Limnology, Austrian Academy of Sciences, Mondseestrasse 9, A-5310 Mondsee, Austria
E-mail: martin.dokulil@oeaw.ac.at

Key words: equilibrium, stable states, urban lake, management, phytoplankton, macrophytes

Abstract
Shallow polymictic lakes are very abundant in many regions of the world, they are more numerous than deep lakes
and are of great importance especially in densely populated areas even when small. Their ecology differs from
stratified lakes in many ways. During the eutrophication process, lakes may change from a clear water, macrophyte
dominated state to an algal dominated state, a process known as the concept of ‘alternative stable equilibria’. The
usefulness of this concept as a descriptor for the eutrophication process and as the basis for rehabilitation of lakes is
tested on long-term data from two contrasting case study sites using a novel multivariate bubble-plot technique. In
one case the concept proved to be useful and successful while it could not be applied to a large shallow turbid lake.
The equilibrium concept is then generalised using further examples. Shallow lakes which are either macrophyte
dominated or light limited because of high inorganic background turbidity have ratios of total phosphorus (TP) to
algal Chlorophyll-a at or below 3:1. Deep oligotrophic or algal dominated lakes in contrast exhibit ratios between
3:1 and 1:1. Alternative stable equilibria of macrophyte or algal domination can occur within the same water-body
at various locations, at different times of the year or during trophic development.

Introduction situations of stable states within phytoplankton as-

semblages or more than two equilibria have attracted
Shallow lakes are more numerous worldwide than much less attention (Scheffer, 1998).
deep lakes and provide humankind with many ser- This paper tries to explore how generally ap-
vices such as nutrition, water for drinking, irriga- plicable this concept is as a tool for restoration of
tion and dilution of pollutants, transportation, recre- eutrophied shallow lakes. The argument is mainly de-
ation and aesthetic enjoyment (Padisák & Reynolds, veloped from results originating from two case study
2003). These services are impaired by exploitation, sites, Alte Donau (Old Danube) and Neusiedler See.
most commonly by nutrient enhancement leading to Generalisation is attempted using published data from
eutrophication. Restoration, rehabilitation and man- a number of other lakes.
agement are therefore needed to ensure sustainable
development.
In contrast to deep lakes, many shallow lakes can The sites
quite abruptly switch between different stable states
representing alternative equilibria, a hypothesis de- Both sites are situated in the Pannonian European Eco-
veloped and established over the past 20 years and region, they are shallow, wind exposed and with sim-
summarised in Scheffer (1998). Lakes can switch from ilar nutrient levels. Morphometric and hydrological
a clear-water macrophyte dominated state to a new features of the two sites are summarised in Table 1.
turbid state dominated by phytoplankton. This bi- Alte Donau is an urban seepage lake of moderate size
stable situation has been intensively investigated (e.g. and has a relatively short retention time. The lake has
Blindow et al., 1993; Scheffer et al., 1993) while been densely covered by submerged macrophytes and
stone wort in the past (Löffler, 1988). As an endorheic
30
Table 1. Morphometric data for Alte Donau and
Neusiedler. See from Dokulil et al. (2000) and Herzig &
Dokulil (2001)

Alte Neusiedler Unit

Donau See

Altitude 157 115.45 m a.s.l.

Area 1.583 315.04 km2
Volume 3.697 367.5 106 m3
Maximum depth 6.8 1.8 m
Mean depth 2.33 1.17 m
Mean theoret. ∼190 days ∼1.5 years
retention time

Figure 1. Long-term development 1987–2001 of macrophyte

dry-weight biomass in tons (hatched bars) and phytoplankton
fresh-weight biomass as mg l−1 (line with filled squares) in Alte
Donau.
lake, the large turbid Neusiedler See has a long re-
tention time, a huge reed-belt and a narrow zone of
submerged vegetation in more sheltered areas (Löffler,
1979). Both lakes have undergone cultural eutrophic- Macrophyte-rich years are separated from algal-
ation followed by a period of oligotrophication as a rich years by the chlorophyll to TP ratio of 1:3
consequence of restoration measures. indicated by the dashed line in Figure 2.
Annual average phytoplankton biomass in Alte
Donau was 1.7 mg l−1 and was composed of a mixed
Case study: Alte Donau algal assemblage in 1987 (Fig. 2). At that time the lake
water was clear with visibility down to the bottom,
Long-term data from Alte Donau show dominance by low chlorophyll concentrations (5.2 µg l−1 ) and dense
macrophytes, low phytoplankton biomass and hence macrophyte cover (Löffler, 1988). Average concentra-
clear water in 1987 (Fig. 1). At that time the entire tions of TP doubled in 1988 and further increased in
water body was covered by submerged macrophytes 1989 but chlorophyll-a levels remained below the crit-
and Chara-beds. Six years later, in 1993, the biomass ical 3 TP to 1 Chl-a line. Although no quantitative data
of under-water vegetation had declined by about 95% on under-water vegetation are available, field notes
while annual average phytoplankton biomass had in- from these years clearly indicate unchanged macro-
creased by more than 1000%. This trend continued phyte stands. The shift from the macrophyte domin-
further in the following year resulting in almost van- ated to the algal dominated state occurred during the
ishing macrophyte beds and a significantly enhanced period 1990–1992. In the first stage TP concentrations
phytoplankton biomass (maximum 81 mg l−1 ). Phyto- declined at constant chlorophyll levels. Thereafter,
plankton biomass was dramatically reduced in 1995 phytoplankton biomass drastically increased, passed
and 1996 as a result of whole lake internal phos- the critical ratio of 3:1 and was accompanied by a shift
phate stripping. In the following years, macrophyte towards cyanobacterial dominance at the expense of
biomass slowly recovered (Pall et al., 1999) while declining macrophyte beds. The situation of vanishing
phytoplankton biomass values remained relatively low under water vegetation and overwhelming plankton
(Fig. 1). biomass finally became evident in the 1992 monitoring
Long-term development is synoptically summar- samples calling for immediate management action.
ised in a multivariate bubble-plot showing the four As a result of the deterioration of this intensively
variables total phosphorus, chlorophyll-a, fresh- used urban lake, a monitoring program covering al-
weight biomass and composition of phytoplankton most all aspects of limnology was initiated early in
(Fig. 2). In this plot, the areas of the circles are pro- 1993 to elucidate the status and trophic situation of
portional to the biomass. Phytoplankton composition Alte Donau, with the prime aim to explore possible
is expressed by the relative contribution of taxonomic remediation measures (Mayer et al., 1997). These in-
groups shown as conventional pie-charts. Trophic vestigations soon revealed cyanobacterial dominance
delineation follows Forsberg & Ryding (1980). (Fig. 2), anoxic sediments at various locations and
 31

Figure 2. Multivariate bubble-plot of total phosphorus (TP) versus phytoplankton chlorophyll-a (Chl-a). The size of the bubbles is equivalent
to the phytoplankton fresh-weight biomass (top right). Bubbles show the relative contribution of algal classes (bottom right). Grey bubbles
indicate years for which no biomass data and algal contributions are available. The 1:1 and the 3:1 TP:Chl-a lines are inserted. Trophic
delineation according to Forsberg & Ryding (1980). Data for 1987 from Löffler (1988), 1989–1990 unpublished data from MA 15, 1991 and
1992 according to BMLF (1994).

vanished under-water vegetation (Dokulil et al., 1995; the amount of chlorophyll is still high and macrophyte
Dokulil & Mayer, 1996). As an immediate short-term biomass has not yet fully recovered (Teubner et al.,
measure, half of the water volume of Alte Donau was 2003).
replaced in December 1993 by water of better quality
from the near-by impoundment New Danube (Dokulil
et al., 1995). The reduction in nutrient concentration Case study: Neusiedler See
brought about by this water exchange significantly im-
proved the water quality in the first five months of In contrast to Alte Donau, submerged macrophytes
1994. This improvement is reflected in a decreased never played a very important role in the open wa-
cyanobacterial contribution and an increased share ter of Neusiedler See because of the high turbidity
of diatoms (pie chart for 1994 in Fig. 2), although and the strong wind exposure. Stands of Potamogeton
the second half of the year again saw an abrupt cy- pectinatus and Myriophyllum spicatum had a distinct
anobacterial bloom outbreak. To improve the nutrient distribution restricted to sheltered bays and the north-
status of the lake and re-establish macrophyte stands western shore which is more protected from the wind
phosphates were flocculated and the sediment oxid- (Schiemer & Prosser, 1976). Macrophyte densities
ised in 1995 and 1996 using the RIPLOX-technique declined in the late 1970s primarily because of lake
(Ripl, 1976). For further details of the action plan level changes (Schiemer, 1979) and the introduction of
consult Donabaum et al. (1999) and Dokulil et al. grass carp Ctenopharyngodon idella (Herzig & Dok-
(2000). As a result, annual average TP concentra- ulil, 2001). Submerged plants recovered again in the
tions declined to about 30 µg l−1 , chlorophyll-a to 1990s. The wax and wane of under water vegetation
less than 15 µg l−1 , biomass dropped to below 3 mg is, however not reflected in the long-term develop-
l−1 , the nitrogen-fixing cyanobacterium Cylindrosper- ment of phytoplankton biomass and related parameters
mopsis raciborskii disappeared (Shafik et al., 2003; (Fig. 3). This is primarily because of the very high
Spröber et al., 2003), the phytoplankton composition inorganic turbidity due to sediment re-suspension. As
changed back to more mixed populations (Teubner et a result, the open water of the lake is continuously light
al., 1999 a, b) and under water vegetation re-appeared limited. Enhanced nutrient input due to eutrophication
(Figs. 1 and 2). Since then, TP and Chl-a further de- is therefore only moderately reflected by an increase in
clined somewhat and reached mesotrophic conditions algal biomass and chlorophyll-a (Fig. 3). All variables
and the critical 3:1 delineation in 2001 (Fig. 2). Al- remain well below the critical limit of 3 TP:1 Chl-a.
though TP concentrations are now lower than in 1987, Macrophytes could therefore occupy the entire lake
32

area but are prevented from doing so because of un- arate lakes with low chlorophyll levels relative to TP
stable sediment surface, high inorganic turbidity and from those with a more direct relationship. Mechan-
large wave action. Given the size, area and depth of the isms for a transition to low chlorophyll levels at high
lake (Table 1), moderate wind speeds of 1 m s−1 are TP include phosphorus partitioning between phyto-
sufficient to re-suspend sediment material (Scheffer, plankton and underwater vegetation, light-limitation
1998). by inorganic turbidity, shadowing by the vegetation,
increased grazing pressure or allelopathic substances
(Hupfer & Dollan, 2003) among others (e.g. Dokulil
Discussion & Janauer, 1993; Herzig & Dokulil, 2001).
Results from a number of lakes are summarised
The theory of alternative stable equilibria explains as TP versus Chl-a in Figure 4. Deep stratifying oli-
shifts between ecological states in ecosystems. These gotrophic to hypertrophic polymictic lakes (shaded
changes usually occur inter-annually as part of a long- sigmoid in Fig. 4) follow the typical pattern expec-
term development. Triggering mechanisms include ted for this relationship. Annual average long-term
increased nutrient loading, changes in water level data from a variety of turbid or macrophyte dominated
and human interference among others (Blindow et al., lakes, marked individually by elliptical envelopes in
1998; Scheffer, 1998). In the case of Alte Donau, Figure 4, somewhat deviate from these expectations
alterations in hydrology and water level have mainly and can be delineated by a line of 3 TP to 1 Chl-a
caused the shift to algal domination (Donabaum, (Fig. 4). The macrophyte periods in both Old Danube
1999) similar to observations elsewhere (Mitchell, (AD) and New Danube (ND), the vegetated and the in-
1989; Blindow et al., 1993). The rehabilitation meas- organic turbid lakes in the Danube Delta (DD) as well
ures applied effectively reduced algal biomass, in- as both the wind exposed turbid lakes Tai Hu (TH) and
creased light penetration and enhanced the grazing Neusiedler See (NS) serve here as examples. Overlap
pressure by zooplankton inducing a back-shift to the to the ‘regular’ pattern or transition to another stable
original macrophyte dominated state (Dokulil et al., state is possible, as seen for instance in Old Danube,
1998, 2000; Teubner et al., 2003). Although po- New Danube, and some of the Danube Delta lakes.
tentially macrophyte inhabited, large shallow wind All these lakes and situations are characterised by low
exposed lakes remain in the algal state mainly because Chl-a:TP ratios as a result of one or the other reason
of frequent sediment re-suspension preventing mac- mentioned above.
rophytes from colonising the lake bottom. Examples Different types of equilibria can occur simultan-
include Neusiedler See in Austria, Tai Hu in China and eously in the same lake. Sheltered bays may be mac-
some of the Danube Delta lakes in Romania (Schiemer rophyte dominated while main parts of the lake are
& Prosser, 1976; Schiemer, 1979; Vadineanu et al., dominated by algal blooms. Examples are, Neusiedler
1992; Oosterberg et al., 2000). See, Tai Hu and some of the Danube Delta lakes.
Lakes which are either turbid or macrophyte Stable equilibria of macrophyte dominance and
dominated are characterised by lower phytoplankton algal pre-dominance may seasonally follow each other
chlorophyll-a relative to total P. Results from Norwe- within a year in some of the Danube Delta lakes. The
gian Lakes, earlier work from New Danube, as well shift is triggered by a multitude of coinciding factors.
as data from reservoirs and shallow lakes in China Filamentous algae over-grow the submerged vegeta-
suggest delineation at around 3 TP:1 Chl-a (Mjelde tion in early summer due to elevated temperatures and
& Faafeng, 1997; Faafeng & Mjelde, 1998; Doku- high nutrient levels. High wind speed then elimin-
lil & Janauer, 2000; Dokulil et al., 2000). A similar ates the macrophytes almost completely from these
relationship can be deduced from a number of invest- lakes giving rise to algal blooms (Vadineanu et al.,
igations in England, the Netherlands and Denmark 1992). Rooted and free-floating plants re-establish in
(e.g. Perrow et al., 1994; Breukers et al., 1997; Schef- the following year because of the prevailing hydro-
fer, 1998, Fig. 5.13; Jeppesen et al., 2002) and can logy bringing clear water in from the River Danube
even be seen in models (e.g. Janse, 1997). The ratio which allows macrophytes to grow and absorb P early
suggested indicates a transition zone between mac- in the year preventing development of algal blooms
rophyte and algal domination (see Fig. 2) within a (Oosterberg et al., 2000).
continuum and shall not be seen as a strict threshold. In some cases two or three levels of stability oc-
It may, however be used as a predictive tool to sep- cur in the same lake at the same time (Van den Berg,
 33

Figure 3. Multivariate bubble-plot to for Neusiedler See. Data for 1990 to 2000 from the Biological Station Illmitz. Legends as in Figure 2.

1998). Tai Hu is good example (Chen, 2003). While

some sheltered bays are full of macrophytes and hence
have clear water, the major part of the lake is tur-
bid having either a mixed algal population or, in
areas of high nutrient content, being dominated by
cyanobacteria.

Conclusions

Alternative stable equilibria can occur within the same

water-body at various locations, at different times of
the year or during trophic development. A critical
level of 3 TP:1 Chl-a has been established separat-
ing macrophyte dominated lakes or lakes with high
background turbidity from algal dominated deep and
shallow lakes. The stable state concept proved to be
helpful for analysing eutrophication events as well as
a conceptual tool for restoration measures in medium
to moderately sized lakes. In large wind-exposed lakes
with high inorganic back-ground turbidity, however
the concept is of no or only limited value.
Figure 4. Conceptual double-log plot of TP versus Chl-a. The
shaded grey sigmoid area covers many deep oligotrophic to shallow
polymictic lakes from Austria and Germany. The elliptical envel- Acknowledgements
opes depict long-term data from the macrophyte dominated period
of Old Danube (AD), the New Danube (ND), the turbid lakes, Tai
Hu (TH) and Neusiedler See (NS), and several vegetated and turbid
We are grateful to the municipality of Vienna (MA
Danube Delta lakes (DD). Data for Tai Hu from Chen (2003), for 45-Wasserbau) for continuous financial support to the
Danube Delta Lakes from Oosterberg et al. (2000) augmented by ‘Restoration project Alte Donau’. We thank the Bio-
own unpublished data. logical Station Illmitz, especially Alois Herzig for
34

continuous data support. Special thanks are extended Donabaum, K., M. Schagerl, K. Teubner & M. T. Dokulil, 1999.
to the technical offices ‘Donabaum & Wolfram’ and Sanierung und Restaurierung der Alten Donau in Wien, Öster-
reich. Deutsche Gesellschaft für Limnologie (DGL) – Tagungs-
‘Systema’ providing data and to the many individu- berichte 1998 (Klagenfurt): 264–268, Tutzing 1999.
als with whom we have worked and co-operated in Faafeng, B. A. & M. Mjelde, 1998.Clear and turbid water in shallow
various projects over the years. Norwegian lakes related to submerged vegetation. In Jeppesen
E., Ma. Søndergaard, Mo. Søndergaard & K. Christoffersen
(eds), The Structuring Role of Submerged Macrophytes in Lakes.
Springer Verlag, New York: 361–368.
Forsberg, C. G. & S. O. Ryding, 1980. Eutrophication.Parameters
References and trophic state indices in 30 Swedish waste water receiving
lakes. Hydrobiologia 89: 189–207.
Blindow, I., G. Andersson, A. Hargeby & S. Johansson, 1993. Long- Herzig, A. & M. T. Dokulil, 2001. Neusiedler See – ein Steppensee
term pattern of alternative stable states in two shallow lakes. in Europe. In Dokulil, M. T., A. Hamm & J.-G. Kohl (eds),
Freshw. Bio. 30: 149–167. Ökologie und Schutz von Seen. Facultas UTB, Wien: 401–415.
Blindow, I., A. Hargeby & G. Andersson, 1998. Alternative stable Hupfer, M. & A. Dollan, 2003. Immobilization of phosphorus
states in shallow lakes: what causes a shift? In Jeppesen E., Ma. by iron-coated roots of submerged macrophytes. Hydrobiologia
Søndergaard, Mo. Søndergaard & K. Christoffersen (eds), The 506–509: 635–640.
Structuring Role of Submerged Macrophytes in Lakes. Springer Janse, J. H., 1997. A model of nutrient dynamics in shallow lakes in
Verlag, New York: 353–360. relation to multiple stable states. Hydrobiologia 342/343: 1–8.
Breukers, C. P. M., E. M. van Dam & S. A. de Jong, 1997. Lake Jeppesen, E., J. P. Jensen & M. Søndergaard, 2002. Response of
Volkerak-Zoom: a lake shifting from the clear to the turbid state. phytoplankton, zooplankton, and fish to re-oligotrophication: an
Hydrobiologia 342/343: 367–376. 11 year study of 23 Danish lakes. Aquat. Ecosys. Health Manag.
BMLF (Bundesministerium für Land- und Forstwirtschaft), 1994. 5: 31–43.
Gütezustand stehender Gewässer in Wien. Wasserwirtschaft- Löffler, H. (ed.), 1979. Neusiedlersee, the Limnology of a Shallow
skataster: 87 pp. Lake in Central Europe / Series: Monographiae Biologicae; v.
Chen, Y., C. Fan, K. Teubner & M. Dokulil, 2003. Changes of nu- 37 Publisher: The Hague; Boston: Dr. W. Junk; Hingham, MA:
trients and phytoplankton chlorophyll-a in a large shallow lake, distribution for the U.S. and Canada, Kluwer Boston, 1979.
Taihu, China: an 8-year investigation. Hydrobiologia 506–509: Löffler, H. (ed.), 1988. Alte Donau. Projektstudie im Auftrag der
273–279. Wasserstraßendirektion. Eigenverlag: 272 pp.
Dokulil, M. T. & G. A. Janauer, 1993. Phosphorus partitioning Mayer, J., M. T. Dokulil, M. Salbrechter, M. Berger, T. Posch, G.
between phytoplankton and macrophytes as a water quality tool Pfister, A. K. T. Kirschner, B. Velimirov, A. Steitz & T. Ul-
in two interconnected impoundments affected by diffuse nutri- bricht, 1997. Seasonal successions and trophic relations between
ent input from the river. In Giussani, G. & C. Callieri (eds), phytoplankton, zooplankton, ciliophora and bacteria in a hyper-
Strategies for Lake Ecosystems Beyond 2000. Proceedings of the trophic shallow lake in Vienna, Austria. Hydrobiologia, 342/343:
5th International Conference of Conservation and management 165–174.
of lakes, Stresa, Italy: 91–94. Mitchell, S. F., 1989. Primary production in a shallow eutrophic
Dokulil, M. T. & G. A. Janauer, 2000. Alternative stable states lake dominated alternately by phytoplankton and by submerged
of macrophytes versus phytoplankton in two interconnected macrophytes. Aquat. Bot. 33: 101–110.
impoundments of the New Danube (Vienna, Austria). Arch. Mjelde, M. & B. A. Faafeng, 1997. Ceratophyllum demersum
Hydrobiol. Suppl. 135 (Large Rivers 12): 75–83. hampers phytoplankton development in some small Norwe-
Dokulil, M. T. & J. Mayer, 1996. Population dynamics and photo- gian lakes over a wide range of phosphorus concentrations and
synthetic rates of a Cylindrospermopsis – Limnothrix association geographical latitude. Freshwat. Biol. 37: 355–365.
in a highly eutrophic urban lake, Alte Donau, Vienna, Austria. Oosterberg, W., M. Staras, L. Bogdan, A. D. Buijse, A. Constantin-
Algol. Stud. 83: 179–195. escu, H. Coops, J. Hanganu, B. W. Ibelings, G. A .M. Menting, I.
Dokulil, M. T., W. Chen & Q. Cai, 2000. Anthropogenic impacts to Nãvodaru & L. Török, 2000. Ecological gradients in the Danube
large lakes in China: the Tai Hu example. Aquat. Ecosys. Health Delta Lakes; present state and man-induced changes. RIZA the
Manag. 3: 81–94. Netherlands, Danube National Institute Romania and Danube
Dokulil, M. T., A. Knoll & J. Mayer, 1995. Impact of diffuse nutri- Delta Biosphere Reserve Authority Romania. RIZA rapport nr.
ent input by polluted ground-water and sediment release to Alte 2000.015.
Donau, the old course of River Danube in Vienna, Austria. Pro- Padisák, J. & C.S. Reynolds, 2003. Shallow lakes: The absolute, the
ceedings of 2nd International IAWQ Conferecne on DIFusePOL relative, the functional and the progmatic. Hydrobiologia 506–
’95: Part II: 462–467. 509: 1–11.
Dokulil, M. T. K. Teubner & K. Donabaum, 2000, Restoration of Pall, K., G. A. Janauer & M. T. Dokulil, 1999. Sanierung der
a shallow, ground-water fed urban lake using a combination of Alten Donau in Wien Entwicklung der Makrophytenbestände.
internal management strategies: a case study. Arch. Hydrobiol. Deutsche Gesellschaft für Limnologie (DGL) Tagungsberichte
Spec. Issues Adv. Limnol. 55: 271–282. 1998 (Klagenfurt): 269–273, Tutzing 1999.
Dokulil, M. T., K. Donabaum, K. Pall & K. Teubner, 1998. Die Perrow, M. R., B. Moss & J. Stansfield, 1994. Trophic interactions
Alte Donau in Wien als Beispiel für die Sanierung eines flachen in a shallow lake following a reduction in nutrient loading: a long
grundwasserabhängigen Stadtgewässers. Int. Seen-Fachtagung term study. Hydrobiologia 275/276: 43–52.
1998: 154–154. Ripl, W., 1976. Biochemical oxidation of polluted lake sediments
Donabaum, K., M. Schagerl & M. T. Dokulil, 1999. Integrated lake with nitrate. A new restoration method. Ambio 5: 112–135.
management to restore macrophyte domination. Hydrobiologia Scheffer, M., 1998. Ecology of shallow lakes. Chapman & Hall,
395/396: 87–97. London: 357 pp.
 35

Scheffer, M., S. H. Hosper, M.-L. Meijer & B. Moss, 1993. Teubner, K., K. Donabaum, W. Kabas, A. Kirschner, G. Pfister, M.
Alternative equilibria in shallow lakes. TREE 8: 275–279. Salbrechter & M. T. Dokulil, 1999b. What are the differential
Schiemer, F., 1979. Submerged macrophytes in the open lake. Dis- consequences on components of a planktonic food web induced
tribution pattern, production and long-term changes. In Löffler, by in-lake restoration of a shallow urban seepage lake? Pro-
H. (ed.), Neusiedlersee. Limnology of a Shallow Lake in Central ceedings of 8th International Conference on Conservation and
Europe. Junk Publisher, The Hague, Boston, London. Mono- Management of Lakes, Lake ’99, S8A-6. Kopenhagen.
graphiae Biologici 37: 235–250. Teubner, K., N. D. Crosbie, K. Donabaum, W. Kabas, K. Kirschner,
Schiemer, F. & M. Prosser, 1976. Distribution and biomass of sub- G. Pfister, M. Salbrechter & M. T. Dokulil, 2003. Enhanced
merged macrophytes in Neusiedlersee. Aquat. Bot. 2: 289–307. phosphorus accumulation efficiency by the pelagic community
Shafik, H.M., L. Vörös, P. Spröber, M. Présing & A.W. Kovács, at reduced phosphorus supply: a lake experiment from bacteria
2003. Some special morphological features of Cylindrospermop- to metazoan zooplankton. Limnol. Oceanogr. (in press).
sis raciborskii (isolated from Lake Balaton, Hungary) in batch Vadineanu, A., S. Christofor & G. Ignat, 1992. Phytoplankton and
and continuous cultures. Hydrobiologia 506–509: 163–167. submerged macrophytes in the aquatic ecosystems of the Danube
Spröber P., H. M. Shafik, M. Présing, A.W. Korács & S. Herodek, Delta during the last decade. Hydrobiologia 243/244: 141–146.
2003. Nitrogen uptake and fixation in the cyanobacterium Cyl- Van den Berg, M. S., H. Coops, M.-L. Meijer, M. Scheffer & J. Si-
indrospermopsis raciborskii under different nitrogen conditions. mons, 1998. Clear water asociated with a dense Chara vegetation
Hydrobiologia 506–509: 169–174. in the shallow and turbid lake Veluwemeer, The Netherlands. In
Teubner, K., K. Donabaum, W. Kabas, A. Kirschner, K. Pall, G. Jeppesen E., Ma. Søndergaard, Mo. Søndergaard & K. Christof-
Pfister, M. Salbrechter & M. T. Dokulil, 1999a. Auswirkung der fersen (eds), The Structuring Role of Submerged Macrophytes in
Sanierung der Alten Donau auf die planktische Nahrungskette lakes. Springer Verlag, New York: 339–352.
und den Makrophytenbestand. Deutsche Gesellschaft für Lim-
nologie (DGL) Tagungsberichte 1998 (Klagenfurt): 274–277,
Tutzing 1999.