The Alte Donau: Successful Restoration and Sustainable Management

Aquatic Ecology Series
Martin T. Dokulil · Karl Donabaum

Katrin Teubner Editors
The Alte Donau:

Successful
Restoration and
Sustainable
Management
An Ecosystem Case Study of a Shallow
Urban Lake
Volume 10
Editor
Jef Huisman
Institute for Biodiversity and Ecosystem Dynamics, University of Amsterdam,
Amsterdam, The Netherlands
More information about this series at http://www.springer.com/series/5637
Martin T. Dokulil • Karl Donabaum
Katrin Teubner
Editors
The Alte Donau: Successful

Restoration and Sustainable
Management
An Ecosystem Case Study of a Shallow
Urban Lake
Editors
Martin T. Dokulil Karl Donabaum
DWS-Hydro-Ökologie GmbH DWS-Hydro-Ökologie GmbH
Wien, Austria Wien, Austria
Katrin Teubner
Dept. of Limnology & Bio-Oceanography
University of Vienna
Wien, Austria

ISBN 978-3-319-93268-2 ISBN 978-3-319-93270-5 (eBook)
https://doi.org/10.1007/978-3-319-93270-5
Library of Congress Control Number: 2018954812
© Springer International Publishing AG, part of Springer Nature 2018

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Preamble
The motivation to write and assemble scientific knowledge on the deterioration and
rehabilitation of an urban lake into this compilation originated from an almost
unique combination of facts. Metropolitan waters are highly diverse, multifunc-
tional ecosystems, usually man-made. Managing these water bodies is challenging
particularly when used for recreation. Such systems often require focused managing
plans and intensive in-lake efforts because several techniques such as complete
draw-down are impossible in such circumstances. Rarely rehabilitation programs
run more than a few years. Here we can report on a 25-year long-term sequence of
measures to return a deteriorated recreational urban lake to good water quality.
During the investigation, we were able to explore almost all levels of the food web
from physico-chemical features via viruses and bacteria to fish and even to water
birds, including the sediment, the littoral zone and the catchment. Several restora-
tion techniques were tested and critically evaluated. The final management plan was
based on bistable theory to reset the lake from turbid and algal dominated to clear,
macrophyte controlled. During the recovery phase, numerous surplus adjustments
had to be implemented to secure sustainable achievement. In the final stage, supple-
mentary monitoring procedures were necessary to maintain good water quality for
recreational activities.
The project was initiated early in 1993 by the municipal authority 45-Water
Management, Vienna (Magistrat 45-Wiener Gewässer), which is responsible for all
waters within the city of Vienna. The monitoring program is still ongoing and will
hopefully persist.
Essential prerequisites for such a huge long-term project were continuous finan-
cial support by the municipality, cooperation with several institutions and a large
number of individual investigators, impossible to name them all. Two key persons
in the early stage of the project were Dipl.-Ing. Gernot Ladinig ( ) who was head of
the water engineering section and Dr. Werner Pekarek ( ) from the hydrology sec-
tion in the municipal authority 45 at that time. Their encouragement, continuous
support and friendship are greatly acknowledged. We also like to express our grati-
tude for the persistent sustenance of the project by the present head of the municipal
authority 45, Dipl.-Ing. Gerald Loew. Furthermore, we recognize that successful
vii
viii Preamble
restoration would not have been possible without the help and advice of Prof. Dr.
Wilhelm Ripl who had developed the RIPLOX-technique which was realized by the
System Institute Aqua Terra (SAT).
We would also like to thank all the people who even participated in the project in
one or the other way throughout the years. We are grateful to everyone who was
directly or indirectly involved in the rehabilitation effort. We further hope that the
compilation of our work in form of this book will help to encourage other groups in
their effort to keep water quality at its best.
Finally we thank the publisher and particularly Judith Terpos for her support,
encouragement and patience during the long-lasting preparation process.
Wien, Austria The Editors

July 2017
Contents
Part I Introduction
1 Concepts for Restoration: Nutrient Loading, Thresholds
and Alternative Stable States�� 3
Martin T. Dokulil and Karl Donabaum
Part II History, Physiography, Chemistry and Climate

2 From a River to an Urban Lake. Historic Development�� 17
Gerhard Nagel
3 Morphology, Water Temperature and Climate�� 27
Georg Kum and Martin T. Dokulil
4 Hydrological Aspects of Alte Donau �� 35
Raimund Taschke
5 Restoration and Lake Management �� 45
Karl Donabaum and Martin T. Dokulil
6 Long-Term Changes of the Physico-chemical Conditions
in Alte Donau�� 55
Karl Donabaum and P. Riedler
7 Sediment�� 69
P. Riedler and Karl Donabaum
Part III Biology, Ecology and Production

8 Wax and Wane of Macrophytes�� 89
Karin Pall
9 Phytoplankton in Alte Donau: Response to Trophic Change
from Hypertrophic to Mesotrophic Over 22 Years �� 107
Katrin Teubner, Wilfried Kabas, and Irene E. Teubner
ix
x Contents
10 Phytoplankton Photosynthesis and Production�� 149

Martin T. Dokulil and Wilfried Kabas
11 Response of Zooplankton to Restoration
and Climate Warming in Alte Donau�� 163
Katrin Teubner, Monika Großschartner, and Irene E. Teubner
12 The Ciliate Assemblage in Alte Donau �� 213
Gerald Pfister
13 Microbial and Viral Loop in Alte Donau: A Case Study�� 227
Branko Velimirov, Ulrike R. Fischer, Alexander K. T. Kirschner,
and Claudia Wieltschnig
14 The Effect of Restoration Measures on the Benthic
Invertebrates of a Danube Backwater (Alte Donau)�� 243
Berthold Janecek, Patrick Leitner, Otto Moog, and Katrin Teubner
15 Fish Assemblages of the ‘Alte Donau’ System:
Communities Under Various Pressures �� 275
Herwig Waidbacher and Silke-Silvia Drexler
16 The Stocks of Waterbirds on the Alte Donau in Vienna �� 313
Rainer Raab and Ulrike Goldschmid
Part IV Recreation, Landscape Planning and Synthesis

17 Hygienic Aspects of a Recreational Water-Body�� 331
Wolfgang Zoufal
18 Restoration of the Littoral Zone�� 337
Karin Pall and Ulrike Goldschmid
19 New Planning Activities in Vienna’s Water-Management�� 355
Brigitte Hozang
20 Eutrophication, Management and Sustainable Development
of Urban Lakes: General Considerations and Specific Solutions
for Alte Donau – A Synthesis�� 373
Martin T. Dokulil, Karl Donabaum, Karin Pall, Georg A. Janauer,
and Katrin Teubner
Subject Index�� 387
Taxonomic Index�� 399

About the Editors
Martin T. Dokulil was born 1943 in Vienna, Austria.

He graduated in Biology from the University of Vienna
in 1970. In the same year, he started his professional
career at the Institute for Limnology of the Austrian
Academy of Sciences as an ecologist in the field of lim-
nology. Research interests in freshwater ecology
focused on the quantitative ecology of freshwater algae,
applied limnology and climate change. He coordinated
and actively worked in several interdisciplinary studies
and projects in Austria and abroad. He became univer-
sity lecturer (docent) at the University of Vienna in
1983 and was awarded the honorary title University
Professor in 1988. His research and teaching activities
were in Asia (Sri Lanka, China, Japan), Latin America
(Chile, Argentina, Uruguay, Costa Rica) and Africa
(Kenya). As guest professor he was teaching at the
University of Buenos Aires, Argentina; the University
of Montevideo, Uruguay; and the University of Girona,
Spain. As a member of several scientific boards, asso-
ciations, boards and journals, he was Austrian National
Representative of the International Association for
Limnology (SIL) from 1984 to 2004 and of the
International Association for Danube Research (IAD)
from 1996 to 2002, among several other advisory func-
tions. He continues teaching as external professor at the
University of Vienna and works as consultant.
xi
xii About the Editors
Karl Donabaum was born in Oberpullendorf, Austria,

in 1963. He received his Ph.D. degree from the
University of Vienna in 1992. In his thesis he investi-
gated changes of the chlorophyll-a content of algal
strains in relation to different environmental factors in
continuous and batch culture experiments. In parallel to
giving lectures in algal taxonomy and courses for algal
physiology at the Institute of Plant Physiology, he
started to work as private consultant and founded a
company in the mid-1990s (today under the name DWS
Hydro-Ökologie, led together with two other managing
directors and approximately 20 employees; www.dws-
hydro-oekologie.at). The focus of his work is placed on
the ecological functioning and integrity of standing
waters. In the past 25 years, he successfully led several
restoration projects in Austria, but also worked as lim-
nological consultant in various foreign countries. The
field of experience ranges from nutrient flocculation,
sediment treatment, hypolimnic aeration and eco-
technological measures to shift food web conditions.
Katrin Teubner born in Leipzig, East Germany, in

1964, is a limnologist focussing on algae in lakes. She
received her Ph.D. degree in Ecology from Humboldt
University of Berlin, Germany, in 1996, and the habili-
tation degree as university lecturer in limnology from
University of Vienna, Austria, in 2004. Her studies are
on lake assessment by algae from eutrophied shallow
urban to pristine deep alpine lakes in Austria, Germany
and China. In view of algal ecology, her main research
interest is on adaptation, acclimation and regulation of
primary producers tracked by their marker pigments,
photosynthesis, climate response, growth response to
ambient nutrient availability and acquisition of short-
Foto ©kunstfotografin.at lived nanomolar phosphorus sources in the field and lab.
Her studies apply to man-made impact on lakes as
eutrophication, soda-mining and recent global warm-
ing. She has been involved in many projects related to
lake restoration, microbial food web and climate
change. She was guest researcher at Freshwater
Laboratory of University of Ulster supported by
Austrian Academy of Sciences and British Royal
Society of Science in 1998 and has been awarded the
About the Editors xiii
Marie-Curie Intra-European research grant at Biological

Sciences at University of Essex, UK, in 2007–2008.
Since 2008, she is a board member and since 2018
General Secretary of International Association for
Danube Research (IAD).
Part I
Introduction
Chapter 1
Concepts for Restoration: Nutrient
Loading, Thresholds and Alternative
Stable States
Martin T. Dokulil and Karl Donabaum
Abstract Shallow groundwater seepage lakes are difficult to manage and to restore
once they are eutrophied. Effective management and rehabilitation of such lakes
need strict and systematic planning based on solid concepts. The model most com-
monly used for eutrophication is the nutrient loading concept which relies on the
limiting nutrient theory. Although not particularly designed for lakes primarily
dependent on groundwater, nutrient loading models can be adapted and used to
investigate the nutrient input and nutrient balance from the aquifer. These models
can also be used to make predictions and to calculate thresholds or limits which
must be reached to improve water quality. The theory of alternative stable states
provides another excellent concept. Clear water, macrophyte dominated stages can
switch to turbid conditions characterized by high algal concentrations. Such for-
ward switches are often associated with anthropogenic pressure or changes in the
hydrological regime. Recreational activities often enhance eutrophication processes
resulting in a system collapse and an almost spontaneous switch to an alternative
state. Backward shift to the original, macrophyte dominated stage can be difficult to
attain. Return times are often prolonged due to hysteresis as a result of resilience.
Concepts are outlined in general and then specified for the ecosystem in question.
Keywords Eutrophication · Cyanobacteria · Blooms · Rehabilitation · Bistable

theory · Shallow lake
1.1 Introduction
Restoration and lake management is a broad and difficult task for any inland water
body. It becomes even more complicated when shallow and/or urban lakes are under
consideration, particularly if these lakes are groundwater seepage lakes with now
surface in- or outflow. The greatest problem is usually posed by human induced
M. T. Dokulil (*) · K. Donabaum

DWS-Hydro-Ökologie GmbH, Wien, Austria
e-mail: martin.dokulil@univie.ac.at; karl.donabaum@dws-hydro-oekologie.at
© Springer International Publishing AG, part of Springer Nature 2018 3

M. T. Dokulil et al. (eds.), The Alte Donau: Successful Restoration
and Sustainable Management, Aquatic Ecology Series 10,
https://doi.org/10.1007/978-3-319-93270-5_1
4 M. T. Dokulil and K. Donabaum
acceleration of nutrient input, commonly termed cultural or anthropogenic eutro-

phication. Other problems include organic matter pollution or siltation.
To effectively manage and rehabilitate a lake, a systematic plan is necessary.
Such a plan must first define the problems within the watershed and then develop a
strategy to correct the lake’s problems. In the case of eutrophication, this is usually
the concentration of the limiting nutrient in the runoff or, in other words, the loading
to the lake. Most commonly the limiting nutrient is phosphorus, in some cases nitro-
gen or a combination of both.
Once the problems have been identified, specific strategic plans must be devel-
oped based on solid scientific principles. For a shallow, urban, seepage lake such as
Alte Donau, two concepts seem appropriate: the nutrient loading concept
(Vollenweider and Dillon 1974; Rast and Thornton 2005; Brett and Benjamin 2008)
and the concept of alternative stable states (Scheffer 1990; Blindow et al. 1997).
These two models will be outlined here with respect to their relevance for the resto-
ration and management of the system in question.
1.2 Nutrient Loading Reconsidered
Eutrophication modelling approaches are applicable only to waterbodies in which

aquatic plant growth is dominated by phytoplankton. This technique cannot be used
to reliably assess planktonic algal chlorophyll response to nutrient loading when
there is substantial aquatic macrophyte or attached algae growth present. A further
general constraint on the application of most eutrophication models is that the
average hydraulic residence time of the waterbody must be 2 weeks or more.
Such a residence time allows algal growth in response to nutrient input.
Measuring groundwater inputs is more difficult and often more expensive than
determining surface water inflows. Defining water and nutrient budget for
groundwater contributing to a lake is not as precise as for surface inflows. The
groundwater basin around a lake is laborious to elaborate. Typically estimations
are based on measurements from a network of wells. Groundwater flow into or
out of a lake is often non-uniformly distributed and may vary considerably from
season to season and year to year depending on water level. Quantification of
subsurface inputs requires experienced professionals and cooperation between
disciplines, particularly when nutrient loading has to be evaluated. Linkages
between groundwater nutrient inputs and cyanobacterial blooms have been
demonstrated by e.g. Jones and Bachmann (1976), Dillon and Rigler (1975) and
summarized in Dokulil and Teubner (2003).
Although the nutrient loading concept (Vollenweider 1968, 1975, 1976) was
never designed for groundwater lakes, fundamentals and formulations can be
adopted to describe and evaluate loading from subsurface aquifers (e.g. Löffler
1988; Riedler et al. 2007). Observed or estimated areal phosphorus loading data can
thus be contrasted to permissible and excessive loading thresholds (Fig. 1.1).
1 Concepts for Restoration: Nutrient Loading, Thresholds and Alternative Stable States 5
10000
Eutrophic Excessive
Phosphorus loading [mg TP m-2y-1]
1000 Permissible
1994
1987
100
10
5
1971-73
max Loading
min Loading
AD Loading Oligotrophic
10
0.1 1 10 100
-1
Hydraulic load qs = z/tw [m y ]
Fig. 1.1 Areal annual total phosphorus loading as dependent on the hydraulic load. Thresholds for
permissible and excessive load are inserted assuming a critical spring overturn phosphorus concen-
tration of 10 μg L−1 (Vollenweider 1975). (Data for Alte Donau (AD) originate from Nachtnebel
and Fürst (1998), Löffler (1988) and Dokulil et al. (1994). The triangle symbols represent maxi-
mum and minimum loading calculated from Table 1 in Jeppesen et al. (2005) for 22 shallow lakes.
The downward arrows indicate the approximate loading levels which must be reached to achieve
chlorophyll-a concentration of 10 or 5 μg L−1 calculated according to Dillon and Rigler (1975))
Since no direct TP measurements are available for 1971–1973, approximate TP

loadings were estimated from data in Nachtnebel and Fürst (1998) assuming similar
in-lake TP concentrations as observed by Löffler in 1987 (Löffler 1988). Loading
rates have increased from about 50–60 mg TP m−2year−1 in the 1970s to 115 mg TP
m−2year−1 in 1987. The increase was attributed to significant alterations in the flow
direction, quantity and dynamic of the groundwater due to newly built water man-
agement structures such as the flood water bypass Neue Donau and the hydropower
dam Freudenau on the Danube, but also to freeway and sewer constructions along
the left shore of the Danube (Nachnebel and Fürst 1998). As a result of the shifts in
the water balance, the hydraulic load of the lake increased from 2.9 to 3.9 m year−1
(Fig. 1.1) as the water residence time shortened from 266 to 195 days (Nachtnebel
and Fürst 1998, Table 7–3). Based on a single observation, Löffler (1988, p. 17)
estimated residence time as 3 month. Recalculation from his data however, reveals
218 days much better fitting to the values mentioned above.
Permissible loading was surpassed in 1993 reaching 160 mg TP m−2year−1 1994
(Fig. 1.1) accompanied by a shift from clear water, macrophyte dominated to turbid
and algal dominated.
Formulations by Dillon and Rigler (1975) allow the calculation of critical phos-
phorus levels which must be reached to achieve desired concentrations of chloro-
phyll-a. To obtain a target value of 10 μg L−1 chlorophyll-a, the annual average TP
concentration must be at or below 30 μg L−1 equivalent to an areal loading of
70 mg m−2 TP. To achieve 5 μg chl-a L−1, the corresponding values are 18.9 μg L−1
and 45 mg m−2 TP respectively. The reduction necessary is also graphically illus-
trated in Fig. 1.1. Following considerations and formulations by Dillon and Rigler
(1975), the theoretical response time to changes in P-loading calculates to about
0.60 years for Alte Donau.
Similar to chlorophyll-a, phytoplankton biovolume (PB) can be estimated from
in-lake TP concentration (Håkanson 1999). From this relation, Håkanson suggested
5 and 10 mm3 L−1 as critical and alarm values respectively for phytoplankton bio-
volume; these figures are roughly equivalent to 25 and 50 μg L−1 chlorophyll-a
assuming a chl-a content in PB of 0.5%. Since these threshold values are created
from a logarithmic relation, uncertainties are rather wide. For the annual mean in-
lake TP concentration of 35 μg L−1 in 1987 (Löffler 1988) for instance, phytoplank-
ton biovolume should be 2.24 mm3 L−1 but could range from 0.44 to 11.22 mm3 L−1.
These figures convert to an approximate mean value of 10.1 μg L−1 chlorophyll
(range 2–50 μg L−1). The true average concentrations of 1.72 mm3 L−1 biovolume
and 5 μg L−1 chlorophyll-a represented 75 and 50% of the expected values, possibly
a reflection of the submersed macrophytes present. Judging from the limited infor-
mation available, phytoplankton biovolume gradually increased in the following
years to finally exceed the alarm value in 1993 and 1994. Chlorophyll-a concentra-
tion remained below the equivalent value because the phytoplankton assemblage
consisted largely of cyanobacteria having low chlorophyll content (0.24%).
A critical point for lake restoration is the amount of phosphorus retained in a
lake, also often used in mass balance models such as the Vollenweider model.
Applying different approaches and formulations to estimate the retention coef-
ficient (Dillon and Rigler 1975; Nürnberg 1998; Brett and Benjamin 2008) to
Alte Donau, unequivocally resulted in coefficients of 0.70–0.78. Phosphorus
retention of 70–78% in the lake nicely agrees with mass balance calculations by
Riedler et al. (2007).
1.3 Stable Equilibria Revisited
The existence of alternative stable states in natural ecosystems was hypothesized

half a century ago. According to Lewontin (1969) a system possesses alternative
stable states if it can return to one or more equilibria after a disturbance. Changes
only occur when the severity of the disturbance exceeds the tolerance level, e.g. the
‘normal’ intensities a species assemblage may typically experience (Sousa 1984).
Regime shifts can be caused by natural or anthropogenic perturbations and may be
gradual or catastrophic (Scheffer et al. 2001a, b; Scheffer and Carpenter 2003; Van
Nes and Scheffer 2005).
Fig. 1.2 Pictorial illustration of the clear water, vegetation dominated and the turbid, algal domi-
nated stable state
The existence of alternative stable states has been supported by ecological mod-
els (Holling 1973; Sutherland 1974), but their presence in the ‘real’ world has been
much debated (Conell and Sousa 1983; Peterson 1984; Jasinski and Asselin 2004).
Experimental evidence shows the alternating presence of persistent communities,
which are often difficult to observe directly (Petraites and Latham 1999; Bertness
et al. 2002; Scheffer et al. 2003).
The presence of alternative equilibria has been demonstrated for a number of
aquatic and terrestrial ecosystems (Knowlton 1992; Hughes 1994; Baker and
Walford 1995; Steele 1998; Hare and Mantua 2000; Van de Koppel et al. 2001;
Bertness et al. 2002; Sedia and Ehrenfeld 2003; Rietkerk et al. 2004; Jasinski and
Payette 2005; Holmgren et al. 2006), recently summarised by Schröder et al. (2005).
In shallow freshwaters, the theory of bi-stability has been widely used, accepted
and modified (Scheffer 1990, 1991, 1998; Scheffer et al. 1993). Moreover, the con-
cept has been adopted and expanded for use in lake management and restoration
(Moss et al. 1996, 1997). Dense stands of submerged plants and clear water or algal
blooms associated with high turbidity (Fig. 1.2) are usually the two different situa-
tions that can exist in lakes (Uhlmann 1980). In deep lakes environmental condi-
tions usually change gradually from one state to the other. In shallow lakes however,
more abrupt switches are common allowing alternative states at almost identical
conditions (Fig. 1.3). The equilibrium trajectory is folded producing an unstable,
never realized region (dashed line in Fig. 1.3), which marks the ‘turning point’ of
the system called ‘bifurcation’.
Two mechanisms are involved: Environmental disturbances partly absorbed by
the resilience of the system gradually modify internal structures (Gunderson et al.
2002), but when the ‘break-point’ region is reached, a small further alteration may
result in a ‘catastrophic’ shift to another stability domain. Strong perturbations can
flip an ecosystem across the unstable region when disturbances are large enough to
exceed resilience.
without vegetation
Forward
Nutrient switch
reduction
Critical
Backward factor
shift
Eutrophication
with vegetation
Fig. 1.3 Conceptual diagram of forward and backward switches between a vegetated and a non-
vegetated stage during nutrient increase and recovery
One reason for a forward switch from clear water and macrophyte domination to
the turbid, algal dominated state can be an increase in nutrient concentration leading
to enhanced growth of planktonic and epiphytic algae which in turn affects the
under-water vegetation, especially when combined with an increase of algivorous
and benthivorous fish species (Scheffer 1998). Massive stocking with carp and
grass-carp can result in abrupt changes in equilibrium conditions leading to a for-
ward switch inducing a turbid algal dominated situation. Grazing by waterfowl dur-
ing summer, however, is considered to be of minor importance while migrating
birds can damage over-wintering macrophytes severely. Grazing by waterfowl may
also strongly affect re-colonization by water-plants. Other factors causing destruc-
tion and loss of macrophytes are disturbances by heavy storms, extreme frost at low
water level or a permanent increase in water depth.
The macrophyte dominated stage is stabilized by the uptake and incorporation of
nutrients by macrophytes and their associated periphytic algae. Both strongly reduce
nutrient availability for primary producers in the pelagic. Such systems often have
greater potentials for top-down control of phytoplankton (Jeppesen et al. 1997).
As has already been mentioned earlier, the shallow urban lake Alte Donau in
Vienna has switched from one to the other stable state and thus serves as an excel-
lent example in a natural environment (Donabaum et al. 1999). Reversal of such a
regime shift can be triggered by natural factors (Blindow 1992; Blindow et al. 1993;
Van Nes et al. 2007) but anthropogenic forcing by restoration measures must be
Fig. 1.4 Total phosphorus 100

versus phytoplankton
chlorophyll-a for Algal
domin. Turbidity
floodplain lakes on the domin.
Chlorophyll-a [Chl-a, mg m-3]

River Danube east of
Vienna, Austria. Waters
dominated by 10
phytoplankton, submersed
macrophytes or inorganic
turbidity are separated by
envelopes. Symbols in the
legend are ordered from
top to bottom that lakes 1 Macrophyte
Side arm 1
Rosskopf 1u
become increasingly domin.
Rosskopf 1o
isolated. The line 5TP:1 Traverse 2u
1
5:
Traverse 2o
Chl-a characterizes Dam
macrophyte domination Old side arm
and separates lakes Oligo Meso Eu Hypertrophic

dominated by 0.1
phytoplankton from turbid 1 10 100 1000
ones. (Modified from
-3
Dokulil et al. 2006) Total phosphorus [TP, mg m ]
considered as an alternative in situations where natural internal switching is unlikely.

In such cases, methods to re-establish macrophyte domination are essential.
From a study on Danish lakes, Jeppesen et al. (1990) concluded that shallow
lakes >3 ha were generally macrophyte-dominated at TP levels <0.05 g P m−3 and
turbid at levels > ca. 0.125 g P m−3, with alternative states possible at intermediate
levels. In accordance, data from several floodplain lakes in Austria scatter around
5TP:1Chl-a, a line characteristic for lakes dominated by submersed macrophytes
(Fig. 1.4). Turbid lakes tend to have higher ratios up to 10:1 while algal dominated
systems are generally below 5:1. This implies that turbid and macrophyte domi-
nated systems support less phytoplankton biomass than the TP concentration would
advocate. Moreover, such ecosystems can have alternative stable states and may
switch if environmental parameters change.
The scheme described in Fig. 1.4, is expanded and generalized for deep and shal-
low lakes in Fig. 1.5. Deep and shallow lakes from Austria and Germany dominated
by phytoplankton are summarized by the grey sigmoid indicating deviation from
linearity at the lower end, because of variable phytoplankton biomass at very low
nutrient supply and because of nutrient saturation and light limitation at the upper
end. All other lakes, here shown as envelopes only, are either turbid and/or macro-
phyte dominated systems. Some of them are spatially diverse (e.g. Tai Hu), others
can switch from algal to macrophyte domination and back (e.g. the Danube Delta
lakes) as alternative equilibrium theory predicts.
Natural ecosystems are never in equilibrium or steady state, because unpredict-
able changes in weather and hydrology, seasonal changes of light and temperature,
and internal mechanisms in populations continuously affect the structure and
100
5:1
1:1
Chlorophyll-a [mg m-3]
TH
NS
10
AD
ND
DF DD
1
1 10 100
Total phosphorus [mg m-3]
Fig. 1.5 Conceptual plot of total phosphorus versus Phytoplankton chlorophyll-a for different
lake types. The shaded area covers deep oligotrophic to hypertrophic lakes from Austria and
Germany. Elliptic envelopes surround data from Alte Donau (AD), Neue Donau (ND), Danube
floodplain lakes (DF) in Austria, the Danube Delta lakes (DD) in Romania, the turbid Neusiedlersee
(NS) in Austria/Hungary, and the hypertrophic Tai Hu in China (TH). Systems dominated by inor-
ganic turbidity and/or submersed macrophytes are separated from deep lakes by the line
5TP:1Chl-a. (Data were extracted from Dokulil et al. (2000a, b, 2006), Jagsch et al. (2002),
Teubner (1996, 2000) and Teubner and Dokulil (2000))
function of freshwater systems. The theory of alternative stable equilibria however

offer strategies to restore shallow, algal dominated lakes to the stage characterised
by macrophytes.
1.4 Conclusions
Regime shifts in shallow lakes as a result of ecohydrological changes can be suc-

cessfully rehabilitated (Hosper 1998; Dokulil and Janauer 2000) if mass balance
models (Vollenweider 1968, 1975, 1976) and alternative equilibrium theory
(Scheffer 1998; Scheffer and Carpenter 2003) are combined. When systems shall be
forced back to an alternative stage, resilience is likely to produce hysteresis and
consequently return times become longer than expected or calculated (Janse et al.
2008; Donabaum et al. 2004; Dokulil and Teubner 2003; Dokulil et al. 2011).
In general, detection and removal of conditions leading to unidirectional switches
require several phases during rehabilitation:
• Reduction of external nutrient loading (catchment restoration)

• Reduction of internal nutrient cycles and fluxes (internal restoration)
• Restructuring of the ecosystem by suitable techniques (e.g. biomanipulation)
• Recovery of water vegetation and restoration of any existing wet-land
• Implementation of unconventional management practices to reach the desired
goal of ecosystem quality might be necessary in certain cases
• Stabilization and management of the rehabilitated ecosystem
In Particular, recovery and backward shift from algal domination to the return of
macrophyte cover in Alte Donau required appropriate planning and suitable
procedures:
• A prime prerequisite is the detailed knowledge of the quantity, flow direction,
nutrient status and potential pollution of the aquifer in this shallow urban lake
primarily influenced by groundwater
• To improve water quality, an indispensable first step is to lower phytoplankton
biovolume and chlorophyll-a concentrations
• To reach a target chlorophyll-a concentration of 10 μg L−1or less, in-lake total
phosphorus (TP) concentration must be reduced to at least 30 μg L−1
• To realize such an in-lake concentration, the areal loading must be lowered to
about 70 mg m−2 TP and internal loading from the sediment must be stopped
• As an outcome of several techniques tested in the field, internal phosphorus floc-
culation with nitrate addition proved to be the optimal technique to attain these
targets
• Application of several options and methods to restructure the ecosystem were
necessary to re-establish the submersed vascular plants
• Continuous environmental monitoring, implementation of a management plan
and permanent adaptation to the changing milieu and anthropogenic require-
ments was eventually essential
Acknowledgments This study was supported by several grants from the Municipal Department –
45 (Water Management – Vienna). We would like to thank all of the numerous collaborators and
the Municipal Department for permission of publication.
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Part II
History, Physiography, Chemistry and
Climate
Chapter 2
From a River to an Urban Lake. Historic
Development
Gerhard Nagel
Abstract The geological setting of the River Danube near Vienna is outlined as it
was before the river was regulated in the nineteenth century. The reasons for the
regulation and how it was done is then described followed by a detailed description
of hydrological management constructions to foster flood protection, to create
hydroelectric power and to protect surface waters from underground pollution origi-
nating from a former, abandoned dump site. The history of urban development and
leisure activities at Alte Donau, the former main channel of the river and now an
oxbow groundwater lake within the city limits of Vienna, is then detailed. Alte
Donau became a famous recreational area already before World War 2 and further
developed after until today. At the end an outlook is presented explaining some of
the future developments and management strategies necessary.
Keywords Urban lake · Historic development · Geology · Flood protection
2.1 T
he Geology of the ‘Wiener Becken’ and the River
Danube Before the Regulation
Vienna is situated at the western rim of a tectonic basin called Wiener Becken which
is filled with sediments of different grain sizes. The river Donau enters the basin
passing through the Flysch mountains which form the eastern branch of the Alps.
During the Quaternary Period the Donau and partly its tributaries have accumulated
gravel horizons which form today’s aquifer. The silty and sandy sediments below
the gravel are a relatively impervious layer.
The development of Vienna has always been strongly related to the River
Danube. In the Wiener Becken the river furcated into a number of main and second-
ary arms. The landscape was dominated by floodplains and riparian forests up to
some kilometers in width. Every bigger flood changed the system considerably,
G. Nagel (*)
City of Vienna, Municipal Department 45 – Water Management, Vienna, Austria
e-mail: gerhard.nagel@wien.gv.at

https://doi.org/10.1007/978-3-319-93270-5_2
18 G. Nagel
created new river arms and filled up others by sedimentation. Therefore no settle-
ments existed nearby the river and the bridges were damaged or destroyed regu-
larly. The historic city of Vienna was situated at the most southern arm of the
Danube which later was named Donaukanal and is now a side arm within the city
(Mohilla and Michlmayr 1996).
The floods of the Donau were a continuous threat to the city and its suburbs. The
biggest flood took place in 1501 with a flow of approximately 14,000 m3 s−1. Today’s
flood protection system is based upon this design flood. The average flow of the
Danube is 1900 m3 s−1 (including the side arm Donaukanal).
The inhabitants of Vienna have made attempts to tame the river for many centu-
ries. The oldest still existing plans date from the seventeenth century. The main
goals of all works were to provide flood protection, create and preserve a navigable
waterway close to the city and construct bridges which could withstand the strain of
the floods. Various constructions were completed, but they all proved not to be resis-
tant. The dikes built between 1780 and 1785, for example, already broke in 1787
during a disastrous flood.
In 1830, a devastating flood revived the discussions to regulate the river bed of
the Danube. Two fundamentally different solutions were debated: one idea was to
stabilize the course of the Donau and provide flood protection by means of dikes;
the other plan was to dig a new bed and unify and straighten the river. Experts were
having heated discussions about this issue for many years.
Most of the proposals included a new straightened riverbed for all river arms
except the Donaukanal. There existed various versions for the location of this cut-
off. The discussion included arguments of hydraulic engineering, but also of urban
planning, economy, military and traffic requirements. Harbours, transshipment
facilities, railway stations, bridges and industrial sites should be constructed together
with the new riverbed.
2.2 The Major Regulation of the Donau
A special commission for the regulation was initiated which organised hearings
with foreign experts among other subjects. Finally, the commission recommended a
cutoff and defined its alignment. The successful construction of the Suez Channel
(1859–1869) strengthened the confidence in the feasibility of such a gigantic proj-
ect. The construction works for the new bed of the Donau took place from 1870 to
1875 and brought a lasting change to the cityscape of Vienna (Fig. 2.1).
The new riverbed had a length of approximately 13 km. It measured 280 m in
width and had an adjoined floodplain of 450 m with a dike on the left bank. The
excavation material was used to fill up one of the big former river arms. Five new
bridges for roads and railways were built together with the regulation, three of them
on dry land. During the World’s Fair in 1873 the construction site was a big attrac-
tion. It could be viewed from a special lookout tower.
2 From a River to an Urban Lake. Historic Development 19
Fig. 2.1 Map of Vienna before the regulation with the alignment of the cutoff (1868). The city
centre at the bottom is situated in the south-west of the Donaukanal. The main arm of the Donau
in the north-east of the cut-through became today’s Alte Donau
Filling up most of the former river arms created new areas for urban develop-
ment. Thus, only few parts of the old system of river arms have survived. One part,
which is still connected to the Donau, has become a national park in the meantime.
The biggest arm that still exists is today’s Alte Donau (the name has the meaning of
‘Old Danube’). It was part of the principle arm of the Danube before the regulation,
what still can be seen in its morphology (Chap. 3).
In 1875 the new cutoff was opened for navigation with a big celebration. But
only a few years later in 1899 a catastrophic flood gave rise to doubts about the
discharge capacity of the new riverbed. Thus, a new discussion of flood protection
requirements in Vienna started a few years later upon completion of this huge
project.
Different versions for an improvement were proposed and some of them even
planned. But the two World Wars and the poor economic situation between the wars
impeded the realisation of those projects. There were plans to dig new bypasses or
to broaden and deepen the cutoff. The construction of hydropower plants was part
of some of the projects. Many of these plans included the Alte Donau: it could have
become part of a new bypass channel, a big harbour or filled up with excavation
material.
20 G. Nagel
2.3 The Neue Donau (‘New Danube’)
A flood in 1954 caused inundations of the city particularly on the right river bank.
It initiated new efforts to improve the flood protection of Vienna. Various proposals
were made again and discussed among experts. The City Administration decided on
a project creating a new flood bypass channel (Neue Donau) along the left embank-
ment and the creation of a flood-free island (Donauinsel) between the new and the
existing river bed, using the excavation material. This project was favoured also
because of its advantages for urban development.
The impoundment Neue Donau was constructed between 1972 and 1988. Its
length is approximately 21 km, its profile width is 210 m. It is divided into two
basins with different water levels. Three weirs were built which regulate the water
levels and control the discharge during floods. Water discharge only takes place dur-
ing high-water periods, most of the year Neue Donau has horizontal, calm water
surfaces similar to a lake. This new flood protection system has shown excellent
performance during big floods, for example in 2002. Besides, Neue Donau and the
Donauinsel have become very important recreation areas in Vienna.
Neue Donau is very important for Alte Donau, because it has changed the hydro-
logical situation substantially. Alte Donau is fed only by groundwater and precipita-
tion, there is no surface inflow. Before the construction of Neue Donau the
groundwater situation between the River Danube and Alte Donau depended on the
often rapidly changing water levels of the Danube. Nowadays the water levels in the
two impoundments of Neue Donau form the constraints for the groundwater fluxes
to Alte Donau.
2.4 The Hydroelectric Power Plant Freudenau
The construction of the hydroelectric power plant Freudenau between 1992 and
1997 caused another change of the hydrological situation. In March 1996 partial
damming of the Danube started, full damming was achieved in November 1997 ris-
ing the mean water level by 8.3 m at the barrage. Damming of the Danube has
increased the groundwater flow through the gravel of the Donauinsel to Neue
Donau. Therefore excessive seepage water is diverted via the weirs of the Neue
Donau and the water levels in both impoundments of the Neue Donau are higher
than before. These water levels are regulated by the weirs and follow an annual
hydrograph curve with amplitude of 0.8 m (upper impoundment). The water level of
the Alte Donau follows this hydrograph subdued and with a time-shift. The average
water level of Alte Donau hence has been raised, but its amplitude has been reduced
substantially (Chap. 4).
The water surplus from Neue Donau allows water management measures for
Alte Donau and other former river arms of the Danube. Since 2006, for example,
Alte Donau has been flushed every year with water of Neue Donau to increase the
calcium concentration and stabilise the pH-value (Chap. 5). Another former part of
Alte Donau, the Wasserpark which is separated by a dam, is flushed regularly with
water of Neue Donau.
2.5 The Contaminated Site ‘Donaupark-Bruckhaufen’
After the major regulation of the Danube the area between Alte Donau and the new
river bed was abandoned land. It included small oxbow lakes, pools and cavities
from digging gravel, which were filled up with different discarded waste materials
during the following decades. Waste from households, industries and construction
waste of unknown composition were deposited in an area of 1.5 km2. The waste
material reached heights of up to 9 m. The overall volume of deposited material was
coarsely estimated to 5 million m3.
In the northern part of the area, dumping ended at about 1930. Afterwards houses
and cottages were built on this area. In 1964 the municipal landfill in the southern
part was closed and the area became a public park.
Problems with subsidence and gas ebullition occurred soon afterwards. In 1986
a comprehensive risk assessment was carried out. During periods of high water level
of the Danube, large areas of the dump site got in contact with the groundwater lead-
ing to groundwater pollution. Due to the groundwater flow towards Alte Donau
potential contamination had to be stopped. A hydraulic barrage was constructed and
parts of the deposited material were removed.
The barrage system consists of 5 wells which extract water from the ground and
are controlled automatically by the groundwater level measured. The target of the
barrage system is to keep the water level within the contaminated site always below
the water level of Alte Donau. The extracted water is cleaned in a pond and dis-
charged to the Danube.
The barrage system was built in 1992/1993 and went into operation in 1994. It is
a crucial hydraulic system to reduce the nutrient input to Alte Donau (Urban and
Ladinig 1996).
2.6 T
he Alte Donau as an Urban Development and Leisure
Area
After the regulation in 1875 many elements of riverine landscape still existed on
Alte Donau and its surroundings such as islands, sandbanks and relicts of old river
arms, some of them connected to Alte Donau. This typical mosaic of land and water
gradually disappeared. Over time the embankments of the Alte Donau were straight-
ened and reinforced, bays and ponds were filled up, lawns for recreation were
created.
22 G. Nagel
The surroundings of Alte Donau were only sparsely populated in the beginning
and served different purposes such as agriculture, industrial sites, landfill, military
use, new – mostly scattered – settlements, sports and leisure activities. Some of
these activities have vanished; others are still prevalent in the area (Ladinig 2000).
2.6.1 Settlements
During and after the First World War people began to settle into the area surround-
ing Alte Donau. Due to housing shortage and bad food supply in those days new
settlements were – partly illegally – established consisting of cottages and small
gardens to grow vegetables. Those settlement areas were situated on the border of
the town without supply and disposal facilities of the municipality. Most houses had
their own wells for water supply, there was no electricity, waste water was collected
in septic tanks or it seeped away. Some of these settlements still exist hence expan-
sion of the sewer system was part of the sanitation program in the surrounding area
of Alte Donau.
2.6.2 First Leisure Time Activities
Leisure activities in the Alte Donau area started soon after the regulation in 1875. In
those days, such activities were restricted to wealthy or rich social classes, because
most people did not have enough money or time for leisure. Furthermore, Alte
Donau was far from the city centre and difficult to reach.
Nevertheless the first boat hire opened in 1880 and in the same year the first row-
ing regatta took place. In 1890 already six boat rentals existed. At the end of the
nineteenth century the first rowing clubs settled at the Alte Donau, later on – at that
time very exclusive – sailing clubs came along. Various gastronomic facilities
opened and Alte Donau soon became a popular destination particularly after an
electric tramway line was completed in 1898.
A big island remained in the southern part of Alte Donau, called ‘Gänsehäufel’.
The name refers to the goose kept there at that time. The island was sandy and partly
covered by trees. Around 1900 the island was popular among bathers who felt
friends of nature. In 1907, the municipality opened the public bath ‘Gänsehäufel’,
the first one of several others along both embankments of Alte Donau. According to
the moral conception of those days it consisted of separated areas for men, women
and families with high fences between them. The Public bath Gänsehäufel was later
enlarged and connected in 1926 by a bridge with the mainland. Till today the
Gänsehäufel has been the biggest and widely known public bath at Alte Donau.
2.6.3 First and Second World War
After the First World War the Austrian Hungarian Monarchy collapsed and Austria
remained as a small country. Its capital Vienna lost importance and the population
decreased. Today Vienna has approximately 1.8 million inhabitants.
In 1918 working hours were limited to 48 per week and big parts of the popula-
tion got more leisure time at their disposal. Vienna was administrated by the social-
ist party, whose conception of life valued health, physical culture and outdoor
sports. This mentality gave an incentive for the leisure time facilities at Alte Donau.
In the 1920s numerous bathes were established, some of them still exist today.
During the Nazi regime plans for city enlargements were made. They included a
big harbour in Alte Donau but were never realised. From 1943 till the end of war
Vienna was a target of bombing raids that struck also the Alte Donau because of the
industrial sites in its surrounding. The Gänsehäufel bath, for example, was com-
pletely destroyed by bombing.
2.6.4 Development Since the Second World War
After the Second World War economic conditions were very poor and priority was
given to the reconstruction of destroyed buildings and facilities. But hand in hand
with the economic recovery leisure time activities started to boom again and Alte
Donau became an important recreational area. The construction of Neue Donau and
the Donauinsel created a new pulse for urban development near the Danube. Office
buildings for the United Nations (1979), called ‘UNO-City’ and a conference centre
(1987) enhanced the importance of the area between the River Danube and Alte
Donau. Completion of the underground line 1 in 1982 made it easy to reach the
recreation area by public transport. Today Alte Donau can be reached by three
underground lines; a ride from the city centre to Alte Donau takes only 10 min.
Till today the region between Neue Donau und Alte Donau has been a focus of
urban development. Skyscrapers with offices and flats, cinemas and fitness centres
were built that now shape the skyline near Alte Donau. On the banks of Neue
Donau many pubs and restaurants have opened which make this area a very lively
and crowded place especially during summer. For these reasons this region has got
a metropolitan atmosphere during the last years. An aerial view of the area is
shown in Fig. 2.2.
24 G. Nagel
Fig. 2.2 Aerial view of Alte Donau area with River Danube, Donauinsel and Neue Donau. (Data
origin: City of Vienna – data.wien.gv.at)
2.6.5 Leisure Time Activities Today
Leisure activities play an important role in our society. Especially during the sum-
mer months many activities focus on water bodies like the Neue and Alte Donau. A
recent scenic view is shown in Fig. 2.3. Today Alte Donau offers a very good infra-
structure for leisure activities:
• Boat rentals offer boats for sailing and rowing, pedal boats and electric powered
boats. Some sport clubs with a long tradition still have their domiciles at Alte
Donau. There are clubs for rowing or sailing and schools for sailing and surfing.
Alte Donau is situated downwind the two most likely wind directions in Vienna
and therefore is very popular for sailors and surfers.
• Of the 10 bathes situated at the Alte Donau, the three largest are operated by the
municipality. These three bathes have about 800,000 visitors in an average year,
Fig. 2.3 The Alte Donau today
in the very hot summer of 2003 there were more than one million. On a summer-
weekend with fine weather up to 42,000 people per day come to the bathes at
Alte Donau. In addition to the bathes there are various sunbathing areas and
shores with public access which are also very crowded on hot summer days.
Numerous restaurants and inns on the waterfront offer drinks and food, most of
them are open all year.
In some parts of the banks settlements with small houses still exist. The houses
on the waterfront stand very close to each other and have their own access to the
water. They are intensely used during the summer months.
Alte Donau is also a fishing ground, which is administrated by a fishing associa-
tion that was founded already in 1880. The fish caught are restocked by the associa-
tion. Alte Donau is stocked mainly with carps and pikes.
In cold winters the Alte Donau is covered completely by ice. If it is thick enough
many people do ice skating ore play ice hockey there.
2.7 Outlook
This survey shows the importance of Alte Donau as a recreation area close to the
city centre of Vienna. Leisure activities cover the whole water body, there are no
secluded or inaccessible parts. Therefore the municipality of Vienna has made big
26 G. Nagel
efforts to achieve and stabilise a good water quality. Today Alte Donau shows a
‘good status’, which is the aim for all water bodies under the Water Framework
Directive of the European Union.
To maintain this good status, continuous management is necessary such as con-
trolled cutting of macrophytes or the lowering of the water level in spring to enhance
groundwater inflow. Accompanying monitoring programs are crucial to provide
information for the evaluation of all measures. Furthermore they show the develop-
ment of the water quality and identify changes early enough to react in time. The
objective of all efforts and management measures is to find stable solutions that
combine the needs of ecology and leisure time activities.
References
Ladinig G (ed) (2000) Die Alte Donau – Menschen am Wasser. Bohmann Druck und Verlags-
GmbH, Vienna
Mohilla P, Michlmayr F (1996) Donauatlas Wien. Österreichischer Kunst- und Kulturverlag,
Vienna
Urban H, Ladinig G (1996) Altlast Donaupark – Alte Donau, Perspektiven, vol 10/1996. Compress
Verlagsges.m.b.H, Kraków
Chapter 3
Morphology, Water Temperature and
Climate
Georg Kum and Martin T. Dokulil
Abstract General features of the urban lake Alte Donau and its surroundings are
described. As a cut-off arm of the former main channel of the River Danube, the
lake has several unique morphometric features. These characteristics are outlined,
tabulated and discussed, including the depth contours and the hydrology.
Development and characteristics of water temperature, as well as the general cli-
matic situation of the city district and the lake area in particular are discussed.
Keywords Morphometry · Hydrology · Temperature · Warming · Climate ·

Continentality
3.1 Introduction
The main characteristics of the water basin, the water temperature regime, the
response to climate warming and the general climatic situation are outlined in this
chapter. A description of specific morphological features and a depiction of the
depth contours are followed by a discussion of developments and trends in air and
water temperature. Effects of climate warming are briefly mentioned. Finally the
general climatic situation of the area is defined and characteristics of the river valley
and the adjacent area are outlined.
Most of the climate data were extracted from Auer et al. (1989). Monthly mean
surface water temperatures originate from the hydrological yearbooks of Austria.
(http://www.bmlfuw.gv.at/wasser/wasser-oesterreich/wasserkreislauf/hydrogra-
phische_daten/jahrbuecher.html)
G. Kum (*) · M. T. Dokulil

e-mail: georg.kum@dws-hydro-oekologie.at; martin.dokulil@univie.ac.at

https://doi.org/10.1007/978-3-319-93270-5_3
28 G. Kum and M. T. Dokulil
Fig. 3.1 Depth contours for Alte Donau
3.1.1 Morphology
As a cut-off arm of the Danube (Chap. 2), Alte Donau has several unique morpho-
metric features. The remaining backwater situated at an altitude of 157 m a.s.l.
(adriatic) has no natural in- or outflow. The water balance entirely depends therefore
on groundwater seepage and precipitation (Dokulil et al. 2011a, b; Chap. 4). The
water body is divided into two main basins, connected by a narrow passage
(Fig. 3.1). The area of the upper basin is 0.55 km2 and has a volume of 1.34 × 106 m3.
The larger lower basin has a surface area of 0.88 km2 and a volume of 2.20 × 106 m3
3 Morphology, Water Temperature and Climate 29
Table 3.1 Morphometric parameters for Alte Donau and the two basins. The appendix
‘Kaiserwasser’ is not included
Parameter Abbreviations Unit Total Upper basin Lower basin
Altitude a. Adria m 156.7 156.7 156.7
Maximum length l m – 2130.0 2370.0
Maximum width b m – 320.0 390.0
Mean width (A0/b) b’ m – 260.1 370.9
Surface area Ao ha 143.3 55.4 87.9
Bottom area As ha 144.1 55.7 88.4
Volume V 103 m3 3548.6 1345.4 2203.2
Maximum depth Zm m 7.0 7.0 5.2
Mean depth (V/Ao) Z’ m 2.5 2.4 2.5
Relative depth Zr % 0.5 0.8 0.5
Reduced depth (4√Ao) Zred 34.6 27.3 30.6
Max. red. depth (Zm/Zred) Zred max 0.2 0.3 0.2
Shore line L km 17.0 5.5 11.5
Shore line development DL 4.0 2.1 3.5
(Table 3.1 and Donabaum et al. 1999). Monthly ground-water input varies between
28 and 80 × 103 m3 in the upper basin and 250–625 × 103 m3 in the lower basin
(Chap. 4). The average hydrological retention time has been estimated to 175 days
(98–246 days) for the upper basin and 375 days (200–900 days) for the lower basin.
The length of the lake is about 5 km bending from NNW to S with an average width
of 0.35 km, a reminiscence of the former fluvial character also indicted by the
shoreline development >2 (Table 3.1). The maximum depth is 7.0 m located at a
single depression in the northern basin (Fig. 3.1). The mean depth of 2.5 m and the
reduced depth of less than 0.5 identify Alte Donau as a shallow polymictic lake
which sometimes stratifies, but never for long periods (Löffler 1988). Water depth is
greater near the eastern shore, the former erosion bank of the river.
3.1.2 Temperature
Monthly mean surface water temperature (SWT) peaks in August at 25.3 °C. Average

monthly maximum temperatures are already reached in June and remain high until
August (Fig. 3.2a). Water temperatures are lowest in winter under the ice cover usu-
ally lasting for about 4 weeks. Long term average SWT is 11.7 °C. Average annual
SWT of Alte Donau depends significantly on mean annual air temperature (AT) and
can be predicted from SWT = 2.99 + 0.83 AT (r2 = 0.61, F = 58.09, p < 0.001). Mean
annual long-term development of SWT since 1973 and AT since 1951 are shown in
Fig. 3.2b, c. A trend shift occurs in both time series around 1985 as indicated by
rescaled adjusted partial sums (Garbrecht and Fernandez 1994) not shown here. No
30 12.0
Alte Donau
A C Wien - Stadlau
1973-2011
25 154 m a.s.l. 1951 - 2011
11.5
160 m a.s.l.
20
Air temperature [°C]

Average SWT [°C]
11.0
15
10.5
LT average
10
10.0
5 max
9.5
Mean
0 min
9.0
AT = - 70.74 + 0.04 Year, n = 26
Jan Feb Mar Apr May Jun Jul Aug Sep Okt Nov Dec 8.5
14 1950 1960 1970 1980 1990 2000 2010
B
Surface water temperature [°C]
13
12
11
10
SWT = -75.64 + 0.04 year, n = 25
r² = 0.25, F = 7.801, p = 0.0103
9
1970 1975 1980 1985 1990 1995 2000 2005 2010 2015
Fig. 3.2 Temperature characteristics of Alte Donau. (a) Monthly minimum, mean and maximum
surface water temperature (SWT) averaged for the period 1973–2011. The dotted line indicates the
long-term annual average SWT. (b) Annual mean SWT 1973–2011. Trend lines and statistics for
the 2nd period are indicated. (c) Annual average air temperature at Wien-Stadlau 1951–2011.
Trend lines and statistics for the 2nd period are indicated
insignificant temperature trend can be observed in the period preceding the shift.
Both SWT and AT increase at a rate of 0.4 °C per decade after 1985 (Fig. 3.2a, b).
Similar rates of increase in SWT of between 0.4 and 0.6 °C per decade are observed
in several Austrian lakes (Dokulil 2013). This increase in water temperature is in
accordance with the significant rise in air temperature (Auer and Böhm 2011).
3.1.3 Climate
The geographic position at about 48° north and 16° east puts Vienna into the warm-
temperate climate. Precipitation can occur at all seasons like in many regions of
Central Europe. The climate in Vienna is characterised by a modest continentality
caused by the absence of a nearby sea which would compensate extreme fluctua-
tions in temperature. The degree of continentality is calculated as the difference of
the average air temperature between the warmest and the coldest month in relation
to the maximum difference worldwide, and expressed as percentage. The resulting
32% put Vienna in the middle between the 10% of islands in south-west Europe and
the 50% near the Ural Mountains, the most continental region of Europe (Auer and
Böhm 2011). The climate in Vienna has a few special local and regional
Fig. 3.3 Climate diagram 30
Average air temperature [°C]

for Vienna-Stadlau (a, b). A Vienna - Stadlau
25 1951 - 1980
(a) Minimum, mean and 160 m a.s.l
maximum monthly mean 10.1°C
20
air temperature for the 559 mm.
Cfb
period 1951–1980. Dotted 15
line indicates the long term LT average
annual average. (b) 10
Monthly average
precipitation for the same 5
max
period as in (a). (c) Mean
0
Average monthly number min
of frost, summer and -5
vegetation days for the B
Average precipitation [mm]
same period as in (a)

60
40
20
0
Jan Feb Mar Apr May Jun Jul Aug Sep Okt Nov Dec
Frostdays Tmin< 0°C
40
Summerdays Tmax>25°C
C Vegetationdays Tmin> 5°C
30
Average days
20
10
0
Jan Feb Mar Apr May Jun Jul Aug Sep Okt Nov Dec
characteristics originating from the landscape, particularly from the location

between the hilly ‘Vienna woods’ and the plains of the ‘Wiener Becken’. The cli-
mate of Vienna can be summarised by a climate diagram (Fig. 3.3a, b). In the course
of the year, highest temperatures are usually reached in July or early August, mini-
mum temperatures occur in February. The absolute maximum temperatures ever
measured are 38.3 °C in July 1957 and −28.8 °C in February 1929. Precipitation is
largest in June and smallest in January. Accumulated precipitation for April to
September (summer half-year) exceeds the winter precipitation by 100 mm.
According to the classification by Köppen (1900) as modified by Geiger (1961), the
N
360
337.5 22.5
315 45
292.5 67.5
-1
km h
W 270 90 E
0 5 10 15 20
247.5 112.5
135
225
202.5 157.5
180
Fig. 3.4 Diagram combining annual wind direction and speed
climate can be characterised by Cfb (Fig. 3.3a), which means moderately warm, no

dry periods and warm summers.
Average 1951–1980 air temperatures reach 20.2 °C ± 10 °C near the Alte Donau
in July (Fig. 3.3a) coinciding with 17.2 summer days defined as days with maxi-
mum temperature greater than 25 °C (Fig. 3.3c). Monthly mean maximum and
minimum temperature exceeds 25 °C in July and − 3.5 °C in January respectively.
Temperatures are generally cooler by 0.1–1.0 °C near the lake than in the city with
largest differences during the cold season. Frost days (minimum temperature below
0 °C) occur in the region from October to April reaching the maximum of almost
24 days in January. On the other hand, each month has at least one vegetation day
with minimum temperature exceeding 5 °C (Fig. 3.3c).
The river valley and the adjacent area to the east have more frost and summer
days but slightly less vegetation days compared to the city. Precipitation varies
between 29 mm in January and 73 mm in June without any dry month (Fig. 3.3b).
Mean total precipitation in the Alte Donau region is 2–15 mm less than in the city,
except for August when rainfall exceeds precipitation in the city on average by
5 mm. Average sum of potential evaporation is 448 mm for April to October and the
30-year period 1957–1986. In the plains east of Vienna evaporation is higher than
closer to the hills in the west.
Fig. 3.5 Distribution of annual average air temperature in Vienna for the period 1951–1980 indi-
cating a corridor of moderate temperatures on either side of the River Danube and associated
waters. Redrawn and modified from Auer and Böhm (2011, p. 92)
Annual average wind speeds and directions are depicted in Fig. 3.4 for the station
Stadlau. Highest average wind speeds between 11.5 and 16.1 km h−1 originate from
westerly directions. Between North and East wind speeds are minimal.
With an annual average air temperature of more than 10 °C Vienna is one of the
warmest regions in Austria. The main reason is the densely covered and populated
area in the city producing a ‘heat island’. This effect is less pronounced at Alte
Donau. The water masses of this ox-bow lake together with the River Danube, the
flood-water protection system ‘Neue Donau’ and other smaller water bodies create
a corridor of more moderate temperatures cutting through the town (Fig. 3.5).
Climate development in Vienna in general indicates that frost days have decreased
to 30 days and summer days increased by about 35 days since 1872. Annual total
precipitation varied between 404 and 988 mm with no trend to more extreme events.
Potential evaporation increased by about 25 mm per decade since 1970 mainly due
to rising global radiation (Kromp-Kolb et al. 2007).
3.2 Summary
Alte Donau, a former branch of the River Danube, is now a narrow shallow poly-
mictic ground-water seepage lake of moderate size (total surface area 1.433 km2).
This urban lake is completely engulfed by the city of Vienna and is situated in a
windy corridor of moderate climate along the River Danube. Annual average long-
term temperature is 10.1 °C for air and 11.7 °C for SWT. Average maximum surface
water temperatures of the urban lake can reach over 25 °C. Both air temperature and
SWT increase at a rate of 0.4 °C per decade since 1985. Precipitation is largest in
June and smallest in January with an annual average of 559 mm.
References
Auer I, Böhm R (2011) Wetter und Klima in Wien. In: Berger R, Ehrendorfer F (eds)
Ökosystem Wien. Die Naturgeschichte einer Stadt. Böhlau Verlag, Wien, pp 88–108.
isbn:978-3-205-77420-4
Auer I, Böhm R, Mohnl H (1989) Klima von Wien. Eine anwendungsorientierte Klimatographie.
Verlag Magistrat der Stadt Wien, Wien, p S. 270
Dokulil MT (2013) Predicting summer surface water temperatures for large Austrian lakes in 2050
under climate change scenarios. Hydrobiologia 731:19–29
Dokulil MT, Donabaum K, Pall K (2011a) Successful restoration of a Shallow Lake: a case study
based on bistable theory. In: Ansari AA, Gill SS, Lanza GR, Rast W (eds) Eutrophication:
causes, consequences and control. Springer, Dordrecht, pp 285–294. https://doi.
org/10.1007/978-90-481-9625-8_1
Dokulil MT, Teubner K, Donabaum K, Seebacher F (2011b) Die Alte Donau. In: Berger R,
Ehrendorfer F (eds) Ökosystem Wien. Die Naturgeschichte einer Stadt. Böhlau Verlag, Wien,
pp 128–132. isbn:978-3-205-77420-4
Donabaum K, Schagerl M, Dokulil MT (1999) Integrated management to restore macrophyte
domination. Hydrobiologia 395/396:87–97
Garbrecht J, Fernandez GP (1994) Visualization of trends and fluctuations in climatic records.
Water Resour Bull 30:297–306
Geiger R (1961) Überarbeitete Neuausgabe von Geiger, R.: Köppen- Geiger/Klima der Erde.
(Wandkarte 1:16 Mill.). Klett-Perthes, Gotha
Köppen W (1900) Versuch einer Klassifikation der Klimate, vorzugsweise nach ihrer Beziehung
zur Pflanzenwelt. Geogr Z 6:593–611
Kromp-Kolb H, Formayer H, Clementschitsch L (2007) Auswirkungen des Klimawandels
auf Wien unter besonderer Berücksichtigung von Klimaszenarien. Studie im Auftrag der
Magistratsdirektion der Stadt Wien – Klimaschutzkoordination, p. 41. http://www.wien.gv.at/
umwelt/klimaschutz/pdf/klimawandel.pdf
Löffler H (ed) (1988) Limnologische Projektstudie – Ökosystem Alte Donau. Bericht im Auftrag
der Wasserstraßendirektion, Wien, p 272
Chapter 4
Hydrological Aspects of Alte Donau
Raimund Taschke
Abstract The hydrological conditions in the Alte Donau and the appendix

Wasserpark depend on the water levels of the Neue Donau and the groundwater
levels in the Hinterland, the climatic water balance (precipitation minus evapora-
tion), and the withdrawals by the barrage of wells situated in the area of the old
dump side between Neue and Alte Donau. These wells protect the Alte Donau from
contaminated groundwater. From the long-term measurement of the groundwater
levels in the area, the flow rates were calculated, modelled and the water balance
determined. Water levels in Alte Donau vary from 156.5 m to 157 m a.s.l. (adriatic)
with an average of 156.7 m a.s.l. and flow rates to and from fluctuate between 100
and 300 L s–1. The quantities of water blocked by the wells and withdrawals from
Alte Donau used as dotation for ox-bow lakes in the Hinterland result in a deficit of
about 45 L s–1 in the water balance of the urban lake Alte Donau. This deficit must
somehow be compensated in the future, possibly by a percolation soil filter cleaning
water coming from the Neue Donau.
Keywords Hydrology · Long-term trend · Numeric model · Nutrient balance ·

Water balance
4.1 Introduction
Urban lakes differ in many aspects from natural or artificial lakes in the landscape
(Birch and McCaskie 1999). Since these lakes are often heavily used for recreation
more people interact with these usually small and shallow water bodies which have
a lower resilience than larger deep lakes. As a consequence, urban lakes are more
sensitive to pollution and eutrophication often enhanced by high shoreline develop-
ment causing additional negative impacts (Naselli-Flores 2008).
R. Taschke (*)
Gruppe Wasser, Vienna, Austria
e-mail: r.taschke@gruppewasser.at

https://doi.org/10.1007/978-3-319-93270-5_4
36 R. Taschke
Fig. 4.1 Network of groundwater profiles from the impoundment Neue Donau to Wasserpark
(WP), Alte Donau and into the Hinterland
Many urban lakes have no in- or outflow. Their water balance solely depends on
groundwater seepage, precipitation and evaporation leading to often unique, site
specific hydrological conditions. In city areas with high human impacts on natural
processes such as recreational activities in urban lakes acquire a special ‘urban’
hydrology. Hydrological sub-processes and data acquisition need to be assessed at
small spatial scales and at high temporal resolution (Niemczynowicz 1999). Data
collection must be site specific depending on the local hydrological boundary
conditions.
The hydrological setting in the surrounding of Alte Donau is rather complex. The
artificial flood-water system Neue Donau with its two impoundments acts as a
boundary but is in turn affected by the nearby River Danube with its hydropower
dam (Fig. 4.1). This specific situation had to be considered when gathering hydro-
logical data.
Improvements in the utilization of the water body, in the ecological state, the
water quality, the nutrient balance, the buffer capacity and the water balance were
the prime aim of this study. As for any lake, the nutrient balance is of crucial
4 Hydrological Aspects of Alte Donau 37
importance for the water quality in Alte Donau basically triggered by the in- and
output of groundwater. The essential parameters are the quantity of groundwater
throughput (water balance of groundwater) and the nutrient concentrations there-in.
It was therefore indispensable to analyse and evaluate the hydrological situation in
the vicinity of Alte Donau.
4.2 Methods
A network of groundwater probes, gauging posts and climate monitoring stations

was installed for the acquisition of the hydrological data from Neue Donau (the
flood water bypass of the Danube), the Wasserpark (a north-western appendix of
Alte Donau) and from Alte Donau (Fig. 4.1). This network was operated by the
municipality of Vienna, to measure and document groundwater levels, rainfall and
evaporation. These records together with the knowledge of the hydrogeology from
bore holes form the basis to describe and model the groundwater flow.
Data were collected in the field by reading water levels at gauging points or in
groundwater bore holes. Flow rates (discharges) were estimated from Pygmy meter
measurements at significant sections (Fig. 4.2).
The hydrological conditions are recorded by a network of monitoring points
measuring precipitation, evaporation and recording temperatures. Water levels were
measured in groundwater bore holes and at staff gauges. Selected locations were
equipped with automatic data collectors. The data collected from the network were
stored in a database for further assessment. Data evaluation was done graphically
with specific computer software as time series charts or plots of groundwater iso-
flow. These plots were used to correct the data if necessary and graphically repre-
sent at the same time the hydrological conditions (Taschke and Voggenberger 2007,
Fig. 4.2 Measure devices from left to right: Weir with staff gauge level for flow determination;
metering a groundwater probe (bore hole); flow measurement with a Pygmy meter
38 R. Taschke
2014; Taschke et al. 2004a, b, 2014). The groundwater flow is calculated using
numerical sub-models as part of the model used to calculate the water balance of
Alte Donau (Taschke et al. 2004a, b, 2014).
The crucial elements to the water balance in Alte Donau are precipitation, evapo-
ration, the groundwater input to the system, and the output into the aquifers of the
Hinterland. The water balance, responsible for changes in the water level of Alte
Donau, can thus be modelled and compared with the observed changes.
4.3 Results
Neue Donau represents the boundary condition for the flow to the Wasserpark situ-
ated slightly lower and further down to Alte Donau. The associated water level dif-
ferences are decisive for the amount of inflow to the Wasserpark and to Alte Donau.
Accordingly, the differences to the Hinterland determine the outflows from the
Wasserpark into the aquifers of the Marchfeld, the region north-east of the system.
The water levels in the upper impoundment of the flood water channel Neue
Donau, in the Wasserpark, in Alte Donau and in the Hinterland are depicted in
Fig. 4.3 for the years 2000–2014.The water level varies seasonally by about a meter
or more in Neue Donau with minima occurring at the end of the year and maxima
during summer (average 157.6 m, maximum 158.1 m, minimum 156.68 a.s.l.). Inter-
Fig. 4.3 Water level changes in Neue Donau, Wasserpark, Alte Donau and the Hinterland for the
years 2000–2014. Water level as meters above adriatic sea level (a.s.l.)
Fig. 4.4 Climatic water balance components of Alte Donau: Precipitation (blue) and evaporation
(red), both in mm for the years 2000–2014
annual variation is negligible. Groundwater levels in Wasserpark and Alte Donau

resemble these variations but annual differences are smaller. Wasserpark and Alte
Donau vary from 156.5 m to 157.3 m a.s.l. (average 157 m a.s.l.) and from 156.5 m
to 157 m a.s.l. (average 156.7 m a.s.l.) respectively (Fig. 4.3). The associated water
level difference between Neue and Alte Donau is on average 0.88 m (range –0.07 to
1.29 m). The outflow to the Hinterland via groundwater varies by more than a meter
from 154.5 m to 155.7 m a.s.l. (average 153 m a.s.l.) and does not correspond to the
level changes in Neue and Alte Donau. Inter-annual differences are more pronounced
and long-term trends are observable. Water level in the Hinterland first decreased
from 2000 to 2003 and then increased till mid-2010. Levels decline again until the
end of 2012 and finally stabilised at 155.2 m a.s.l. in 2013.
Precipitation and evaporation, the other significant components of the water bal-
ance, are depicted in Fig. 4.4 as climatic water balance. Withdrawals, also affecting
the water balance, are not shown. Precipitation varies considerably between years.
Average long term precipitation of 2.5 mm (daily) was equal to mean evaporation.
Maximum rainfall of 57.5 mm occurred on 22nd of June 2009 while maximum
evaporation was measured on 29 June 2014 (Fig. 4.4).
A flow model for groundwater was used to calculate the inputs and outflows of
the Wasserpark and the Alte Donau. This model accounted also for the barrage of
wells withdrawing contaminated groundwater from the waste deposits at Donaupark
and Bruckhaufen (see Chap. 2), providing a hydraulic barrier against input of dan-
gerous substances into Alte Donau (Fig. 4.5).
Flow rates were calculated for selected areas and dates. In each area water level
differences to Alte Donau were calculated from a reference station. Water level dif-
ferences were related to flow rates by regression analysis allowing calculation of
in- and outflows from level differences (Taschke et al. 2004a, b, 2014).
40 R. Taschke
Fig. 4.5 Contour lines of groundwater levels calculated with the numerical model. The five black
rings represent the wells to protect Alte Donau from pollution seepage (see text)
Fig. 4.6 Relation of water level difference (m) to inflow rate (L s–1) for the Wasserpark. The
regression equation and the variance are indicated
Such a relationship is exemplified in Fig. 4.6 for the inflow to the Wasserpark.

The influx to the Wasserpark depends on the water level difference between the
Wasserpark and the upper impoundment of Neue Donau. The contour lines of
groundwater flow derived from this relation is shown in Fig. 4.7.
The numerical model puts together the in- and outflow quantities of the ground-
water, the inputs to and the withdrawals from the Alte Donau, and the climatic
components to derive a volume balance. The modelled output must match the water
level changes measured within the observation period (e.g. one week). The mea-
sured and calculated volume changes and the resulting volume balance are shown in
Fig. 4.8. All calculations and simulations must consider any water management
attempts such as water level draw down in spring in Alte Donau to promote macro-
phyte growth, water introduced from Neue Donau into Alte Donau, and water
drained from the Wasserpark into the left Danube sewer to improve limnochemical
conditions particularly to stabilize the pH value (Chaps. 5 and 6).
Fig. 4.7 Contour lines of groundwater level and groundwater flow as L s–1 derived from the
groundwater model for the area Wasserpark
Fig. 4.8 Comparison of the volume change measured and determined with the water balance
model (both as 103 m3). The total withdrawals and the input into the upper basin (both as L s–1 at
Kagraner Brücke are also indicated
42 R. Taschke
Fig. 4.9 Scheme of current groundwater flow rates in the surrounding of Alte Donau: Red arrows
indicate the average direction and the range of flow rates (as L s–1) of the groundwater derived from
weekly data 2008–2012. Black dots indicate the wells (Sperrbrunnen)
The current situation of the groundwater in- and outflow is generalised and sche-
matically shown in Fig. 4.9. As can be seen, the quantities reaching Alte Donau
from Neue Donau vary considerably in the length of the water body. The barrage of
wells (black dots in Fig. 4.9) protect Alte Donau from an input of 23–65 L s–1 con-
taminated ground-water from the old dump side creating a back flow of 0–35 L s–1
from Alte Donau. This is compensated within Alte Donau by 1–48 L s–1 flowing
from the south-eastern basin to the north-western (upper) basin. Flow rates to the
Hinterland increase from north to south.
4.4 Discussion
Early records of water level changes in the region have indicated a highly dynamic
system which was at that time not permanently separated from the main river. Water
levels changed seasonally by up to 1.20 m with minimum levels in winter and maxi-
mum height during summer (e.g. Pesta 1928; Oberzill 1941; Grohs 1943). These
and previous studies also demonstrated that water level changes in back-waters are
closely coupled to the level changes in the river and follow those within 2–3 days
(Steuer 1901; Brunnthaler 1907). This dynamic linkage is characteristic for flood-
plain lakes and back-waters in general and has persisted with minor modifications
in Alte Donau until the mid-1970th (Chap. 5).
The grade of connectivity largely determines the range of seasonal changes in the
water levels of floodplain lakes and their dependence from the river (Dokulil 2003;
Rezabek et al. 2003). Accordingly ground water levels close to a river are different
and retarded close to oxbow lakes (e.g. Kim et al. 2009; Lewandowski et al. 2009).
The present situation of the average groundwater flow rates in Alte Donau
(Fig. 4.9) are a result and a reflection of the present complex boundary conditions.
Over the past decades newly built water management constructions have added to
the complexity of the groundwater flow in the vicinity of Alte Donau. These struc-
tures include, among others, the flood water bypass Neue Donau, a hydro-power
dam in the Danube (Michelmayr 2005) and the pumping wells protecting Alte
Donau from underground pollution input (Prohaska 2005). These structures signifi-
cantly altered the water balance, the water level and the original dynamic of the
ecosystem in the second half of the twentieth century. The water balance analysed
for June 1987 (Löffler 1988) already indicated similar flow directions and rates as
in the present study. The ground-water flow behaviour of Alte Donau nowadays is a
mixture of oxbow and seepage lake conditions. Differences of inflow and outflow
side are typical for many water bodies solely fed by ground-water (e.g. Swancar
et al. 2000; Baron 2007).
4.5 Conclusions
Although the water balance model cannot include all relevant variables into the
water budget, the match is satisfactory. The methodology used sufficiently describes
the water budget of the Alte Donau (Taschke et al. 2014). Close investigation of the
water balance indicates a water deficit in the upper basin of Alte Donau which is
compensated by water from the lower basin through the bottle-neck at Kagraner
Brücke.
The deficit in water balance is caused by the wells to secure the dump site
Donaupark and Bruckhaufen. The wells not only prevent the influx of polluted
water to the upper basin of Alte Donau, they also collect water from there. Further
water management measures, such as the water dotation of the backwaters in the
flood-plain Lobau and other withdrawals, result in an annual average deficit of
about 45 L s–1 in Alte Donau.
The water budget model can simulate water levels at different amounts of inputs
to and withdrawals from Alte Donau. The annual average water level in Alte Donau
will rise by 15–20 cm if the deficit of 45 L s–1 is compensated. This deficit may be
compensated in the near future by introduction of water from Neue Donau which
must first percolate through a soil filter to remove phosphorus and to add calcite.
Acknowledgemnts I would like to thank all staff members of the engineering bureau Gruppe
Wasser for their significant contributions in collecting and analysing the hydrographic data. Special
thanks go to Katrin Teubner for her support in finalising this chapter.
44 R. Taschke
References
Baron U (2007) Hydrogeologie der “Vier bunten Seen” im Muskauer Faltenbogen. Bandenburger
geowissenschaftlichen Beiträge 14:75–86
Birch S, McCaskie J (1999) Shallow urban lakes: a challenge for lake management. Hydrobiologia
395/396:365–377
Brunnthaler J (1907) Die Algen und Schizophceen der Altwässer der Donau bei Wien.
Verhandlungen Zoologisch-Botanische Gesellschaft Wien 57:170–223
Dokulil MT (2003) Horizontale und vertikale Interaktionen im Mega- und Mikro-Ökoton einer
Flussaue: Phytoplankton und Phytobenthos. In: Janauer GA, Hary N (eds) Ökotone Donau-
March, Veröffentlichungen des Österreichischen MaB-Programmes, vol 19. Universitätsverlag
Wagner, Innsbruck, pp 113–155
Grohs H (1943) Limnologische Untersuchungen zweier Donaualtwässer bei Wien. Arch Hydrobiol
39:369–402
Kim KH, Yun ST, Choi BY, Chae GT, Joo Y, Kim K, Kim HS (2009) Hydrochemical and multivari-
ate statistical interpretations of spatial controls of nitrate concentrations in a shallow alluvial
aquifer around oxbow lakes (Osong area, central Korea). J Contam Hydrol 107(3):114–127
Lewandowski J, Lischeid G, Nützmann G (2009) Drivers of water level fluctuations and hydrologi-
cal exchange between groundwater and surface water at the lowland River Spree (Germany):
field study and statistical analyses. Hydrol Process 23(15):2117–2128
Löffler H (1988) Alte Donau. Projektstudie im Auftrag der Wasserstrassendirektion. 272 p
Michelmayr F (2005) Gegen den Strom. Die Regulierung der Donau. In: Brunner K, Schneider P (eds)
Umwelt Stadt, Geschichte des Natur- und Lebensraumes Wien. Böhlau Verlag, Wien, pp 307–317
Naselli-Flores L (2008) Urban lakes: ecosystems at risk, worthy oft the best care. In: Proceedings
Taal 2007: the 12th world lake conference, pp 1333–1337
Niemczynowicz J (1999) Urban hydrology and water management ± present and future chal-
lenges. Urban Water 1:1–14
Oberzill W (1941) Biologisch-chemische Untersuchungen des Tritonwassers im Gebiet der Alten
Donau bei Wien. Arch Hydrobiol 37:533–577
Pesta O (1928) Berichte zur Limnologie der „Alten Donau“bei Wien. Arch Hydrobiol 19:301–317
Prohaska W (2005) Altlastensanierung. In: Brunner K, Schneider P (eds) Umwelt Stadt, Geschichte
des Natur- und Lebensraumes Wien. Böhlau Verlag, Wien, pp 294–295
Rezabek HR, Hladej M, Halbritter T, Stritzinger W, Stepan C (2003) Hydrologische
Randbedingungen für Ökotone in Donauauen. In: Janauer GA, Hary N (eds) Ökotone Donau-
March, Veröffentlichungen des Österreichischen MaB-Programmes, vol 19. Universitätsverlag
Wagner, Innsbruck, pp 309–319
Steuer A (1901) Die Entomostrakenfauna der „alten Donau“bei Wien. Zoologische Jahrbücher.
Abt. Systematik, Geographie und Biologie der Tiere 15:1–155 + 26 Abb. im Anhang
Swancar A, Lee TM, O´Hare TM (2000) Hydrogeologic setting, water budget, and preliminary
analysis of ground-water exchange at Lake Starr, a seepage lake in Polk County, Florida.
U.S. Geological eurvey, water resources investigation report 00.4030, Tallahassee, Florida,
p. 55 + Appendix p. 65
Taschke R, Voggenberger C (2007). Alte Donau – Anhebung des Puffervermögens mittels
Wassertausch – Beweissicherung Bericht 2006 – Teil B Hydrologie
Taschke R, Voggenberger C (2014) Alte Donau – Anhebung des Puffervermögens mittels
Wassertausch – Beweissicherung 2007 – 2011 Teil B Hydrologie
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Wasserwirtschaftlicher Versuch – Beweissicherung 2004 bis Beweissicherung 2014. 15
Jahresberichte an die MA 45
Taschke R, Luza T, Voggenberger C, Schwillinsky S, Prantl M (2004b) Wasserhaushalt Alte Donau
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Taschke R, Voggenberger C, Leonhartsberger I, Wögerer C (2014) EU-Life + Urban Lake Alte
Donau Life 12, Action B2, Aktualisierung der hydrologischen Bilanz
Chapter 5
Restoration and Lake Management
Karl Donabaum and Martin T. Dokulil
Abstract At the beginning of the 1990ies Alte Donau was characterized by severe
eutrophication problems, leading to a deterioration of water quality. The former
macrophyte-dominated state changed by a catastrophic shift to a turbid state domi-
nated by high biomass of filamentuous cyanobacteria, associated with a significant
reduction in transparency. A holistic lake management concept was developed to
restore macrophyte domination. Integrated restoration plans included internal and
external measures. Improvements in the catchment aimed to minimize the input of
nutrients from contaminated groundwater, from sewage and from storm waters.
Internal measures included aeration, water exchange, chemical flocculation and
nitrate oxidation of the sediments (Riplox treatment). The main measures were fol-
lowed by biomanipulation actions like selective fish stocking, planting of macro-
phytes and weed management after recovery. Technical measures like the simulation
of low water levels to enhance the light supply for the macrophytes in early spring
or partially water exchange with water from the nearby impoundment Neue Donau
to stabilize the buffer capacity of the system to compensate biogenic calcium pre-
cipitation are still in progress. Restoration was successfully. Full recovery of Alte
Donau was achieved in 2004, when intense macrophyte growth became apparent,
consequently triggering the switch to a clear water phase. Since then Alte Donau is
in a macrophyte dominated stable state, characterized by high water quality with
low nutrient and chlorophyll-a concentrations.
Keywords Restoration · Lake management · Alternative stable states · Water

management measures · Resilience
K. Donabaum (*) · M. T. Dokulil

e-mail: karl.donabaum@dws-hydro-oekologie.at; martin.dokulil@univie.ac.at

https://doi.org/10.1007/978-3-319-93270-5_5
46 K. Donabaum and M. T. Dokulil
5.1 Eutrophication History
Evidence from the literature suggests that phytoplankton dominated in Alte Donau
at the beginning of the twentieth century (Brunnthaler 1907; Schiller 1929), where
high abundances of bloom forming species have been reported. In the time period
between 1960 and 1985, the water body was characterized by high transparency,
low chlorophyll-a concentrations and extensive macrophyte growth. The dominant
species in the lake were Myriophyllum spicatum L. and Potamogeton pectinatus L..
Charophytes like Nitellopsis obtusa (DESV. in LOIS) J. GROVES, Chara tomen-
tosa L. and Chara hispida WOOD were also abundant (Löffler 1988). A similar
macrophyte composition was reported from the swedish lakes – Lake Takern and
Lake Krankesjon - which have shifted several times between a clear – water state
and a turbid state (Blindow et al. 1993).
In the early 1990ies a dramatic change to a phytoplankton dominated state was
observed. Significant numbers of the blue-green algae Limnothrix redekei VAN
GOOR and Cylindrospermopsis raciborskii (WOLOSZ.) SEENAYYA ET SUBBA
RAJU were recorded in water quality samples in the year 1992 (Chap. 9). Average
annual chlorophyll-a concentrations increased substantially and secchi depth
decreased (Dokulil 1994; Mayer et al. 1997, Chap. 6). Associated with these
changes was a remarkable decline of submerged macrophytes (Chap. 8). Within
2 years only remnants of macrophyte and charophyte stands were left (Dokulil and
Janauer 1995).
Several reasons were responsible for the shift from a clear water state to a turbid
state in Alte Donau. Substantial nutrient influx from non point sources, such as leak-
ing septic tanks, inclompletion of the sewage network and elution from a nearby
former dump site were believed to be responsible for excess nutrient loading.
Nutrient inputs originating from exretion of waterfowl and from recreation activi-
ties additionally contributed to eutrophication. Estimation of phosphorus addition
by 1.2 Mio visitors in a typical summer period amount to approximatley 114 kg P.
Investigations of the fish community in 1996 (Waidbacher et al. 1996) revealed a
substancial dominance of bleak (Alburnus alburnus L.). The cyprinids roach
(Rutilus rutilus L.), rudd (Scardinius erytrophthalmus L.), bream (Abramis brama
L.), white bream (Blicka bjoerkna L.) and tench (Tinca tinca L.) were abundant.
Common Carp (Cyprinus carpio L.) was the main fish stocked. Perch (Perca fluvia-
tilis L.) was also abundant in Alte Donau and the piscivores pike (Esox lucius L.) as
well as pike-perch (Stizostedion lucioperca L.) were stocked. An extensive grazing
pressure on zooplankton was assumed by the fish community, especially from bleak.
Confirmation came from zooplankton studies in 1994 (Dokulil 1994, Chap. 11).
Abundances and species composition were typical for eutrophic conditions. Among
the crustaceans, small species like Eubosmina coregonii BAIRD showed higher
abundances than larger species like Daphnia cucculata G. O. SARS. These results
coincide with investigations in 1987 (Löffler 1988), but they are in contradiction to
earlier reports, where high species diversity amongst the cladocerans was reported
(Pesta 1928).
5 Restoration and Lake Management 47
Fig. 5.1 Water level fluctuations in Alte Donau from 1964–2011. Min. & Max. values – white
area, black line – mean level, dashed line – mean level of periods
Investigations on benthic communities (Waidbacher et al. 1996) showed a loss of

characteristic species and poor diversity of macroinvertebrates in comparison to ear-
lier studies (Löffler 1988) and to similar backwaters. Lack of habitates in the littoral
zone and loss of macrophytes were recognized to be responsible (Chaps. 8 and 18).
One of the triggering factors for the shift to a phytoplankton dominated state was
the water level dynamics in the second half of the twentieth century. Water level
fluctuations for the period from 1958 to 1993 are shown in Fig. 5.1. Reduced water
level dynamics in combination with a higher water level can be seen from the late
70’s onwards. The mean water level calculated for the period from 1978 to 1993
was about 37 cm higher than in the period before. Similarly the range of water level
fluctuations decreased by 1 m. The sharp transition in hydrological dynamics in
1978 are basically due to the nearby construction of a highway and the impound-
ment Neue Donau.
These changes seem to be correlated to the severe decline of the macrophytes
similar to observations elsewhere (Scheffer et al. 1993; Blindow et al. 1993). The
decisive environmental variable for macrophyte decrease is the altered underwater
light climate (Chambers and Kalff 1985). In addition, macrophytes were exten-
sively removed by the local authorities as a management practice for many years.
Grazing by high densities of water fowl contributed to the disappearence of sub-
merged macrophytes in Alte Donau. According to Steiner (1986), water fowl grazed
about 194 tons (wet weight) of macrophytes per year. The vanishing of macrophytes
was accelerated through light limitation when phytoplankton began to dominate.
Fig. 5.2 Striking yellow colour in Alte Donau during blooms of cyanobacteria, Aerator on a plat-
form before installing
As a consequence of increased nutrient input, macrophyte decrease and grazing

pressure on zooplankton, cyanobacteria became abundant in 1993 and 1994. The
water showed a striking yellow colour and transparency decreased to 30 cm (Fig. 5.2).
5.2 Restoration Measures
Integrated lake management strategies included external and internal measures.

Rapid expansion of the sewerage network was initiated when the problem became
apparent. To keep off contaminated groundwater from the lake pumping wells were
constructed between the former dump site and the lake. Since 1995 they keep off the
inflow of nutrient enriched ground water from the area of the dump site.
As a first internal measure, artificial aeration combined with vertical mixing was
performed in 1993 in areas where thermal stratification and hence oxygen depletion
near the sediment were expected to create problems. Substantial nutrient release
from the sediment under anoxic conditions, shown by laboratory experiments, could
thus be avoided (Dokulil 1994).
Half of the water volume of the lake was exchanged by water low in nutrient
concentrations from the impoundment Neue Donau in December 1993 (Dokulil
Riplox-Treatment
1995: 997 t 1995: 303 t

1996: 980 t 1996: 200 t
39% FeCl3 50% Ca(NO3)2
water surface
-1 -
1995: 69 mg l Cl
36 mg l-1 Fe3+
1996: 68 mg l-1 Cl-
36 mg l-1 Fe3+
1995: 7 mg l-1 NO3--N

1996: 5 mg l-1 NO3--N
1995: 295 g m-2 Fe3+ 1995: 12 g m-2 NO3--N

1996: 290 g m-2 Fe3+ 1996: 8 g m-2 NO3--N
Sediment
Fig. 5.3 Sheme of Riplox treatment in Alte Donau
and Gasser 1994). This treatment resulted in a moderate phytoplankton develope-

ment in spring 1994, consisting mainly of diatoms and chlorophyceans. However,
in summer cyanobacteria reappeared reaching the highest biomass ever recorded
(Chaps. 6 and 8).
After careful consideration of most of the restoration techniques commonly
used, the RIPLOX-method was selected as the main restoration measure (Ripl 1976,
1978, 1986, 1995; Ripl and Lindmark 1978; Ripl and Wolter 1993a, b). It was
implemented in the spring periods of 1995 and 1996 (Fig. 5.3). The method consists
of two steps which are performed in chronological sequence. In the first step, FeCl3
buffered with limestone was added to remove phosphorus and suspended material
by chemical and mechanical flocculation. The FeCl3 acts as a barrier against phos-
phorus release from sediments. In a second step, Ca(NO3)2 was added to the sedi-
ments to enhance nitrate oxidation. Nitrate is reduced to elementary nitrogen by
anaerobic denitrification. Organic mud is oxidized to carbon-dioxide and water.
Consequently oxygen deficit caused by heterotrophic metabolism decreases. Thus
prolonged anoxic conditions near the sediment surface can be avoided and internal
loading reduces.
Immediatly after the Riplox treatment periods transparency significantly

increased and concentrations of TP and chlorophyll-a dropped down (Chap. 6). The
composition of the phytoplankton assemblage shifted from cyanobacteria towards
diatoms and green algae (Chap. 9), accompanied by an increase in the relative bio-
mass contribution of heterotrophic bacteria (Kirschner et al. 1998) and zooplankton
(Chap. 11). For short periods cryptophyceans, chrysophyceans, dinoflagellates and
coccal cyanobacteria were also abundant. The shift in the phytoplankton composi-
tion may be due to the artificially altered N/P ratio. Among other factors, summa-
rized by Varis (1993), low N/P ratios seem to favour cyanobacterial growth (Smith
1986; Varis 1993). The restructured plankton community had an enhanced phospho-
rus accumulation efficiency and acted as a sink for phosphorus in a reduced total
pool size (Teubner et al. 2003). The period of intermediate nutrient concentrations
from 1997 to 1999 was characterized by the continued abundance of filamentous
cyanobacteria (Chap. 9), poor growth of macrophytes (Chap. 8) and slow increase
in the abundance of larger species of zooplankton (Chap. 11). This time period can
be seen as the resilience phase of the ecosystem during recovery. It may also be seen
as an unstable situation where shifts in any direction could have been possible.
To push the system further in the desired direction, biomanipulation was
attempted.
Attempts were made to concentrate most of the water fowl in a small appendix
of the north-eastern basin which has been cut off from the rest of the lake. Collecting
eggs of the ducks was proposed as a measure to reduce the high stock of water fowl
(Steiner 1986). In addition, people were asked not to feed the birds.
Great efforts were made by the local authorities in planting a reed belt to give
more structure to the littoral zone in the southern basin and to enhance sedimenta-
tion of organic material. Structuring and revitalisation of the shore line is still in
progress (Chaps. 18 and 19).
For several reasons, large scale fish removal was not possible in Alte Donau. But
after carrying out top down and bottom up experiments in an appendix of Alte
Donau (massive stocking with pikeperch) as well as mesocosm experiments to esti-
mate the feeding rates of predators (Chap. 15), stocking with predators like Aspius
aspius L. and pike perch (Stizostedion lucioperca L.) was intensified to reduce bleak
(Alburnus alburnus L.) and other planktivorus fish to enhance the development of
larger cladocerans. After the recovery of macrophytes also the stocking with pike
(Esox lucius L.) was promoted.
Several experiments using enclosures and cages were performed in view of sup-
porting the recovery of macrophytes. Macrophytes were taken from the impound-
ment Neue Donau and transferred into experimental areas in Alte Donau. The
influence of light (artificial illumination), carbon supply (CO2-gassing) and the
absence of fish on the growth rates of different macrophyte species were investi-
gated (Chap. 8). Finally light supply was identified as the most limiting factor for
recovery of macrophytes. A technical measure was introduced to improve the under-
water climate for macrophyte growth – the simulation of low water levels in spring
(Chap. 4). From 2002 onwards the water level of Alte Donau was artificially low-
ered from march to may in a range of approximately 25 cm. Within only 2 years this
20
18
Macrophyte dry weight biomass [MB, t]
700 16
14
12
Phytoplankton biomass [mg L-1]

400
4
300
200
2
100
0 0
Year
Fig. 5.4 Biomass of Macrophytes and Phytoplankton (white section of bar corresponds to the
amount of cyanobacteria) in the time period from 1987 to 2012
treatment resulted in a dramatic increase of the macrophyte stands, mainly consist-

ing of Myriophyllum spicatum L. (Fig. 5.4, Chap. 8). Beginning with the year 2004
a management plan for mowing was established to ensure activities like swimming
and boating. The removed dry weight of macrophytes differs per season but it is in
the range of 100–200 to per year.
The intensified macrophyte growth revealed an unexpected problem, which
became apparent in the year 2005. The pH-values highly increased (up to max val-
ues of 10) in the northern basin of Alte Donau due to biogenic calcium precipitation
and low ground water influxes (Chaps. 4 and 6). To overcome this problem a techni-
cal management measure was introduced in 2006. At high pH values (9.5) and
defined minimum levels of Ca and HCO3, Alte Donau is flushed with water from the
nearby impoundment Neue Donau for a few weeks. The exchange rate is in the
range of 1.5 Mio m3 and the measure is stopped when the target values in the north-
ern basin are reached (Ca > 40 mg L1, pH < 8.5).
To ensure sustainability of macrophyte domination, recently the focus was set on
measures like enhancing species diversity and planting of charophytes. In addition to
this, protected areas have been defined, where no mowing is allowed (Chaps. 8 and 19).
In the last decade stable mesotrophic conditions could be observed, with TP yearly
mean values ranging from 12 to 14 μg L–1 and yearly mean chlorophyll-a values rang-
ing from 3 to 4.5 μg L–1, corresponding to Secchi depth values between 3.5 and 4 m.
Phytoplankton became dominated by chrysophytes and cryptophytes in spring and a
mixture of green algae, dinophytes and coccal cyanobacteria in summer (Chap. 9).
The zooplankton composition reflects the mesotrophic conditions (Chap. 11).
5.3 Conclusion
Big efforts were made to restore the backwater system of Alte Donau after the
occurence of sever eutrophication problems at the beginning of the 1990ies. The
case study of Alte Donau has once more verified the theory of alternative stable
states (Chap. 1). An integrated lake management plan, based on the theory of alter-
native stable states was developed and implemented over the past decades
(Table 5.1). Monitoring of recovery showed that this water management plan has
been fully successful. Moreover, long term investigations indicated significant resil-
ience of the system against changes and hysteretic behaviour during recovery phase.
In addition to this the long term data set and the permanent monitoring of water
quality is the basis for finetuning of water management measures which are neces-
sary to keep the ecosystem in a stable macrophyte dominated state.
Table 5.1 Sequence of restoration and rehabilitation measures

Year(s) Measure Result
1989– Planting reed and marsh plants Enlargement of shore zones (Chap. 18)
1994
1993 Water exchange (winter period) Water quality improved for 6 months
1994/1995 Aeration Effect on oxygen, no effect on water quality
Sedimentrehabilitation Effect on oxygen, no effect on water quality
P-Flocculation effect on all water quality parameters
Sediment aeration No significant effects on sediment and water
(details see this chapter)
1995 1st Riplox treatment Significant improvement of water quality
1996 2nd Riplox treatment Stabilisation of clear water (Chaps. 6, 9, and
10)
1996/1997 First macrophyte plantings Successful, but stands were effected by grass
carps and other fish; as a result fish free areas
were planed
1998 Fish-Exclosure Rapid macrophyte growth in fish free area
(unpublished result)
1998 Biomanipulation with fish in the effect on Zooplankton biomass and size,
small appendix ‘Kaiserwasser’ improvement of water quality (Chap. 15)
1999– Macrophyte plantings for testing Experiments showed that enhanced light
2000 growth factors supply was the key factor for stimulating
macrophyte growth (Chap. 8)
Since Temporary water level draw-down in Significant and rapid proliferation of
2002 spring to enhance light supply in macrophyte stands (Chap. 8); stabilisation of
spring period good water quality (Chaps. 6, 9, and 11)
Since Macrophyte management (mowing, To ensure utilization; to establish type-
2004 stone wort planting) specific vegetation, to enlarge resilience of
macrophyte vegetation, to remove P bound in
macrophytes
Since Periodically water exchange (with To avoid high pH-values (this chapter)
2006 water from Neue Donau) associated with fotosynthesis and
Ca-preciptitaiton
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Smith VH (1986) Light and nutrient effects on the relative biomass of blue-green algae in lake
phytoplankton. Can J Fish Aquat Sci 43:148–153
Steiner E (1986) Untersuchungen zur Dynamik des Wasservogelbestandes an der Alten Donau und
dessen Bedeutung für die Ökologie dieses Gewässers. Bericht an die Wiener Magistratsabteilung
22, 42 pp
Teubner K, Crosbie N, Donabaum K, Kabas W, Kirschner A, Pfister G, Salbrechter M, Dokulil
MT (2003) Enhanced phosphorus accumulation efficiency by the pelagic community at
reduced phosphorus supply: a lake experiment from bacteria to metazoan zooplankton. Limnol
Oceanogr 48(3):1141–1149
Varis O (1993) Cyanobacteria dynamics in a restored Finnish lake: a long term simulation study.
Hydrobiologia 268:129–145
Waidbacher H, Moog O, Janecek B, Kummer, Mann H, Schmidt-Kloiber A, Nesemann H, Graf W
(1996) Endbericht zu fischökologischen und benthosbiozönotischen Aufnahmen im Rahmen
der „FeCl3 Behandlung“ in der Alten Donau. Jahresbericht an die Wiener Magistratsabteilung
45. Wasserbau, 102 pp
Chapter 6
Long-Term Changes of the
Physico-chemical Conditions in Alte Donau
Karl Donabaum and P. Riedler
Abstract The long-term development of physico-chemical changes will be

described here from the early twentieth century till recent years with the main focus
on the past three decades. Four periods can be distinguished: the eutrophication
phase dominated by phytoplankton meaning high contents of phosphorus and low
water transparency. By applying the RIPLOX treatment (restoration) algal biomass,
particulate matter and nutrients dropped down, but also some chemical parameters
temporary changed, like chloride, calcium and nitrate. When macrophytes recov-
ered pH values raised significantly, moreover influenced by low buffering capacity.
To encounter this process flushing from the impoundment Neue Donau was carried
out periodically, representing a further impact on physico-chemical development.
Currently Alte Donau indicates stable mesotrophic conditions with low annual
mean concentrations of TP and chl-a and high transparency.
Keywords Long-term changes · Physico-chemical coniditions · Nutrients ·

Chlorophyll-a · Transparency · Nutrient balances
6.1 Introduction
The physico-chemical conditions of Alte Donau, which are associated with environ-
mental factors like morphology, hydrology, geology and urban development reflect
the variable history of Alte Donau which was marked by severe eutrophication
problems in the 1990ies, triggered by human and environmental impacts.
K. Donabaum (*)
e-mail: karl.donabaum@dws-hydro-oekologie.at
P. Riedler
DWS-Hydro-Ökologie GmbH, Technisches Büro für Ökologie und Landschaftsplanung,
Wien, Austria
e-mail: patricia.riedler@dws-hydro-oekologie.at

https://doi.org/10.1007/978-3-319-93270-5_6
56 K. Donabaum and P. Riedler
Actually, the good physico-chemical status and the outstanding optical proper-
ties of the water body can be seen as the outcome of numerous restoration measures
set in the past decades, eventually leading to a drop down of the eutrophication level
and a shift back from phytoplankton to macrophyte domination.
6.2 The History of Investigations in Alte Donau
The first investigations in Alte Donau were made by Brunnthaler (1907) and Schiller
(1929) at the beginning of the twentieth century. They described the species composi-
tion of the phytoplankton community and gave evidence of the formation of temporary
algal blooms. It can be assumed that at that time Alte Donau has been a typical back-
water of the river Danube with similar hydrochemical properties as the river itself.
The onset of intensive investigations of Alte Donau was given in the 1980ies with
the study of Löffler (1988). This was followed by a period with more scarce data
material up to 1992. From 1992 up to now a continuous and extensive monitoring
was performed in Alte Donau. Long-term changes can be presented, showing the
variable history and fluctuations of the physico-chemical conditions as well as
changes in ecological and trophic status (Dokulil et al. 2000; Dokulil and Teubner
2000; Donabaum et al. 2004).
The time span of intensive investigations in Alte Donau covers the following four
periods:
• Eutrophication period (1987–1994)
• Restoration period (1995–1999)
• Recovery of Macrophytes (2000–2006)
• Stable conditions with macrophyte domination (2007–2014)
6.3 Methods
The determination of the physical-chemical and chemical parameters was carried

out according to standard methods (see Donabaum et al. 2004). An overview of the
applied methods is found in https://www.bmwfw.gv.at/TechnikUndVermessung/
Akkreditierung/Documents/testing%20laboratories.pdf and http://www.dws-
hydro-oekologie.at/leistungsspektrum.htm
6.4 L
ong Term Changes – Physico-chemistry and Main
Constitutes
Like in most shallow lakes temperature regime of Alte Donau is strongly related to
air temperatures, which in Vienna show the typical trends of the temperate zone.
Duration of ice covering in winter is usually 4–8 weeks, highest water temperatures
in summer can reach 30 °C (Chap. 3). In general Alte Donau is polymictic. Two
6 Long-Term Changes of the Physico-chemical Conditions in Alte Donau 57
eutrophication restoration recovery stable conditions

10.0 phytoplankton
domination
macrophyte
domination
9.5
9.0
pH [-log[H+]]
8.5
8.0
7.5
7.0
87 88 89 90 91 92 93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
years
Fig. 6.1 pH-values (points) and running average (line) in Obere Alte Donau during the four peri-
ods between 1987 and 2014 (see description 6.2)
sites with depths above 5 m show thermal stratification for a few weeks in summer.
Area and volume of the stratified water body is small, so apart from local effects no
main influence to the physico-chemical conditions of Alte Donau is resulting from
stratification processes. The long-term changes in temperature indicate an increase
of the water temperatures in the last years (Chap. 3).
Apart from the physical exchange processes at the surface, oxygen content in the
euphotic zone is largely dependent on primary production. In the recent phase of
evolvement (2007–2014) only moderate oversaturation with oxygen near the surface
was observed in Alte Donau. The oxygen supply of the deeper zones and near the
bottom is sufficient throughout the year, the profiles show only a slight decrease with
increasing depth. One exception is a small area in the deepest part of the main basin
of Obere Alte Donau where thermal stratification (see above) and oxygen depletion
for a few weeks in summer are common. Consequently redissolving of phosphorus
and enrichment of ammonium can be observed but only to a minor extent. In the
eutrophication phase when phytoplankton was dominant (1992–1999) more massive
oversaturation with oxygen occurred in the surface water. Oxygen supply near the
bottom was neither a problem in the recolonisation phase with macrophytes nor in
the eutrophication phase. Whereas Löffler (1988) reported unremarkable oxygen
content in the surface water, but oxygen depletion in larger areas of the bottom within
stands of charophytes and in relationship to zones of organic rich sediments.
According to the geochemical characteristics and the buffer capacity, pH of Alte
Donau is slightly or moderate alkaline (7.6–8.8). The mean values of the period in
the 1980ies resemble probably the most type specific conditions (Fig. 6.1). In all
other periods the pH-values reflect various events like chemical treatment, a decreas-
ing buffer capacity in combination with biogenic effects (excessive macrophyte
growth) and other management measures like the flushing with water from the
Table 6.1 Mean, minimum Mean Minimum-maximum

and maximum values of main
Calcium [mg L–1] 38.0 13.3–83.0
ions in the Alte Donau in the
period between 1996 and Magnesium [mg L–1] 16.6 9.70–21.9
2014 Natrium [mg L–1] 16.8 8.88–29.7
Potassium [mg L–1] 2.89 1.50–4.22
Alkalinity [mmol L–1] 2.49 1.06–3.38
Sulfate [mg L–1] 12.2 7.20–35.5
Chloride [mg L–1] 30.9 12.3–115
impoundment Neue Donau. A dramatically increase in pH with values up to 10 in

the summer period was observed in the years from 2003 to 2005, when macrophyte
growth was intensive. A reduced ground water inflow especially in the area of Obere
Alte Donau (Chap. 4) and the biogenic calcium carbonate precipitation were the
reasons. To prevent the fish population from damages, buffer capacity was increased
by periodically flushing with water from the impoundment Neue Donau. Neue
Donau is similar to Alte Donau compared by nutrients but shows higher contents of
buffering substances like calcium (Ca++) and Alkalinity (HCO3-). With this measure
pH-values could be stabilized in the range of 8.2 up to maximum values of 9.0. In
the 1980ies when macrophytes also were prominent, pH-values showed a high sea-
sonal fluctuation range, but because of a higher ground water inflow buffer capacity
was sufficient, values which could be harmful to fish were not observed in this
period.
The long-term trend of conductivity covers the range of 320–550 μS cm–1, show-
ing a decrease in the period of macrophyte recolonisation (2000–2006), which is
related to the same processes as explained for the increase of pH. Conductivity was
slightly higher (20–50 μS cm–1) in the basin of Obere Alte Donau in most of the
periods. The long-term trend shows also a weak increase of conductivity in both
basins over the last years, corresponding to the chloride values (see above).
The main constitutes and dominant cations are calcium (Ca++) and magnesia
(Mg++), the main anions are HCO3–, sulfate (SO43–) and chloride (Cl–). An overview
about the mean concentrations of some ions and the alkalinity is given in Table 6.1.
The total salinity is about 8 mval L–1.
The long-term records for calcium show dramatic changes, which are related to
the chemical treatment in 1995 and 1996 when the values were highest (up to
80 mg L–1) as a consequence of adding Calcium carbonate (CaCO3) as buffering
agent to the acidic ferric chloride (FeCl3) solution (Fig. 6.2). The treatment with
FeCl3 was followed up by the the addition of Ca(NO3)2 to promote oxidation of sedi-
ments. The rapid decrease of calcium and the minimum values (17 mg L–1) in the
period between 2000 and 2006 are related to biogenic decarbonisation and reduced
groundwater fluxes. According to the biogenic activities calcium values show a dis-
tinctive seasonality. Stabilisation of calcium content on a yearly mean level of
30–35 mg L–1 could be achieved by flushing with water from Neue Donau. The
long-term trend of alkalinity shows a similar pattern like the calcium content, with
the main difference in the years 1995–1996 where the values dropped down as a

90
phytoplankton macrophyte
domination domination
80
70
calcium [mg L-1]
60
50
40
30
20
10
87 88 89 90 91 92 93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
years
Fig. 6.2 As Fig. 6.1 but for concentrations of calcium

120
100
chloride [mg L-1]
80
60
40
20
0
87 88 89 90 91 92 93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
years
Fig. 6.3 As Fig. 6.1 but for concentrations of chloride
consequence of the treatment with FeCl3. Related to this, values for chloride were
highest in the chemical treatment period (up to 100 mg L–1), followed by a rapid
decrease down to the range of 20–30 mg L–1 (Fig. 6.3). Cl– values show a slight
increase since the year 2004 which can be linked to an increase in the river Danube
and the impoundment Neue Donau which can be seen as the hydrological
constraint.
6.5 L
ong Term Changes – Nutrients, Chlorophyll-a
and Transparency
Total nitrogen (TN) was high in the eutrophication and restoration phase. In the
eutrophication phase the TN content (up to 4 mg L–1) was largely determined by the
particulate fraction. The anorganic compounds like nitrate (NO3–), ammonia (NH4+)
and nitrite (NO2–) played a minor role. In the course of the chemical treatment, after
addition of CaNO3, the two TN peaks mainly comprised of NO3–, which was effec-
tively incorporated by phytoplankton and relocated by bacteria in the denitrification
process. When this measure was terminated TN values rapidly dropped down to a
range between 500 and 600 μg L–1. A further decrease was observed after recoloni-
sation with macrophytes (300–400 μg L–1). In the last years, beginning with 2006,
TN is slightly increasing, showing short-term peaks in the single years. Tis peaks
are due to flushing events with water from Neue Donau, where the main constitute
of TN is represented by NO3–. Like in many shallow lakes, where denitrification
processes play an important role, Alte Donau is characterised by extremely low
nitrate availability for phytoplankton. These are conditions which typically favour
the growth of nitrogen fixing blue greens. Therefore the increase in NO3– by flush-
ing is seen as measure which favours the growth of other algal groups like green
algae and diatoms. The shift from blue greens to other algal groups, especially to
green algae, triggered by stocking the NO3– content was clearly observed after the
addition of Ca(NO3)2 in 1995 and 1996.
Although Alte Donau was dominated by macrophytes in 1987, the total phospho-
rus (TP) values (18–35 μg L–1) gave a first indication of the onset of an eutrophica-
tion process (Figs 6.4 a, Chaps. 5 and 20). In 1993 and 1994 eutrophication
culminated in extraordinary high TP values (110 μg L–1). Between 1987 and 1994
macrophytes completely vanished and phytoplankton blooms, dominated by fila-
mentous blue greens progressed. Chlorophyll-a (chl-a) increased dramatically and
reached the maximum concentration of 140 μg L–1 in year 1993. Accordingly the
Secchi transparency decreased rapidly to a minimum of 0.3 m (Figs 6.4b, c).
Successful RIPLOX treatment resulted in a remarkable decline of TP and chl-a and
in an increase of transparency. The period after the treatment was associated with
moderate TP and decreasing chl-a concentrations. Furthermore, in the year 2000
macrophytes started to recolonise the ecosystem and progressively reestablished in
the years 2003 and 2004. After switch back of Alte Donau to macrophyte domina-
tion, TP and chl-a shifted further downwards accompanied by a significant increase
in transparency. The latest period indicates stable mesotrophic conditions with low
annual mean concentrations of TP (10–14 μg L–1) and chl-a (3.5–6 μg L–1) and with
high transparency (2–4 m). In accordance with decreasing TP and chl-a, the concen-
tration of particulate matter shows a decreasing trend over decades.
Within the TP pool, the predominant fraction is particulate phosphorus. After
chemical treatment the values for soluble unreactive phosphorus (SUP) were in the
range of 5–8 μg L–1. After recolonisation with macrophytes SUP values dropped
further down to yearly means of 2–4 μg L–1. The decline of this fraction of soluble
restoration recovery stable conditions

a 120
eutrophication
total phosphorus [µg L-1]
100
80
60
40
20
0
b 160
120
chlorophyll-a [µg L-1]
60
40
20
0
c
0
1
secchi depth [m]
5
87 88 89 90 91 92 93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
years
Fig. 6.4 As Fig. 6.1 but for Phosphorus (a), chlorophyll-a (b) and Secchi transparency (c)
phosphorus, which can only be incorporated by algae after enzymatic digestion, can
be seen as an indicator of increasing phosphorus limitation (Teubner et al. 2003).
Values for the directly incorporable phosphorus fraction – soluble reactive phospho-
rus (SRP) – have been in the range of the detection limit (1 μg L–1) within the last
15 years.
Soluble reactive silica (SRSi) is an important nutrient for diatoms and some
chrysophyte species. If tributaries are missing its supply depends on the groundwa-
ter inflow and on the redissolution of diatom frustules in the sediments. Alte Donau
is characterized by Si concentrations in the range of 500–2500 μg L–1. But if diatom
biomass is high, silica can be completely depleted. The long-term records show a
decline during the periods of restoration and macrophyte recovery but an increase in
the most recent period of ‘stable conditions’. The increase in the last years is at least
partially related to the flushing measure with water from Neue Donau, where the
basic content of SRSi is higher than in Alte Donau.
Monitoring of dissolved organic carbon (DOC) started in 1994. DOC reached
maximum values of about 8 mg L–1 at the time of phytoplankton blooms during
‘eutrophication’ and decreased after chemical treatment. Since then yearly means
are between 2.8 and 3.5 mg L–1. Apart from the clear dropdown after the eutrophica-
tion phase there is no really conclusive pattern in the long-term trend.
6.6 Nutrient Balances
The nutrient balance is based on the hydrological balance of Alte Donau (Taschke
et al. 2014) covering a five years period from 2008 to 2012. In a first step, a balance
for chloride was calculated to validate the hydrological balance by comparing all
in- an output elements with volume based loads. Subsequently some further ion and
nutrient balances were modelled but the focus here is set on the phosphorus
balance.
The following list shows the elements of the chloride balance. If an element was
only relevant for one section of Alte Donau, upper and/or lower basin (Untere Alte
Donau = UAD, Obere Alte Donau = OAD, it is indicated in brackets1:
precipitation – Meteorological Station Illmitz
groundwater inflow
introduction from Neue Donau (ND) during water exchange (UAD)
groundwater outflow
discharge via Sperrbrunnen (OAD)
discharge via Wasserpark during water exchange (OAD)
discharge in the backwaters Lobau during waterlevel-decrease (UAD)
pass over from UAD in OAD at “Kagraner Brücke”
Temporal resolution was one month. The point of temporal changes in concen-
tration was fixed by end of month.
1
During water exchange, water flows from Neue Donau at marker 9.75 in Untere Alte Donau, goes
upstream in Obere Alte Donau and leaves the system via Wasserpark. It is possible to forward part
of the inflow to the backwaters Lobau. In times when no management measures are proceeded
water continuously passes over from the UAD to the OAD.
After preparing the phosphorus balance with the same level of detail as the chlo-
ride model, some further input elements had to be considered: bathers, waterbirds,
foliage and baiting by anglers. All internal processes, like uptake by macrophytes,
sedimentation and dissolution or any transformations in the water column could not
be treated separately because of complexities and estimation uncertainties. Numbers
for concentrations in the groundwater and the water body were collected in the
course of various monitoring programs.
Information on the numbers of bathers was given by the Municipality of urban
baths (MA 44) and Hozang (2008). Phosphorus input by swimmers was taken from
Schulz (1981), who estimated a daily input of 94 mg P per person. The stock of
waterbirds is based on counts by Raab (2002) and Dvorak (2003). Mandl and Sampl
(unpubl. 2004) determined an average P-Input per waterbird of 0.15 kg a–1.
Statements to the extent of baiting by fisherman were provided by the Österreichische
Fischereigesellschaft. Fishing regulations for Alte Donau say, that „for baiting max-
imal two hands of predominantly plant products is allowed“, this corresponds about
0.25 kg. Herbal products contain approximately 5 g phosphorus kg–1 (Maly,
unpubl.). The phosphorus input due to foliage could only be roughly estimated
because of contradictory data in the literature (LFU 1996). Because of obstruction
of the banks and treeless bath meadows only 0.5% of the area of Alte Donau is leaf
covered. One square meter foliage is equivalent to 200 g dry mass, which in turn
contains 2.5 mg phosphorus, leaching up to 5%. Finally this process occurs mainly
in October and November.
Groundwater inflow and the intro from Neue Donau during water exchange
dominate the chloride input in Alte Donau (Fig. 6.5). In contrast the atmospheric
entry is negligible (<1%). On these paths the monthly chloride load of UAD is
between 6.6 and 19.8 to. During water exchange the load sharply increases up to 44
to. The chloride input to OAD is determined by groundwater (2/3) and by the pass
over from the UAD (1/3). The monthly mean is in the range of 8.5 to. The monthly
discharge of chloride from UAD varies from 6.6 to 16.3 to. During water exchange
it can rise up to 53.3 to. The water-level-decrease of Alte Donau (discharge of sur-
face water) determines the chloride output in spring. In summer during water
exchange the pass over to OAD dominates the output, whereas in autumn the
groundwater outflow becomes more important. In periods with no water exchange
the monthly discharge of chloride by groundwater outflow from OAD is in the range
of 2.0–5.2 to.
The OAD receives annually 76 kg phosphorus, 40% of it by swimmers during
the period of May to September (Fig. 6.6). Of equal importance is the passover from
UAD with 34%, which is significantly enhanced by water exchange. Considerably
smaller portions come from waterbirds, groundwater inflow and precipitation
(5–8 kg respectively 6–11%). Baiting results only in a low phosphorus load (1 kg or
2%) and the input by foliage, focused only on a few weeks per year, covers 0.1%.
With regard to the volume relation of 1:2 (OAD: UAD), the annual input of 186 kg
phosphorus to UAD is comparable to the load of OAD. But unlike in OAD, in UAD
the input from groundwater represents the major part with 85 kg or 45%. In second
place follows the input by bathers (54 kg or 29%) concentrated on the summer
Fig. 6.5 Comparisons of the montly chloride input and output [to] of the OAD (above) and the
UAD (below) considering the main elements of the balance and complimented by a correction
quantity.
months. Also temporary is the intro from Neue Donau during water exchange, con-
tributing 26 kg (14%) to the phosphorus load of UAD. Bird droppings are respon-
sible for an annual input of 12 kg (7%) phosphorus and the portion of wet deposition
is around 8 kg (4%). By contrast baiting and foliage are negligible (<1.5 kg or 1%)
for UAD. The monthly phosphorus output of UAD by groundwater outflow was
estimated with 0.7 kg. 1.3 kg of phosphorus is discharged to the backwaters of the
Lobau in periods of water level decrease or water exchange. When water exchange
is done (e.g. in the years 2008–2011) the discharge to OAD amounts 19–39 kg P a–1.
Without water exchange measure, as in the season 2012, only 7 kg P a–1 pass over
groundwater
200 8.4 precipitation
discharge/pass over 6.5 foliage
bathers discharge/pass over 0.1
34.0
175 feeding feeding
1.6
waterbirds
waterbirds
foliage 10.7
precipitation
150
groundwater
OAD
125
Phosphorus [kg a-1]
bathers
38.7
100
groundwater
45.4
75
discharge/pass over
14.1
precipitation
50 4.4
foliage
0.1
25 UAD feeding
0.7
waterbirds
0
6.6
OAD UAD
bathers
28.7
Fig 6.6 Annual load (left) and relative share (right) of phosphorus in the OAD and UAD based on
the period 2008–2012.
to OAD. The monthly phosphorus export from OAD by groundwater is in the range

of 0.3 kg. In periods with water exchange the monthly phosphorus discharge from
OAD is between 2 and 16 kg.
In both basins of Alte Donau the modelled phosphorus input exceeds the output
by far (Fig. 6.7).
6.7 Summary
Like in most shallow lakes temperature regime of Alte Donau is strongly related to
air temperatures. Duration of ice covering in winter is usually 4–8 weeks, highest
water temperatures in summer can reach 30 °C. In general Alte Donau is polymictic,
thus oxygen supply of the deeper zones and near the bottom is sufficient throughout
most of the year. According to the geochemical characteristics pH should be slightly
to moderate alkaline (7.6–8.8), but during phases of excessive macrophyte growth,
an enormous increase took place. By flushing with water from the impoundment
Neue Donau pH values could be stabilized under 9.0. The long-term trend of con-
ductivity covers the range of 320–550 μS cm–1, mainly influenced by the dominant
cations Calcium (Ca++) and Magnesium (Mg++) and the main anions HCO3–, Sulfate
(SO43–) and Chloride (Cl–). In accordance with pH, Ca dropped between 2000 and
2006 down to a minimum of 17 mg L–1, due to biogenic decarbonisation and reduced
groundwater fluxes. Stabilisation of Ca++ content on a yearly mean level of
40
Phosphorus in/output [kg]
30
20
10
0
UAD
OAD
-10
J F MAM J J A S ON D J F MAM J J A S ON D J F MAM J J A S ON D J F MAM J J A S ON D J F MAM J J A S ON D
2008 2009 2010 2011 2012
Fig. 6.7 Monthly balance of the weekly phosphorus input and output [kg] of the UAD an OAD by
considering groundwater inflow and outflow, precipitation, intro from Neue Donau, discharge in
the backwaters Lobau via Wasserpark, passover from the OAD in the OAD as well as foliage, bird
droppings, baiting and bathing.
30–35 mg L–1 could be achieved by flushing with water from Neue Donau. Total
nitrogen (TN) was high during the eutrophication and restoration phase, whereas
first the particulate fraction (phytoplankton) dominated and after chemical treat-
ment NO3–. A decrease of nitrate was observed after the recolonisation with macro-
phytes, favouring the growth of green algae and diatoms instead of blue-greens.
Between 1987 and 1994 macrophytes completely vanished and phytoplankton
blooms, dominated by filamentous blue greens progressed culminated in extraordi-
nary high TP values (110 mg L–1). Chlorophyll-a increased (140 μg L–1), accord-
ingly the Secchi transparency dropped rapidly to the the minimum of 0.3 m.
Successful RIPLOX treatment resulted in a remarkable decline of TP, particulate
matter and chl-a and in an increase of transparency. Since 2004, after the switch
back of Alte Donau to macrophyte domination, this trend continues. The latest
period indicates stable mesotrophic conditions with low annual mean concentra-
tions of TP (10–14 μg L–1) and chl-a (3.5–6 μg L–1) and with high transparency
(2–4 m). Based on a phosphorus balance Alte Donau receives annually 260 kg of
phosphorus. In UAD the main source for phosphorus input is the groundwater
inflow and in OAD bathing is the most important source for the load. In both basins
the modelled phosphorus input exceeds the output by far, due to processes like sedi-
mentation and uptake by macrophytes.
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Chapter 7
Sediment
P. Riedler and Karl Donabaum
Abstract Lake sediments play an important role for the nutrient cycling, especially
in shallow, unstratified water bodies. Phosphor release by the sediments is often a
main source for internal loading in eutrophicated shallow lakes. First investigations
of the sediments of Alte Donau were done by Löffler (Alte Donau, Projektstudie im
Auftrag der Wasserstrassendirektion, 272 pp, 1988). More detailed research was
done prior and after the restoration of Alte Donau. The results of the investigations
by Ripl (Sediment treatment. In: Eiseltová M (ed) Restoration of lake ecosystems –
a holistic approach, vol 32. IWRB Publication, Slimbridge, pp 75–81, ISBN
0-9505731-5-9, 1994) were used as the basis to establish the treatment process.
Considerably low iron content of the sediment and a high proportion of organic
content were of importance to use the Riplox method (Ripl W, Sediment treatment.
In: Eiseltová M (ed) Restoration of lake ecosystems – a holistic approach, vol 32.
IWRB Publication, Slimbridge, pp 75–81, ISBN 0-9505731-5-9, 1994) as the main
restoration measure in Alte Donau. The aim of the investigations in 2004 was to
evaluate the sustainability of the restoration measures and to create a data base rec-
ognizing undesirable trends like an enrichment of the organic content in an early
stage. In this chapter the focus is set on the results of the investigations in 2004.
Keywords Sediment · Interstitial water · Phosphor binding capacity · Iron content ·

Apatite
P. Riedler (*)
DWS-Hydro-Ökologie GmbH, Technisches Büro für Ökologie und Landschaftsplanung,
Wien, Austria
e-mail: patricia.riedler@dws-hydro-oekologie.at
K. Donabaum
e-mail: karl.donabaum@dws-hydro-oekologie.at

https://doi.org/10.1007/978-3-319-93270-5_7
70 P. Riedler and K. Donabaum
7.1 Introduction
The Riplox treatment (Ripl 1994; Wolter 1994) was used as the main measure for
restoration of Alte Donau in the years 1995 and 1996 (Chap. 5). In a first step, FeCl3
was added to enhance flocculation and precipitation of phosphorus from the water
column. CaCO3 was used as buffer reagent for the acidic FeCl3 solution, to prevent
a drop down of pH-values. To reduce phosphorus release and internal loading from
the sediments, the amount of iron added was calculated to raise the iron content of
sediments from 8–14 to 28–34 mg g−1. The addition of FeCl3 was followed by the
insertion of Ca(NO3)2 to enhance denitrification for the decomposition of organic
material and to reduce oxygen consumption of the sediments. It was assumed that
the sustainability of positive effects would last for a period of 10 years (Ripl 1997).
After the Riplox treatment and after the implementation of several additional
restoration measures water quality improved significantly from year to year. In 2004
after full recovery of macrophytes the system switched to a clear water state.
However, measurements of bacterial production in the sediments gave evidence for
an increase of the organic content on the sediment surface (Donabaum et al. 2003;
Velimirov et al. 2004). According to this observation, sediment investigations were
done in 2004 to assess the possible risks of an internal loading.
7.2 Methods
Sediment sampling was done with a modified Kajak corer throughout a three days
period in August 2004 (Fig. 7.1 left and center). Sampling sites were selected allow-
ing the comparison with investigations between 1994 and 1996 (Ripl 1997).
Sampling was done on seven sites in the main basin of Obere Alte Donau (OL1-OL7)
and on nine sites in the two basins of Untere Alte Donau (Fig. 7.2). UL1-UL5 were
sampled in the left branch of Untere Alte Donau, four sites (UR1-UR4) in the right
branch of Untere Alte Donau and one additional sample was taken from the so
called Kaiserwasser (KW), which is an appendix of Untere Alte Donau in the south
west. An analysis of the main constitutes and the water content was done in three
depth layers (0–5, 5–10 and 10–15 cm). Analyses of phosphor components and
fractions, loss on ignition (all 0–5 cm) and chlorophyll (0–0.5 cm) was done in the
upper sediment layer.
The water content of the sediment was estimated by weighing the wet sediment,
drying it by 95 °C to constant weight and reweighing. The dry sediment was com-
busted at 450 °C and weighed again to determine ash free dry weight and loss on
ignition. Fractions of the samples were dried at 50 °C and ground with a vibration
mill determining the main constitutes like silicon (Si), titanium (Ti), aluminium
(Al), iron (Fe), manganese (Mn), calcium (Ca), magnesium (Mg), sodium (Na),
potassium (K), phosphor (P) and sulfur (S). Elements were analysed as oxides by
7 Sediment 71
Fig. 7.1 Modified Kajak corer (left), sediment core (center), scheme of a “sedimentturbo” (right)
means of wavelength dispersive X-ray fluorescence technique (RFA – Philipps

PW2400).
Phosphor compounds of the sediment were determined by fractionation accord-
ing to the method of Psenner et al. (1984).
Chlorophyll-a in the upper layer of the sediments was extracted in 90% Aceton
and analysed according to Lorenzen (1967), using the spectral absorption coeffi-
cient of Jeffrey and Humphrey (1975). Additionally, chlorophylls and accessory
pigments were separated by reversed-phase HPLC (Wright et al. 1991), to deter-
mine roughly the proportions of algal classes present (Schagerl and Donabaum
2003; Schagerl et al. 2003; Greisberger and Teubner 2007). Investigations of the
interstitial water were done on the same sites as described above, but in October
2004. Insertion and removal of the sampling equipment for the analysis of the inter-
stitial water (“sedimentturbos”) was done by diving (Fig. 7.1 right). The
“sedimentturbos” were subdivided in 5 separate compartments, which could be
filled with deionised water. After this, the polyethylen bars were covered with a
dialysis membrane. They were placed for 48 h in the sediment. Samples of the inter-
stitial water of three layers (0–5 cm, 5–10 cm und 10–15 cm depth) and of the
sediment-water contact zone were further processed for chemical analyses (SRP,
NH4-N, NO3-N, SO4-S, Cl, DOC).
Fig. 7.2 Map of sediment sampling sites in 2004
7.3 Main Constitutes and Iron-Content
Water content of sediment samples of Alte Donau ranged from 26 to 90%, showing
on most of the sampling sites a decrease with depth. The mean water content of all
samples from the layer 0–5 cm was 79%, the means for the deeper layers were 69%
(5–10 cm) and 67% (10–15 cm). The decrease of the water content with sediment
depth is a consequence of consolidation and compaction (Hupfer 2001).
The proportion of loss on ignition of the sediment dry weight ranged from 3%
to 16% for the surface layer 0–5 cm. The highest percentage of organic material
(16%) was observed in the deepest area of the main basin of Obere Alte Donau
(Station OL2). The mean value for Obere Alte Donau was 10% and for Untere Alte
Donau 11%. 96% of the variability of the organic content could be explained by the
water content of the sediments. In general, the relative low organic content in the
upper layer of the sediment is indicative for moderate primary production in the
7 Sediment 73
20 40
16
loss of ignition [mass-%]
30
iron content [mg g-1]

12
20
10
4
0 0
1994-96 2004 1995/96 2004
Fig. 7.3 Comparison of loss of ignition and iron content in the sediment of Alte Donau in the
years 1994–1996 and 2004
water body and/or for the effective mineralisation (see also benthic invertebrates in
Chap. 14).
There was no considerable difference of organic content compared to the results
of the investigations in 1986 (6–13% n = 4; Löffler 1988) and 1993, prior to restora-
tion (range: 4–22%, mean 10%; Dokulil et al. 1993) as well as to the investigations
in the period from 1994–1996 (mean value Untere Alte Donau – 12%, mean value
Obere Alte Donau 9%; Fig. 7.3 left).
Calcium is the main constitute of the sediment in Alte Donau, the proportion of
CaO is in the range of 29–45 mass%, followed by carbon (organic and inorganic –
loss on ignition at 1000 °C), which comprises 35 mass% of the sediment. Together
with SiO (18%) these three fractions resemble around 90 mass% of the sediment.
The proportion of Fe2O3− and Al2O3− is in the range of 3%.
Typical for Alte Donau is a very uneven distribution of the sediments (Ripl and
Wolter 1995), which is due to the origin of Alte Donau, when in former times high
velocities formed an uneven riverbed with scours.
In 1994, Ripl determined iron concentrations between 8 (Obere Alte Donau) and
14 mg g−1 (Untere Alte Donau) in the upper layer (0–10 cm) of the sediments (Ripl
and Wolter 1995). According to Mackereth (1966) the estimated concentrations are
in the range of the lithospheric mean. Ripl and Wolter (1995) assumed that a big part
of the iron is bound to minerals and therefore not available for P-fixation. To create
a buffer and to enhance the P-binding capacity of the sediment of Alte Donau the
aim was to stock the sediment surface with 20 mg g−1 of iron, which finally should
give iron concentrations in the range of 28 (Obere Alte Donau) to 34 mg g−1 (Untere
Alte Donau). After implementation of Riplox treatment in 1995 and 1996 sediment
analyses (layer 0–10 cm) showed mean iron contents between 14 (Obere Alte
Donau) and 19 mg g−1 (total range 8–37 mg g−1). Investigations revealed big spatial
differences in iron content of the sediment on a large scale as well as on a small

scale, which is possibly due to wind induced pooling. Most of the added iron was
accumulated in the upper 7 cm of the sediment. In Fig. 7.3 (right) the iron contents
measured immediately after the Riplox treatment are compared with the iron con-
tents found in 2004. The sites which were marked by a high iron content after the
treatment in 1995/1996 (>20 mg g−1) showed in 2004 a significant decrease, whereas
sites with moderate iron stocking (10–15 mg g−1) showed almost no changes when
analysed 10 years later. In 2004 the mean iron content in Alte Donau was 12 mg g−1.
For the period between 1978–1995 Ripl and Wolter (1995) assumed a sedimenta-
tion rate of 10 mm a−1. Due to the improvement of the water quality in the period
after the treatment it can be presumed that the accumulation rate of sediment has
decreased and was in the range of 5 to max. 10 mm a−1. It can be concluded that
sediment accumulation between 1995 and 2004 has been in a magnitude of 5–10 cm.
So the highest iron concentration should be found in the upper 10 cm of the sedi-
ment. However, a comparison of the three upper layers (0–5 cm, 5–10 cm, 10–15 cm)
showed no significant differences and might possibly mirroring bioturbation activi-
ties, which would trigger the relocation of the sediment surface.
No significant differences could be found when comparing the mean iron con-
tents of the sediment in Alte Donau prior to, immediately after the Riplox treatment
and then years after. Thus it can be concluded that the main effect of the FeCl3 addi-
tion was given by flocculation and coprecipitation of phytoplankton organisms that
intercept primary production and internal metabolic cycling.
In absolute numbers the stocking of the sediment with Fe was from 8 to 14 mg g−1
in Obere Alte Donau and from 14 to 19 mg g−1 in Untere Alte Donau, which corre-
sponds to an increase of 75 and 36%. The role of Fe as an important P-binding
partner and as a buffer to prevent internal P-loading from the sediment is question-
able. According to the results of sediment P-fractionation calcium seems to be the
most important P-binding partner (see Sect. 7.5).
7.4 Phosphor Components in the Sediment
Phosphor is chemically bound in the sediment in many ways. Only a small propor-
tion of phosphor is directly available for biogenic uptake and other processes like
diffusion into the water-sediment contact zone or reversible bindings triggered by
water-sediment interactions.
The “water soluble fraction” of the sediment phosphor corresponds to a large
extent to the solved phosphorus in the interstitial water (see Sect. 7.5). This fraction
is a good indicator for the internal loading capacity. Concentration of soluble reac-
tive phosphorus (SRP) in the sediment layer 0–5 cm was between 0.2 and 1.6 mg kg−1
dry weight. With a percentage of 0.3% the direct available water soluble fraction
plays a minor role in comparison to the total phosphorus (TP) (423–1270 mg kg−1;
Fig. 7.4). However, the SRP fraction can be seen as a main nutrient source for the
macrophytes. According to P-mass budgets assessed for the near impoundment
7 Sediment 75
Fig. 7.4 Total phosphorus 800

and portion of different
phosphorus fractions in the
sediment of Alte Donau in
2004 600
[µg g-1]
400
refractory P
apatite-P
200 organic P
aluminium-P
reductively soluble P
water soluble P
0
2004
Neue Donau the input of phosphorus from groundwater influx is not sufficient to
explain the standing crop of macrophytes (Wolfram et al. 2001). The authors assume
that macrophytes release phosphor into the sediments during the winter period
where it is chemically bound, whereas in spring the macrophytes seem to utilize
SRP from the interstitial water. Their conclusion is that assimilation of SRP is shift-
ing the dissociation terms, leading to a re-dissolution of bound fractions, which then
are available for the uptake by macrophyte roots.
The P-fraction, which can be extracted from the sediment under reductive condi-
tions resembles phosphor that is bound to iron- or manganese hydroxides. The
amount of reductive soluble phosphorus was in the range of 20–45 mg kg−1. Unlike
the water soluble SRP fraction, the iron- and manganese bound phosphor is not
directly available for metabolism. This fraction is mainly mobilized under anoxic
conditions in the sediment-water contact zone. The estimated proportion of reduc-
tive soluble phosphorus (6%) was found to be typical for meso- and eutrophic con-
ditions. Measurements made prior to implementation of Riplox treatment (Dokulil
et al. 1993) were marked by a high range of fluctuation (0.5 bis 175 mg kg−1), show-
ing a mean value of 43 mg kg−1, corresponding to 3.4% of the total sediment phos-
phorus content. No significant differences could be observed comparing the
investigations prior in 1993 and 10 years after treatment with FeCl3 in 2004. As
mentioned already in Sect. 7.3, the largest part of sediment phosphor is deposed and
fixed as Ca bound apatite-phosphate. Assuming a total release of the reductive
P-fraction from the sediment into the water phase, the calculated P-concentration in
Alte Donau would be 225 μg L−1. The highest TP concentrations ever documented
for Alte Donau were measured in 1993 and 1994, where the TP values were around
100 μg L−1. Internal P-load by the release of a big portion of the iron and manganese
bound fraction could have happened only if large areas of the water-sediment con-
tact zone of Alte Donau would have been anoxic. But this was not the case, neither
in the eutrophication phase in the early 1990ies nor in recent years. Oxygen deficits
near the sediment occured only locally, in the deepest parts of Alte Donau and the
duration of anoxic phases was never exceeding a few weeks. Furthermore Ripl
(1997) supposed, that re-dissolution of phosphor from the sediments might play a
minor role in Alte Donau, because the calcium rich sediments allow the fixation of
phosphorus even under anoxic conditions.
According to Psenner et al. (1984) the third fraction (eluted with cold NaOH)
includes organic P-compounds like sugar phosphates, nucleins, nucleoproteids and
complex bindings like humic acids and fulvic acids. Moreover, this fraction includes
further inorganic phosphor like aluminium phosphate and adsorbed phosphates. The
entire fraction is independent from redox reactions but is activated at high pH-
values (Psenner 1984). It is in exchange with the quote of soluble reactive P in the
interstitial water. The estimated amount of inorganic aluminium bound P was in the
range of 27 and 138 mg kg−1, on average it corresponds to 9% of TP in the sediment.
Because of the high P binding capacities and because of the low solubility the alu-
minium salts are frequently used for P-precipitation. Iron, however, is less poison-
ing than aluminium.
Organic phosphates comprised 23% of the sediment TP (38–442 mg kg−1). There
was a clear linear relationship between organic phosphor and loss on ignition
(r2 = 0.64, p = 0.000).
Apatite phosphate and calcium and magnesium phosphates are determined in the
fourth elution step. In Alte Donau apatite phosphate was the dominant phosphor
compound, comprising 45% of the sediment TP. Estimated concentrations of apatite
phosphor ranged from 213 to 483 mg kg−1 of sediment dry weight. Numerous stud-
ies underline the importance of autochthonous calcite precipitation in the context of
the immobilisation of P in the sediments in calcareous lakes because of low dissolu-
tion capacity and independency of redox reactions (Boström et al. 1988; Istvanovics
et al. 1989; Jäger and Röhrs 1990; Gonsiorczyk et al. 1998; Golterman 2001; Prairie
et al. 2001). From the results of the sediment investigations in Alte Donau it becomes
evident that calcium plays a more important role as a phosphor fixing agent than
iron. Calcium is delivered in remarkable amounts by the ground water influx and it
was stocked in the course of the Riplox treatment as CaCO3 and Ca(NO3)2
(Donabaum and Riedler 2004). Although no data are available concerning the cal-
cium richness of the sediments from the 1980ies or prior Riplox treatment, it is
plausible that the portion of apatite phosphate in the sediment has increased because
of the treatment.
In the last extraction step non-degradable and refractory phosphates are eluted.
These P-compounds play almost no role for re-dissolution and metabolism activi-
ties. There are almost no exchange processes with the interstitial water. A small
portion of this refractory P consists of inorganic P-compounds, the main part of this
fraction is organic P. The portion of refractory phosphates in the sediment of Alte
Donau in relation to sediment TP was in the range of 8 und 22% (mean 99 mg kg−1).
There were no significant differences between the sampling sites.
TP was calculated by summation of the single P-fractions, giving phosphorus
values between 0.4 and 1.3 g kg−1 dry weight. Here, spatial differences could be
observed within the two basins of Alte Donau. These differences were largely
related to the allocation of organic P-compounds and therefore were also correlated
7 Sediment 77
to loss on ignition. There is, however, no substantial difference within the sediment
TP values of 2004 in comparison to the results of the investigations in 1993 (0.7–
2.4 g kg−1) or in comparison to 1995/1996 (around 0.7 g kg−1; see Dokulil et al.
1993; Ripl 1997).
7.5 Interstitial Water
The pore water of the sediment of Alte Donau was first analysed by Ripl and Wolter
(1995) before the implementation of Riplox treatment. Their investigations revealed
SRP concentrations between <0.1 mg L−1 and 1 mg L−1. Measurements were
repeated after the addition periods in 1995/1996. The majority of the analysis
showed low concentrations of SRP in the interstitial water (<0.1 mg L−1; Ripl 1997).
Because of these findings Ripl concluded, that internal P-loading from the sediment
played a minor part in the eutrophication process of Alte Donau, also considering
the fact that the calcium rich sediments of Alte Donau are marked by a high P bind-
ing capacity even under reductive conditions.
Measurements in 2004 showed a substantial decrease of the SRP concentrations
in the interstitial water in comparison to the results of 1995/1996. The mean of all
sites and over depth (water-sediment contact zone, 0–5 cm, 5–10 cm and 10–15 cm)
was about 30 μg L−1. In general, the SRP concentrations increased with sediment
depth (mean of vertical profiles: 9 − > 15 − > 40 − > 73 μg L−1). Highest SRP val-
ues, up to 0.2 mg L−1 were found in the north western part of the main basin in
Obere Alte Donau (Fig. 7.5 left). In this area Alte Donau is fed by nutrient enriched
3.0 20 80
2.5
2.0
1.5
16
60
0.8
NH4-N [mg L-1]
SO4-S [mg L-1]

SRP [µg L-1]
12
0.6 40
8
0.4
20
4
0.2
0.0 0 0
1995/96 2004 1995/96 2004 1995/96 2004
Fig. 7.5 Comparison of SRP, ammonia and sulphate in the pore water of the sediment in Alte
Donau in 1995/1996 and 2004
ground water (see Chap. 4). The incoming ground water of Obere Alte Donau
showed SRP concentrations in the range of 40 μg L−1, whereas the values for Untere
Alte Donau were considerably lower (15 μg L−1; Donabaum and Riedler 2004). The
magnitude of the observed SRP concentrations in the interstitial water of Alte
Donau, which has been found in 2004, is typical for oligo- and mesotrophic condi-
tions (Hupfer 2001).
Nitrogen cycle in the sediment is triggered by deposition and decomposition of
organic material. Under oxygen, the basic process is given by nitrification, where
ammonia is oxidized to nitrite and finally to nitrate, whereas ammonification takes
place under reductive conditions, with ammonia as final product. Ammonia can be
adsorbed to particles or it can be released to the pore water or the water body. The
analyses of NH4-N in the compartments of the sedimentturbos showed an increase
with depth. NH4-N concentrations of 0.3 mg L−1 have been found in the sediment-
water contact zone, showing an increase to 1.7 (0–5 cm) and furthermore to 3 mg L−1
(5–10 und 10–15 cm) in the deeper sediment layers. Absolut numbers for NH4-N
and the observed gradient are in the range which has been found in oligo- to meso-
trophic lakes (Hupfer 2001). The values also correspond to measurements in
groundwater stations along the shore line of Alte Donau (1–15 mg L−1; Donabaum
and Riedler 2004). In general, a decrease of NH4-N concentrations in the pore water
could be observed in comparison to investigations in 1995/1996 (Fig. 7.5 center).
NO3-N typically permeates only the topmost layer (1–2 mm) of the sediment
(Hupfer 2001). No NO3-N could be found in the interstitial water of Alte Donau in
2004. Accordingly the NO3-N concentrations in the water-sediment contact zone
were low, corresponding to the very low nitrate concentrations in the water body
(<0.05 mg L−1; Donabaum et al. 2005).
In anoxic sediment layers sulfate is reduced by dissimilative sulfate reduction to
H2S. Consequently, the SO4-S concentrations show a typical vertical profile in the
sediment. Mean concentrations of 9 mg L−1 were measured in the water-sediment
contact zone, followed by concentrations decreasing from 4 to 2 mg L−1 with
increasing sediment depth (0–5 cm, 5–10 and 10–15 cm). The sulfate concentra-
tions of the open water body (mean value for 2004: 12 mg L−1, Donabaum et al.
2005) were slightly higher than in the water sediment contact zone. The mean value
for sulfate in ground water measuring stations around Alte Donau is given by
11 mg L−1 (Donabaum and Riedler 2004). Formation of H2S and of sulfides like FeS
is important for the immobilisation of heavy metals. Apart from fixation of sulfur as
sulfides, organic sulfur compounds can be formed or deposed in the sediment.
Because of the numerous fixation processes, the sediments play an important role as
a sink for sulfur and sulfate.
In 1995 the sulfate concentrations in the pore water have been 10–15 fold higher
than in 2004 (Fig. 7.5 right). The high sulfate concentrations found in 1995 can be
seen as a result of the Ca(NO3)2 addition to the sediment. An increase of oxygen in
the sediment can activate processes like the formation of sulfate by sulfur bacteria
and chemical oxidation of reduced sulfur components to sulfate.
Chloride concentrations show a similar decrease like sulfate concentrations with
increasing sediment depth. The mean value for Cl in the sediment water contact was
7 Sediment 79
29 mg L−1, followed by a decrease to 21 (0–5 cm) and furthermore to 18 (5–10 cm)

and 15 mg L−1 (10–15 cm) with depth. The Cl concentrations in the open water body
are comparable to the concentrations that have been measured in the water sediment
contact zone (Donabaum et al. 2005). Significant differences were observed in the
two main basins of Alte Donau. In Obere Alte Donau the mean Cl concentrations of
the water sediment contact zone was in the range of 33 mg L−1, whereas in Untere
Alte Donau the mean value for Cl was 26 mg L−1. The Cl concentrations in ground
water measuring stations (Cl mean: 27 mg L−1) were slightly higher than the con-
centrations in the pore water (Donabaum and Riedler 2004).
The DOC concentrations measured in the pore water were in the range of 0.9–
4.7 mg L−1. Highest DOC values were found in the 5–10 cm sediment layer.
Considerable differences were observed between the sampling sites in Alte Donau.
With a mean DOC value of 1.5 mg L−1 the right branch of Untere Alte Donau was
significantly different (two-sample KS-Test, p < 0.000) to the left branch of Untere
Alte Donau (DOC mean: 2.6 mg L−1). Mean DOC values of 2.2 mg L−1 were found
in Obere Alte Donau. The DOC concentrations in the pore water were higher than
in ground water measuring stations (DOC mean: 1 mg L−1; Donabaum and Riedler
2004) but lower than in the open water body of Alte Donau (DOC mean: 3 mg L−1;
Donabaum et al. 2005). The decrease in the interstitial water is due to mineralisation
processes as fixation in the biofilms on sediment (Kreuzinger and Matsché 2000).
The right branch of Untere Alte Donau is marked by a high ground water exchange
rate. The inflowing ground water is oxygenised, which enhances mineralisation pro-
cesses and DOC consumption and overturn (Wolfram and Humpesch 2003). This is
consistent with observed secondary gross production rates in Alte Donau (Velimirov
et al. 2005). Lowest bacterial production rates were detected in Obere Alte Donau
(mean: 4605 μg C L−1 h−1). Untere Alte Donau was marked by generally higher
secondary gross production rates with 6129 μg C L−1 h−1 on average in the left
branch and with 7421 μg C L−1 h−1 highest mean values in the right branch.
7.6 Phytobenthic Biomass and Algal Classes
In 2004 chlorophyll-a on the sediment surface was considerably lower than in

1995/1996, the period immediately after Riplox treatment (mean 1995: 12.9 μg cm−2,
1996: 15.4 μg cm−2, 2004: 1.5 μg cm−2, Fig. 7.6; see also relative high abundance of
benthic invertebrates after Riplox in 1995&1996 in Chap. 14). In 1995/1996 growth
of phytobenthos was enhanced by sufficient light supply which was initiated by the
chemical treatment and the lack of competing macrophytes (Dokulil et al. 1997).
Shading effects by high stands of macrophytes were responsible for the significant
decrease of phytobenthos organisms in 2004. As expected, in any of the investiga-
tion periods, diatoms were dominating the benthic algal biomass with portions
between 59% und 95% of total biomass in 2004 (Fig. 7.7) and with 44% in previous
years 1995/1996 (Dokulil et al. 1997). The higher proportion of diatoms in 2004
40
1995
1996
2004
30
chlorophyll-a [µg cm-2]
20
10
0
O7 UL2 UL5 UR4
Fig. 7.6 Comparsion of chl-a content on the sediment surface of Alte Donau in 1995, 1996 and
2004. Sampling sites are shown in Fig. 7.2
UAD UAD
OAD left branch right branch KW
100
80
algal classes [%]
60
40
Dino
20 Cyano
Chloro
Crypto
Baci
0
UR3
UR4
UR1
UR2
UL3
UL4
UL5
KW
UL1
UL2
O7
O4
O5
O6
O1
O2
O3
Fig. 7.7 Proportion of different algal classes in the benthic community of Alte Donau, HPLC
analysis in 2004. OAD Main basin of Obere Alte Donau, UAD Main basin of Untere Alte Donau –
left and right branch, KW Kaiserwasser
7 Sediment 81
was due to a decrease of the proportions of chlorophytes (26% in 1995/1996) and

blue greens (16% in 1995/1996). The shift in the benthic algal composition might
be due to nutrient reduction and the alteration of the underwater light climate.
7.7 Conclusions
The first investigations of the sediment of Alte Donau by Löffler (1988) comprises
estimations for the particle size, water content, loss on ignition and heavy metals. In
his study especially the values for iron or calcium showed a high scattering and
some of the given numbers seem not plausible. Therefore a comparison of this study
and later investigations is difficult. In 1988 the values for the calcium content of the
sediment ranged from 0.3 to 75.2 mg g−1, whereas Ripl (1997) analysed contents of
about 240 mg g−1 and in 2004 the investigations revealed contents of 90–230 mg g−1.
According to the geochemical atlas (1988) the calcium content for carbonate rocks
is in the range of 300 mg g−1. Values below 10 mg g−1 as given by Löffler (1988) are
unrealistic.
In 1993 in view of designing the restoration measures, Dokulil made an assess-
ment of the status of the open water body of Alte Donau and of the sediments.
Sediment samples were taken from four profiles, analysing a similar parameter set
as Löffler (1988) but without heavy metals. Additionally, Dokulil made experiments
for re-dissolving of phosphorus and analysed soluble sediment P compounds and
nitrogen content (Dokulil et al. 1995).
Several investigations of the sediments were made by Ripl and Wolter in the
course of the combined treatment with FeCl3 and the insertion of Ca(NO3)2. Both
reports contain comprehensive data material regarding sediment- and pore water
analyses during the period from 1994 to 1996 (Ripl and Wolter 1995; Ripl 1997). In
a first step, Riplox treatment was tested in an experimental area in Alte Donau. The
results were used as a basis to calculate the amounts of FeCl3, CaCO3 and Ca(NO3)2
for the whole lake treatment (see Tables 7.1 and 7.2). Nevertheless a mass balance
of the compounds is difficult to establish, because there were some uncertainty
about the amounts that have been finally used for the treatment (Ripl 1997). Another
problem is the heterogeneous spatial dispersion, due to wind induced transportation
Table 7.1 Key numbers for Addition of 39% FeCl3-Solution 1977 t

the Riplox-treatment (Dokulil et al. 1997)
1995/1996 and the added iron
Fe added in total 265 t
Treated sediment area (Ripl and 1,078,000 m2
Wolter 1995, Ripl 1997)
Fe added per m2 sediment 246 g m−2
Volume of wet sediment in 0–5 cm 50,000 cm3 m−2
Dry weight per cm3 of sediment 0.25 g cm−3
Fe added to the sediment per g 20 mg g−1
dry weight
Table 7.2 Fe-balance (Riplox-treatment)

Sources Fe [mg g−1]
Fe-content measured by Löffler (1988) 0.1–7.1
Fe-content measured by Ripl and Wolter (1995) – Obere Alte 8.3
Donau
Fe-content measured by Ripl and Wolter (1995) – Untere Alte 14.3
Donau
Fe-added + 20
Predicted Fe-content (based on numbers by Löffler 1988) = 20–27
Predicted-Fe-content (based on numbers by Ripl and Wolter = 28–34
1995)
Measured Fe-content by Ripl (1997) = 17.2
Measured Fe-content by Donabaum and Riedler (2004) = 6.1–27.5 (MW 13.3)
and difficulties with the addition per boat. Despite these uncertainties, in Tables 7.1
and 7.2 an attempt is made to compare expected and estimated values for iron in the
sediment.
The amount of iron in the sediment has only slightly changed in the period
between the treatment 1995/1996 and 2004. Thus it can be concluded that the P
binding capacity of the sediment is still high enough to prevent from phosphorus
release. The question arises if the importance of iron as P fixing binding partner is
overestimated in case of Alte Donau. From numerous reports in the literature it
becomes evident that iron plays an important role for the P cycle in the sediments
(Boers et al. 1992, 1994; Jäger 1994; Donabaum and Wolfram 2001; Hupfer 2001;
Riedler et al. 2002). The complex chemical exchange processes show a strong
dependency to physico-chemical reactions like the redox potential and to biological
constraints. It is known that mobilisation of considerable amounts of phosphorus is
also possible in iron rich sediments (Roden and Edmonds 1997; Kjellberg-
Christensen 1997; Christensen 1999). It can be concluded that the main function of
FeCl3 is to be seen in its property as flocculation agent, removing plankton organ-
isms and phosphor from the water body, leading to a temporary but severe distur-
bance of primary production activities in the ecosystem. Analyses of phosphor
compounds in the sediment revealed a domination of calcium bound apatite phos-
phate and showed that the iron bound phosphorus compounds play a minor role
(Sect. 7.4). Calcium is thus the dominant constitute for sustainable immobilisation
of phosphor in the sediment of Alte Donau.
As already mentioned data for the calcium content of the sediment given in
Löffler (1988) are too low and questionable. Unfortunately there are also no data
available for the calcium content from the period of 1989–1994. In this context the
data base for the calcium balance given in Table 7.3 and Table 7.4 might be uncer-
tain. Under the assumption, however, that the calcium content at the beginning of
the 1990ies was in the same range as found in 2004 (mean: 180 mg g−1), the stock-
ing with 58 mg g−1 matches the measured value of 1997 with 243 mg g−1.
7 Sediment 83
Table 7.3 Key numbers to CaCO3 added (Ripl and Wolter 1995) 1784.4 t
addition of Ca in the course
Ca(NO3)2 solution (50%) added 503 t
of Riplox-treatment in
1995/1996 Ca added in total 776 t
Treated sediment area (Ripl and Wolter 1,078,000 m2
1995, Ripl 1997)
Ca added per m2 sediment 720 g m−2
Volume of wet sediment in 0–5 cm 5000 cm3 m−2
Dry weight per cm3 of sediment 0.25 g cm−3
Ca added to the sediment per g dry weight 58 mg g−1
Table 7.4 Ca-balance Sources Ca [mg g−1]

Löffler (1988) 0.3–75.2
Ca-added + 58
Predicted Ca-content 1997 = 58–133
Measured Ca-content by Ripl (1997) = 243
Measured Ca-content by Donabaum = 91–235 (MW 181)
and Riedler (2004)
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Part III
Biology, Ecology and Production
Chapter 8
Wax and Wane of Macrophytes
Karin Pall
Abstract From 1993 to 2014 extensive investigations of the macrophyte vegetation

in the Alte Donau were carried out. They included a yearly detailed mapping of water
vegetation as well as periodical surveys of the submerged plant biomass. At the begin-
ning of the investigations, an enormous decrease of the macrophyte vegetation could
be registered compared to previous studies. The breakdown of the macrophyte vege-
tation was caused by the poor water quality at that time and reached its climax in 1995
when the Alte Donau was nearly free from macrophytes. To improve the situation a
set of management measures was realized by order of the city of Vienna. This study
shows the response of the macrophyte vegetation with regards to the development of
the submerged biomass as well as to the observed changes in species composition.
Keywords Aquatic vegetation · Lake restoration · Macrophyte planting · Weed

cutting (=underwater vegetation mowing) · Submerged plant biomass ·
Macrophyte experiments · Underwater floodlight · Plant mass index · Charophyte ·
Myriophyllum spicatum
8.1 Introduction
The Alte Donau in Vienna was a product of the regulation of the Danube River in the
late nineteenth century (Chap. 2). The former highly dynamic tributary developed
from an oxbow to form an inner urban lake without connection to the main river
itself. Since the beginning of the twentieth century, the Alte Donau and the sur-
rounding area were a very popular place for recreation (Chaps. 2 and 19). Along the
shoreline many swimming areas developed and many weekend houses were built.
For a long time, the Alte Donau could preserve its good water quality in spite of
its intensive use as a swimming, boating, sailing and fishing area. Until the middle of
K. Pall (*)
Systema GmbH, Vienna, Austria
e-mail: karin.pall@systema.at

https://doi.org/10.1007/978-3-319-93270-5_8
90 K. Pall
the 1980s, the Alte Donau was characterised by widespread stands of submerged
macrophytes and thus low phytoplankton densities as well as high water transparency
(Chaps. 6 and 9). In the following 10 years, a strong deterioration of the water quality
took place and resulted in a nearly totally breakdown of the macrophyte vegetation.
For this reason, the city authorities of Vienna immediately started extensive
investigations of the water body. The results led to a series of management mea-
sures, which were carried out within the last 20 years (Chap. 5), accompanied by a
limnological and an ecological monitoring program (Chaps. 6, 7, 8, 9, 10, 11, 12,
13, 14, 15 and 16), including a comprehensive surveillance of the development of
the macrophyte vegetation (this chapter).
In addition to the results of macrophyte investigations described in this chapter,
the methods for planting of reed and marsh plants (helophyte vegetation), that were
carried out to enlarge shore zones covered by reed communities from 1984 to 1994,
are explained in Chap. 18.
8.2 Methods
The periodical investigations of the macrophyte vegetation began in 1993 and con-
sisted of two parts. On the one hand, the development of the aquatic plant biomass,
i.e. from year to year, was recorded. On the other hand, a yearly mapping of the
species spectrum and the abundances of the single species took place. In order to
detect the driving factors for plant growth in the Alte Donau, additionally lake
experiments with underwater floodlight were performed.
8.2.1 Determination of Plant Biomass
The recording of the plant biomass took place at least once a year. It was carried out
by a combination of echo-sounding and scuba diving as described in Janauer et al.
(1989). As a first step, the submerged plant stands were recorded by echo-sounding
along transects, situated every 100 m perpendicular to the shoreline. Using these
data a model of the three-dimensional extension of the submerged vegetation could
be derived. In a second step, four samples (25 cm × 25 cm) of plant biomass from
all transects with remarkable macrophyte growth were collected by scuba divers.
The plant material was carefully washed and dried at 93 °C. The extrapolation of the
measured dry weight to the three-dimensional vegetation model allowed the estima-
tion of the plant biomass of the entire water body.
8.2.2 Mapping Procedure
Every summer (July and August), the entire macrophyte species spectrum of the
Alte Donau was mapped. The shoreline was divided into 100 m long sections.
Mapping was done by boat with the help of an aqua-scope and a rake in the
8 Wax and Wane of Macrophytes 91
combination with random sampling by scuba diving. The abundance of each species
was estimated according to a five-level scale in every 100 m long section, reaching
from the shoreline to the middle of the water body. Following Kohler (1978) the
different levels of the “Plant Mass Index” (PMI) can be interpreted as follows:
1 = very rare, 2 = rare, 3 = common, 4 = abundant, 5 = very abundant, in masses.
8.2.3 M
acrophyte Lake Experiment with underwater
Floodlight
In order to test the interrelationship of light conditions and plant growth, some
underwater floodlights were installed in the Alte Donau. They were adjusted at dif-
ferent depths between 2.5 and 3.5 m, in each case 1.5 m above the ground. Spectral
range and intensity of the light were chosen in such a way, that the light climate
under the floodlights was very similar to the natural conditions in 1.5 m water depth
of the Alte Donau.
The floodlights were switched on at 6 o’clock in the morning and switched off at
6 p.m. Just under the floodlights a circular area with a diameter of 1 m was planted
densely with about 10 cm high plants of Myriophyllum spicatum. As reference in
the same depth, equal areas were planted with the same plant material, but not lit.
Additionally some areas were lit without any plantings below.
The experiment took place in a time-span of 2 years, from June to September in
1999 and from May to September in 2000. The development of the planted and the
spontaneous growing vegetation was investigated by scuba divers every 2–4 weeks.
The monitoring each time included measuring the plant height and estimating the
plant cover in the experimental lake areas.
8.2.4 Further Evaluations
The Relative Plant Mass (RPM) after Pall and Janauer (1995) was used to analyse
the quantitative composition of the macrophyte vegetation and the dominance struc-
ture of the different species. The RPM-value of a species gives the percentage of the
plant mass of this species of the entire plant mass of all occurring species in a cer-
tain water body. The calculation of RPM refers to the following formula:
Formula 8.1 Calculation of the Relative Plant Mass (RPM) after Pall and Janauer
(1995)
∑(M 3
i )
⋅ Li ⋅ 100
RPM [%] = i =1
k
 n 
∑ ( )
 ∑ M ji ⋅ Li 
3
j =1  i =1 

92 K. Pall
RPM = relative plant mass of a species

Mi = for a section i estimated plant mass of this species
Li = length of section i
j = current index of different species
The Mean Mass Index (MMT) after Janauer et al. (1993) describes the total plant
mass of a species in the investigated area. It is a length-weighed mean of the esti-
mated plant masses (PMIs) of all investigated sections. MMT is calculated accord-
ing to following formula:
Formula 8.2 Calculation of the Mean Mass Index (MMT) after Janauer et al.
(1993)
3
∑M 3
i ⋅ Li
MMT = i =1
L
MMT = mean mass index of a species over all investigated sections
Mi = plant mass of a species in section i
Li = length of section i
L = total length of investigated area sections
8.3 Results and Discussion
8.3.1 Macrophyte Biomass
Figure 8.1 shows the development of the macrophyte biomass from the beginning of
our investigation in the year 1993 until 2001. Additionally, the corresponding value
from the year 1987 (Löffler 1988) is given. According to Löffler (1988) the Alte
Donau was in a mesotrophic state at that time. The whole bottom was covered with
dense macrophyte stands, dominated by charophytes, and their overall biomass was
about 721 t dry mass. In the following years, the macrophyte biomass decreased dra-
matically. In 1993 we recorded only 34 t and in 1994 only 2 t dry mass. By 1995 the
macrophytes had nearly totally disappeared from the lake (less than 0.5 t dry mass).
During 1995 and 1996 for the first time massive external and internal lake man-
agement measures were taken (Chap. 5). The application of the RIPLOX-treatment
(Fig. 5.3 in Chap. 5) caused an enormous increase of the water transparency due to
the removal of phosphorus from the water body (Fig. 6.1 in Chap. 6). The sub-
merged macrophyte vegetation thus started to grow again (see short-term response
of phytoplankton production and the composition of phytoplankton and zooplank-
ton during this period in the Chaps. 9, 10, 11 and 12). In 1996 the macrophyte bio-
mass was 26 times higher than in 1995. A further increase of about 20% followed
until 1997. In this year 10 t dry weight of macrophyte biomass were recorded. In the
721
Biomass
Macrophyte dry weight [t]
40
30
R R
20
10
0
1987 1993 1994 1995 1996 1997 1998 1999 2000 2001
Year
Fig. 8.1 Development of the macrophyte biomass from 1993 to 2001 (summer surveys) in com-
parison with 1987 (Löffler 1988). All values in t dry weight (dry mass). R: years with chemical
phosphate precipitation by RIPLOX-treatment
following years the increasing rates became lower and showed a decreasing ten-
dency due to the lack of temporarily clear water conditions. Until 2001 the biomass
of the submerged vegetation in the Alte Donau had increased only to 15 t dry mass.
It seemed that larger increasing rates of macrophyte biomass could be found only
in case of periods with high water transparency in the previous year. The strong cor-
relation between availability of light and macrophyte growth could also be derived
from the spatial distribution of the macrophyte vegetation in the Alte Donau.
Figure 8.2 shows the vegetated areas in the year 1987 (Löffler 1988) and 2001
(results of echo sounding).
While in 1987 nearly the whole lake bottom was vegetated, by 2001 some mac-
rophytes could be found only in a narrow belt along the shoreline. The submerged
vegetation ended in a depth of about 2.1 m. This value corresponded well with the
theoretical depth limit of macrophyte growth in the Alte Donau, which could be
derived from the average Secchi depth (method see Chambers and Kalff 1985).
In order to verify further this relationship for the Alte Donau and to make it
usable for management purposes, experiments with underwater floodlights were
carried out. The floodlights were installed in different water depths. It was thus pos-
sible, to have light conditions comparable with the natural situation in 1.5 m water
depth, even in the non-vegetated zones between 2.1 and 3.5 m. Figures 8.3 and 8.4
show the development of planted stands of Myriophyllum spicatum in a water depth
of 3 m with and without lighting.
In the first year the lit site shows, after a short reduction of the cover in the middle
of July, a high vegetation density until the end of September (site: area 1, Fig. 8.3).
The plant height increased rapidly from 10 cm after planting during the summer and
reached its maximum (150 cm) in late August. In contrast, the plant cover decreased
consecutively in the non-lit site (site: area 2, Fig. 8.4) and was only 30% by the end
of September. The remaining plants grew very slowly and the maximum height,
94 K. Pall
Fig. 8.2 Vegetated area in the years 1987 (left picture, data from Löffler 1988) and 2001 (right
picture). 1987 dark grey: stands of Myriophyllum spp. and Potamogeton spp., light grey: stone-
worts (mainly Nitellopsis obtusa). 2001 grey: nearly exclusively Myriophyllum spicatum
Area 1
Plant stands under floodlight in a depth of 3 m
200 100
mean plant height [cm]
180 90
160 80
140 70
cover [%]
120 60
100 50
80 40
60 30
40 20
20 10
0 0
plant hight plant cover
Fig. 8.3 Development of planted stands of Myriophyllum spicatum under floodlights at a water
depth of 3 m (macrophyte experiment in 1999)
reached in late August, was only about 20 cm. The non-planted but lit site showed
no plant growth at all in the first year (site: area 3, not shown in a figure).
During winter the plants degraded at all sites. In the second year, the plants in area
1 started in May with a cover of 20% and a plant height of about 10 cm. The maximum
cover reached in this year was 50% and the maximum plant height (middle of August)
Area 2
Plant stands without floodlight in a depth of 3 m
200 100
180 90
160 80
140 70
cover [%]
120 60
100 50
80 40
60 30
40 20
20 10
0 0
plant hight plant cover
Fig. 8.4 Development of planted stands of Myriophyllum spicatum without floodlights at a water
depth of 3 m (macrophyte experiment in 1999)
Area 1
Development of the 1999 planted plant stands
200 100
180 90
160 80
140 70
cover [%]
120 60
100 50
80 40
60 30
40 20
20 10
0 0
mean plant height plant cover
Fig. 8.5 Development of planted stands of Myriophyllum spicatum under floodlights at water
depth of 3 m in the second year of macrophyte experiments (2000)
was 120 cm (Fig. 8.5). In site 2, no plants could be found in the second year (site: area
2, without figure). However, surprisingly in site 3 (no planting, but lighting) some
plants (Myriophyllum spicatum) spontaneously started to grow (site: area 3, Fig. 8.6).
By the end of September, they reached a cover of 30% and a plant height of 25 cm.
96 K. Pall
Area 3
Sponaneous plant growing in the second year
200 100
180 90
160 80
140 70
cover [%]
120 60
100 50
80 40
60 30
40 20
20 10
0 0
mean plant height plant cover
Fig. 8.6 Spontaneous plant growth at a water depth of 3 m in a non-planted but lit area in the
second year of macrophyte experiments (2000)
These results provided evidence for the planning of a temporally limited water-
level-lowering scheme. By lowering the water level, light can reach deeper zones of
the lake. In the case of the Alte Donau with an average depth of 2.3 m (morphometry
see Chap. 3) and a current vegetation limit of 2.1 m, even a little lowering of the
water level would result in large additional areas for potential plant growth. In 2002
a first test was initiated. Since 2003, the water level of the Alte Donau has been
artificially lowered yearly from the ice melt until the end of April. A detailed
description of the long-term lake management strategy including measures for re-
establishing macrophytes is given in Table 5.1 in Chap. 5.
Figure 8.7 shows the response of the macrophyte biomass to temporal water level
lowering. In 2002, the waxing rate (a plus of 33%) of the macrophyte biomass
showed an increasing tendency for the first time since 1996. From 2002 to 2003, the
plant biomass grew 6-fold up to 125 t dry mass. In the following year with 370 t a
3-fold wax was recorded. Since then the wax of the macrophyte biomass slowed
down. From 2004 to 2005, the growing rate was only 4%, which finally led to 385 t
dry mass.
The reduced waxing rates in the latter years were partly the result of an intensive
weed cutting operation in the Alte Donau, due to the enormous increase of
Myriophyllum spicatum. This high-growing species impeded recreational activities
in a wide area. This is the reason why future management measures will have to aim
at a change of the dominance structure of the macrophyte vegetation, supporting the
low-growing macrophytes at the expense of high-growing species (see also Chap.
20). The development of the species spectrum and the quantitative composition of
the macrophyte vegetation are described in the following sections.
721
Biomass
400
Macrophyte dry weight [t]
350
300
250
200
150
100
50
0
87 93 94 95 96 97 98 99 00 01 02 03 04 05
Year
Fig. 8.7 Development of the macrophyte biomass following the temporal water level lowering.
2001: test phase, 2002–2005: normal operation. L: Water level lowering
8.3.2 Species Spectrum
Table 8.1 provides a list of all species found from 1993 to 2005. Additionally,
the species mentioned by Löffler (1988) are listed. In the second column,
abbreviations used later on are given. All in all, there are 39 different macro-
phyte species (only hydrophytes): 32 of them occur submerged and seven build
the floating-leaf-vegetation. The occurrence of 11 different charophytes is
remarkable. Some important macrophyte species in Alte Donau are shown in
Fig. 8.8.
Table 8.2 shows the annual occurrence and frequency of the different species in
the Alte Donau. Due to other sampling methods no frequency data for 1987 and
1993 are available. For this reason, only the occurrence is given. It is obvious that
from 1987 to 1993 many species disappeared from the Alte Donau, of which some,
as most charophytes, occurred again later, but others, as Chara vulgaris, Elodea
canadensis, Hippuris vulgaris and Myriophyllum verticillatum, disappeared from
the lake.
8.3.3 Dominance-Structure of the Vegetation
Figure 8.9 shows the dominance-structure of the macrophyte vegetation in the Alte
Donau from 1993 to 2005. There are the RPM-values of four different plant groups,
i.e. the charophytes, the low-growing spermatophytes, the high-growing spermato-
phytes and the floating-leaf-plants. For each group separately the RPM-value of the
dominant species is given.
98 K. Pall
Table 8.1 Species spectrum Species Abbr.

and list of abbreviations
Ceratophyllum demersum L. Cer dem
Chara aspera Detharding ex Cha asp
Willdenow
Chara contraria A. Braun ex Kützing Cha con
Chara delicatula Agardh Cha del
Chara globularis Thuillier Cha glo
Chara hispida L. Cha his
Chara intermedia A. Braun Cha int
Chara tomentosa L. Cha tom
Chara vulgaris L. Cha vul
Eleocharis acicularis var. fluitans Döll Ele aci
Elodea canadensis Michaux fil. Elo can
Elodea nuttallii (Planchon) St. John Elo nut
Hippuris vulgaris L. Hip vul
Lemna minor L. Lem min
Myriophyllum spicatum L. Myr spi
Myriophyllum verticillatum L. Myr ver
Najas marina L. Naj mar
Najas minor Allioni Naj min
Nitella mucronata (A. Braun) Miquel Nit muc
Nitellopsis obtusa (Desvaux in Nit obt
Loiseleur) J. Groves
Nuphar lutea (L.) J.E. Smith Nup lut
Nymphaea alba L. Nym alb
Nymphoides peltata (S.G. Gmelin) Nym pel
O. Kuntze
Polygonum amphibium L. Pol amp
Potamogeton x cooperi (Fryer) Fryer Pot coo
Potamogeton crispus L. Pot cri
Potamogeton friesii Ruprecht Pot fri
Potamogeton lucens L. Pot luc
Potamogeton nodosus Poiret Pot nod
Potamogeton pectinatus L. Pot pec
Potamogeton perfoliatus L. Pot per
Potamogeton pusillus L. sec. Dandy et Pot pus
Taylor
Ranunculus circinatus Sibthorp Ran cir
Stratiotes aloides L. Str alo
Tolypella intricata R. D. Wood Tol int
Utricularia vulgaris L. Utr vul
Veronica anagallis-aquatica L. Ver ana
Veronica catenata Pennell Ver cat
Zannichellia palustris L. Zan pal
Fig. 8.8 Important macrophyte species in Alte Donau. 1: Nymphoides peltata, 2: Trapa natans, 3:
Nymphaea alba, 4: Myriophyllum verticillatum, 5: Myriophyllum spicatum, 6: Potamogeton pecti-
natus, 7: Chara aspera, 8: Najas marina, 9: Chara tomentosa
100 K. Pall
Table 8.2 Frequency of the different species in the Alte Donau: x = general occurrence, =
occurrence in one 100 m-section, = 2 to 5 sections, = 6 to 25sections, = 26 to 50
sections, = 51 to 100 sections and = more than 100 of a total of 298 sections; Abbreviations
of species names in Table 8.1.
Year 87 93 94 95 96 97 98 99 00 01 02 03 04 05
Cha asp x
Su b merg ed sp ecies, Ch aro p h y tes
Cha con x
Cha del
Cha glo x x
Cha his x
Cha int x
Cha tom x x
Cha vul x
Nit muc
Nit obt x x
Tol int
Cer dem x x
Ele aci
Elo can x
Elo nut
Hip vul x
Myr spi x x
Myr ver x
Naj mar x x
Naj min
Pot coo
Su b merg ed sp ecies, Sp eramto p h y ts
Pot cri x
Pot fri
Pot luc
Pot pec x x
Pot per
Pot pus
Ran cir
Utr vul x
Ver ana
Ver cat
Zan pal
Lem min
Flo atin g -leaf-p lan ts
Nup lut
Nym alb x x
Nym pel
Pol amp
Pot nod
Str alo x
. Number 19 8 16 14 20 22 23 25 25 29 25 31 34 29
Nitellopsis obtusa other charophytes

Najas marina other low-growing spermatophytes
Myriophyllum spicatum other high-growing spermatophytes
Nymphaea alba other floating-leaf-plants
100
90
80
70
60
50
40
30
20
10
0
1993 1994 1995 1996 1997 1998 1999 2000 2001 2002 2003 2004 2005
Fig. 8.9 Relative Plant Mass (RPM in %) of different plant groups and their dominating species
in the Alte Donau from 1993 to 2005
At the beginning of the monitoring program in 1993 about 90% of the entire
macrophyte plant mass was represented by Nitellopsis obtusa. The remaining 10%
were shared by Myriophyllum spicatum and Nymphaea alba. Other species did not
play a remarkable role. Until 1995, Nitellopsis obtusa and other charophytes disap-
peared completely, while Myriophyllum spicatum became the dominant species.
Najas marina and Nymphaea alba had a remarkable part of the entire plant biomass,
too. In the following years, Myriophyllum spicatum eventually became the overall
dominating species, while the proportion of all other spermatophytes declined. The
only plant group, which could, slowly but continually, enlarge its proportion were
the charophytes – interestingly enough not only Nitellopsis obtusa but more impor-
tantly other charophyte species.
8.3.4 Absolute Plant Mass
The RPM gives information about the proportions and dominances structures
between different macrophyte groups or species. As it is a relative dimension it
has no relevance for describing the real plant masses. In order to show the devel-
opment of the absolute plant mass of the different plant groups examined, the
MMT was calculated. Figure 8.10 shows the development of MMT from 1993
to 2005.
102 K. Pall
MMT of different plant groups of the submerged and floating-leaf

vegetation in the Alte Donau from 1993 to 2005
1993
Charophytes 1994
1995
1996
1997
1998
1999
2001
other small plants 2002
2003
2004
2005
high-growing plants
floating-leaf-plants
00 1 2 3 4 5
MMT
Fig. 8.10 Absolute Plant Mass (MMT) of different plant groups in the Alte Donau from
1993 to 2005
In 1993, charophytes built the highest absolute plant mass. They reached an
MMT-value of 3, which corresponds to abundance class “common”. All other plant
groups were only “rare”. The macrophyte breakdown reached its climax in 1995.
Only the floating-leaf plants survived. In the following years the absolute plant mass
of the submerged macrophytes increased again, especially after improving the light
availability by management measures. The high-growing plants started to grow first.
In 2005 they reached an MMT-value between 4 and 5, which corresponds to “abun-
dant” to “very abundant, in masses”. The mass increase of the charophytes only
started later, but until 2005 they did not equal the status at the beginning of the
investigations.
8.3.5 F
urther Development of the Macrophyte Vegetation
in the Following 10 Years
As regards to the fact that the Alte Donau is an important place for recreational
activities, it is essential, which plant group dominates the vegetation. With regard to
this a quite high proportion of charophytes and other low-growing species would be
ideal. Such low growing vegetation (especially charophytes) can preserve a good
water quality (low nutrient concentrations and high transparency) in the Alte Donau
equally well as the currently dominating high-growing vegetation.
Intensive weed cutting became necessary in 2003 due to the masses of high-
growing plants, especially Myriophyllum spicatum. On the one hand, the aim of
weed cutting was to ensure the usability of the Alte Donau for e.g. swimming and
boating, whereas on the other hand, conditions for favouring the low-growing spe-
cies should be established by means of a well-organized weed-cutting plan.
After successfully establishing dense macrophyte vegetation, mainly by means
of water level lowering, the focus now turned on optimising the species composi-
tion. Even with regard to this, the periodical water level lowering provides some
benefits: The inflowing ground water during the phases of lowered water levels has
especially positive effects on the charophyte vegetation. Furthermore, in general the
low-growing vegetation can benefit from the high water transparency during the
process of water level lowering.
From 2005 onwards, the yearly measured biomasses were around 350 t dry mass
(Fig. 8.11). Only 2014 a bigger increase of biomass, up to nearly 500 t dry mass,
could be registered. This resulted from the resettlement of macrophytes in an up to
that time not vegetated part of the Alte Donau. From this we can conclude that with
the recapturing of the whole potential growth areas in the Alte Donau by macro-
phytes a biomass-value similar to that measured in 1987 will be reached again.
Since 2004 (27 submerged species) from year to year a decreasing trend in the
number of submerges species could be registered. The minimum with only 14 spe-
cies was reached in 2012. In order to enrich the species spectrum and to strengthen
the resilience of the macrophyte vegetation, extensive planting measures with
submerged – preferably low-growing species – were set. The plant material used
consisted exclusively of species which could be found in the Alte Donau in former
years. The planting measures, carried out by divers, showed satisfactory results as
most of the introduced species could establish considerable new plant stands. In
2014 the number of submerged species which could be registered during the routine
mapping reached 24 again.
In 2005 the overall dominant plant group in the Alte Donau were high-growing
species (predominantly Myriophyllum spicatum, see Fig. 8.8). In spite of intensive
efforts in planting low-growing species, especially charophytes, until 2014 the dom-
inance structure of the macrophyte vegetation in the Alte Donau could be changed
only to a small extent. Still the group of high growing species contributes about 90%
to the overall plant mass of the submerged vegetation.
104 K. Pall
800
Biomass
Macrophyte dry weight [t] 700
600
500
400
300
200
100
0
1987 2005 2006 2007 2008 2009 2010 2011 2012 2013 2014
Year
Fig. 8.11 Development of the macrophyte biomass from 2005 to 2014
Especially the re-establishing of charophytes turned out as to be a great chal-

lenge. As well-documented by a purpose-built automatic underwater video-trap fish
hindered effectively the growth of charophytes by grazing (amur [Ctenopharyngodon
idella], rudd [Scardinius erythrophtalmus]) or digging out (bream [Abramis brama],
carp [Cyprinus carpio]) the new planted stoneworts.
8.4 Conclusions
The measures set in the Alte Donau, especially the temporally limited water-level-
lowering, led – after the nearly completely break-down in the 1990th – to an enor-
mous wax of the macrophyte vegetation. The aim of re-establishing dense
macrophyte stands could be attained within only a few years. However, one problem
remained, the former species-rich system turned to a system dominated nearly
exclusively by one species – Myriophyllum spicatum.
The further challenge will be to achieve a change in the species composition,
steering away from the “monoculture” of high-growing Myriophyllum spicatum and
leading towards a predominantly ground covering vegetation (preferably dominated
by type-specific charophytes). First steps with low-growing spermatophytes could
effectively enrich the species spectrum but, until now, not really change the domi-
nance structure of the vegetation.
In order to achieve this aim on the one hand the temporarily limited water level
lowering should be continued as it provides positive effects especially on charo-
phytes. On the other hand, as well the weed-cutting as the fish-stock would have to
be optimised.
Since 2017 in wider areas of the Alte Donau a temporarily removal of non-
predatory fish and – in return – a stocking of predatory fish took place. From these
areas Myriophyllum spicatum was removed and, instead of it, charophytes planted.
First results show a great potential of this approach with regard to re-establishing a
type-specific charophyte-vegetation, as it existed in the Alte Donau in former times.
Acknowledgments This study was funded by the City of Vienna, Municipal Department 45
(Water Management). We would like to thank for permission of publication. Thanks furthermore
to the team of the systema company, namely Veronika and Stefan Mayerhofer, Gregor Hoheneder,
Irene Teubner and Sascha Pall.
References
Chambers PA, Kalff J (1985) Depth distribution and biomass of submersed aquatic macrophyte
communities in relation to Secci depth. Can J Fish Aquat Sci 42:701–709
Janauer GA, Wychera U, Dirry P (1989) Biomasse und Phosphormanagement von
Wasserpflanzen im Jahresverlauf. Grundlagen für ein Phosphormanagement in donaunahen
Oberflächengewässern, 150 pp
Janauer GA, Zoufal R, Christoph-Dirry P, Englmaier P (1993) Neue Aspekte der Charakterisierung
und vergleichenden Beurteilung der Gewässervegetation. Ber Inst Landschafts-
Pflanzenökologie Univ Hohenheim 2:59–70
Kohler A (1978) Methoden der Kartierung von Flora und Vegetation von Süßwasserbiotopen.
Landschaft + Stadt 10(2):73–85
Löffler H (1988) Alte Donau. Limnologische Projektstudie – Ökosystem Alte Donau. Endbericht,
72pp
Pall K, Janauer GA (1995) Die Makrophytenvegetation von Flußstauen am Beispiel der Donau
zwischen Fluß-km 2552,0 und 2511,8 in der Bundesrepublik Deutschland. Archiv für
Hydrobiologie, Supplement 101, Large Rivers 9/2:91–109
Chapter 9
Phytoplankton in Alte Donau: Response
to Trophic Change from Hypertrophic
to Mesotrophic Over 22 Years
Katrin Teubner , Wilfried Kabas, and Irene E. Teubner
Abstract The long-term phytoplankton study in groundwater-seepage lake Alte

Donau, a former side-arm of the Danube River in Vienna, covers four main lake
treatment periods (1–4) from 1993 to 2014. During hypertrophic conditions with
annual total phosphorus (TP) concentrations of 50–70 μg L−1 and mean summer
phytoplankton biovolume of 18–24 mm3 L−1 before restoration (1), the filamentous
cyanobacterium Cylindrospermopsis raciborskii was the main taxon in association
with Limnothrix redekei. The drastic phosphorus reduction by chemical RIPLOX-
precipitation was repeated twice (2a/b, 1995 and 1996) and resulted in a prompt
drop of summer phytoplankton to 4.6 mm3 L−1 in 1995 and 1.7 mm3 L−1 in 1996.
Non-filamentous cyanobacteria contributed here only moderately while relative
high peak contributions of chlorophytes occurred. After years of re-establishment of
macrophytes (3), the summer phytoplankton biovolume remained low during the
period of sustained ‘stable conditions’ (4) with values between 0.5 and 1.5 mm3 L−1.
In the long-term, phytoplankton was responding to low annual total phosphorus
(10–11 μg L−1) which finally indicated a mesotrophic state close to oligotrophic
conditions according to the lake classification scheme. The long-term median of
chlorophyll-a (chl-a) content was 0.50% of wet weight phytoplankton biomass. As
the phytoplankton composition shifted from a cyanobacteria dominated assemblage
to a phytoplankton assemblage that was composed of taxa of various taxonomic
affiliations, the chl-a content varied considerably. Chl-a content reached its lowest
median value of 0.19% when cyanobacteria formed blooms contributing 77% to
K. Teubner (*)
Dept. of Limnology & Bio-Oceanography, University of Vienna, Wien, Austria
e-mail: katrin.teubner@univie.ac.at
W. Kabas
Vienna, Austria
I. E. Teubner
Department of Geodesy and Geoinformation, Faculty of Mathematics and Geoinformation,
Vienna University of Technology, Vienna, Austria
e-mail: irene.teubner@geo.tuwien.ac.at

https://doi.org/10.1007/978-3-319-93270-5_9
108 K. Teubner et al.
total phytoplankton (period 1) and was highest with 0.83% during the peak develop-
ment of chlorophytes which contributed 18% to total biovolume (period 2b). The
relationship between phytoplankton chl-a and TP is more robust than between phy-
toplankton biovolume and TP for indicating the lake’s trophic state, although both
response curves are statistically significant and provide roughly the same main pic-
ture of an ecosystem shift from hypertrophic in 1993 to mesotrophic in 2000 and the
persistence of mesotrophic conditions for the 15 recent years. Trophic shifts were
also indicated by the phytoplankton assemblage metric when comparing phyto-
plankton species composition between the lake treatment periods. The main picture
of seasonal development of phytoplankton taxa and functional phytoplankton
groups indicated that assemblages either prevailed in winter to spring or summer to
autumn. Annual phytoplankton development thus seems primarily distinctive
between the two half-year-cycles, namely the winter-spring and the summer-autumn
period, rather than between the four seasons. While the seasonal development of
phytoplankton follows the lake phenology commonly observed in temperate lakes,
long-term compositional shifts of phytoplankton especially responded to the sus-
tained reduction of TP forced by lake treatment measures in Alte Donau.
Keywords Oxbow lake · Lake restoration · Lake recover · Lake biomanipulation ·

Riplox · Algae · Cyanobacteria · Cylindrospermopsis raciborskii · Seasonality ·
Trophic classification · Phosphorus · Chlorophyll-a · Chlorophyll:TP ·
Biovolume:TP · Phytoplankton assemblage metric
9.1 Introduction
Rising awareness of eutrophication and advances in the management of eutrophied

aquatic systems have been made since the late 1960s. The OECD study (OECD
1982) in the 1980s and the initiative by the EU Water Framework Directive (2000)
in the 2000s provided the two milestones in the scientific understanding of the
mechanisms of eutrophication and consequently launched the search for aquatic
biota, which indicate the specific reference status of each lake type. Since massive
phytoplankton growth corresponds most notably to nutrient enrichment in lakes and
as these primary producers play a key role in the food chain, phytoplankton is a key
biotic parameter assessing lakes. Among the biotic parameters monitored in the
oxbow lake Alte Donau, phytoplankton (this chapter) and zooplankton (Chap. 9)
provide with 22 and 19 years, respectively, the longest records.
Alte Donau is popular for recreation including swimming and fishing (Teubner
et al. 2015). During the 22-year lake restoration, fish were only removed by angler
sport. The number of fish catches and fish stocking, however, decreased with years
of phosphorus reduction towards a lower trophic level (time series of fish see Fig.
15.1 in Chap. 15, fish biomass versus chl-a in Fig. 20.3 in Chap. 20). During all
years, both mainly carp and predatory fish were added. In the long term, the local
9 Phytoplankton in Alte Donau: Response to Trophic Change from Hypertrophic… 109
fishery increasingly forced the stocking of predatory fish. Some aspects of both
bottom-up and top-down control on plankton development are discussed in view of
main compositional shifts from a cladoceran-rotifer-rich to a copepod-rotifer-rich
of zooplankton assemblage along the lake management periods in Chap. 11.
Phytoplankton development in this chapter focuses on the bottom-up control with
main emphasis on phosphorus. This main nutrient element is commonly known to
limit phytoplankton growth in freshwaters. Accordingly, the main target of restora-
tion and lake management in Alte Donau was to reduce phosphorus availability for
phytoplankton growth and aimed at increasing water transparency and thus to
enhance water quality. The emphasis of phosphorus-resource control on phyto-
plankton is the cause for plotting the trophy classification scheme applying Austrian
standards ÖNORM M6231 (2001) or using a phytoplankton metric based on an
Austrian Trophic State Index (see Sect. 9.2.4) in this chapter. The phosphorus-
phytoplankton response also remains relevant when applying functional phyto-
plankton groups (Reynolds et al. 2002; Padisák et al. 2009), as was done for Alte
Donau, since these groups describe the co-occurrence of algae not only along sea-
sons but also along spatio-temporal gradients of nutrient source availability.
Freshwater phytoplankton surveys with a focus on eutrophication commonly
describe the phytoplankton shifts in view of resource-driven control. These studies
describe a unique phytoplankton signature along a trophic gradient across lakes
from shallow to deep water bodies (Rojo and Alvarez-Cobelas 1994; Teubner 1996;
Teubner et al. 1999, 2003a, 2004; Naselli-Flores and Barone 2003; Stefaniak et al.
2005; Nixdorf and Deneke 1997; Søndergaard et al. 2005), from flushed riverine to
stratified systems (Krienitz et al. 1996; Teubner 1996; Teubner et al. 1999; Köhler
et al. 2000) or from tropical to temperate climate zone (Chen et al. 2003; Bouvy
et al. 2006; Burford et al. 2006, 2016; Liu et al. 2011, 2016; Deng et al. 2016). In all
these field surveys, the changes in phytoplankton assemblages associated with
eutrophication or restoration are mainly discussed by species alterations among or
within four predominant taxonomic affiliations: cyanobacteria, diatoms, golden
algae and green algae.
The aim of this chapter is to track the impact of lake restoration measures on
compositional shifts of phytoplankton described for four main lake treatment peri-
ods (1–4) including the chemical phosphorus flocculation and the re-establishment
of submerged vegetation. We further distinguish two sub-periods 2a and 2b con-
cerning the first and second chemical phosphate precipitation by RIPLOX-treatment
(Ripl 1976; Donabaum et al. 1999). The comparison of these four main periods by
other biota is described for zooplankton in Chap. 11 and for macrozoobenthos in
Chap. 14. Besides the compositional shift of phytoplankton due to phosphorus
reduction along restoration and management measures in Alte Donau, we analysed
the general phytoplankton pattern of seasonal cycles and pigment relationships and
its impact on the assessment by trophic classification.
9.2 Methods
9.2.1 Phytoplankton Sampling During Four Treatment Periods
Alte Donau, a former stretch of the Danube River, is an urban polymictic lake with
a maximum depth of 7 m and mean depth of 2.5 m (relative depth 0.52 calculated
according to Kõiv et al. 2011) and a surface area of 1.43 km2. The shallow oxbow
lake consists of two main impoundments, the south basin (in other chapters of this
book also called lower basin, ‘Untere Alte Donau’ or UAD) and the north basin
(also called upper basin, ‘Obere Alte Donau’ or OAD) with their main sampling
sites ‘AD1’ and ‘AD4’, respectively. According to Löffler (1988) and Mayer et al.
(1997), the retention time in the 1980s was roughly estimated by about 20 days to a
few months. With the construction of the impoundment Neue Donau (1970) and a
hydro power plant in the river Danube (1997) the water level fluctuations markedly
decreased step by step as the ground water fluxes changed dramatically (Chap. 2,
Fig. 5.1 in Chap. 5; see also Donabaum et al. 2004). A more detailed calculation for
the recent years revealed a much longer hydrological retention time fluctuating from
98 days to about 900 days, with a mean of 365 days for south and 175 days for north
basin (Chap. 4). The morphology and hydrology of these two basins are further
described in Chaps. 3 and 4, respectively.
Samples of phytoplankton and chlorophyll-a (chl-a) were taken with a
5 L-Schindler sampler at 0.2 m depth in the two main impoundments at biweekly
(to monthly) intervals from April 1993 to December 2014.
Phytoplankton biovolume was estimated from phytoplankton abundance and
size measurements using the sedimentation technique and light microscopy.
Biovolume was calculated for individual phytoplankton species according to the
basic geometric shape of the cell bodies or the aggregation of cells (Rott 1981).
Different from the 19-year record of zooplankton samples, which were analysed by
a single person, the microscopical phytoplankton counting over the 22 years was
conducted by a number of colleagues. The authors of this chapter (K.T., W.K.) con-
tributed to the phytoplankton counting for several years before and during the resto-
ration period. Taxonomic references reported in AlgaeBase [http://www.algaebase.
org; searched on 9 August 2017] were used for determining the phytoplankton taxa.
The 22-year time series covers all periods of the lake treatment described in
Chap. 5. For this phytoplankton chapter, the periods were slightly modified analo-
gous to zooplankton in Chap. 11. The periods are as follows: 1 – ‘eutrophication’
(before April 1995), 2 – ‘restoration including the first RIPLOX-treatment in April
1995 and the second RIPLOX-treatment in April 1996 (chemical phosphate precipi-
tation see Fig. 5.3 in Chap. 5, further Chap. 6, methods Chap. 11; details in Ripl
1976; Donabaum et al. 1999) and further restoration measures (April 1995–1999),
3 – ‘macrophyte re-establishment’ (2000–2006) and 4 – ‘stable conditions’ (2007–
2014). An earlier study comparing the pelagic community in the first and second
RIPLOX-year revealed different pathways of prompt responses of plankton organ-
isms to phosphorus reduction (Teubner et al. 2003b). Accordingly, we split the
r estoration period in two sub-periods, i.e. (2a) April 1995 to March 1996 and (2b)
April 1996 to 1999.
9.2.2 Data Treatment and Statistics
Phytoplankton shown separately for the two impoundments is based on the original
data set (Fig. 9.5). As phytoplankton, chl-a and total phosphorus (TP), however,
were not consistently sampled at exactly 2-week intervals, we interpolated the
observations at daily resolution (Livingstone 2003; Sapna et al. 2015) and averaged
these over 2 weeks, a month, a summer season or a year (Figs. 9.2, 9.3, 9.4 and 9.6,
9.7, 9.8, 9.9, 9.10, 9.11.). This data treatment is analogous to the data preparation
for the zooplankton analysis in Chap. 11.
We calculated net change rates of chl-a (kchl-a, hereafter referred to as net
growth rate of chl-a) at monthly time intervals as:
kchl - a  ln chl - at 2  ln chl - at1 / t (9.1)

where chl-at2 is the chl-a concentration at time t = t2, chl-at1 the chl-a at time t = t1
and Δt the time span in number of days (e.g. Teubner et al. 2003b). Likewise, we
analysed the persistence of phytoplankton composition at monthly time intervals
expressed by the standardised Bray-Curtis similarity index (0–100) using the soft-
ware package PRIMER 5. The higher this index, the higher is the resemblance of
the phytoplankton composition between successive samples as indicated by species
or functional phytoplankton groups. The seasonal variability of this Bray-Curtis
index and of chl-a and biovolume data was estimated using the coefficient of varia-
tion (CV = standard deviation/mean).
Prior to the statistical analysis, we tested data for normal distribution. According
to the non-parametric Shapiro-test (Dunn and Clark 1974), not all parameters fol-
lowed a normal distribution, in particular not those of single phytoplankton taxa,
which were rare in at least one lake treatment period. Also, chl-a and biovolume
data for the whole study period did not follow normal distribution. We conducted
statistical analyses to identify the differences among sampling sites (AD1, AD4)
and restoration periods using R (R i386 Version 2.15.2). We applied Kruskal-Wallis
H-tests to determine differences between annual median values for both impound-
ments and for the five restoration periods (Figs 9.7 and 9.8). To identify homoge-
neous subsets when comparing the phytoplankton assemblages between the two
impoundments (see text for difference between site in Fig. 9.5) and during the five
restoration periods, we further conducted pairwise Mann-Whitney U-tests with a
Bonferroni correction (Figs 9.7 and 9.8). The temporal agreement between chl-a
and biovolume was assessed using the non-parametric Spearman rank correlation.
The biovolume of higher ranks of phytoplankton taxa are displayed as notched box-
whisker plots using SYSTAT 10 (SPSS Inc.) (Figs 9.7 and 9.8). The boxes are
notched at the median; the length of the notches indicates the 95% confidence
interval.
9.2.3 Assessment by Functional Groups
Phytoplankton species were categorised by their co-occurrence during seasonal suc-

cession in similar habitats as ‘functional associations’ according to Reynolds et al.
(2002) modified by Padisák et al. (2009). We tried to avoid using taxa on genus level
instead of species level as suggested by Padisák et al. (2009) and further made sure
that the functional group categories comprise more than one species. For this rea-
son, we allocated Melosira varians C.Agardh to ‘P’ and not as a single species to
template ‘TB’ as nominated in Padisák et al. (2009). The individual taxa were allo-
cated to 20 functional groups as follows: cyanobacterial Chroococcales and
Synechococcales of the genera Woronichinia naegeliana (Unger) Elenkin and
Microcystis aeruginosa f. aeruginosa Kützing, M. viridis (A.Braun)
Lemmermann, M wesenbergii (Komárek) Komárek ex Komárek (Lm),
Aphanocapsa and Aphanothece (K), filamentous nitrogen fixing genera of
Nostocales as Dolichospermum and Aphanizomenon (H1) and Cylindrospermopsis
(SN) and of non-nitrogen fixing Oscillatoriales as Limnothrix, Planktolyngbya,
Planktothrix and Pseudanabaena (S1), unicellular centric diatoms (A), pennate dia-
toms as mainly Diatoma and Tabellaria (B) and Asterionella, Navicula and Nitzschia
(C), Aulacoseira granulata (Ehrenberg) Simonsen, Fragilaria cf. construens,
F. crotonensis Kitton, Melosira varians C.Agardh (P), cyanobacteria and dinofla-
gellates of the genera Ceratium, Gomphosphaeria, Gymnodinium, Merismopedia,
Snowella, Woronichinia, and Peridinium (Lo), cryptophytes mainly of the species
Cryptomonas curvata Ehrenberg C. erosa Ehrenberg, C. marssonii Skuja (Y),
small-cell taxa of Chlorophyta, Euglenophyta and Charophyta as e.g., Ankyra,
Carteria, Crucigenia, Elakatothrix, Koliella, Monoraphidium, Schroederia,
Tetraedron, Tetraselmis, Tetrastrum and Trachelomonas (X1), mainly colonial
Trebouxiophyceae and Chlorophyceae of the genera Botryococcus, Coelastrum,
Dictyosphaerium, Lagerheimia, Oocystis, Quadricoccus, Radiococcus and
Tetrachlorella; Radiocystis (F) and Pediastrum and Scenedesmus (J), large cell bod-
ies or filaments mainly of Charophyta and Chlorophyta as Closterium, Cosmarium,
Mougeotia, Planktonema, Staurastrum and Zygnema (N), flagellates mainly of
Ochrophyta and Cryptophytes as Chrysomonas, Ochromonas and Rhodomonas
(X2) and of other species of Ochrophyta of the genera Dinobryon, Mallomonas and
Synura (E) and Uroglena and two further groups of lower biovolume (U; names for
the functional groups are given in brackets and are based on the nomenclature of
Reynolds et al. (2002) and Padisák et al. (2009)).
9.2.4 A
ssessment of Trophic State by ÖNORM
and Phytoplankton Assemblage Metric Modified
from Brettum Index
The assessment of phytoplankton and chl-a along the TP gradient in the water col-
umn follows the lake’s trophic classification recommended by the ÖNORM M6231
(2001). According to this description of Austrian standards, we display the annual
concentration of TP versus summer chl-a and summer phytoplankton biovolume,
respectively. TP ranges are described for five trophic levels from oligotrophic to
hypertrophic including a sub-category for the eutrophic state with moderately eutro-
phic and highly eutrophic. For chl-a, the sub-categories for the eutrophic state are
missing in the ÖNORM M6231 (2001) and, therefore, we used an empirical value
calculated from the chl-a-TP response curve plotted for Alte Donau. Furthermore,
although rough numbers of phytoplankton biomass for some trophic levels are pro-
vided in the ÖNORM M6231 (2001), we used a more detailed calculation for phy-
toplankton biovolume for describing the five trophic levels. We calculated the class
limits for phytoplankton biovolume by converting the chl-a thresholds to biovolume
using a constant ratio that was empirically determined for Alte Donau (median
value, 0.50% chl-a of wet weight biomass of phytoplankton assuming a density of
1 mg biomass per 1 mm3 biovolume, 515 observations, details in Fig. 9.9). All num-
bers for the class limits describing the five trophic levels are displayed in Fig. 9.10.
In addition to the trophic states by ÖNORM M6231 (2001), we applied a phyto-
plankton assemblage metric to infer the ecological integrity of Alte Donau without
referring to the individual species again. The Austrian phytoplankton assemblage
metric is modified from the Brettum index (Brettum 1989), which was originally
developed for lakes in Norway and refers to a calibration data set over seven ranks.
The Austrian metric is based on a multi-year calibration data set of 167 phytoplank-
ton taxa observed in 29 lakes (2429 phytoplankton samples including TP measures,
Dokulil et al. 2005), which are mainly deep, oligo- to mesotrophic lakes and pre-
dominantly from the pre-alpine and alpine regions in Austria. To cover the full range
of trophic states as suggested by ÖNORM, ten mainly shallow and eutrophied lakes
in Germany were included in the data set (data of 8 meso- to hypertrophic lakes
from Teubner 1996 and of mesotrophic pre-alpine Ammersee from Teubner et al.
2004; Teubner 2006 in Dokulil et al. 2005). The Austrian phytoplankton assemblage
metric displays five ranks according to the five states recommended by ÖNORM
M6231 (2001).
1. oligotrophic
2. mesotrophic
3. moderately eutrophic
4. highly eutrophic
5. hypertrophic
In the calibration data set, the weighted species scores reflect the frequency dis-
tribution of individual phytoplankton species along these five ranks. The weighting
among species, which may differ in their biovolume yield by more than one order of
magnitude, counterbalances the contribution of many less abundant phytoplankton
species against few highly abundant, blooming species, thus giving a reliable lake
assessment. The advantage of the Austrian phytoplankton assemblage metric
described in Dokulil et al. (2005) is that it includes all species observed in a phyto-
plankton sample regardless of the species-specific phosphorus range. Thus, biased
interpretation using few, a priori selected single indicator species is avoided.
Assessing an actual phytoplankton lake sample by the Austrian assemblage metric,
the biovolume of each phytoplankton species is multiplied by its species-specific
weighted score from the calibration data set. Finally, the scores over all observed
lake species are summed up and weighted among the five TP ranks. The rank with
the highest total score indicates the highest probability of the trophic state.
Phytoplankton assessment methods to identify ecological integrity differ among
countries and regions as for example recently summarized by Pasztaleniec (2016).
Habitat scores in general, however, have a long tradition in ecology and are com-
monly retrieved from a meta-analysis of empirical observations of various biotic
community structure or other phenomena discovered across individual habitats
(Karr 1998; Hofmann 1993; Moog and Chovanec 2000; Moog 2002; Henderson
2003; Crossetti and Bicudo 2008, see also chironomid score in Chap. 14). In case of
phytoplankton, such methods are used to assess lake ecology in accordance with the
European Framework Directive (e.g., Padisák et al. 2006; Dokulil and Teubner 2006;
Solimini et al. 2008; Nõges et al. 2009; Poikane et al. 2011; Pasztaleniec 2016).
9.3 Results
9.3.1 P
hytoplankton Species Composition, Functional Groups
and Their Seasonal Pattern
Photographs of some cyanobacteria and algae which were observed in Alte Donau
are shown in Fig. 9.1. These photosynthetic organisms vary in their size from small-
celled pico-plankton as e.g. Aphanocapsa spp. (Fig. 9.1b) to macroscopically visi-
ble, large cells as e.g. Cosmarium botrytis (Fig. 9.1i) or may form large aggregates,
such as Microcystis aeruginosa, M. flos-aquae, M. novacekii, M. viridis, M. wesen-
bergii (Fig. 9.1a), Uroglena volvox (Fig. 9.1c), Dinobryon divergens (Fig. 9.1d),
Botryococcus braunii (Fig. 9.1g) and Coenochloris spec. (Fig. 9.1h). In terms of
taxonomy, common freshwater species of all taxonomic groups were observed with
the exception of the generally rare freshwater Rhodophyta. Considering the 17 main
Fig. 9.1 Phytoplankton species from small-sized picoplankton (b) to large netplankton species
(e.g., c) in Alte Donau; (a and b) Cyanobacteria, (a) – Microcystis wesenbergii, (b) – Aphanoscapsa
spec.; (c and d) Ochrophyta, Chysophyceae, c – Uroglena volvox Ehrenberg, (d) – Dinobryon
divergens O.E.Imhof; (e and f) Bacillariophyta, Bacillariophyceae, (e) – Encyonema triangulum
(Ehrenberg) Kützing, (f) – Fragilaria spec.; (g and h) Chlorophyta, (g) – Trebouxiophyceae,
Botryococcus braunii Kützing, (h) – Chlorophyceae, Coenochloris spec., (i and j) Charophyta,
(i) – Conjugatophyceae, Cosmarium botrytis Meneghini ex Ralfs, (j) – Hyalotheca dissiliens
Brébisson ex Ralfs, k Euglenophyta, Euglenophyceae, Euglena acus (O.F.Müller) Ehrenberg.
Scale bar for (a, b, h): 10 μm; for (d, e, g, i, j): 20 μm; for (f, k): 30 μm; for (c): 50 μm
win spr sum aut win spr sum aut win spr sum aut win spr sum aut
0.04 Bacill: 0.6 Bacill: 0.15 Dino: Bacill:
Astfor UniCen 0.10 Cerhir Aulspp
0.002
0.02 0.3
0.05
0.00 0.0 0.00 0.000

Bacill: 0.10
Bacill: 0.08 Cyano: Ochro:
0.007 Fracon Frauln Micwes 0.005 Urovol
0.05 0.04
0.000 0.00 0.00 0.000

Bacill: 0.008 Bacill: 0.003 Cyano: Cyano:
0.0010
0.002 Nitaci Fracro 0.002 Placir Cycrac
0.004
0.001 0.0005
Biovolume [mm3 L-1]
0.000 0.000 0.000 0.0000

0.002 Bacill: 0.15 Ochro: Cyano: 0.8 Cyano:
0.0010
Tabflo 0.10 Dindiv Snolac Plasub
0.001 0.0005 0.4
0.05
0.000 0.00 0.0000 0.0
Bacill: Chloro: 0.008 Chloro: 0.04 Cyano:
0.002
Navspp Kollon Coeast Psecat
0.0002
0.004 0.02
0.000 0.0000 0.000 0.00

Crypto: Crypto: Chloro: 0.0008 Chloro:
0.02 0.12
0.08 Rholac Coeret Schrob
0.01 0.06 0.0004
0.04
0.00
Cryero 0.00 0.00 0.0000
0.004 Chloro: 0.006 Chloro: Charo: Charo:
0.02 Cloacu
Tetspe Carpse 0.04 Stagra
0.002 0.003 0.01
0.000 0.000 0.00 0.00

DJF MAM JJA SON DJF MAM JJA SON DJF MAM JJA SON DJF MAM JJA SON
Months Months Months Months
Fig. 9.2 Seasonal development of 28 phytoplankton taxa as long-term monthly averages from
1993–2014. Black areas indicate the season with the highest mean seasonal biovolume. The verti-
cal line separates biovolumes of winter-spring from summer-autumn
Abbr: winter (DJF): Astfor – Asterionella formosa HASSAL, Fracon – Fragilaria construens
(EHRENB) GRUNOW, Nitaci – Nitzschia acicularis W.SMITH, Tabflo – Tabellaria flocculosa
(ROTH) KÜTZ., Navspp – Navicula spp., Cryero – Cryptomonas erosa EHRENB., Tetspe –
Testraselmis spec, spring (MAM): UniCen-unicellular centric diatoms, Frauln – Fragilaria ulna
(KÜTZ.) LANGE-BERTALOT, Fracro – Fragilaria crotonensis KITTON, Dindiv – Dinobryon
divergens O.E.IMHOF, Kollon – Koliella longiseta (VISCH.) HIND., Rholac – Rhodomonas
lacustris PASCHER et RUTTNER, Carpse – Carteria pseudomultifilis PETERFL L., summer
(JJA): Cerhir – Ceratium hirundinella (O.F.MÜLLER) DUJARDIN, Micwes – Microcystis wesen-
bergii (KOM.) KOM., Placir – Planktolyngbya circumcreta, Snolac – Snowella lacustris (CHOD.)
KOM. et HIND., Coeast – Coelastrum astroideum DE-NOT., Coeret – Coelastrum reticulatum
(DANG.) SENN, Stagra – Staurastrum gracile RALFS, autumn (SON): Aulspp – Aulacoseira
spp., Urovol – Uroglena volvox EHRENBERG, Cylrac – Cylindrospermopsis raciborskii, Plasub –
Planktolyngbya subtilis (W.WEST) ANAGN. Et KOM., Psecat – Pseudanabaena catenata
LAUTERB., Schrob – Schroederia robusta KORS., Cloacu – Closterium acutum (LEMM) W.
KRIEG; Bacill – Bacillariophyta, Crypt – Cryptophyta, Chloro – Chlorophyta, Ochro – Ochrophyta,
Dino – Dinoflagellata, Cyano – Cyanobacteria, Charo – Charophyta
functional phytoplankton groups for Alte Donau (see method), five groups refer to
cyanobacteria, four to chlorophyte and three to diatom taxa.
The monthly development of single species in Alte Donau is given in Fig. 9.2.
Most species with pronounced development in winter and spring were bacillario-
phytes (diatoms) of Tabellariales, Bacillariales and Fragilariales (needle-shaped
diatoms, such as e.g., Asterionella formosa, Nitzschia acicularis, Fragilaria con-
struens) and Stephanodiscales (a number of species categorised as unicellular cen-
tric diatoms). Some cryptophytes as Cryptomonas erosa and Rhodomonas lacustris
and few ochrophytes (e.g. Dinobryon divergens) and chlorophytes (Tetraselmis
spec. and Carteria pseudomultifilis) also developed their main biovolume in winter
to spring. Many species, such as e.g., Asterionella formosa, Cryptomonas erosa,
Rhodomonas lacustris and Tetraselmis spec., established large biovolumes through-
out these both seasons with the main peak or main seasonal average neither in win-
ter or spring. Typical phytoplankton species blooming in summer (Fig. 9.2) were
cyanobacteria (Microcystis wesenbergii, Planktolyngbya circumcreta), chloro-
phytes (Coelastrum astroideum, C. reticulatum), dinoflagellates (Ceratium hirundi-
nella) and charophytes (Staurastrum gracile) with high biovolumes lasting to
autumn. Other species with a biovolume peak in autumn reached already had high
yields in summer (e.g., cyanobacteria Cylindrospermopsis raciborskii,
Planktolyngbya subtilis). The majority of species developed their main yield either
in winter and spring or in summer and autumn. Hence, annual phytoplankton devel-
opment seems primarily distinctive between the two half-year-cycles, namely the
winter-spring and the summer-autumn period, rather than between the four
seasons.
Analogous to Fig. 9.2., the seasonality of biovolume of four phytoplankton asso-
ciations is shown in Fig. 9.3. As found for single species, functional phytoplankton
groups built pronounced biovolumes lasting either in winter and spring (functional
group ‘Y’ with species of Cryptomonas and ‘P’ of mainly pennate diatom species)
or in summer and autumn (functional group ‘Lo’ with species of cyanobacteria of
Genera Woronichinia, Snowella, Gomphosphaeria, Merismopedia and dinoflagel-
lates and ‘Lm’ of Cyanobacteria, e.g. the genus Microcystis). The monthly net
growth rate of chl-a (Fig. 9.4a) and changes of phytoplankton composition
win spr sum aut win spr sum aut win spr sum aut win spr sum aut
Biovol [mm3L-1]
1.00 1.00 1.00 1.00

Lm
0.75 Y 0.75 P 0.75 0.75
0.50 0.50 0.50 0.50
0.25 0.25 0.25 0.25
Lo
0.00 0.00 0.00 0.00
DJF MAM JJA SON DJF MAM JJA SON DJF MAM JJA SON DJF MAM JJA SON
Months Months Months Months
Fig. 9.3 Seasonal development of biovolume for four functional phytoplankton groups (Y, P, Lo,
and Lm; species allocation see method) as long-term monthly averages from 1993–2014
A
Net growth rateschl-a [d ] 0.6
-1
0.4
0.2
0.0
-0.2
-0.4
-0.6
J-F F-M M-A A-M M-J J-J J-A A-S S-O O-N N-D D-J
Monthly intervals
B
0.30
Bray-Curtis similarity for phytoplankton [%]
simil
85
CV
80 0.20
CV
75
0.10
70
phyto species
0.10
85
80 0.08
CV
75
0.06
70
phyto fct groups 0.04
J-F F-M M-A A-M M-J J-J J-A A-S S-O O-N N-D D-J
winter spring summer autumn winter
Fig. 9.4 Seasonal distribution pattern (a) of net growth rates of chl-a and (b) of compositional
shifts of phytoplankton (standardised Bray-Curtis similarity) for 28 phytoplankton species (phyto
species) shown in Fig. 9.3 and 13 dominant phytoplankton functional groups (phyto fct groups)
displayed as long-term mean (1993–2014) at monthly intervals from January to December. A simi-
larity value of 100% indicates an identical composition of two successive phytoplankton samples,
0% a totally different composition. The coefficient of variation (CV) illustrates the variability of
the Bray-Curtis similarity
(Fig. 9.4b) illustrate the seasonal pattern of phytoplankton development. In the

long-term average, the net growth rates of chl-a are positive in the first three monthly
intervals in the year revealing an increase of phytoplankton biovolume. From April
to May and May to June these net growth rates are slightly negative indicating the
break down of the spring phytoplankton bloom during the transition from spring to
summer. According to the long-term lake phenology analysed in Chap. 11, the
clear-water phase varied between the 95th (early April) and the 145th (late May)
day in the year (1994–2014, Fig. 11.7). The following strong increase of net growth
rates of chl-a in June to July stands for a rapidly progressive development of sum-
mer phytoplankton. Later in summer, the net growth rates of chl-a become moderate
positive and are weakest then successively decreasing to lowest and negative values
from October to November, i.e. from autumn to winter. With the winter season, net-
growth rates of chl-a increase again. The long-term seasonal change of phytoplank-
ton composition is expressed by the Bray-Curtis similarity between two successive
monthly samples (Fig. 9.1b). As indicated by the CV, the similarity index varies
most at particularly low values during the transition from spring to summer (from
April to May and May to June, respectively), which coincides with low net growth
rates of chl-a. It corresponds to a most pronounced species shift after the break
down of spring bloom and a new growth of summer plankton. A second but less
pronounced shift is found from October to November. Both species shifts during the
transition from spring to summer and autumn to winter reflect the development in
winter-spring or summer-autumn for many single species described before in
Fig. 9.2. This general pattern of seasonal phytoplankton phenology is also seen
when assessing compositional shifts by biovolume of phytoplankton functional
groups (Fig. 9.4b).
9.3.2 L
ong-Term Development of Taxonomic Phytoplankton
Groups
Time series of taxonomic groups are shown in Fig. 9.5 for both basins. When com-
paring the phytoplankton development year by year between the south (AD1) and
the north basin (AD4) (Fig. 9.5), we found statistically higher total biovolumes in
AD1 only in 2004 (H-test, p < 0.001) and 2010 (H-test, p < 0.05), while higher total
biovolumes in AD4 occurred in 2011 (H-test, p < 0.005) and 2012 (H-test, p < 0.05).
The biovolume of chlorophytes never differed between sites and built relatively high
biovolumes before chemical phosphorus precipitation and modest biovolumes in
years following the chemical treatment (Fig. 9.5). Analogous, the biovolume of cya-
nobacteria stayed similar between both basins with the exception of 3 years with
peak biovolumes of more than 40 mm2 L−1 in the eutrophied years 1993 and 1994
before chemical restoration. Large differences between sites were observed for bac-
illariophytes (H-test; p < 0.05) with statistically higher values in the south basin
from 2003 to 2010 and statistically lower values in 2011, both during the lake treat-
ment of re-establishment of macrophytes and stable conditions. Bacillariophytes
reached highest yields of 15 mm3 L−1 in years 1993 and 1994 before the chemical
phosphorus precipitation. In later years of lake treatment periods, biovolumes did
not exceed 3 mm3 L−1. Similar to the bacillariophytes, the ochrophytes were signifi-
cantly higher in the south basin from 2004 to 2006 (late years of the macrophyte
re-establishment) but higher in the north basin from 2011 to 2013 (during ‘stable
conditions’). Their biovolume was relatively low and did not exceed 2 mm3 L−1 bio-
volume (Fig. 9.5). Cryptophytes and euglenophytes developed only modest biovol-
umes (Fig. 9.5), but often had statistically significant higher biovolumes in the north
basin than in the south basin in some years from 2006 onwards. These differences in
phytoplankton structure, mainly from 2004 onwards only, mirror the spatial patchi-
ness of submerged vegetation but are of minor importance when describing the main
picture of the long-term response of phytoplankton to lake restoration. For this rea-
son, results in the following graphs are shown as averages for both lake basins of
Alte Donau summarizing aspects in the water body as a whole.
North basin (AD4) South basin (AD1)

50 50
40 40
30 30
10 10
Cyanobacteria cyanobact
Biovolume of taxonomic algal groups [mm L ]
5 5
-1
0
3
0
15 15
10 Bacillariophyta
10 bacillario
5 5
0 0
0.2 0.2
Euglenophyta eugleno
0.0 0.0
6 6
3 Chlorophyta 3 chloro
0 0
1.2 1.2 desmid
Charophyta
0.6 0.6
0.0 0.0
2 Cryptophyta 2 crypto
0 0
2 2
Ochrophyta 1 chryso
0 0
4 4
2 Dinoflagellata 2 dino
0 0
93 94 95 96 97 98 99 00 01 02 03 04 05 93 94 95 96 97 98 99 00 01 02 03 04 05
1 2a 2b 3 1 2a 2b 3
Years Years
Fig. 9.5 Time series of original data for monthly phytoplankton development at the sampling sites
AD1 (south basin) and AD4 (north basin) from 1993 to 2005 comprising the first three periods of
lake treatment: before restoration (1), restoration with chemical RIPLOX treatment (2a, 2b) and
re-establishment of macrophytes by periodical water level drawdown (3)
A
50
Phyto [mm3 L-1] 40 total biovolume

30
20
10
0
93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
B 100
80
% Cyanobacteria
60
40
Relative biovolume contribution of taxonomic phytoplankton groups
20
100
80 % Bacillariophyta
60
40
20
100
80
60
40 % Charophyta
20
100
80
60 % Chlorophyta
40
20
100
80
60 % Cryptophyta
40
20
100
80 % Ochrophyta
60
40
20
100
80
60
% Dinoflagellata
40
20
93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
1 2a 2b 3 4
Years
Fig. 9.6 22-year time series of phytoplankton development for total biovolume (a) and the contri-
bution of higher ranks of phytoplankton taxa (b) comprising five treatment periods: before restora-
tion (1), restoration with chemical RIPLOX treatment (2a, 2b), re-establishment of macrophytes
by periodical water level drawdown (3) and ‘stable conditions’ (4). Data are displayed as averages
of the two main impoundments shown in Fig. 9.5
The total biovolume as average for both basins was peaking during the eutro-
phied years 1993 and 1993 with peak values more than 50mm3 L−1 before chemical
precipitation by RIPLOX-treatment was carried out (Fig. 9.6a). Under these nutri-
ent rich conditions, filamentous cyanobacteria contributed up to 98% to phytoplank-
ton (Fig. 9.6b). As described in detail in former studies, Cylindrospermopsis
raciborskii, a taxon of the Nostocales, was the main cyanobacterium at that time and
occurred in association with Limnothrix redekei (Dokulil and Mayer 1996; Mayer
et al. 1997). With various measures of lake treatment, the biovolume contribution of
cyanobacteria decreased successively but still could contribute remarkable biovol-
umes with peak summer values from 20 to 50% (Fig. 9.6b). The cyanobacterial
composition, however, changed considerably when RIPLOX-treatment was con-
ducted in 1996 and 1997 and further lake treatment measures were carried out in
onward years. C. raciborskii almost disappeared and was replaced by various other
cyanobacterial taxa such as Chrococcales and Synechococcales (genera
Chroococcus, Gomphosphaeria, Limnothrix, Microcystis, Pseudanabaena,
Radiocystis, Snowella, Woronichinia). Bacillariophytes and chlorophytes reached
an intermediate importance in building up phytoplankton biovolume during chemi-
cal restoration (period 2), charophytes during the re-establishment of macrophytes
(period 3). Ochrophytes and cryptophytes contributed most to biovolume during
re-establishment of macrophytes and stable conditions (period 3 and 4).
9.3.3 C
omparison of Phytoplankton Composition and Chl-a
Content Between the Five Lake Treatment Periods
Besides the time series of the 22-year phytoplankton development, the box-plots in
Fig. 9.7 depict the distribution pattern of total biovolume and the chl-a concentra-
tion during the five periods of lake management (1, 2a & b, 3 and 4). For reference
of the trophic situation, the total phosphorus is displayed in this figure accordingly.
With the first chemical phosphorus precipitation in the RIPLOX-treatment year
1995, the average concentration during the eutrophied period (1, Fig. 9.7) dropped
to less than a half for TP (45% mean, 38% median), to about 20% for chl-a and to
about 15% for phytoplankton biovolume. This drastic reduction of phosphorus
availability for phytoplankton growth led to statistically significant differences in
the median value between the treatment periods (Fig. 9.7 Kruskal-Wallis test, a:
H = 305.15, df = 4, p < 0.001, b: H = 149.78, df = 4, p < 0.001, c: H = 143.38, df = 4,
p < 0.001) with a significant reduction of all three parameters from period 1 (sub-
group ‘a’) to 2a. The subsequent decrease of TP seems to be very closely associated
with chl-a as both these parameters follow almost the same statistically relevant
allocation of subgroups displayed by boxes, namely a less distinct phase in the first
RIPLOX-year followed by a slight recovery of the trophic level from the first to the
Fig. 9.7 Box-whisker plots of (a) TP concentration (in μmol L−1 and μg L−1), (b) phytoplankton
biovolume (mm3 L−1) and (c) chl-a concentration (μg L−1) during the five treatment periods. All
data are averages of AD1 and AD4. Statistically significant differences between homogeneous data
subsets are represented by letters above each box as determined by pairwise Mann-Whitney U tests
with Bonferroni correction. 2a, 2b, 3 and 4 indicate the treatment periods as in Fig. 9.6 (see
methods)
second RIPLOX-year and a further reduction during the re-establishment of macro-

phytes (period 3, subgroup ‘c’) and the period of stable conditions (period 4, sub-
group ‘d’). Different to chl-a, the biovolume decreased gradually as subgroups were
statistically less distinctive during periods 2a to 4. The response of single phyto-
plankton groups to lowered TP is described in more detail for phytoplankton chl-a
and biovolume in Figs 9.8 and 9.9.
Analogous to the time series in Fig. 9.6, the compositional changes of phyto-
plankton can be described by shifted proportions among taxa of various taxonomic
affiliations (Fig. 9.8). The distribution pattern of chlorophyll content (Fig. 9.8a) and
phytoplankton groups (Fig. 9.8b–h) is displayed for the five lake treatment periods.
The most significant compositional change is caused by the reduction of cyanobac-
teria (Fig. 9.8b). The eutrophied period 1 (‘subgroup a’) does not overlap with later
lake treatment periods. After a strong reduction in the first RIPLOX-treatment
(period 2a), the biovolume contribution of cyanobacteria slightly increased in the
second RIPLOX-treatment (period 2b) before decreasing in onward periods s howing
the lowest contribution in the period of stable conditions (period 4, subgroup ‘c). An
almost opposite pattern is found for ochrophytes and cryptophytes (Fig. 9.8c, d). A
prompt increase of their relative biovolume contribution was observed in the period
of the first RIPLOX-year of phosphorus precipitation (period 2a) followed by a
slight decrease during the second RIPLOX-treatment and a subsequent increase in
phytoplankton biovolume in periods of macrophyte re-establishment (period 3) and
stable conditions (period 4). The biovolume contribution of chlorophytes and bacil-
lariophytes (Fig. 9.8e, f) revealed an intermediate increase during both RIPLOX-
a cd b c d a bc b b c a b a c d a ab a b c
2. 4 100 100 100
A B C D
Chlorophyll-a [%]
Cyanobacteria [%]
80 80 80
Cryptophyta [%]
1. 8
Ochrophyta [%]
60 60 60
1. 2
40 40 40
0. 6
20 20 20
0. 0 0 0 0
1 2a 2b 3 4 1 2a 2b 3 4 1 2a 2b 3 4 1 2a 2b 3 4
Period Period Period Period
a bc b b c a b b c c a b c c d a a a a a
100 100 100 100
E F G H
Dinoflagellata [%]
80 80 80 80
Bacillariophyta [%]
Chlorophyta [%]
Charophyta [%]
60 60 60 60
40 40 40 40
20 20 20 20
0 0 0 0
1 2a 2b 3 4 1 2a 2b 3 4 1 2a 2b 3 4 1 2a 2b 3 4
Period Period Period Period
Fig. 9.8 Box-whisker plots of (a) chl-a content of phytoplankton as % of wet weight phytoplank-
ton biomass (see method 9.2.4) and (b–h) relative contribution of taxonomic phytoplankton
groups. All data are averages of AD1 and AD4. Statistically significant differences between homo-
geneous data subsets for each parameter are represented by letters above each box as determined
by pairwise Mann-Whitney U tests with Bonferroni correction. 2a, 2b, 3 and 4 indicate the treat-
ment periods as in Fig. 9.6 (see methods)
years, but a successive decline in onward periods. The contribution of charophytes

and dinoflagellates (Fig. 9.8g, h) remained low during all five periods. Charophytes
reached their highest relative biovolume in the period of macrophyte re-establish-
ment. For dinoflagellates, no significant differences between the treatment periods
were observed (Kruskal-Wallis test, H = 9.18, df = 4, p = 0.057). The chl-a content
of total phytoplankton (Fig. 9.7a) varied also among lake treatment periods. Its
median value was lowest with 0.19% (mean 0.20%) for period 1 when cyanobacte-
ria were mainly contributing to total phytoplankton (median 77%, mean 64%). The
chl-a content is highest for period 2b (median 0.83%, mean 0.84%) and coincides
with a high contribution of chlorophytes during period 2b (median 18%, mean 21%)
and a large contribution of bacillariophytes (median 33%, mean 35%) while the
contribution of cyanobacteria was moderate (median 16%, mean 21%).
When looking at biweekly data over the whole study period, phytoplankton chl-a
concentration and biovolume shows a high temporal agreement (Spearman rank,
r = 0.85 with p < 0.01; Fig. 9.9). The range of variation, however, is different for
both parameters. With a CV of 1.5 for phytoplankton chl-a but of 2.4 for phyto-
plankton biovolume, the chl-a values are less variable than those of phytoplankton
biovolume estimating the biweekly phytoplankton yields. The median percentage of
chl-a is 0.50% of wet weight phytoplankton biomass (see methods). All data points
2.5
1
2a
2.0
2b
Chlorophyll-a [LOG µg L ]
-1
3
1.5 4
1.0
0.5
0.0
-0.5
-1.0 -0.5 0.0 0.5 1.0 1.5 2.0
3 -1
Phytoplankton biovolume [LOG mm L ]
Fig. 9.9 Relationship between phytoplankton chl-a concentration and phytoplankton biovolume,
both displayed on a logarithmic scale (LOG10). The solid line indicates 0.50% chlorophyll of phy-
toplankton biomass wet weight (median of biweekly averages of AD1 and AD4 over 22 years, see
methods); Spearman rank correlation, r = 0.85, p < 0.001, n = 513; 1, 2a, 2b, 3 and 4 indicate the
lake treatment periods as in Fig. 9.6
describing the eutrophied period before the first chemical phosphate precipitation in
April 1995 are below the 0.50% line while most data points of period 2b are above
this value.
9.3.4 A
ssessment of Trophic State and Phytoplankton Habitat
Score Index
The trophic classification according to Austrian standards is described by the rela-

tionship of summer phytoplankton chl-a concentration (Fig. 9.10a) or summer phy-
toplankton biovolume (Fig. 9.10b) versus annual TP concentration. As shown by
22 years passing five lake treatment periods, Alte Donau went through an ecosystem
shift from hypertrophic to mesotrophic, and finally reached a relative poor nutrient
status close to oligotrophic under ‘stable conditions’. This main picture is observed
regardless of assessing chl-a or phytoplankton biovolume versus TP. The equivalent
assessment by chl-a and biovolume, however, is not valid throughout all five lake
treatment periods. Large differences in the assessment of lake nutrient status by
chl-a or biovolume were found for period 1 and 2b. During years before chemical
restoration (period 1, 1993 and 1994), summer chl-a values and annual TP concen-
trations indicate a hypertrophic situation (Fig. 9.10a). The values of chl-a and TP
A B
1.8 1.4
hyper
1.6 35 µg L-1 1.2
Chlorophyll-a [LOG µg L-1]
highly-eutr
Biovolume [LOG mm3 L-1]

24 µg L-1
1.4 1.0 3 -1 hyper
7.0 mm L
mod-eutr
1.2 -1 0.8 4.7 mm3 L-1 highly-eutr
12 µg L
mes o mod-eutr
1.0 0.6 2.4 mm3 L-1
1
0.8
4 µg L
-1 0.4 mes o
2a
0.8 mm3 L-1
0.6 0.2 2b
oligo -1
-1
-1
-1
-1
-1
-1
-1
60 µg L oligo
10 µg L
20 µg L
40 µg L
60 µg L
10 µg L
20 µg L
40 µg L
0.4 0.0 4
0.2
0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0 0.6 0.8 1.0 1.2 1.4 1.6 1.8 2.0
TP [LOG µg L-1] TP [LOG µg L-1]
Fig. 9.10 Scheme of trophic lake classification: trophy classes (oligo oligotrophic, meso mesotro-
phic, mod-eutr moderately eutrophic, highly-eutr highly eutrophic, hyper hypertrophic) are shown
as response curve of summer phytoplankton chl-a concentration (a) and summer phytoplankton
biovolume (b) to annual TP concentration displayed on a logarithmic scale (LOG10); Solid lines
and respective numbers of class limits refer to Austrian standards (ÖNORM M6231 2001). Dashed
lines and numbers for class limits between highly and moderately eutrophic for chl-a and biovol-
ume are calculated to distinguish the five TP classes accordingly (see methods); data are averages
of AD1 and AD4; 1, 2a, 2b, 3 and 4 indicate the lake treatment periods as in Fig. 9.6 (see
methods)
are both close to the limit between hypertrophic and highly eutrophic. When assess-
ing the trophic classification by summer phytoplankton biovolume and annual TP
concentration, the biovolume exceeds the class limit of biovolume towards hyper-
trophic conditions much more than those of TP (Fig. 9.10b). In addition, trophic
class assignment differs for the period with repeated RIPLOX-treatment and fol-
lowing years of restoration (period 2b), i.e. summer chl-a still demonstrates a mod-
erately eutrophic level while summer biovolume already indicates a mesotrophic
level. According to the compositional shifts of phytoplankton, the chl-a content
changed during the treatment periods as described for Figs. 9.8 and 9.9. The pre-
dominance of cyanobacteria during hypertrophic summer phytoplankton develop-
ment was associated with a relatively low chl-a content per phytoplankton biomass.
In turn, the chl-a content in period 2b was particularly high when the contribution
of cyanobacteria was low but that of chlorophytes considerably high. Comparing
the trophic classification scheme of Fig. 9.10a, b, double logarithmic scaled data
points for chl-a versus TP follow a linear relation more closely (Spearman rank,
r = 0.86 with p < 0.01; Fig. 9.10a) than those of biovolume versus TP (Spearman
rank, r = 0.51 with p < 0.05; Fig. 9.10b).
The trophic classification assessed by the Austrian phytoplankton assemblage
metric are displayed for the five lake treatment periods separately Fig. 9.11. The
bars mirror the distribution pattern of species in assemblages occurring from oligo-
to hypertrophic. The black bars indicate the most probable trophic classification
during each lake treatment period. According to this metric, Alte Donau shifts from
Fig. 9.11 Phytoplankton 100

habitat quality index for
the five lake treatment 1
periods. The phytoplankton
assemblage metric refers to 0
five trophic states:
1-oligotrophic 100
(TP < 10 μg L−1),
2-mesotrophic (TP 2a
10–20 μg L−1),
0
Species assemblage score

3-moderately eutrophic
(TP 20–40 μg L−1), 100
4-highly eutrophic (TP
40–60 μg L−1), and 2b
5-hypertrophic state (TP
>60 μg L−1), The black bar
0
indicates the highest
probability of trophic state; 100
1, 2a, 2b, 3 and 4 indicate
the lake treatment periods 3
as in Fig. 9.6 (see methods)
0
100
0
1 2 3 4 5
Habitat quality index
hypertrophic to mesotrophic along the five lake treatment periods which is in agree-
ment with the main picture of the trophic classification shown in Fig. 9.10. With the
first RIPLOX-treatment, the metric indicates the pronounced step by prompt species
response from an assemblage typically found in hypertrophic lakes (mainly built up
by the cyanobacterium Cylindrospermopsis raciborskii) to an assemblage shared by
species of various taxonomic affiliations commonly found in moderately eutrophic
environments. This result thus mirrors the drastic change indicated by the relation-
ship of chl-a versus TP more closely than by phytoplankton biovolume versus
TP. According to this metric, the second RIPLOX treatment and the measures dur-
ing this period 2b manifest this species composition of a moderately eutrophic lake.
With period 3 and 4 more species are referring to a mesotrophic reference, which is
mainly due to cyanobacteria taxa other than C. raciborskii and a further mixture of
species allocated to the ochrophytes and cryptophytes (see results for Fig. 9.8
before) commonly observed in mesotrophic lakes.
9.4 Discussion
Total biovolume of phytoplankton did not significantly differ for the two impound-
ments with the exception of 4 years in the 22-year time series. The similar phyto-
plankton development in the two impoundments is found despite the large spatial
heterogeneity of the banks and basins of the former Danube river branch (Table 3.1
and Fig. 3.1 in Chap. 3; see also Chaps. 18 and 19). For example, macrophyte stand-
ing crop was about 30% lower in the south basin than in the north basin during the
period from 2005 to 2008, while it was 20% higher in the south basin from 2009 to
2012; in onward years the distribution was more even (Karin Pall, pers. comm.). In
addition, annual phosphorus loading was more than twice as high in the south than
in the north basin in the recent years 2008–2014 (Fig. 6.6 in Chap. 6) which is asso-
ciated with a longer retention time (Chap. 3). The even distribution of total phyto-
plankton in both basins, however, is in line with the strong consistency of zooplankton
(Chap. 11) and ciliate biomass (Chap. 12) in both impoundments. When looking at
particular taxonomic affiliations of phytoplankton, such as bacillariophytes, ochro-
phytes, cryptophytes and euglenophytes, no statistically significant differences were
found before 2002, i.e. the 10 years before underwater vegetation was re-established.
In onward years, a frequent alternation of low and high biovolume of different taxo-
nomic affiliations was observed for the two impoundments which arose from the
strong re-growth of underwater vegetation (mainly built of Myriophyllum spicatum,
Fig. 8.9 and Table 8.2 in Chap. 8, Fig. 20.5 in Chap. 20). During that period under-
water vegetation strongly increased from a very low biomass in 2002 to a 6-fold
higher yield of 125 t dry mass in 2003 and then remained stable at about 300–500 t
dry weight (see time series of macrophyte development Figs. 8.7 and 8.11 in Chap.
8). As total phytoplankton biovolume did not significantly differ in years with mas-
sive underwater vegetation yield, it can be assumed that biovolume among taxa of
different taxonomic affiliations was counterbalanced as e.g. statistically higher bio-
volumes were found for bacillariophytes in the north basin and for cryptophytes in
the south basin. This might display the substitution of benthic-pelagic species ver-
sus exclusively pelagic species, as many bacillariophytes are benthic species and
might have been additionally found as phytoplankton in water column samples near
patchy underwater vegetation, while areas with less underwater vegetation might
have stimulated the growth of exclusively planktonic species, such as e.g. the cryp-
tophytes. The shift of mainly tychoplanktonic pennate diatoms towards mainly
planktonic solitary centric diatoms and the further shift towards e.g. cryptophytes is
commonly described for ecosystems along the transition from lentic to lotic habitats
in association with modified light and nutrient exposure (e.g., Bahnwart et al. 1998;
Nicklisch 1998; Nicklisch et al. 2007; Centis et al. 2010; Wu et al. 2010; Zohary
et al. 2010; Naselli-Flores and Barone 2011; Abonyi et al. 2012; Shatwell et al.
2012).
The species composition of the phytoplankton community found in Alte Donau
is similar to that found in the river-floodplain system of the River Danube (e.g., Kiss
1987; Schmidt 1994; Schagerl and Riedler 2000; Mihaljević et al. 2010; Dokulil
and Donabaum 2014). All species reported for Alte Donau were further common in
shallow lakes as, e.g., documented for North Germany by Täuscher (2014).
9.4.1 T
he Success of Filamentous Cyanobacteria
Cylindrospermopsis raciborskii in the Nutrient Rich
Oxbow Lake
The awareness of the unhealthy state of the ecosystem Alte Donau increased with
the ongoing eutrophication in 1993 and 1994, when progressive turbidity of the
water body was mirrored by a Secchi depth of approximately 0.5–1.5 m which was
caused by cyanobacterial blooming (Chap. 6, see also Mayer et al. 1997; Donabaum
et al. 1999; Dokulil and Teubner 2003). Effective lake restoration and management
measures were deemed necessary in this urban lake when realising that the main
planktonic cyanobacterium was Cylindrospermopsis raciborskii since strains of this
cyanobacterium can potentially produce several cyanobacterial toxins, saxitoxins
and cylindrospermopsin (e.g., Fastner et al. 2003, 2007; Kurmayer and Christiansen
2009). During the mass development in Alte Donau, however, no evidence of such
substances was recorded by symptoms of bathing people, which confirms other field
studies of non-toxic strains building up C. raciborskii blooms (Alster et al. 2010).
Although no measurements of toxins were carried out in Alte Donau and hence no
chemical information is available on the presence of toxins attributed to strains of
C. raciborskii, it can be assumed that such toxic substances were not a major issue
in this popular bathing lake in 1993 and 1994 even though being aware of the scien-
tific challenge in predicting toxic cyanobacterial blooms (e.g. Bukowska et al.
2017). This cyanobacterium was originally commonly observed in the subtropical
and tropical region (e.g., Bouvy et al. 2006; Figueredo and Giani 2009). It is now
more increasingly found as an invasive species in the temperate zone and often seen
to be growth stimulated by global warming in nutrient rich shallow lakes (Dokulil
and Mayer 1996; Padisák 1997; Isvánovics et al. 2000; Mischke 2003; Nixdorf et al.
2003; Stüken et al. 2006; Fastner et al. 2007; Kling 2009; Kaštovský et al. 2010;
Aubriot and Bonilla 2012; Bonilla et al. 2012; Paerl and Otten 2013; Burford et al.
2016; Dokulil 2016; Kokociński et al. 2017). C. raciborskii belongs to the Nostocales
among the Cyanobacteria which are known to be able to fix N2 in heterocytes during
periods of nitrogen limitation (e.g. see trait description for this taxon in the phyto-
plankton functional group by Mantzouki et al. 2016). N2-fixation by nostocale cya-
nobacteria, however, is not necessarily a dogma for acquiring nitrogen, even if
among vegetative cells also heterocytes are grown by these prokaryotic species
(Teubner 1996; Teubner et al. 1999). The preferred and hence more likely sources
for utilising nitrogen are ammonium and further nitrate if these N-fractions are
available for cyanobacteria (e.g., Dudel and Kohl 1991; Burford et al. 2006). The
cyanobacterial filaments of C. raciborskii were the most important for building up
phytoplankton in 1993 and 1994 when total biovolume yielded a summer maximum
of more than 50 mm3 L−1 (original description in Dokulil and Mayer 1996 and
Mayer et al. 1997, reinforced analysis of this dataset in Dokulil 2016). C. racibor-
skii developed highest biovolumes at a water surface temperature of 21.4 °C under
well-mixed, hypertrophic conditions in the polymictic Alte Donau (Dokulil and
Teubner 2000; hydrological retention time and hydrological aspects including the
ground-water flow attributed to the oxbow and groundwater-seepage lake see meth-
ods and Chaps. 3 and 4). Nixdorf and Deneke (1997) compared a variety of eutro-
phied lakes in North Germany with deep stratified and shallow well mixed water
columns during summer and stated that at least ‘very shallow’ lakes were the most
efficient systems in utilising phosphorus due to the favourable proportion between
euphotic depth and mixing depth. An example for this category is Lebbiner See with
a maximum depth of 4 m, a contribution of summer cyanobacteria by more than
90% to total biovolume, an occurrence of C. raciborskii among other filamentous
cyanobacteria, and a summer chlorophyll peak value of 71 μg L−1, indicating three
aspects of a phytoplankton situation that is comparable to the condition of Alte
Donau before restoration. According to a field survey in The Netherlands referring
to more than 400 ‘lake years’ from 80 lakes, which were mainly shallow and eutro-
phied, cyanobacteria contributed with 46% the most to phytoplankton (Schreurs
1992). Schreurs emphasized that the comparison of annual yields in ‘Oscillatoria‘,
‘Aphanizomenon‘ and ‘Microcystis‘-dominated lakes shows that the highest con-
centrations of chlorophyll-a occurred in the ‘Oscillatoria lakes‘ with a long-lasting
dominance especially of e.g. Planktothrix agardhii. The success of building up an
extremely high phytoplankton yield in ‘typical‘ Oscillatoria- years could be verified
by the flushed riverine lake ‘Langer See’ in the early 1990s in Germany (Teubner
1996; Teubner et al. 1999), which built up blooms with solitary filaments of
Planktothrix agardhii in association with solitary filaments of Limnothrix redekei as
opposed to alternatively blooming associations forming scum mainly composed of
two colonial forms of cyanobacteria, Aphanizomenon flos-aquae (bundles of fila-
ments) and Microcystis spp. (coccal cell aggregates). In Alte Donau, the massive
bloom of C. raciborskii was associated with L. redekei (Dokulil and Mayer 1996;
Mayer et al. 1997; Dokulil 2016), which would support the finding by Schreurs
(1992) that in particular solitary filaments of cyanobacteria are successfully build-
ing up high yields in nutrient-rich shallow lakes.
One aspect in describing the successful development of cyanobacteria in a highly
fluctuating environment in terms of light and nutrient availability is analysing the
maximum growth rate for optimal growth in comparison to modulated nutrient and
light conditions. Such experiments under different light and nutrient exposure are
aimed at mimicking a regime of a turbulent turbid water column and could show
that cyanobacteria forming solitary filaments (Nicklisch 1998; Nicklisch and Fietz
2001; Nicklisch et al. 2007; Shatwell et al. 2012; Deng et al. 2016) can cope well
with continuous or fluctuating light and nutrient availability. Lab experiments and
field assays specifically with Cylindrospermopsis raciborskii underpin the finding
on the successful growth in well mixed turbid environments for this solitary fila-
mentous cyanobacterium (Isvánovics et al. 2000; Aubriot and Bonilla 2012; Bonilla
et al. 2012; Amaral et al. 2014; Dokulil 2016). Other aspects for prosperous devel-
opment of cyanobacteria in general or of C. raciborskii in the specific case indicated
by long-lasting blooms in particular in summer are a higher temperature optimum
for growth (Nicklisch et al. 2007; Joehnk et al. 2008; Kokociński et al. 2017), a
highly adaptive strategy of phosphorus acquisition even of ephemeral nutrient-
source patches (e.g., Falkner and Falkner 2003; Aubriot and Bonilla 2012; Amaral
et al. 2014) and storage behaviour (e.g. Isvánovics et al. 2000).
As maximization of phosphorus uptake is a function of cell surface (in addition
to enzyme activity and related capacity of transport incorporating phosphorus, see
e.g. Finkel et al. 2009), the long thin filaments of solitary trichoms, as e.g. found for
Cylindrospermopsis raciborskii and Limnothrix redekei (their filament diameter is
only 1.8–2.4 μm and 3.05–3.8 μm, respectively; unpublished data Teubner), might
provide a further aspect of ecological relevance mirrored by field studies. Comparing
surface to volume ratios of species of various taxonomic affiliations in meso- to
hypertrophic lakes, the Oscillatoriales together with filaments of Synechococcales
(former Oscillatoriales) represent the second highest surface to volume ratios
(median 1507 mm2 L−1 per mm3 L−1, n = 259), while those of filamentous Ulotrichales
were much higher (4071 mm2 L−1 per mm3 L−1, n = 126) and of pennate diatoms
slightly lower (1374 mm2 L−1 per mm3 L−1, n = 357, dimensions measured by
Interference reflection microscopy, data from Fig. 23 in Teubner 1996; empirical
measures of surface and volume for algae of various taxonomic affiliations see also
Morabito et al. 2007).
During the ecosystem shift from the hypertrophic to mesotrophic, the relative
contribution of cyanobacteria to total biovolume in summer remained relatively
high with about 20% although the TP concentration and total phytoplankton dropped
drastically. A closer look at the species shift within the cyanobacteria, however,
reveals that cyanobacteria, such as blooming Cylindrospermopsis raciborskii disap-
peared while other cyanobacteria such as Microcystis, Aphanocapsa taxa became
typical cyanobacterial summer taxa even in the period of ‘stable conditions’ (period
4). Many cyanobacteria found in period 4, however, were already documented for
mesotrophic conditions in 1986 and 1987. In a multi-site phytoplankton field survey
during that time, Donabaum (1988) reported cyanobacterial taxa (Chroococcales
and Synechococcales) such as Microcystis aeruginosa, M. flos-aquae and
Planktolyngbya limnetica in addition to ochrophyte and charophyte taxa (Uroglena
spp., Closterium aciculare T.West).
The growth of cyanobacteria is described for a variety of ecotypes from oligo- to
hypertrophic and occurs under particular underwater light climate at surface water
or deep metalimnetic layers from mixed to stratified water columns, from shallow
to deep lakes, from lakes of the tropical and subtropical to the temperate zone (e.g.,
Jewson 1977; Schreurs 1992; Mur et al. 1993; Dokulil and Teubner 2000; Teubner
et al. 2004; Teubner 2006; Paerl and Otten 2013; Qin et al. 2013). The maximum of
about 20% peak contribution of cyanobacteria in summer is thus not critical for our
conclusion of successful restoration in mesotrophic Alte Donau as (1) cyanobacte-
rial taxa mirror a high ecological plasticity and (2) their portion in phytoplankton is
in the same range as for diatoms and green algae, while only those for cryptophytes
and ochrophytes are a bit higher. The success of restoration in view of phytoplank-
ton composition thus might be seen in the balanced mixture of taxa of many taxo-
nomic affiliations (e.g. Bouvy et al. 2006) and points in the direction of enhanced
biotic diversity and richness stabilising a healthy ecosystem (e.g., Costanza and
Mageau 1999; Dokulil and Teubner 2010). This is also in agreement with the phy-
toplankton assemblage metric applied for Alte Donau (Dokulil et al. 2005), which
indicates a shift from phytoplankton taxa commonly found in hypertrophic lakes
observed before restoration to taxa commonly occurring in mesotrophic lakes
described for the ‘stable’ condition of the restored Alte Donau. This result verifies
that phytoplankton assemblage metrics are useful tools for analysing the trophic
status of lakes and rivers, which is often used to assess the ecological status of an
aquatic ecosystem (e.g., Dokulil et al. 2005; Dokulil and Teubner 2006; Padisák
et al. 2006; Lepistö et al. 2006; Crossetti and Bicudo 2008; Solimini et al. 2008;
Nõges et al. 2009, 2010; Marchetto et al. 2009; Rimet et al. 2009; Poikane et al.
2011; Napiórkowska-Krzebietke et al. 2012; Katsiapi et al. 2016; Lobo et al. 2016;
Pasztaleniec 2016; Vadrucci et al. 2017).
9.4.2 The Seasonal Pattern of Phytoplankton Development
The bloom of Cylindrospermopsis raciborskii and Limnothrix redekei is not only

observed for summer periods (it is commonly agreed that cyanobacteria are typi-
cally blooming during the hot season; e.g., Dokulil and Teubner 2000; Cao et al.
2008; Liu et al. 2011; Paerl and Otten 2013), but is lasting also until autumn (Dokulil
and Mayer 1996; Mayer et al. 1997; Dokulil 2016). Such a ‘warm assemblage’ last-
ing during summer and autumn (Komárková et al. 2003) is dedicated to the growing
season established after spring overturn and lasting to autumnal mixing and are
commonly studied when describing phytoplankton succession. The importance of
‘cold assemblages’ lasting from winter to spring (Komárková et al. 2003) became
more relevant in recent years with increasing awareness of over-wintering phyto-
plankton (e.g., Rodhe 1955; Adrian et al. 1999; Weyhenmeyer et al. 1999; Teubner
2000; Morabito et al. 2002; Barone and Naselli-Flores 2003; Naselli-Flores and
Barone 2003; Anneville et al. 2005; Dokulil and Herzig 2009; Hampton et al. 2017).
It’s impact on vernal bloom is most relevant for lake assessment and climate
research. This pattern of main compositional shifts that occur only twice a year, i.e.
first during the transition from spring to summer establishing a ‘warm assemblage’
lasting from summer to autumn, and secondly during the transition from autumn to
winter building up a ‘cold assemblage’ lasting from winter to spring, was also found
in Alte Donau and verifies earlier studies from shallow to deep lakes (using dis-
criminant function analysis in Teubner 2000 and Bray-Curtis similarity in Teubner
et al. 2003a). This seasonal pattern is a cornerstone of understanding phytoplankton
development in view of lake phenology in the temperate climate zone, whether in
view of the succession of phytoplankton species (e.g. plankton groups ecology by
Sommer et al. 1986, 2012; De Senerpont Domis et al. 2013) or the description of
seasonally co-occurring taxa of various affiliations by functional groups (e.g.,
Reynolds et al. 2002; Padisák et al. 2009; Crossetti and Bicudo 2008; Nõges et al.
2010; Vadrucci et al. 2017). The transition from spring to summer, i.e. ‘clear-water
phase’, is a key parameter when studying lake phenology (Sommer et al. 1986; De
Senerpont Domis et al. 2013). In Alte Donau this short period of increased water
transparency occurred from early April to late May (Chap. 11) stimulating the ver-
nal growth of submerged plants (see Sect. 8.3.1 in Chap. 8) which is most important
for the recovery of aquatic vegetation (e.g. Scheffer et al. 2001; Wu et al. 2014). The
timing of the clear-water phase is in accordance with phenology for other shallow
polymictic lakes (Teubner et al. 1999; Straile and Adrian 2000; Huber et al. 2010;
Shatwell et al. 2008) and different from an often later timing for deeper lakes
(Müller-Navarra et al. 1997; Winder and Schindler 2004; Straile 2002). The trend
analysis revealed a time shift by about 11 days advance per decade and is associated
with climate change (details in Sect. 11.3.5 of Chap. 11). According to our analysis,
the phenology of the vernal clear-water phase can be described for two distinct
periods with regard to the phosphorus decline, i.e. a more pronounced shift of
33 days per decade earlier during the rapid decline of TP from 1994–1999 (period 1
and 2) and a shift of only 7 days earlier per decade for years of moderately low TP
(period 3 and 4). According to empirical analysis including lakes with long-term
external reduction of phosphorus loading, the phenology of the vernal clear-water
phase should be interpreted with caution as time-shifts cannot be attributed to cli-
mate change only but are a result of the interplay of climate forcing and trophic
changes in the ecosystem (e.g. Jeppesen et al. 2003).
9.4.3 T
he Success of Restoration in Terms of Controlling
Phytoplankton Yield, Assessed by Chl-a And Biovolume
During the late 1980s large water level fluctuations in Alte Donau (Fig. 5 in Chap. 5,
see further Chap. 6, Donabaum et al. 2004) were associated with a considerable
nutrient input from contaminated groundwater and sewage. During this period, high
nutrient concentrations were observed in the Danube River (TP 238 μg L−1, total
nitrogen 2.53 mg L−1) which is in the close vicinity of Neue Donau and Alte Donau
(Dokulil and Janauer 1990). The small lake surface, the low ‘relative depth’ and
large inputs of contaminated groundwater characterize Alte Donau as groundwater-
seepage fed ecosystem acting similar to those of riverine lakes as highly productive
lakes under hypertrophic conditions (Teubner 1996; Teubner et al. 1999). Such
lakes are known for their low phosphorus retention (Brett and Benjamin 2008; Kõiv
et al. 2011). As reported in detail in the eutrophication history in Chap. 5, an inte-
grated lake management plan included external (e.g. completion of remediation
wells and sewage network for allotment gardens and other arrears in close neigh-
bourhood in 1995) and internal measures (e.g., initial chemical RIPLOX-treatment
in 1995 and 1996, Fig. 5.3 in Chap. 5) to reduce phosphorus loading from external
and internal sources.
According to a multi-lake study by Jeppesen et al. (2005), the internal phospho-
rus load recovery is about 10–15 years when internal abiotic and biotic lake pro-
cesses are shifting towards an efficient phosphorus cycling system and at the same
time are diminishing the phosphorus availability for phytoplankton growth. The
phytoplankton composition analysed from another long-term multi-lake recovery
survey shifted after 15 years from an assemblage mirroring a phosphorus enriched
environment to an assemblage occurring in an environment of low phosphorus pool
(Anneville et al. 2005). Although the phosphorus pool has already decreased the
phytoplankton yield response is often delayed by a few years during internal lake
recovery (Dokulil and Teubner 2005, see also hysteresis in Fig. 20.4 in Chap. 20).
In Alte Donau, mainly two in-lake treatment measures, the chemical phosphorus
flocculation associated with nitrate oxidation of the sediment by RIPLOX-treatment
(RIPLOX-scheme see Fig. 5.3 in Chap. 5, sediment analysis in Chap. 7) and the re-
establishment of the reed-belt and submerged vegetation (Chaps. 8 and 18), were
aimed at accelerating the internal load recovery in Alte Donau, which shifted this
urban lake from a hypertrophic state in year 1994 to a moderately eutrophic state in
years 1995/96, and to a mesotrophic state in year 2000 (total phosphorus loading as
function of the hydraulic load see Fig. 1.1 in Chap. 1). After this five-year in-lake
management (1995–1999), the lake remained mesotrophic and reached even lower
levels of moderate mesotrophic conditions.
Triggered by chemical phosphorus precipitation, the total phosphorus concentra-
tion of about 70 μg L−1 (2.37 μmol L−1) before the restoration (hypertrophic)
decreased to about 30 μg L−1 (0.89 μmol L−1) in 1995 and 20 μg L−1 (0.6 μmol L−1)
in 1996 (moderately eutrophic). Forced by this drastic decline of TP, the planktonic
community adjusted promptly to efficiently utilising the remaining phosphorus
sources. According to an earlier study by Teubner et al. (2003b) for the years 1994–
1996, planktonic biota acted as a sink for phosphorus at lowered TP pool in two
ways: a) with the tighter coupling between planktonic food and consumer organ-
isms, a larger portion of phosphorus was bound by biota at the expense of the dis-
solved P-fractions, and b) with the compositional shifts from an N-rich towards a
P-rich biota when assessing stoichiometry of plankton community (Teubner et al.
2003b). The zooplankton to phytoplankton ratio markedly increased after the
RIPLOX treatment and stands for a close coupling of zooplankton and their food
(Fig. 11.6 in Chap. 11, see also Teubner et al. 2003b). The increase of this ratio was
mainly due to the low biovolume of phytoplankton under P-limited conditions as
many short-lived primary producers often respond promptly to drastic nutrient
reduction (e.g., Jeppesen et al. 2000; Ibelings et al. 2007). Such an increase of zoo-
plankton relative to phytoplankton is commonly observed in lakes or particular lake
sites at declining nutrient levels (e.g., Padisák 1993; Teubner et al. 2003b, Jeppesen
et al. 2000, 2005; Moustaka-Gouni et al. 2006; Wu et al. 2007; He et al. 2017).
A spontaneous growth of submerged macrophytes was stimulated by enhanced
light availability at low phytoplankton yield after past RIPLOX treatment (see time
series of phytoplankton and macrophyte yield in Fig. 20.2 in Chap. 20). In addition,
the successful re-planting of the reed belt and submerged macrophytes (Chaps. 8
and 18) contributed further to reduce the phosphorus availability for phytoplankton
growth from year 2000 onwards. According to the monograph on Alte Donau
1986/87 by Löffler (ed, 1988), the TP plant tissue content varied between sub-
merged species and sampling sites and ranged for individual measurements from
0.52 to 3.6 mg FW g−1 (mean values: 1.54 mg FW g−1 for Myriophyllum spicatum,
1.51 mg FW g−1 for species of Characeae). The long stems of M. spicatum, that are
close to the water surface, are removed by moving which is necessary in some rec-
reational areas used for swimming and boating. Mowing thus further contributes to
some extent to eliminate phosphorus from Alte Donau. The impact of accidentally
removing fish through mowing is described in Chap. 15. In a longer perspective, the
aim is to further increase the standing crop of macrophytes while successively
replacing tall-growing macrophyte species (e.g. Myriophyllum, Figs. 11.8 and 8.8,
plant biomass in Fig. 8.10) by short-growing macrophytes inhabiting deeper layers
(species of Characeae, Fig. 8.8) (see Chaps. 8, 18 and 20). The recovery of macro-
phytes, whether by exclosures or cage planting or by whole-lake stimulated growth
due increased underwater light availability forced by biomanipulation, is seen as a
successful restoration tool in addition to or instead of common fish manipulation for
stabilising a high water quality in shallow to deep lakes (e.g., Ozimek et al. 1990;
Jeppesen et al. 2000; Qiu et al. 2001; Ibelings et al. 2007; Hilt et al. 2010; Wu et al.
2014; Phillips et al. 2016). Beside the discussed two main in-lake management mea-
sures, the more balanced water fluctuation and the longer retention time (methods,
see also Chaps. 2, 3, 5 and 6) might have contributed to stabilizing the low phospho-
rus availability and subsequently the successful re-establishment of macrophytes in
recent years. The persistence of mesotrophic conditions for the 15 recent years of
our long-term study Alte Donau stands for the sustained shifting towards a healthy
ecosystem (Costanza and Mageau 1999; Dokulil and Teubner 2010).
The dimensionality of ecosystem response is often reduced to the relationship
between phytoplankton yield and concentration of the total pool of phosphorus, the
most limiting nutrient element in freshwaters. This relationship is commonly used
to develop a trophic classification scheme for identifying the overall success of res-
toration measures or the trophic state in lakes (e.g., Vollenweider 1968; Carlson
1977; Forsberg and Ryding 1980; McCauley et al. 1989; Reynolds 1992; Watson
et al. 1992; Teubner and Dokulil 2002; Dokulil and Teubner 2003; Lepistö et al.
2006; Søndergaard et al. 2011; Qin et al. 2013). TP vs Chl-a graphs are also shown
in other chapters to introduce (Chap. 1) or summarize the main idea of restoration
(Chap. 20) in Alte Donau although data were treated differently than in this phyto-
plankton chapter (see methods). According to our analysis, the relationship between
phytoplankton chl-a or phytoplankton biovolume versus TP roughly provides the
same main picture of ecosystem shift from a hypertrophic state in 1993 to a meso-
trophic state in 2014. Taking into account the high variability of chl-a content in
phytoplankton biomass, it might be a rather counterintuitive result that both the
chl-a:TP relationship (p < 0.01) and biovolume:TP relationship (p < 0.05) are statis-
tically significant when analysing the empirical data on a double logarithmic scale.
Chl-a is a ubiquitous photosynthetic pigment occurring in phytoplankton of all tax-
onomic affiliations (e.g., Tolstoy 1979; Bricaud et al. 1995; Richardson et al. 1983;
Donabaum 1992) and is thus used as a proxy for phytoplankton yield. The median
of chl-a wet weight phytoplankton biomass for the whole data record in Alte Donau
confirms the average chlorophyll content of 0.50% wet weight phytoplankton bio-
mass reported for a large data set across trophic gradient by Kasprzak et al. (2008)
and a large number of cultures including cyanobacteria and eukaryotic taxa (chloro-
phytes, cryptophytes, bacillariophytes and ochrophytes) by Donabaum (1992).
Fluctuations in the chl-a content per biomass or biovolume occur along different
time scales of acclimation and adaptation (Geider and MacIntyre 2002).
Photoacclimation often results in a relative increase of chl-a yield per cell or per
carbon (also measured by an increase of both the size and number of photosynthetic
units) for algae that are adjusted to a low light environment while light saturation
commonly leads to the opposite (e.g., Geider et al. 1997; Felip and Catalan 2000;
Teubner et al. 2001; Schagerl and Müller 2006; Grant and Louda 2010; McKew
et al. 2013). Furthermore, chl-a content and light absorption by chl-a increase in
response to fluctuating light when compared with constant light (e.g. for cyanobac-
teria see Nicklisch 1998; Nicklisch and Fietz 2001; Shatwell et al. 2012). According
to allometric rules, the cellular chl-a content per biomass is relatively high for small
cell dominated assemblages compared to large cells (e.g., Richardson et al. 1983;
Bricaud et al. 1995; Vörös and Padisák 1991; Kalchev et al. 1996; Woitke et al.
1996; Felip and Catalan 2000; Teubner et al. 2001). The impact of adaptation is
most relevant when species composition changes as the chl-a content per phyto-
plankton biomass depends on the photosynthetic apparatus that is different among
taxa of various affiliations. The elevated chl-a content during periods of relative
high contribution of chlorophytes (0.83% wet weight phytoplankton biomass) in
Alte Donau relies on the especially high contribution of this photosynthetic pigment
in chlorophyte species (e.g., Donabaum 1992; Greisberger and Teubner 2007). In
turn, cyanobacteria with photosynthetic phycobilines, which are contributing to
light harvesting in addition to chl-a, are well known to rely on low chl-a content per
biomass or biovolume yield (Feuillade and Davies 1994; Donabaum 1992; Schagerl
and Müller 2006; Schagerl and Donabaum 2003; Greisberger and Teubner 2007),
which thus corresponds to lowest chl-a content (0.19% wet weight phytoplankton
biomass) during cyanobacterial blooms in Alte Donau (time series of chl-a content
per wet weight of phytoplankton see also Dokulil et al. 2007). In this view, biomass
phytoplankton yields that are composed of large-celled cyanobacteria and exposed
to saturating light conditions might tend to be underestimated by chl-a measure-
ments. Likewise, the biomass yield of small-sized chlorophytes inhabiting depth
layers of low light availability might usually be over estimated by chl-a. Facing the
different time scales effecting alterations of chl-a (Geider and MacIntyre 2002),
namely rather short-term acclimation due to an adjustment within minutes to hours
lasting for hours to days or long-term adaptation by species shift due to growth
within larger time-sales from 1 day to weeks, a change of chl-a might not be neces-
sarily linked to biovolume increase of phytoplankton. Felip and Catalan (2000)
describe the decoupling between the peak chl-a concentration and the peak biovol-
ume during seasonal succession of phytoplankton. When plotting the compositional
shift between phytoplankton species (e.g. expressed by a similarity index as shown
in Fig. 9.4b) versus the net growth of chl-a (see Fig. 9.4a), the data points follow a
dome-shaped frequency distribution. The common pattern is that the high persis-
tence of species in planktonic assemblages is related to almost zero net changes of

biomass yield, while pronounced changes in species composition are associated
with high values of positive or negative net growth rates (see Fig. 7 in Teubner et al.
2003b, Fig. 2d in Sonntag et al. 2006, the same is valid for Alte Donau but is not
graphically shown). Furthermore, highest stability of phytoplankton species com-
position is often associated with chl-a net change rates ranging between zero and
slightly positive values and thus indicate that small growth success best secure the
stability of species composition (multi-lake study by Teubner, unpublished). Both
examples illustrate that chl-a and phytoplankton biovolume emphasize different
aspects of organism behavior. While the yield of biomass or biovolume stands for
growth, the chl-a concentration mirrors both the phytoplankton yield and the adjust-
ment of primary producers to their environment via photoacclimation. In this view,
chl-a might mirror a more complex response of biota adjusting to their environment
than just phytoplankton biovolume. These arguments are in agreement with our
empirical results from Alte Donau suggesting that the chl-a vs TP relationship
seems more robust than biovolume vs TP (see different levels of significance of both
correlations). This finding might be confirmed by other studies that commonly
choose phytoplankton chl-a (Vollenweider 1968; Carlson 1977; Forsberg and
Ryding 1980; McCauley et al. 1989; Reynolds 1992; Watson et al. 1992; Teubner
and Dokulil 2002; Chen et al. 2003; Dokulil and Teubner 2003; Lepistö et al. 2006;
Søndergaard et al. 2011; Qin et al. 2013) over phytoplankton biovolume (e.g.
Dokulil and Teubner 2005) for the response variable to the total phosphorus pool.
Many studies emphasize the logistic shape of the Chl-a to TP relationship (e.g.,
McCauley et al. 1989; Watson et al. 1992; Dokulil and Teubner 2003; Donabaum
et al. 2004). This sigmoid curve is mirroring an exponential growth at low TP con-
centrations but saturates at high TP. The latter is often discussed in view of a co-
limitation, i.e. that an environmental stimulus other than phosphorus such as light or
further macronutrient elements (nitrogen and silica) are not sufficiently available
for enabling further growth at relatively high TP levels (e.g., Vollenweider 1968;
Forsberg and Ryding 1980; McCauley et al. 1989; Teubner and Dokulil 2002). In
case of Alte Donau, the log-log-transformed Chl-a:TP or biovolume:TP relationship
does not follow the shape of a sigmoid curve and is thus different from multi-lake
studies covering a much wider range of trophic levels (McCauley et al. 1989;
Watson et al. 1992; Dokulil and Teubner 2003).
9.5 Conclusions
The main compositional shift from cyanobacterial dominance under hypertrophic

conditions to a balanced mixture of cyanobacteria, diatoms, green algae, golden
algae and cryptophytes under mesotrophic conditions seems to be mainly driven by
bottom-up control (see integrated lake management plan including external and and
internal measures; Chaps. 5, 8 and 18, 19, 20). The drastic decline in phytoplankton
yield responded to a sustainable reduction of the total phosphorus pool stimulated
by (1) the initial phosphorus precipitation in association with sediment oxidation by
RIPLOX-treatment and (2) the persistent low phosphorus availability in subsequent

years due to the successful re-establishment of underwater vegetation. The hyper-
trophic situation in 1994 was characterised by 35 μg L−1 mean phytoplankton chl-a
in summer, 63 μg L−1 annual mean for TP, 0.85 m annual mean for Secchi depth
(0.69 m summer Secchi depth), about 3 t dry weight of submerged macrophytes
(Fig. 8.1 in Chap. 8), 6157 kg total fish stocking (40% predatory fish) and 5599 kg
total fish catch (37% predatory fish, fish data see further Chap. 15). The period of
sustained lake management is mirrored by the stabilised situation during the last
8 years of observation (2007–2014) with 5 μg L−1 mean phytoplankton chl-a in
summer, 10 μg L−1annual mean for TP, 3.7 m annual mean Secchi depth (3.0 m
mean summer Secchi depth), about 365 t annual dry weight of submerged macro-
phytes (Fig. 8.11 in Chap. 8), 4347 kg annual fish stocking (21% predatory fish) and
4314 kg annual fish catch (31% predatory fish, fish data see further Chap. 15 and
Fig. 20.7 in Chap. 20). The control of phytoplankton yields effected biota from
bacteria to macrozoobenthos as described in other chapters of this book. The sup-
pression of filter-feeding herbivorous cladocerans after reduced algal food supply
and the dominance of mainly selective-feeding omnivorous and herbivorous cope-
pods, which survived under mesotrophic transparent-water conditions, is one exam-
ple of how strongly declined phytoplankton yields impacted planktonic assemblage
(Chap. 11). Different from many other lake restoration projects described in litera-
ture, fish was not removed as cyprinid-fishing is most popular in this urban lake.
Alte Donau thus provides an example of successful and sustained lake restoration
mainly accomplished by bottom up (resource-driven) control rather than by top
down (consumer-driven) control.
Acknowledgments We thank David Livingstone and Susanne Wilhelm for valuable comments
on methods for data interpolation and analyzing time series records during European Union proj-
ects REFLECT (http://www.ife.ac.uk/reflect/) and CLIME (http://clime.tkk.fi/) that were useful
for data treatment in Alte Donau. We thank all of the numerous collaborators and the Municipal
Department for permission of publication. We further want to thank the ‘Wiener Fischereiausschuss’
(Austrian Fishery Association) for providing long-term fish catch records, Franz Wagner and
Adrian Boland-Thoms for helpful comments. The long-term lake measurements were financially
supported by Municipal Department – 45 (Water Management - Vienna). ‘Österreichisches
Komitee Donauforschung, Internationale Arbeitsgemeinschaft Donauforschung’ partly funded
data assessment (K.T.). Further data evaluation (I.T.) was partly funded by the TU Wien Science
award 2015 received by Wouter Dorigo (EOWAVE).
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Chapter 10
Phytoplankton Photosynthesis and
Production
Martin T. Dokulil and Wilfried Kabas
Abstract Productivity of aquatic vegetation determines the trophic level of any

freshwater ecosystem. Phytoplankton photosynthetic rates are particularly relevant.
Results are reported here on photosynthetic rates, primary productivity and associ-
ated parameters of phytoplankton from a polymictic, groundwater seepage lake in
an urban environment before, during and after restoration measures. In addition, a
simple regression model is presented to approximate daily column production from
column integrated chlorophyll-a measurements. Calculated and estimates phyto-
plankton annual lake production is compared to production by submerged vascular
plants. Results indicate that macrophytes played an essential role during the clear
water phase preceding the eutrophication phase associated with intense algal pro-
ductivity and vanishing submersed plant production. Internal restoration measures
led to rapidly decreasing phytoplankton production and slowly re-appearing macro-
phytes. The rehabilitation phase following this period was characterized by declin-
ing phytoplankton productivity and re-establishing of macrophyte production. Total
lake production as the sum of phytoplankton and macrophyte production declined
from 626 t C to 186 t C during 1993–1996 mainly as a result of declining plankton
production. During rehabilitation phytoplankton production further declined and
macrophyte production regained importance. The interannual variability was
ascribed to changes in the winter North Atlantic Oscillation Index. The index sig-
nificantly correlated with lake production explaining 48% of the variability.
Keywords Productivity · Model · Algae · Submersed production · Vascular plants ·

Phytoplankton
M. T. Dokulil (*)
e-mail: martin.dokulil@univie.ac.at
W. Kabas
Wien, Austria

https://doi.org/10.1007/978-3-319-93270-5_10
150 M. T. Dokulil and W. Kabas
10.1 Introduction
Productivity of aquatic vegetation determines the trophic level of any freshwater

ecosystem. Of particular importance is phytoplankton, which usually contributes
the most to the total production of an ecosystem (Falkowski and Raven 2007).
Nutrient enrichment, during the process of eutrophication, enhances productivity
and, as a consequence, algal growth in the pelagic ultimately leads to algal blooms
(Dokulil 2014b). In shallow lakes, the growth balance between phytoplankton and
higher vegetation is central to the understanding of such systems (Chap. 1, see also
Dokulil et al. 2006, 2011). Annual production of submerged higher plants can be
approximated from a single census of standing crop. Productivity of micro-algae is
usually estimated from short-term measurements of photosynthetic rates because of
the much higher turnover (Dokulil 2009). Results are reported here on photosyn-
thetic rates, primary productivity and associated parameters of phytoplankton from
a polymictic, groundwater seepage lake in an urban environment before, during and
after restoration measures (schedule of restoration measure see Chap. 5). In addi-
tion, a simple regression model is presented to approximate daily column produc-
tion from column integrated chlorophyll-a measurements. Calculated and estimates
phytoplankton annual lake production is compared to production by submerged
vascular plants.
10.2 Methods
Here we summarize results of studies on primary productivity conducted between

1993 and 2011 in the shallow urban lake, Alte Donau. Data were extracted from
Rivas (1993), Roschitz (1996), and Kabas (1997, 2004) which contain also all meth-
odological details. Photosynthetic rate profiles were obtained either directly from in
situ incubations or calculated from laboratory simulations (for details see Kabas
2004). All productivity data are reported as carbon uptake. Productivity measure-
ments for the years 1993–1995 were derived from the oxygen technique and con-
verted to carbon uptake applying a factor of 3 (assuming a PQ of 1.12). The derived
photosynthetic parameters alpha (α), a measure of energy efficiency and Ek, the
light parameter describing the onset of saturation were derived from P/E curves
relating chlorophyll-specific rates (P*) to photosynthetic active radiation (PAR).
Vertical profiles were integrated over the euphotic zone and multiplied by the factor
PARday/PARinc to calculate water column production per day. These daily rates were
then integrated over a year to derive annual primary production (Dokulil 1984;
Dokulil et al. 2005; Dokulil and Kaiblinger 2009). Chlorophyll-a was estimated
spectrophotometrically after extraction in hot 90% ethanol (ISO 10260 1992). The
year 1993 was omitted from inter-annual analysis because of an incomplete data set.
All data were tested for normality, constant variance and power before running sta-
tistical tests using SigmaStat, Version 3.11. Definitions of variables and their inter-
pretation followed Sakshaug et al. (1997).
10 Phytoplankton Photosynthesis and Production 151
10.3.1 V
ertical Profiles of Photosynthetic Rates and P/E
Curves
Two vertical profiles of photosynthetic rates were selected to characterise the envi-
ronmental conditions before and after the regime shift (Fig. 10.1). Both were
obtained approximately at the same time in August under similar radiation condi-
tions. During the algal dominated stage, exemplified by the profile from 1993
(Fig. 10.1a), light was rapidly attenuated. Accordingly photosynthetic activity was
restricted to the top two meters. The maximum rate (Pmax) occurred at 0.5 m and was
A) P [mg C m-3 h-1] B) P [mg C m-3 h-1]

0 50 100 150 0 50 100 150
0
E E
1
Depth [m]
3
1993 1998
Pmax = 3.32 Pmax = 1.39
3 EK = 200 EK = 220
P* [mg C mg chl-a-1 h-1]
a = 0.017 a = 0.006
0
0 500 1000 1500 0 500 1000 1500
C) E [ µmol m-2 s-1] D) E [ µmol m-2 s-1]
Fig. 10.1 Examples of vertical photosynthetic rate profiles as carbon uptake during summer
before restoration (a, 1993) and after restoration (b, 1998). Vertical light energy profiles are
inserted into the upper panel. Lower panel: Chlorophyll-a specific carbon uptake versus light
energy (P/E curves) for 1993 (c) and 1998 (d)
six times higher than the rate in 1998 (176 and 29 mg C m−3 h−1 respectively). In the
years following the phosphorus stripping, the euphotic zone extended down to the
bottom (Fig. 10.1b), because chlorophyll levels were drastically reduced.
Photosynthetic rates became smaller and the maximum rate occurred at greater
depth (1.8 m in Fig. 10.1b). Accordingly, chlorophyll specific rates of carbon uptake
(P*) and derived parameters changed significantly (Fig. 10.1c, d). Maximum spe-
*
cific rate Pmax decreased by 60% while the saturation parameter Ek remained near
200 μmol photons m−2 s−1. The light utilization efficiency α decreased from 0.017 to
0.006 mg C (mg Chl-a)−1 (μmol m−2 s−1)−1, which suggests switch from low light
acclimation to assemblages better coping with higher light intensities due to greater
transparency.
10.3.2 Inter-annual Variation
Seasonal cycles of chlorophyll-a (Chl-a), maximum photosynthetic rates (Pmax),

chlorophyll specific productivity (P*) and daily column production (ΣΣP) all
decreased during the investigation period (Fig. 10.2). Highest rates occurred in the
pre-treatment years 1993/1994 (data for 1993 are not shown). The inter-annual wax
and wane of Chl-a concentrations was closely reflected in the Pmax-values. In turn,
daily production in the water column was highly dependent on maximum rates.
Primary productivity per unit chlorophyll-a (P*) was lower in 1996 and 1997 (aver-
age 1.3 and 1.5 mg C (mg Chl-a)−1 h−1 respectively) compared to all other years
where values ranged from 2.2 to 2.5 (Fig. 10.2c). Statistical data on measured vari-
ables are reported in Table 10.1. Mean and ranges of Chl-a, Pmax and ΣΣP clearly
indicate eutrophic to mesotrophic conditions typical for temperate lakes (Dokulil
2014a; Dokulil and Kaiblinger 2009; Dokulil et al. 2005; Lewis 2011).
The light saturation parameter Ek generally decreased from 1994 to 2001
(Fig. 10.3a, see also Table 10.2). Photosynthesis saturated on average at 105 μmol
photons m−2 s−1 (Table 10.1). Light utilisation efficiency α was highest in 1994
(average 0.074) indicating light acclimation to low radiation when cyanobacteria
were blooming (Fig. 10.3b, see also Table 10.2). The dominant cyanobacterium in
1994 was Cylindrospermopsis raciborskii contributing to the ‘self-shading effect’
of phytoplankton. The adjustment to low light intensities, however, enabled this
cyanobacterium to form dense blooms with a peak biovolume of more than
50 mm3 L−1 in Alte Donau (Dokulil 2015). An adjustment to low underwater light
climate was also found for phytoplankton in other lakes facing high inorganic tur-
bidity caused by plumes, e.g. from tributaries or industrial effluents (e.g., alpine
Traunsee in Teubner et al. 2001; Teubner and Dokulil 2002). In Alte Donau, mean
α-values dropped to 0.016 during the restoration phase in the years 1995 and 1996
corresponding to the greater light availability in the water column. The recovery
years from 1997 to 2001 were less variable and characterised by an average α of
0.028 and an Ek of 89. Descriptive statistics for α and Ek are summarised in
Table 10.2 for the three periods.
120
A
100
Chl-a [mg m-3]

80
20
0
B
250
Pmax [mg C m h ]
-1
200
-3
150
100
50
6
0
P* [mg C mg Chl-a h ]
C
-1
5
-1
4
3
2
1
0
D
4
SSP [g C m d ]
-1
-2
0
1994 1995 1996 1997 1998 1999 2000 2001
Fig. 10.2 Long-term seasonality for chlorophyll-a (a), maximum hourly photosynthetic carbon
uptake rates (b), chlorophyll-a specific productivity (c) and daily water column production (d) for
the years 1994–2001
Table 10.1 Summary statistics for the variables and major derived parameters for 1994–2001
Variable Size Mean C.I. Range Max Min Median
SWT [°C] 145 15.19 1.14 25.60 26.40 0.80 16.50
Eo´ [μmol m−2 s−1] 145 960 90 1817 1870 53 935
ΣE [Mol m−2 d−1]_ 145 31 3 63 64 1 31
Chl-a [mg m−3] 145 28.92 5.90 257.95 259.54 1.58 19.19
Pmax [mg C m−3 h−1] 145 12.42 1.84 107.07 108.19 1.12 9.41
P* [mg C (mg Chl-a)−1 h−1] 2.09 0.16 5.13 5.25 0.11 2.02
ΣΣP [g C m−2 d−1] 145 0.64 0.10 4.60 4.60 0.003 0.54
Ek [μmol m−2 s−1] 145 105 11 367 378 11 89
α [mg C (mg Chl-a)−1 E−1] 145 0.0292 0.0040 0.1890 0.1900 0.0012 0.0222
400
A
Ek [µmol photons m s ]
-1
-2
300
200
100
0
Alpha [mg C mg Chl (µmol m s )]
-2 -1
0.05
-1
0.10
0.15
B
0.20
1994 1995 1996 1997 1998 1999 2000 2001
Fig. 10.3 Long-term development of the light saturation parameter Ek and the light utilization
efficiency α for 1994–2001. Dotted lines separate the eutrophic year 1994 from the restoration
years 1995/1996 and from rehabilitation years (post 1996)
Table 10.2 Descriptive statistics for light saturation Ek (μmol photons m−2 s−1) and the efficiency
parameter α [mg C (mg Chl-a)−1 E−1] separated for the three periods (94 before, 95–96 during and
97–01 after restoration)
Variable Size Mean C.I. Range Max Min Median
Ek 94 12 123 58 312 324 12 111
Ek 95–96 34 146 31 360 378 18 139
Ek 97–01 99 89 9 240 251 11 78
α 94 12 0.0744 0.0306 0.1770 0.1900 0.0136 0.0595
α 95–96 34 0.0160 0.0039 0.0512 0.0534 0.0022 0.0138
α 97–01 99 0.0282 0.0033 0.1160 0.1170 0.0012 0.0251
c.i. = confidence interval of the mean
10.3.3 Factors Controlling Photosynthetic Rates
The maximum photosynthetic rate Pmax, was actively controlled by three main fac-
tors. Variation in chlorophyll-a concentration explained 60% of the variability of the
maximum photosynthetic rate (Fig. 10.2 and Table 10.3). Lake surface temperature
and the daily integral of photosynthetic active radiation (PAR) impinging on the
lakes surface explained 35% and 29% respectively (Fig. 10.4a, c). A linear combi-
nation of the three variables increased total explained variability in Pmax to 77%
(Table 10.3). In conclusion, temperature and radiation are largely responsible for
Table 10.3 Statistical data for the relations of variables shown in Fig. 10.4 and for maximum
photosynthetic rate (Pmax) versus chlorophyll-a (Chl-a) and the multiple linear regression of Pmax
versus Chl-a, PAR (Daily photosynthetically available radiation) and WT (average column water
temperature). All variables tested for normality and constant variance
Equation r2 n F p
lg Pmax = 0.745 + 0.0344 WT 0.352 145 77.5 <0.001
lg Pmax = 0.955 + 0.00993 PAR 0.291 145 76.6 <0.001
lg Pmax = 0.509 + 0.390 lg Ek 0.192 145 66.2 <0.001
lg α = −0.515 − 0.581 lg Ek 0.261 145 50.1 <0.001
lg Pmax = 0.191 + 1.098 lg Chl-a 0.594 145 209.1 <0.001
lg Pmax = −0.0600 + 0.948 lg Chl-a + 0.00327 PAR + 0.0195 WT 0.769 145 156.4 <0.001
the increment in chlorophyll concentration which ultimately leads to higher photo-

synthetic carbon-uptake.
Maximum rates were also weakly correlated with the light saturation parameter
Ek (r2 = 0.19, Fig. 10.4b) demonstrating a tendency of photosynthesis to saturate at
higher irradiance when Pmax rises. As a consequence, the two photosynthetic param-
eters Ek and α were negatively correlated (Fig. 10.4d) indicating less efficient light
harvesting at low light intensities when photosynthetic rates saturate at higher light
intensities. In other words, light acclimation in general changes from low light to
higher irradiance when maximum rates of photosynthesis increase. Statistical data
on these relations are summarised in Table 10.3.
10.3.4 Annual Primary Production
Annual primary production (APP) was calculated for the years from 1993 to 2001
from column integrated rates shown in Fig. 10.2d assembled from data in Kabas
(1997, 2004) and Roschitz (1996) and augmented by data from Rivas (1993).
Similarly, yearly chlorophyll-a values were derived by integrating euphotic zone
concentrations over time. From these two parameters annual chlorophyll-specific
photosynthetic rates (P*) were calculated. Results, differentiated for the various
periods, are depicted in Fig. 10.5.
During the eutrophic phase, annual chlorophyll-a concentration under a square-
metre of lake surface exceeded 30 g in 1993. Water exchange in winter 1993/1994
(comp. Chap. 5) brought the yearly value down to 24 g in 1994. Correspondingly,
primary production dropped from 429 to 352 g C m−2 y−1. These values indicated
eutrophic conditions regardless which classification was used (see Dokulil 2014a,
Tables on p. 106; see also washout effect discussed in Chap. 11). Annual chlorophyll-
specific production, however, increased from 13.0 to 14.7 g C (g Chl-a)−1 as a result
of the changes in species composition of phytoplankton (Chap. 9) and light condi-
tions during the first 6 months of 1994. Restoration procedures in 1995 and 1996
drastically reduced all parameters, but the 2 years strongly differed (Fig. 10.5). As a
result of the halving of Chl-a and APP in 1995, P* remained at 12.4 g C (g Chl-a)−1.
A r² = 0.35 B r² = 0.19
100 p <0.001, n = 145 p <0.05, n = 144
h ]
Pmax [mg C m-3 h-1]
-3 -1
100
Pmax [mg C m
10
10
1 1
0 5 10 15 20 25 30 10 100 1000
Temperature [°C] Ek [µmol photons m-2 s-1]
C r² = 0.29
Alpha [a, mg C mg Chl-a-1 (µmol m-2 s-1)] D r² = 0.26
p <0.001, n = 145 p <0.001, n = 144
Pmax [mg C m-3 h-1]
0.1
0.01
0.1 0.001
0 10 20 30 40 50 60 70 10 100 1000
-2 -2 s-1]
PAR [mol photons m d-1] Ek [µmol photons m
Fig. 10.4 Dependence of maximum photosynthetic rates (Pmax) on (a) surface water temperature,
(b) photosynthetic active radiation (PAR) and (c) saturation parameter Ek. The relation of Ek to the
light utilisation efficiency α is shown in (d)
Improvements in under-water light attenuation (compare with Fig. 10.1) and altera-

tions in the composition of the phytoplankton assemblage in 1996 (Chap. 9) led to
an almost further halving in APP to 128 g C m−2 y−1 corresponding to meso- or
meso-eutrophic conditions depending on which classification is used (Dokulil
2014a) Chl-a did not concomitantly drop. Consequently P* declined to 6.7 g
(C g Chl-a)−1. In contrast to the characteristics of primary production, the bacterial
secondary production has hardly changed (see Sect. 13.2.1 in Chap. 13, see also
Teubner et al. 2003). The primary production during the post-restoration years from
1997 to 2011 are characterised by ranges of 19–21 g Chl-a, 67–164 g C APP, and
4.9–12.6 P*. These years classify as mesotrophic. The large yearly fluctuations in
P*, cumulating in 2000 (Fig. 10.5c), are mainly driven by the larger variation in
Chl-a (35%) relative to APP (10%). The between-year variability of chlorophyll-a
can be ascribed to alterations in algal taxon contribution varying in their chlorophyll
content.
35
A
Chlorophyll-a [g Chl-a m y ]
-2 -1
30
25
20
15
10
0
Primary production [g C m-2 y-1]
B
400
300
200
100
0
C
14
12
P* [g C Chl-a y-1]
10
-1
8
6
4
2
0
1994 1996 1998 2000 2002 2004 2006 2008 2010 2012
Fig. 10.5 Annual column chlorophyll-a (a), annual column primary production as carbon uptake
(b), and annual chlorophyll-a specific carbon uptake (c) for the years 1993–2011. Indicated are the
following periods: dark gray – before restoration, light gray – restoration RIPLOX-years, hatched –
after restoration; all as direct measurements. Years in black are also for the period after restoration
but values are calculated from a simple model. For further information, refer to the text
For the years 2002–2011 for which no direct measurements of photosynthetic

rates are available, a simple regression model was developed to derive estimates of
annual production from chlorophyll-a. Annual values of Chl-a and APP for the
years 1993–2001 derived from the PP-measurement site presented in Fig. 10.5 were
first linearly related to each other (Fig. 10.6a). The resulting equation
ΣΣP = −74.973 + 14.937 ΣΣChl-a (r2 = 0.76, p < 0.001, n = 9) was used in a second
step to estimate APP from annual chlorophyll calculated individually for measure-
ments from three other stations. The mean and standard deviation of both parame-
ters and their relation is depicted for 1996–2006 in Fig. 10.6b. Values for 1996–2001
allowed comparison to real measurements. The equation derived from actual mea-
1993-2001 measured A B
Primary production [g C m-2 y-1]
SSP = -74.973 + 14.937 SSChl 140

500 r² = 0.76, p < 0.001, n = 9
120
400
100
300 80
1996-2006 calculated
60
200 SSP = 54.366 + 3.856 SSChl
r² = 0.66, p < 0.001, n = 11
40
100 1993-2006 combined equation
20 SSP = -103.528 + 15.497 SSChl
r² = 0.76, p < 0.001, n = 20
0 0
5 10 15 20 25 30 35 5 10 15 20 25 30 35
-2 -1
Chlorophyll-a [g Chl-a m y ]
Fig. 10.6 (a) Measured annual primary production related to annual chlorophyll-a concentration
1993–2001. (b) Calculated annual primary production related to measured annual chlorophyll-a
concentration 1996–2011. Standard deviation is indicated for both variables. Note the different
scales on the Y-axis
surements (Fig. 10.6a) overestimated direct APP calculations on average by 7%

(range − 10 to +32%). The regression based on the calculated values (Fig. 10.6b)
underestimated APP on average by 24% (range − 9 to −34%). Using all data, the
combined equation ΣΣP = −103.528 + 15.497 ΣΣChl-a (r2 = 0.76, p < 0.001, n = 20)
underestimated real values by 6% (range − 24 to +19%).
To stay conservative, the combined equation was used to calculate estimates
of APP for 2002–2011 (Fig. 10.5b). Annual column chlorophyll-a concentrations
declined in the in the period 2002–2011 from almost 16–11.4 g Chl-a m−2 y−1.
Consequently, calculated APP also declined from 143 to 73 g C m−2 y−1, and P*
decreased from 8.9 to 6.4 g C Chl-a−1. In the following years from 2007 to 2011
all three parameters varied and reached higher values in 2009 and 2010
(Fig. 10.5a, b and c).
10.3.5 Lake Primary Production
In a bistable system such as Alte Donau, phytoplankton and macrophyte production

are of equal importance. An estimation of annual primary production for the lake
must include both parameters in identical units. Macrophyte dry-weight was con-
verted to cell carbon assuming a C-content of 35% (Esteves and Suzuki 2010).
Annual phytoplankton carbon production per square meter was amplified using the
lake area and assuming an average euphotic depth of 1 m (Table 10.4). These
approximations of net-production were then used to derive the relation of phyto-
plankton to macrophyte production, and to calculate total primary production for
the lake (Table 10.4).
Table 10.4 Annual lake production for phytoplankton and submersed macrophytes both converted
from biomass to carbon, and the relation of phytoplankton carbon to macrophyte carbon for the
census 1987 (Löffler 1988) and the period 1993–2011
Phyto C Makro C Phyto C/ Annual LPP

Year [t] [t] Makro C C [t]
1987 9 252.4 0.3 321.4
1993 615 12.0 51.2 626.8
1994 504 0.8 649.2 505.2
1995 314 0.1 2488.9 313.7
1996 183 3.4 53.9 186.3
1997 214 4.1 52.7 218.2
1998 230 4.3 53.0 234.2
1999 215 4.9 44.3 220.1
2000 235 5.1 46.2 239.7
2001 230 5.3 43.8 235.2
2002 203 6.8 30.1 210.1
2003 205 43.8 4.7 248.5
2004 127 125.7 1.0 252.9
2005 128 134.4 0.9 262.0
2006 105 124.3 0.8 229.3
2007 96 119.0 0.8 214.9
2008 68 126.0 0.5 194.2
2009 129 131.3 1.0 260.5
2010 129 129.5 1.0 258.7
2011 107 126.0 0.8 233.0
Total annual lake primary production (LPP) as the sum of phytoplankton and macrophyte carbon
is given in the last column. Biomass data for macrophytes derived from Chap. 8
The data from the census in 1987 (Löffler 1988) evidently show the dominance
of submerged macrophytes. The carbon quotient of 0.3 for phytoplankton versus
macrophytes indicates that macrophyte production was at least three times higher
than that of plankton algae. Total carbon accumulation for 1987 was estimated as
321.4 tons carbon. The situation completely reversed when the system switched to
the turbid stage in 1992. Phytoplankton carbon production exceeded carbon accu-
mulation in macrophytes 51 times in 1993 and further increased to almost 2500 in
1995 (Table 10.4). This overwhelming plankton production was the result of rapidly
declining macrophyte biomass despite improvements in algal concentration and
decreasing plankton production due to the restoration measures (Chap. 5).
The annual lake production gradually declined from 626 t C to 186 t C during the
years 1993–1996 mainly as a result of declining plankton production since macro-
phyte contribution was negligible (Table 10.4, see also Fig. 8.1 in Chap. 8). The
post-restoration period between 1997 and 2003 was characterised by a stabilization
of annual planktonic primary production around 219 ± 12.7 C. At the same time,
submerged macrophyte carbon production gradually increased from 1.9 tons in
1997 to 6.9 tons in 2002, reaching 43.8 t C in 2003 corresponding to 17.6% contri-
bution to total lake production. This year marked the break-through in macrophyte
development. In the years to follow, planktonic production further declined while
300
Lake Carbon Production [t C]
250 Lake
200
Phytoplankton
150
100
50
Macrophytes
0
1997 1999 2001 2003 2005 2007 2009 2011
Fig. 10.7 Post-restoration lake primary production for phytoplankton, submerged macrophytes
and the lake; as tons carbon per year
macrophytes gained importance with carbon production varying around 127 ± 4.7 t

C comprising on average 54% of total annual production (Fig. 10.7). Total lake
production varied between 194 and 262 tons carbon (average 234 ± 20.1 t C). These
year to year differences in lake production have been attributed to the mean North
Atlantic Oscillation (NAO) for January and February, used here as a general cli-
matic parameter. The index significantly correlated with lake production explaining
48% of the variability (r2 = 0.48, p = 0.004, n = 15, see also climate response in Sect.
11.3.5 of Chap. 11).
10.4 Conclusions
Phytoplankton photosynthetic rates were characteristic and indicative of the various

phases during regime shift, restoration and rehabilitation in a shallow polymictic
urban lake. Photosynthetic rates were responding to the trophic level. Light accli-
mation to low radiation during the turbid state was indicated by high α-values and
the associated Ek. The greater light availability in the water column during the res-
toration phase led to a drop in light harvesting efficiency. Both parameters were less
variable during the rehabilitation years. The inter-annual variability in Pmax closely
resembled Chl-a concentrations. Daily water-column production in turn was depen-
dent on maximum rates. Therefore daily column production could be predicted
from column integrated chlorophyll-a concentration alone contrary to findings by
Gervais and Behrendt (2003) which had to add radiation to their predictions
equation. It must be emphasised however, that radiation usually is essential (e.g.,

Dokulil 1984, 2014a).
Photosynthetic rates measurements and their associated parameter proved to be
excellent predictors of switches in trophic level as a result of regime shift as well as
indictors for recovery following restoration in an urban lake.
Acknowledgments The authors acknowledge the continuous support from the Municipal

Department – 45 (Water Management – Vienna) and the University of Vienna for providing labora-
tory space. Moreover thanks to the many collaborators during this long-term study.
Appendix
List of Abbreviations:
Abbreviation Description Unit

SWT Surface water temperature °C
WT Average column water temperature °C
PQ Photosynthetic quotient dimensionless
PAR Photosynthetic available radiation μmol m−2 time−1
PARday/PARinc Ratio PAR incubation to PAR daily dimensionless
E o´ Subsurface PAR μmol m−2 s−1
ΣE_ Daily incoming PAR mol m−2 d−1
Chl-a Chlorophyll-a concentration mg m−3
Pmax Maximum photosynthetic rate mg C m−3 h−1
P* Chlorophyll specific rate mg C (mg Chl-a)−1 h−1
ΣΣP Daily euphotic zone production g C m−2 d−1
APP Annual primary production g C m−2 y−1
Ek Light saturation parameter μmol m−2 s−1
α [alpha] Light utilization efficiency mg C (mg Chl-a)−1 E−1
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Oceanogr 48(3):1141–1149
Chapter 11
Response of Zooplankton to Restoration
and Climate Warming in Alte Donau
Katrin Teubner , Monika Großschartner, and Irene E. Teubner
Abstract Lake restoration is commonly assessed by changes in water transpar-

ency, nutrients and biomass of phytoplankton, while information about changes in
zooplankton that is triggered by lake management is often missing. In our 19-year
study we used rotifers and crustaceans to document the effect of restoration mea-
sures on zooplankton in the oxbow lake Alte Donau, a former side-arm of the
Danube River which is most popular for recreation and angler (cyprinid-dominated
shallow water). The record covers four management periods: the period before res-
toration, the restoration (including years of chemical phosphate precipitation by
Riplox treatment), the re-establishment of macrophytes and the sustained ‘stable
conditions’. We found the highest abundance of all zooplankton in the first Riplox-
year, with decreasing zooplankton abundance in following periods associated with
the decline of phytoplankton. In the long term, the main compositional change
related to a shift from a cladoceran-rotifer-rich to a copepod-rotifer-rich zooplank-
ton assemblage. Thus, the large-bodied zooplankton shifted from a community
composed of mainly filter-feeding herbivorous cladocerans under eutrophic algal-
turbid conditions to mainly selective-feeding omnivorous and herbivorous cope-
pods under mesotrophic transparent-water conditions. While the carbon ratio
between zoo- and phytoplankton increased significantly during the first three peri-
ods and remained high under ‘stable conditions’, the mean body size of zooplankton
did not exhibit a long-term trend. Short-term increases of large-bodied zooplankton
coincided with an intermittent increase of calanoid copepod abundance (Eudiaptomus
K. Teubner (*)
M. Großschartner
DWS-Hydro-Ökologie GmbH, Vienna, Austria
e-mail: monika.grossschartner@dws-hydro-oekologie.at
I. E. Teubner
Department of Geodesy and Geoinformation, Faculty of Mathematics and Geoinformation,
Vienna University of Technology, Vienna, Austria
e-mail: irene.teubner@geo.tuwien.ac.at

https://doi.org/10.1007/978-3-319-93270-5_11
gracilis) during the chemical treatment concomitant with a drastic phytoplankton

biomass reduction and the occurrence of large-bodied cladocerans (Simocephalus
vetulus and Sida crystallina) in some years with re-established underwater vegeta-
tion. Besides the main response of zooplankton to the ‘bottom up’ control that was
triggered by the reduction of phytoplankton food supply by one order of magnitude,
we studied the zooplankton response to climate change. The impact of climate
warming was evident from intra-annual coincidence of the climate signal (NAODJFM)
and water temperature (WT) in winter and early spring, the increase of surface
water temperature (SWT) by 1.52 °C per decade in April and the prolongation of the
warm period (SWT > 22 °C) by 10.5 days per decade in summer. This prolongation
of the warm season seemed to support the summer development of the medusa stage
of freshwater jellyfish (Craspedacusta sowerbii). During the transition from spring
to summer, the progressively earlier clear-water phase followed two trends. The first
period with a 33-day earlier clear-water phase per decade coincided with pro-
nounced ecosystem changes from a high to a low eutrophic state created by chemi-
cal restoration measures. The second period with a moderate earlier progression of
7 days per decade was accompanied by a further, slight TP decrease associated with
the re-establishment of macrophytes. When comparing rotifers, cladocerans, cala-
noid and cyclopoid copepods, the latter group benefits most from seasonal tempera-
ture increases and climate warming.
11.1 Introduction
Urban lakes are often altered by humans as they are located in densely populated
areas. Both open water bodies and banks are affected by pollution and artificial bank
stabilisation due to human activities and are thus commonly ‘heavily modified’ or
degraded ecosystems. More recently, however, awareness has increased that urban
ecosystems can, in fact, generate valuable services for urban life (Bolund and
Hunhammar 1999). This awareness has led to a shift in urban planning from a focus
on water exploitation to a focus on ecosystem health, satisfying both nature conser-
vation and the use of waters for recreational and cultural activities. Concepts in
sustainable landscape planning and biomanipulation used as a lake restoration tool
indicate that healthy freshwaters gain supra-regional importance by their efficient
retention of carbon and other nutrient elements (Ripl 1995; Søndergaard et al. 2001;
Reichstein et al. 2013). Lake restoration prevents runoff of nutrients from degraded
urban areas through rivers into the ocean, from where it is difficult to get nutrients
back into terrestrial ecosystems (Ripl 1995). Accelerated circulation of matter
becomes increasingly important during lake restoration, i.e. when a nutrient-rich
lake is facing a drastic reduction of the pool size of essential nutrients (Teubner
et al. 2003). Zooplankton, which is the focus of this chapter, facilitates rapid cycling
of matter in nutrient-poor lakes (Capblancq 1990) and therefore contributes sub-
stantially to retaining nutrients in a healthy ecosystem (Loreau 1995).
11 Response of Zooplankton to Restoration and Climate Warming in Alte Donau 165
Alte Donau in Vienna faced these different aspects of an urban lake mentioned
before. The habitat for cold-water fish assemblages in the Danube River shifted to a
habitat for warm-water fish assemblages after the cut-off from the main river chan-
nel in 1870–1875 (see fish in Chap. 15, history in Chap. 2). During the mesotrophic
period in the 1980s, the oxbow lake was cyprinid-dominated with extensive dense
underwater vegetation (Löffler 1988). In the 1990s, Alte Donau has undergone a
strong nutrient enrichment (Dokulil et al. 2010a). This was accompanied by the
development of large cyanobacterial blooms and a consequently low water transpar-
ency. The highest phytoplankton biomass was measured in 1994. The phytoplank-
ton was dominated by the filamentous cyanobacterium Cylindrospermopsis
raciborskii (Dokulil and Mayer 1996) and reached a chlorophyll-a concentration of
about 140 μg L−1. Concomitantly, the water transparency was lowest with a Secchi
depth of 0.28 m in September 1994 (see Fig. 6.4 in Chap. 6, phytoplankton compo-
sition in Chap. 9). The awareness of the value of urban ecosystems led to actions to
restore Alte Donau (Chap. 5). The restoration comprised different treatments such
as the use of Riplox (1995/1996) to limit the growth of planktonic cyanobacteria
and algae (Chap. 9) and the re-establishment of submerged vegetation for the pur-
pose of sustained restoration by biomanipulation (Chap. 8). The lake restoration
included both the open water bodies and the banks (Chap. 18). Furthermore, a mas-
ter plan of urban planning was elaborated to generate the surrounding recreational
area in order to create a better quality of urban life in Vienna (Chap. 19).
Besides the impact of eutrophication, the recent climate change and its impact on
ecosystems have also led to a growing public awareness of the state of aquatic envi-
ronments. Early climate change studies have focused on identifying and quantifying
the impact of climate change on freshwater and marine ecosystems at local and
global scale. More recent follow up studies have led to the development of future
management strategies for mitigating the risks posed by climate change (IPCC
2014). The vulnerability of ecosystems to climate change is amplified by man-made
degradation of landscape that is most relevant for an urban oxbow lake such as Alte
Donau. Awareness of the risks associated with climate change is here seen in com-
bination with the damage done to ecosystem health. The main threat for ecosystems
is thus not only the direct impact of temperature increase created by global warm-
ing, but also the deterioration of the water cycle, soil and vegetation including habi-
tat loss that are caused and enhanced by anthropogenic eutrophication and landscape
alterations (e.g., Blenckner 2005; Ripl et al. 2007; Jeppesen et al. 2010a; Dokulil
and Teubner 2011; Reichstein et al. 2013). In this view, rivers and lakes are under-
stood as sentinels for landscape degradation and for climate change (e.g., Karr
1998; Verburg et al. 2003; Adrian et al. 2009; Schindler 2009). This chapter focuses
on the change of zooplankton community structure following restoration as well as
on the potential impact of global warming.
Many limnological aspects were studied to understand the long-term develop-
ment of biota in Alte Donau. The zooplankton (this chapter) and the phytoplankton
(Chap. 9) are the two biological plankton parameters monitored for the longest time
in this book. The analysis of the 19-year record of zooplankton aims at describing
the long-term dynamics in species composition and comparing the community
structure between the two large lake basins. Here, we compare the zooplankton
assemblages between four main periods of the ecosystem-based lake management.
These periods cover the eutrophication period before the first chemical treatment in
April 1995, the ‘restoration’ period encompassing the first and the second chemical
treatment including further restoration measures, the period of re-establishment of
macrophytes and the period of ‘stable conditions’. In addition, we explore if the
impact of restoration on the development of zooplankton of Alte Donau was super-
imposed by global warming.
11.2 Methods
11.2.1 Sampling During the Four Periods of Lake Treatment
Alte Donau is a polymictic oxbow lake (7 m maximum depth, 2.5 m mean depth),
with a water volume of 3.54 × 106 m3 and a surface area of 1.43 km2 (Dokulil et al.
2010a, for lake history see also Chap. 2). The water body consists of two main
impoundments. The morphology and hydrology of these two basins are described in
Chaps. 3 and 4, respectively.
Zooplankton samples were taken at three depths (0.5 m, 2.5 m and near the bot-
tom at 4.5 m) with a 5 L Schindler sampler in the south basin (AD1) and the north
basin (AD4) and were integrated over depth. The samples were filtered through a
44 μm mesh size net, resuspended in 50 mL lake water and fixed with formalde-
hyde. The zooplankton time series presents a consistent data set as determination of
species and counting of samples was conducted by the same person (M.G.) for the
whole 19-year period. We calculated the biomass of the crustaceans by length to
weight regressions according to McCauley (1984) and Bottrell et al. (1976), assum-
ing a dry to wet weight ratio of 0.15. We derived the biomass of rotifers from mea-
surements of all three dimensions following Ruttner-Kolisko (1977). To compare
the biomass of zooplankton and the biovolume of phytoplankton, we translated the
mass related measures of both plankton groups into carbon content by using the
volume conversion of Behrendt (1990).
Zooplankton samples were taken at biweekly intervals during the observation
period from April 1994 to November 2012. This 19-year time series thus covers the
four main periods of the lake treatment specified in Chap. 6, which are slightly
modified in this chapter: 1 – ‘eutrophication’ (before April 1995), 2 – ‘restoration
including the two years with chemical phosphate precipitation’ (April 1995–1999),
3 – ‘macrophyte re-establishment’ (2000–2006) and 4 – ‘stable conditions’
(2007–2012).
The restoration period covers the treatment with FeCl3 and limestone for phos-
phate precipitation followed by addition of Ca(NO3)2 to enhance nitrate oxidation of
the sediment (Ripl 1976; Donabaum et al. 1999). This two-step chemical treatment
(Riplox) was applied twice, in April 1995 and April 1996 (Chap. 5). An earlier study
on the ecosystem response comparing the ecosystem before the restoration with the
situation during the first and the second Riplox-year revealed different pathways of
prompt responses by the plankton community. It showed that the short-term impact
of chemical treatment in the first year was quite different from that observed during
the second year (Teubner et al. 2003). For this reason, we split the restoration period
in two, i.e. 2a) April 1995 to March 1996 and 2b) April 1996 to 1999. One main
focus of this book chapter is the long-term impact on zooplankton during the five
restoration periods (1, 2a, 2b, 3 and 4).
11.2.2 D
ata Treatment, Statistics and Graphical
Representations Identifying Development Patterns
upon Restoration
Figures 11.1, 11.2, 11.3 and 11.4 are based on the original dataset. However, as
phytoplankton and zooplankton were not always sampled by exactly 2-week inter-
vals, we interpolated the available data at daily resolution (Livingstone 2003; Sapna
et al. 2015) and averaged these over a two-week period. These data were then used
to calculate the zooplankton to phytoplankton carbon ratios and the net change rates
of zooplankton at regular 2-week time intervals (Figs. 11.5 and 11.6). The net
change rate of zooplankton biomass (kzoo, hereafter referred to as net growth rate
of zooplankton) is calculated as
kzoo  ln Zoot2  ln Zoot1 / t (11.1)

where Zoot2 is the zooplankton biomass at time t = t2, Zoot1 the biomass at time
t = t1 and Δt the time span in number of days (e.g., Teubner et al. 2003). The net
growth rate of phytoplankton is calculated analogous to Eq. 11.1 (see also Chap. 9).
The net growth rates of zooplankton and phytoplankton integrated over time are the
cumulative growth rates of the respective plankton group. The timing of the clear-
water phase was defined by the coincidence of low phytoplankton biovolume after
the spring bloom and enhanced Secchi disc depth, indicating high water transpar-
ency during the spring-summer transition.
Prior to the statistical analysis, data were tested for normality. According to the
non-parametric Shapiro-test (Dunn and Clark 1974), not all parameters followed a
normal distribution, in particular not those of single species. We therefore ln-
transformed these data (using natural logarithm) to normal distribution. For better
graphical illustration, we also plotted data on the LN-scale in Figs. 11.2, 11.3, 11.6a
and 11.8A1, B1, C1 and D1. Data that contained the value zero were transformed
by ln(x + 1).
We conducted statistical analyses to identify the differences among sampling

sites (AD1, AD4) and restoration periods using R (R i386 Version 2.15.2). We
applied Kruskal-Wallis tests to determine differences between annual median values
of both time series (indicated by grey bars in Fig. 11.1) and of the five restoration
periods (Figs. 11.3 and 11.4). To identify homogeneous subsets when comparing
the zooplankton assemblages during the five restoration periods, we further con-
ducted pairwise Mann-Whitney UT tests with a Bonferroni correction (Figs. 11.3
and 11.4). The abundances of single species and of taxonomic zooplankton groups
are displayed as notched box-whisker plots using SYSTAT 10 (SPSS Inc.) (Figs. 11.2
and 11.3). The boxes are notched at the median; the length of the notches indicates
the 95% confidence interval. Additionally, we show the change in zooplankton com-
munity structure in triangular plots, assuming that the three large taxonomic groups
comprise the total zooplankton of large and medium-sized species (Fig. 11.4). The
triangular diagrams (Gibbs 1878) depict here the portions among the three zoo-
plankton groups and have the advantage of graphically displaying the points in one
plane, different from a three-dimensional space of a x-y-z Cartesian coordinate sys-
tem. To avoid overlay of data points, we displayed the results in separate triangles
for each restoration period. Application and interpretation of the triangular graphs
in limnology are described in detail in Teubner and Teubner (1998), Teubner and
Dokulil (2002) and Teubner et al. (2003) for nutrient stoichiometry and phytoplank-
ton composition.
11.2.3 Data Treatment, Statistics to Identify Climate Response
Graphs related to the climate response are displayed in Figs. 11.7, 11.8 and 11.9.
The analyses of long-term trends in zooplankton community development, the
potential responses of zooplankton to water temperature and climate signals super-
imposing restoration efforts are based on biweekly data retrieved from linearly
interpolated data as described above (Figs. 11.7 and 11.9). All graphs in Fig. 11.7,
which are related zooplankton abundance, are analysed for the 19-year period
(1994–2012) and then related to the timing of the clear-water phase for 21 years
(1994–2014) and to water temperature for 22 years (1993–2014).
We used the North Atlantic Oscillation (NAO, Hurrell et al. 2001) Index as cli-
mate signal since it is a common proxy for studying the climate impact on aquatic
ecosystems in the temperate zone (NAO station-based, from Hurrell eds. 2015).
We have chosen the NAO signal for the period December – March (NAODJFM, see
Fig. 11.7) as the winter climate signal does not vanish as fast as the NAO signal of
later months (see, e.g., Blenckner et al. 2007; Dokulil et al. 2010b); thus, it is suit-
able for unravelling the climate response in late spring-early summer in Alte Donau.
In accordance with this, we analysed if warming occurs during the main season of
zooplankton development by selecting two dates in late April and early July to
reveal temperature trends during the 19-year period. We further calculated the num-
ber of ‘warm’ lake water days (surface water temperature, SWT, above 22 °C) to
analyse the length of the warm period in summer (Fig. 11.7b). In order to trace the
climate response related to this summer period, we used the annual NAO. Additionally,
we analysed the extreme hot summer period by determining the number of days
exceeding the threshold of 25 °C, as this temperature was the highest integer of the
biweekly averages of SWT (Fig. 11.7b). According to suggestions regarding the
optimal growth temperature for the polyp (19–25 °C) and the medusa development
of Craspedacusta sowerbii derived in culture experiments (Folino-Rorem et al.
2016), we calculated the annual periods with 19–25 °C SWT and above 21 °C SWT
in number of days per year in Alte Donau. All data in Fig. 11.7 (with the exception
of the skewed distribution for the number of extreme hot days above 25 °C SWT)
and Fig. 11.9 satisfy normal distribution. Despite the normal distribution for data in
Fig. 11.7, the respective trends of the year-to-year variation are calculated as robust
lines by non-parametric fitting according to Theil (1950) as described in Helsel and
Hirsch (2002). These robust trend lines are applied to calculate the reliable slope of
the year-to-year trends, which are robust against outliers of unusual high or low
values in the first or last year of observation. The statistical significance of the trends
is calculated by Mann-Kendall tests using R (McLeod 2015). Regarding the year-
to-year variation of the clear-water phase (Fig. 11.7d), a rapid decrease occurred in
the beginning and was followed by a slower decrease in later years. For this reason,
the time series was tested for breakpoints using Davies-test in R (Muggeo 2008,
2015), but the results were not significant (p-value>0.05). Nevertheless, data were
separated into two phases based on the a priori assumption of different stages of
the lake restoration. The segmented two regression lines are shown in Fig. 11.7d.
The first period refers to the rapid phytoplankton biomass decrease occurring in the
years prior to treatment initiation and the restoration treatment period including the
phosphate precipitation (period 1 and 2), while the second period covers the subse-
quent years where only small changes in the phytoplankton yield occurred, i.e. the
periods with macrophyte re-establishment and stable conditions (period 3 and 4;
periods see in methods Sect. 11.2.1 above). In addition, the regression line for the
whole time series is displayed.
11.3 Results
The long-term development of the zooplankton abundance and biomass is shown in

Fig. 11.1a, b for the south (AD1) and the north basin (AD4). This 19-year develop-
ment comprises the five periods of ecosystem-based management (1, 2a/2b, 3 and
4) covering the periods before, during and after chemical phosphorus precipitation
(for a description of these periods see introduction and methods in Sect. 11.2.1).
Zooplankton
6000
5500 North Basin (site AD4) A
Abundance [ind L-1]
3500 South Basin (site AD1)

3000
2500
2000
1500
1000
500
0
2500 North Basin (site AD4)

South Basin (site AD1)
B
Biomass [µg L-1]
2000
1500
1000
500
0
Rot [ind L-1] Cop [ind L ] Clad [ind L-1]
400
Cladocera C
200
100
0
300 Copepoda D
-1
200
100
0
4000
3000 Rotatoria
2000 E
1000
0
30 Cop
Clad F
Number of species
25
Rot
20
15
10
5
1 2a 2b 3 4
0
94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12
Year
Fig. 11.1 Long-term series of zooplankton development comprising five periods: before restora-
tion (1), restoration with chemical Riplox treatment (2a, 2b), re-establishment of macrophytes by
periodical water level drawdown (3) and ‘stable conditions’ (4). Abundance (a) and biomass (b) of
total zooplankton are shown for both sampling sites, the south basin (AD1) and the north basin
(AD4). The abundance of individual taxonomic groups (c: cladocerans, d: copepods, e: rotifers)
and the comparison of species numbers (f) represent averages of AD1 and AD4. Years with signifi-
cant differences between AD1 and AD4 (H-test, p-values. See text) are indicated by dark grey hori-
zontal bars for graphs (a–e)
11.3.1 C
omparison of Zooplankton Abundance and Biomass
in the South and North Basin
When comparing the zooplankton structure year by year between the north and the
south basin, we only found statistically significant differences in the year 1994
(Fig. 11.1a, b; bars indicate years with significant differences between sites; H-test,
p < 0.05 for rotifer abundance, all other p < 0.001). The annual average of zooplank-
ton abundance in the north basin was 2027 individuals per litre in 1994, which cor-
responds to a zooplankton biomass of 899 μg L−1. Zooplankton abundance in the
south basin was about six times lower than in the north basin with annual mean
values of 324 individuals L−1, which corresponds to a biomass of 152 μg L−1.
Furthermore, we found significantly lower values in the south basin for the indi-
vidual taxonomic groups, i.e. cladocerans, copepods and rotifers, in the year 1994
(see abundance in Fig. 11.1c, d, e). These statistically significant differences are
probably mainly due to the lake treatment by water exchange in 1993 which aimed
at improving the water quality in Alte Donau; relatively nutrient-poor water from
Neue Donau (New Danube) was introduced to flush the eutrophied water body of
Alte Donau. The water from the channel Neue Donau entered Alte Donau in the
south end of the south basin (site ‘Seestern’). Thus, we assume that the north basin
rather than the south basin mirrors the state of the zooplankton community during
the eutrophication period in 1994 before the chemical treatment started in 1995. As
these site differences correspond to a different magnitude of zooplankton abun-
dance and biomass, but not to a significantly different species pattern, we did not
treat the data for the two basins separately. Moreover, we did not find between-site
differences in abundance or biomass for either total zooplankton or for individual
zooplankton groups during the subsequent 5-year ‘restoration’ period from 1995 to
1999 (see period 2 in Fig. 11.1). Also, in the next 13 years that included the period
of ‘macrophyte re-establishment’ and ‘stable conditions’, we did not find any statis-
tically significant between-site differences for total zooplankton abundance or bio-
mass. Statistically significant abundances were only detected for some individual
zooplankton groups in particular years. These significant differences are indicated
by grey bars for years 2000, 2001, 2006, 2009 and 2010 in Fig. 11.1c, d, e (H-test,
p < 0.05 for rotifers 2000, p < 0.05 for all others).
The water exchange using water of the neighbouring channel, Neue Donau, in
1993 aiming at improving the water quality in Alte Donau did not influence the
whole water body but only one basin. Furthermore, this water exchange had only a
short-term effect on the zooplankton in 1994, waning in the following years.
Therefore, we conclude that between-site differences in Alte Donau are of minor
importance in describing the main picture of the long-term response of the zoo-
plankton to the lake restoration. For this reason, results are summarised for the
water body as a whole.
11.3.2 Species Composition of Zooplankton
The zooplankton abundance is mainly dominated by small-sized rotifers (Fig. 11.1f,

see also list of species in Table 11.1). On average, we identified 10 rotifer species
with a maximum number of 18 and a minimum number of 3 species during the
19-year investigation period. About 29% of the identified rotifers are known to be
algivorous, another 29% bacterivorous/algivorous and 26% omnivorous. Another
9% of the rotifer species are known to be bacterivorous. Only about 7% of these
microzooplankton are classified as carnivorous. The seven most abundant rotifers
are Keratella cochlearis, Polyarthra dolichoptera-vulgaris, Kellicottia longispina,
Brachionus angularis, Collotheca mutabilis, Gastropus stylifer and Conochilus uni-
cornis (Fig. 11.2).
Larger-sized zooplankton species are cladocerans and copepods which are found
at an average of five and four species, respectively (maximum 8 and 6, Fig. 11.1f;
see species list in Table 11.1). Most of the cladocerans are classified as omnivorous
Table 11.1 List of zooplankton species found from 1993 to 2012

Taxa
Rotifera Anuraeopsis fissa Gosse, Ascomorpha ecaudis Perty, A. ovalis Bergendal, A.
saltans Bartsch, Asplanchna priodonta Gosse, Brachionus angularis Gosse, B.
calyciflorus Pallas, B. diversicornis Daday, B. quadricornis Meissner,
Cephalodella sp., Collotheca mutabilis Hudson, Colurella uncinata Müller,
Conochilus unicornis Rousselet, Encentrum sp., Euchlanis sp., Filinia longiseta
Ehrenberg, F. terminalis Plate, Gastropus stylifer Imhof, Hexarthra mira
Hudson, Kellicottia longispina Kellicott, Keratella cochlearis Gosse, K.
quadrata Müller, Lecane closterocerca Smarda, L. cornuta Müller, L. elongata
Harring & Myers, L. luna Müller, L. lunaris Ehrenberg, Lepadella sp.,
Lophocharis sp., Macrochaetus subquadratus Perty, Monommata sp., Mytilina sp.,
Notholca acuminata Ehrenberg, N. jugosa Gosse, N. squamala Müller,
Pleosoma hudsoni Imhof, Polyarthra dolichoptera-vulgaris Idelson/Carlin,
Pompholyx solcata Hudson, Squatinella sp., Synchaeta pectinata Ehrenberg, S.
tremula-oblonga Ehrenberg, Testudinella patina Hermann, Trichocerca birostris
Minkiewicz, T. capucina Wierzejski et Zacharias, T. cilindrica Imhof, T.
insignis Herrick, T. pusilla Lauterborn, T. similis Wierzejski, Trichotria
pocillum Müller, T. tetractis Ehrenberg
Cladocera Acroperus harpae Baird, Alona affinis Leydig, Alonella nana Baird, Bosmina
longirostris O.F. Müller, Camptocercus rectirostris Schoedler; Ceriodaphnia
quadrangular O.F. Müller, Chydorus sphaericus O.F. Müller, Daphnia cucullata
Sars, D. hyalina Leydig, Diaphanosoma brachyurum Lievin, D. orghidani
Negrea, Disparalona rostrata Koch, Eubosmina coregoni Baird, Eurycercus
lamellatus O.F. Müller, Graptoleberis testudinaria Fisher, Leptodora kindtii
Focke, Pleuroxus truncatus O.F. Müller, Polyphemus pediculus Linne,
Pseudochydorus globosus Baird, Scapholeberis mucronata O.F. Müller, Sida
crystallina O.F. Müller, Simocephalus vetulus O.F. Müller
Copepoda Calanoida: Eudiaptomus gracilis Sars
Cyclopoida: Cyclops vicinus Ulyanin, Eucyclops serrulatus Fischer, Megacyclops
viridis Jurine, Mesocyclops leuckarti Claus, Thermocyclops oithonoides
G.O. Sars, T. crassus Fischer
Eubo core Diap brac Chyd spha Ceri quad

6 4 4 5
4
3
2 2
2
1
0 0 0 0
Lept kind Daph cucu Eudi grac Bosm long
5 5
crustaceans
3
1 3 3
1
1 1
0 0 0 0
Abundance (LN individuals L )
-1
Ther cras Cycl vici Meso leuc Ther oit h
3 3
1.8 3.0
0.9 1.5
1 1
0 0 0 0
Brac angu Aspl prio Coll muta Gast st yl
5 5 5
3
3 3 3
1
1 1 1
0 0 0 0
Kera coch Kell long Poli doli 3 Trich capu
8 6 7
rotifers
6 5 2
4
4
3 1
2 2
1
0 0 0 0
Leca luna Trich pusi Cono unic Trich insi
5
2
1 1 3
1
1
0 0 0 0
1 2a 2b 3 4 1 2a 2b 3 4 1 2a 2b 3 4 1 2a 2b 3 4
Category of treatment period
Fig. 11.2 Box-whisker plots of the abundance of individual species of cladocerans, copepods and
rotifers during the five treatment periods. Cladocerans: Eubo core – Eubosmina coregoni, Diap
brac – Diaphanosoma brachyurum, Chyd spha – Chydorus sphaericus, Ceri quad – Ceriodaphnia
quadrangular, Lept kind – Leptodora kindtii, Daph cucu – Daphnia cucullata, Bosm long –
Bosmina longirostris, Calanoid copepods: Eudi grac – Eudiaptomus gracilis, Cyclopoid copepods:
Ther cras – Thermocyclops crassus, Cycl vici – Cyclops vicinus; Meso leuc – Mesocyclops leuck-
arti, Ther oith – Thermocyclops oithonoides, Rotifers: Brac angu – Brachionus angularis, Aspl
prio – Asplanchna priodonta, Coll muta – Collotheca mutabilis, Gast styl – Gastropus stylifer,
Kera cochl – Keratella cochlearis, Kell long – Kellicottia longispina, Poly doli – Polyarthra
dolichoptera-vulgaris, Trich capu – Trichocerca capucina, Leca luna – Lecane luna, Trich pusi –
Trichocerca pusilla, Cono unic – Conochilus unicornis, Trich insi – Trichocerca insignis. Black
boxes indicate the highest median abundance of a species; 1, 2a, 2b, 3 and 4 refer to the treatment
periods (see Fig. 11.1)
(45%). Also, high percentages of primarily bacterivorous (25%) or rather bacterivo-

rous/algivorous (21%) cladocerans are found. Carnivorous species only constitute
8%. The most abundant cladocerans are Eubosmina coregoni, Ceriodaphnia quad-
rangular, Daphnia cucullata, Bosmina longirostris, Diaphanosoma brachyurum
and Chydorus sphaericus (Fig. 11.2). The adult stages of copepods are mainly
omnivorous (67%). The remaining recorded species are classified as algivorous
(33%). The main copepod species are Eudiaptomus gracilis, Thermocyclops
crassus, T. oithonoides and Mesocyclops leuckarti, while Cyclops vicinus is less
abundant (Fig. 11.2). In addition to the more common species displayed in Fig. 11.2,
the relatively rare littoral cladocerans, Simocephalus vetulus, Sida crystallina and
Acroperus harpae, were found.
11.3.3 C
omparison of Zooplankton Composition
Between the Five Treatment Periods
In addition to the time series on the 19-year development displayed in Fig. 11.1, the
comparison of zooplankton structure during the five periods of lake management (1,
2a, b, 3 and 4) is shown in Figs. 11.2, 11.3 and 11.4. Fig. 11.2 depicts the distribu-
tion pattern of the most abundant rotifers and crustaceans. None of these species
revealed a quasi-evenly distributed occurrence across all treatment periods. The cla-
doceran Eubosmina coregoni and the rotifer Brachionus angularis were the two
most abundant species during the eutrophic period before the Riplox treatment
(period 1). Furthermore, Eubosmina coregoni, which had the highest peak abun-
dance during the 19-year study, were only dominant in the eutrophic period
(Fig. 11.2) and did not occur from late autumn 2003 until the end of the study. The
rotifer Keratella cochlearis was the only species showing the highest abundance
during the first year of the chemical phosphorus precipitation (Fig. 11.2, period 2a).
For many other rotifers, median abundances increased within the restoration period,
from the first to the second year of the Riplox treatment in 1995/1996 and the fol-
lowing years of restoration until 1999, reaching the highest abundances in period
2b. The cladocerans which were also most abundant in period 2b were Diaphanosoma
brachyurum, Chydorus sphaericus, Daphnia cucullata, Eudiaptomus gracilis,
Cyclops vicinus and Mesocyclops leuckarti. Only few crustacean species reached
peak abundances in the subsequent periods, i.e. the period of ‘macrophyte re-
establishment’ (Fig. 11.2, period 3, Bosmina longirostris) and in the successive
years of ‘stable conditions’ (Fig. 11.2, period 4, Ceriodaphnia quadrangular,
Thermocyclops oithonoides). It is worth noting that also some rarely found cladoc-
erans, such as Simocephalus vetulus and Sida crystallina, occurred during the re-
establishment of macrophytes (not shown in Fig. 11.2). These cladocerans are very
large-sized species such as with a body length of about 2000 and 2500 μm for
Simocephalus vetulus and Sida crystallina, respectively (for size comparison, the
length of Daphnia cucullata varies between 300 and 1200 μm). These species were
Cyclopoida Calanoida all zooplankton
Abundance [LN indiv. L ]

-1
9 ab
1mm a ab c
1mm 8 b
Cladocera Rotatoria 7
6
5
4
1mm 30µm
1 2a 2b 3 4
Cyclopoida Calanoida
a

a
-1
-1
a a b
6 a 5 ab b
b a
5
4
4
3
3
2 2
1 1
1 2a 2b 3 4 1 2a 2b 3 4
Cladocera Rotatoria
a abc
-1
-1
7 b 9 ab c
c b 8
a
6 c b
7
5 6
4 5
3 4
3
2 2
1 2a 2b 3 4 1 2a 2b 3 4
Treatment period Treatment period
Fig. 11.3 Box-whisker plot of the abundance of all zooplankton, copepods (cyclopoids and cala-
noids), cladocerans and rotifers during the five treatment periods. According to Kruskal-Wallis
tests, statistical differences among lake treatment periods were found for all zooplankton groups
(p < 0.01) with the exception of cyclopoids (p = 0.814). Furthermore, statistically significant dif-
ferences between homogeneous data subsets for calanoids, cladocerans, rotifers and all zooplank-
ton are represented by letters above each box as determined by pairwise Mann-Whitney U tests
with Bonferroni correction. 2a, 2b, 3 and 4 indicate the treatment periods as in Fig. 11.1
very low in abundance and were only found in the late period of re-establishment of
macrophytes and under stable conditions (from April 2005 and onwards).
Simocephalus vetulus reached a maximum abundance of 0.75 individuals L−1 and a
biomass of 5.6 μg L−1, and Sida crystallina an abundance of 0.5 individuals L−1 and
a biomass of 2 μg L−1. Both species were only found in 1–2.5% of the plankton
Cop
10
10
0
0
0
0
25
25
75
75
Cop
1
50
50
50
50
75
75
25
25
Clad Rot
0
0
10
10
0
0
0 25 50 75 100 0 25 50 75 100
Clad Rot
10
0
10
0
0
0
25
25
75
75
2a 2b
50
50
50
50
75
75
25
25
0
0
10
10
0
0
0 25 50 75 100 0 25 50 75 100
10
0
10
0
0
0
25
75
25
75
3 4
50
50
50
50
75
25
75
25
0
0
10
10
0
0 25 50 75 100 0 25 50 75 100
Fig. 11.4 Zooplankton structure displayed as the relative contribution of copepods (Cop), cladoc-
erans (Clad) and rotifers (Rot) to total zooplankton for the five treatment periods by triangular
diagrams. Filled blue circles denote abundance, open circles biomass. 1, 2a, 2b, 3 and 4 are the
treatment periods as depicted in Fig. 11.1. The top left triangle is a scheme illustrating how to read
the zooplankton graphs. All points in the sector ‘Cop’ indicate a contribution of at least 50% cope-
pods to the total zooplankton. Analogously, the sector ‘Clad’ represents ≥50% cladocerans and
sector ‘Rot’ ≥50% rotifers. Points in the middle area indicate that no single zooplankton group
exceeded 50% of total zooplankton (more information on triangular diagrams in limnology can be
found in the Methods Sect. 11.2.2). According to the Kruskal-Wallis tests, the relative abundance
and the relative biomass of all three zooplankton groups were statistically different between the
five lake treatment periods (see text)
samples taken during period 3 and 4. Another littoral but smaller bodied cladocer-
ean species, Acroperus harpae with a body length of 700 μm, was also found from
the period of macrophyte re-establishment onwards. A. harpae occurred more regu-
larly during periods 3 and 4 than the above-mentioned two littoral, large-bodied
cladocerans (first occurrence of A. harpae recorded in April 2003, present in about
10% of the samples taken in period 3 and 4).
The rotifer Kellicottia longispina seemed to be the only rotifer exhibiting a similar
distribution pattern as that of the abundant crustacean zooplankton. Unlike the crus-
taceans, many rotifer species showed their highest abundance in the long-term period
after chemical restoration (13 years from 2000 to 2012). The rotifer species with the
highest abundance in period 3 were Collotheca mutabilis, Polyarthra dolichoptera-
vulgaris and Lecane luna, and in period 4 Gastropus stylifer, Trichocerca capucina,
Trichocerca pusilla, Conochilus unicornis and Trichocerca insignis (Fig. 11.2).
When looking at the individual species (see Fig. 11.2), we found at least one rotifer
and one crustacean species with peak abundances in each of the five periods. For
some species, the peak abundance was far from being statistically significant.
Nevertheless, the species-specific comparison provides a rough overview of the per-
sistence pattern of the individual species during this long-term period.
The statistical difference between the five periods of ecosystem-based lake res-
toration management is shown in Fig. 11.3 for the abundances of all zooplankton
species and for the four taxonomic zooplankton groups. For ‘all zooplankton’, we
found significant differences between the median abundance of the five periods
(Kruskal-Wallis test, H = 64.76, df = 4, p < 0.001). The highest abundance of all
zooplankton occurred in the first year of the Riplox treatment (period 2a). At the end
of the ‘restoration‘ (period 2b), total zooplankton abundance decreased significantly
(subgroup ‘b’). The period of macrophyte re-establishment seemed to be accompa-
nied by a recovery of zooplankton abundance (overlap of subgroups a and b).
Finally, during the latter part of lake restoration (period 4 of ‘stable conditions’),
total zooplankton abundance demonstrated a further significant decrease (homoge-
nous subgroup ‘c’) and the lowest abundance in a comparison between all periods
(1, 2a, 2b, and 3).
The four taxonomic groups revealed a different pattern of succession during the
periods of lake restoration. The abundance of calanoids (Kruskal-Wallis test,
H = 17.28, df = 4, p < 0.01) did not differ statistically when comparing the eutrophic
period with the periods of ‘macrophyte re-establishment’ and ‘stable conditions’
(all three periods belong to subgroup ‘b’). The abundance of calanoids increased
only shortly during the first year of Riplox treatment (overlap between subgroups a
and b) and reached an intermediate peak in the second year of the chemical treat-
ment (subgroup ‘a’). During these 2 years, median calanoid copepod contributions
relative to copepods were 21% and 23%: the long-term median wasonly shortly
during the first year of Riplox treatment (overlap between subgroups a and b) and
reached an intermediate peak in the second year of the chemical treatment (sub-
group ‘a’). During these 2 years, median calanoid copepod contributions relative to
copepods were 21% and 23%: the long-term median was 19%. For cyclopoids, the
main group of copepods, no statistically significant differences were observed
between the five periods (Kruskal-Wallis test, H = 1.57, df = 4, p = 0.814). The
abundance of cladocerans showed pronounced differences among lake treatment
periods (Kruskal-Wallis test, H = 60.87, df = 4, p < 0.001). These crustaceans were
mainly abundant in the eutrophication period (1) with a strong intermediate decrease
in the first year of the Riplox treatment (2a), with a rising abundance during period
2b and a decrease in the subsequent periods. The low abundance during the final
‘stable’ period of restoration management is similar to the low abundance achieved
after the abrupt decrease with the first Riplox-treatment. Rotifer abundance showed
also significant differences (Kruskal-Wallis test, H = 69.97, df = 4, p < 0.001) and
tended to increase in the first year of the Riplox treatment (2a), reaching peak abun-
dance compared with all other periods. In the following years, the abundance of
rotifers decreased again. The lowest rotifer abundance, found in the last period, i.e.
‘stable conditions’ (subgroup c), was similar to the initial abundance under eutro-
phic conditions before application of chemical restoration.
Figure 11.4 shows the proportional shifts of the zooplankton groups over the five
periods of ecosystem-based management. The triangle graphs illustrate the relative
contribution of the three main zooplankton groups, the two crustaceans and the
rotifers. All three zooplankton groups revealed highly significant differences
(Kruskal-Wallis test, df = 4, p < 0.001; copepods: H = 129.7, cladocerans: H = 62.65,
rotifers: H = 60.77) when comparing abundances among the five treatment periods
(see filled blue circles in Fig. 11.4). The most pronounced shift of relative abun-
dance during the treatment periods was found for cladocerans vs copepods and the
least pronounced, but still highly significant change, emerged for rotifers (see also
Sect. 11.3.5 about the climate impact). Cladocerans and rotifers each contributed at
least 50% to the total zooplankton abundance during the eutrophic period (Fig. 11.4
1) and were thus the two most abundant groups before the restoration was initiated.
In later periods of the lake restoration treatment, the cladocerans did not reach an
abundance larger than 50% again (Fig. 11.4 2–4). Instead, copepods became abun-
dant and reached more than 50% in period 3 and 4 (Fig. 11.4 3–4). In contrast, the
relative contribution of rotifers was high in all periods and peaked in period 3 when
macrophytes were re-established (Fig. 11.4 3). In the final period of stable conditions,
relative rotifer abundance slightly declined and copepod abundance further
increased. During the 19-year investigation period, an abundance shift occurred
from a rotifer-cladoceran-rich community during the eutrophication period to a
rotifer-copepod-rich community in the later periods of lake restoration.
The relative contribution of the three main taxonomic groups to total zooplank-
ton biomass demonstrated significant differences (Kruskal-Wallis test, df = 4,
p < 0.001, copepods: H = 75.26, cladocerans: H = 52.60; p < 0.01, rotifers: H = 17.02,
see open circles in Fig. 11.4). Again, the change of rotifers during the treatments
was less pronounced when compared to copepods and cladocerans. Nevertheless,
the contribution of the relative biomass reveals a different perspective, as the here
investigated zooplankton covers a larger size-spectrum of animals from small-
bodied rotifers to large-bodied copepods and cladocerans. Even rotifers, which
often contributed much more than 50% to total abundance, never exceeded a bio-
mass contribution of 50% during any time of the 19-year investigation (Fig. 11.4
1–4). The zooplankton biomass was mainly dominated by cladocerans during the
eutrophic period. In the later years of the lake management, in particular during
macrophyte re-establishment and stable conditions (period 3 and 4 in Fig. 11.4
3–4), copepods contributed at least 50% to the zooplankton biomass. Thus, we can
conclude that in terms of biomass proportion the zooplankton is dominated by crus-
taceans and shifted from a cladoceran-rich community in the eutrophic period to a
copepod-rich community throughout the periods of the ecosystem-based lake
restoration.
11.3.4 L
ong-Term Development in Net Growth Rate
of Zooplankton Biomass, Size Structure and the Carbon
Ratio of Zooplankton to Phytoplankton
The time series of the net growth rate of zooplankton, the zooplankton size and
related measures are shown in Fig. 11.5. The net growth rate of zooplankton bio-
mass is depicted in Fig. 11.5 a. Positive rates (n = 219, mean positive rate is
0.0185 day−1) were less common than negative rates (n = 227, mean negative rate is
−0.0182 day−1). The total sum of the positive rates over the whole study period from
1994 to 2012 was 4.05 day−1 and − 4.13 day−1 for the negative rates. Thus, the zoo-
plankton biomass decrease was nearly compensated by the zooplankton biomass
increase over the 19 years. As to the net growth rate of zooplankton abundance, the
numbers differed only slightly (positive changes: n = 205, mean = 0.0191 day−1,
total sum = 3.92 day−1; negative changes n = 241, mean = −0.0165, total
sum = −3.97 day−1) and thus do not describe a substantially different situation than
that for the long-term dynamics of the net growth rates of zooplankton biomass
(graph for the net growth rate of zooplankton abundance is not shown). In accor-
dance with this, the cumulative net growth of zooplankton biomass was relatively
balanced throughout the five periods of ecosystem-based management. The only
period with clearly limited zooplankton development in the growing season occurred
in 1996, i.e. the year of the second chemical treatment (period 2b). In the successive
2 years within the 2b restoration period, zooplankton biomass recovered to the orig-
inal level and then remained stable following a rather regular seasonal pattern with
intra-annual fluctuations (Fig. 11.5b). The fairly balanced long-term dynamic of the
net growth rates of zooplankton biomass was, however, quite different from the
dynamics of the net growth rates of phytoplankton (Fig. 11.5b). In the year of the
first chemical phosphate precipitation (period 2a), the phytoplankton biomass
decreased as a response to the reduced concentration of the main nutrient, phospho-
rus (Fig. 11.5b, d). The drastic loss of phytoplankton biomass could not be compen-
sated by positive net growth rates afterwards due to the persistence of low phosphorus
availability. In turn, this led to a stabilised high water transparency (Fig. 11.5d,
transparency is displayed as Secchi depth; for long-term dynamics of nutrients, phy-
toplankton and related measures see also Chaps. 6 and 9).
Annual average of mean zooplankton body size over the 19-year study is depicted
in Fig. 11.5c. The long-term development did not show a particular trend during the
0.08
kzoo
Net growth rate [d ]
-1
0.04
0.00
-0.04
-0.08
0.1
Cummulative net growth rate [d-1]
0.0
-0.1
-0.2
-0.3
cum kzoo
cum kphyto
Size [µg L : ind L ]
-1
1.5
animal sizezoo
1.0
-1
0.5
0.0
2.0 TP 5
TP [µmol L-1]
1.5 Secchi
Secchi [m]
1.0 3
2
0.5
1
0.0 0
93 94 95 96 97 98 99 00 01 02 03 04 05 06 07 08 09 10 11 12 13 14
1 2a 2b 3 4
Year
Fig. 11.5 Time series of net change rates of zooplankton (A, 1994–2012), cumulative net change
rates of zooplankton and phytoplankton (B, period for zooplankton as in A, for phytoplankton
1993–2014), annual averages of zooplankton body size (C, calculated as the ratio of zooplankton
biomass [μg L−1] to zooplankton abundance [number of individuals L−1], 1994–2012), and annual
averages of total phosphorus concentrations (TP, molar) and Secchi depth (Secchi) (D, 1993–
2014). A and B show net change rates at biweekly intervals (see method); 1, 2a, 2b, 3 and 4 are
treatment periods as in Fig. 11.1
A B
e0 0.28
Czoo : Cphyto [µmol L-1 : µmol L-1]

Czoo : Cphyto [µmol L-1 : µmol L-1]
1
e-1 2a 0.24
2b
3 0.20
e-2 4
0.16
e-3
0.12
e-4
0.08
e-5 0.04
e-6
e-2 e-1 e0 e1 e2 1 2a 2b 3 4
-1
TP [µmol L ]
Fig. 11.6 Relationship between the carbon of zooplankton and phytoplankton for the five treat-
ment periods from 1994 to 2012. (a) carbon ratio (molar) versus total phosphorus concentration
(TP, molar) displayed on a logarithmic scale (LN), (b) carbon ratio for individual treatment periods
as Box-whisker plot. 1, 2a, 2b, 3 and 4 indicate the treatment periods as in Fig. 11.1
treatment periods but displayed two distinct increases, namely in the successive
years 1998/1999 and in year 2004. The first peak coincided with the naturally re-
covered underwater vegetation after the second chemical phosphate precipitation.
The second short-term increase of animal size occurred during the re-planting of
submerged vegetation (macrophyte biomass in Fig. 8.7 in Chap. 8).
According to the large phytoplankton blooms seen before the restoration (period
1), the carbon ratio of zooplankton to phytoplankton was significantly low when
concentrations of total phosphorus were high (Fig. 11.6a, b). The remarkable
increase of this carbon ratio in the following three periods of lake management
(periods 2a, b and 3) was mainly due to the reduced phytoplankton biomass and not
enhanced zooplankton biomass. However, during the final period, the ‘stable condi-
tions’, the carbon ratio of zooplankton to phytoplankton decreased compared with
the period of macrophyte re-establishment. This significant decrease of the ratio
coincided with the decline in total zooplankton abundance and biomass (Figs. 11.1a,
b and 11.3; see here the statistical significance of reduced abundance for total zoo-
plankton in period 4), while phytoplankton biomass remained the same.
11.3.5 I mpact of Climate Warming on Seasonality and Long-

Term Development of Zooplankton
The response to climate driven water warming is illustrated in Fig. 11.7. Figure 11.7a

displays the correlation between the North Atlantic Oscillation index for the winter
period from December to March (NAODJFM, see method) and the water temperature
integrated over depth (WT). The winter climate index significantly correlates with
0.6
p < 0.05
0.4
0.2
R 0.0
-0.2
-0.4
p < 0.05
-0.6 A NAODJFM vs WT
J F M A M J J A S O N D
Biweekly interval
25
20
SWT [°C]
15
10
B SWT first half July
SWT second half April
100 SWT > 22°C
90 SWT > 25°C
Number of days [d]
80 Freshw Jellyfish
70
60
50
40
30
20
10
0 C
150 1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014
140
Julian day
Years
130 CWP
120
110
100 D
90
1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014
1 2 3 4
Year
Fig. 11.7 Impact of climate warming. A: Correlation between the climate index NAODJFM and the
water temperature integrated over the water column (WT) for all biweekly intervals from January
(J) to December (D) during 21 years (1994–2014). Lines in the correlogram indicate the threshold
Fig. 11.8 Freshwater
jellyfish (Craspedacusta
sowerbii) is a commonly
appearing species during
hot summer periods in Alte
Donau. The photo shows
its size compared with the
water plant Spiked
Water-milfoil
(Myriophyllum spicatum).
(Photo taken on August 15,
2015)
water temperature in the beginning of year, from late January to early March. The
impact of this climate signal then abates from late March and onwards.
The surface water temperature (SWT) measured in late April significantly
increased by 1.52 °C per decade (p-value <0.05, Fig. 11.8b). The SWT increase in
early July was much lower (0.9 °C per decade with p-value >0.05, Fig. 11.8b) and
thus indicates less pronounced warming in summer than in spring. During the inves-
tigation period 1993 to 2014, the number of days when SWT exceeded 22 °C varied
between 31 and 87 days (Fig. 11.8c), the number of days significantly increasing
during the investigation period, by 10.5 days per decade (p-value <0.05, Fig. 11.8c).
This lengthening of the warm period is mainly due to a progressively earlier onset
of the warm period (slope = −0.76, p-value <0.05; not shown in graphs). The first
day in the year on which the temperature exceeded the threshold of 22 °C thus
shifted by about 7 days to an earlier date per decade (6.84 days per decade, varying
Fig. 11.7 (continued) of the coefficient of correlation (R) with statistical significance

(p-value<0.05) B: Year-to-year variation of surface water temperature (SWT) measured in spring
(biweekly means of the second half of April, 1994–2014) and summer (biweekly means of the first
half of July, 1993–2014). The lines refer to robust trends with a slope of 0.17 for SWT (tau = 0.39,
p-value = 0.014) in April and of 0.1 in July (tau = 0.27, p-value = 0.08). These trends refer to an
increase of SWT of 1.52 °C per decade in late April and 0.9 °C per decade in early July. C: Year-
to-year variation of the number of days when SWT is above 22 °C and above 25 °C (1994–2014).
The slope of the robust trend line for the number of days with SWT > 22 °C is 1.17 (tau = 0.31,
p-value = 0.049), indicating an increase in the number of hot days in summer (10.5 days per decade
with a SWT higher than 22 °C). Stars denote years of abundant freshwater jellyfish in summer
reported by local newspapers or observed by the authors. D: Year-to-year variation of the timing of
the clear-water phase. The single robust trend (line 1), covering all years from 1994–2014, has a
slope of −10.2, indicating a time shift by about 11-day advance in the occurrence of the clear-water
phase per decade (10.8 days; tau = −0.26, p-value = 0.1). The robust trend line for the 6-year time
segment from 1994 to 1999 (line 2, slope = −3.67, tau = −0.47, p-value = 0.25) refers to treatment
periods 1 and 2 and the 15 years from 2000 to 2014 (line 3, slope = −0.8, tau = −0.14, p-value = 0.5)
for treatment periods 3 and 4
between end of May to end of June). In contrast, the lengthening of the warm sum-
mer period was less pronounced (slope = 0.20, p-value >0.05) and resulted in a later
offset of about 2 days per decade only (1.8 days per decade, varying from the end of
August to mid-September). While the progressively earlier onset is significantly
negatively correlated with the annual NAO index (r = −0.49, p-value <0.05), the
delayed trend of the offset is positively, but not significantly, correlated with the
climate signal (r = 0.31, p-value >0.05, not shown in graphs).
When considering the extremely warm days with SWT above 25 °C (see Methods
Sect. 11.2.3), we found that this threshold was exceeded for at least 10 days in
7 years (1994, 1998, 2002, 2003, 2006, 2012 and 2013), while in the remaining
years the extremely hot summer period only lasted less than 1 week (5 years) or did
not appear at all (9 years, Fig. 11.8c).
During summer periods of extreme warm water, the medusa stage of
Craspedacusta sowerbii Lankester was observed in Alte Donau as documented
in local reports (Fig. 11.7c). This freshwater jellyfish had been sporadically observed
from July to August as from the 1970s in Neue Donau (Waidbacher, personal com-
munication), and repeatedly mentioned in the local news (e.g., 28 July 2006:
‘Süsswasser-Quallen wieder da’) and observed by the authors (2014 and 2015). The
medusa of this freshwater jellyfish is often observed macroscopically in Alte Donau
(Fig. 11.8) but may easily be overlooked in regular samplings as they often emerge
for only a few days in small areas in the elongated lake basin (spatial patchiness).
The abundance of this freshwater jellyfish is therefore not included in the regular
zooplankton estimations in Figs. 11.1a, b. The number of days with a water tem-
perature range from 19 to 25 °C, offering optimal growth conditions for the polyp
stage of C. sowerbii (see method), varies between 58 and 137 in Alte Donau and
exhibits an increasing trend (the slope of robust trendline is 0.4, p-value>0.05, data
not shown). Counting only the number of days where the seasonal threshold of
25 °C was reached, the period varied between 20 and 104 days, which is, on aver-
age, 62% of total length in the period with temperatures between 19 and 25 °C
period. The period with SWT above 21 °C, assuming to promote the development
of mature medusa (see method), varies between 60 and 108 days in Alte Donau
(starting at Julian Day 140 to 180, i.e. late May to the end of June; ends at Julian
Day 232 to 261, i.e. late August to mid-September). Again, the duration of this
period demonstrated an increasing trend (slope of robust trendline is 0.71,
p-value>0.05, data not shown). Thus, the prolongation of the period with an opti-
mum temperature range supporting the medusa stage was more pronounced than for
the polyp development in Alte Donau.
The seasonally low phytoplankton development in spring indicated presence of
a clear-water phase in Alte Donau. The timing of this phase varied between the 95th
(early April) and 145th day (late May) in a year during the 21-year investigation
period (Fig. 11.7d). The robust trend line covering the whole period from 1994 to
2014 has a slope of −10.2, suggesting an earlier occurrence of the clear-water phase
of about 11 days per decade (Fig. 11.7d, p-value >0.05). Considering the two main
treatment periods of different orders of phytoplankton yield separately (see Method

Sect. 11.2.3), the year-to-year variation still showed earlier timing for both time
segments. The robust trend line for 1994–1999, covering the 2 years before restora-
tion and the ‘restoration period’ (period 1 and 2), exhibited a 33-day earlier onset
per decade of the clear-water phase. This shift is much more pronounced towards
early spring than the shift indicated by the second robust line for 2000–2014. The
second robust line is describing only a shift of 7 days earlier per decade (‘re-
established macrophytes’ and ‘stable conditions’, period 3 and 4).
The seasonality of the four taxonomic groups of zooplankton (cyclopoid and
calanoid copepods, cladocerans and rotifers) and the relationship between these
zooplankton groups and WT are shown in Fig. 11.9. Considering the long-term
average of the biweekly abundances throughout the seasons, the seasonal distribu-
tion pattern of cyclopoid copepods closely resembled a unimodal distribution.
The abundance peaks occurred in May and an even higher peak appeared in July,
with a temporary short-time depression in June (Fig. 11.9a3). Due to the weak
summer depression, the annual cycle of cyclopoid abundances coincided closely
with the temperature peak in Alte Donau. When displaying these long-term data
as a scatter plot in Fig. 11.9a1, the statistically significant relationship between
the abundance of cyclopoids and WT becomes obvious (Fig. 11.9a1, P = 0.83,
p0 < 0.001). Fig. 11.9a2 again shows the correlation between the abundance of the
cyclopoids and WT but for each biweekly interval throughout the year in a
correlogram.
The cyclopoids were positively related with temperature at almost all times of
the year (Fig. 11.9a2). The relationship was strongest during the cold season and
culminated in a statistically significant relationship during four biweekly intervals
from December (R = 0.67 and 0.66, p-value<0.05) to January (R = 0.62 and 0.69,
p-value<0.05, Fig. 11.9a2). Thus, cyclopoids, which commonly show peak abun-
dance during the summer season, generated particularly high abundances when WT
was unusually high during winter. In contrast to the cyclopoid copepods, the less
dominant calanoid copepods exhibited a pronounced bimodal seasonal develop-
ment pattern (Fig. 11.9b3) and an inverse relationship with seasonal WT in late
summer to autumn (Fig. 11.9b2). During the autumn peak from September to early
October, the abundance of calanoids seemed to be stimulated in years with rela-
tively low temperatures compared with the situation in warm years (p-value>0.05,
Fig. 11.9b2). This inverse relationship between abundance and warming in autumn
on the one hand and the general suppression of calanoid development for months
during the growing season on the other (midsummer decline from June to August in
Fig. 11.9b3) weakened the long-term relationship between calanoid abundance and
water temperature as shown in Fig. 11.9b1 (P = 0.22, p-value <0.001). In case of the
cladocerans (Fig. 11.9d), the seasonal abundance was suppressed again from June
to August (Fig. 11.9d3). In contrast to the copepods, no significant inverse relation-
ship was found between the long-term warming trend and the long-term develop-
ment of the cladocerans (Fig. 11.9d2). The relationship traced in the long-term data
e6 e5
Abundance [in L-1]
Abundance [in L-1]

e5 A1 cyclopoids e4 B1 calanoids
e3
e4 e2
e3 e1
e2 e0
e-1
e1 P = 0.22, p<0.001
P = 0.83, p<0.001 e-2
e0 e-3
e0 e1 e2 e3 e4 e0 e1 e2 e3 e4
Water temp [°C] Water temp [°C]
0.6 0.6
p<0.05 p<0.05
0.4 0.4
0.2 0.2
0.0 0.0
R
R
-0.2 -0.2
-0.4 -0.4
-0.6 A2 p<0.05
-0.6 B2 p<0.05
25 140 25 30
A3 B3
Abundance [ind L-1]

Abundance [ind L-1]
Water temp [°C]
Water temp [°C]

20 120 20
25
100
15 15
80
20
10 60 10
40 15
5 5
20
0 0 0 10
J F M A M J J A S O N D J F M A M J J A S O N D
Biweekly interval Biweekly interval
e9 e7
Abundance [in L-1]
Abundance [in L-1]
e8 C1 rotifers e6 D1 cladocerans
e7 e5
e4
e6
e3
e5 e2
e4 e1
e3 e0
P = 0.58, p<0.001 P = 0.61, p<0.001
e2 e-1
e0 e1 e2 e3 e4 e0 e1 e2 e3 e4
Water temp [°C] Water temp [°C]
0.6 0.6
p<0.05 p<0.05
0.4 0.4
0.2 0.2
0.0 0.0
R
-0.2 -0.2
-0.4 -0.4
-0.6 C2 p<0.05
-0.6 D2 p<0.05
25 800 25 120
C3
Abundance [ind L-1]
D3
Abundance [ind L-1]
Water temp [°C]
Water temp [°C]
20 20 100
600
80
15 15
400 60
10 10
40
200
5 5 20
0 0 0 0
J F M A M J J A S O N D J F M A M J J A S O N D
Biweekly interval Biweekly interval
Fig. 11.9 Relationship between water temperature (WT) and the abundance of the four main
taxonomic zooplankton groups (a–d). a1: Correlation between WT and the abundance of cyclopoid
copepods reflected by long-term data (P = Pearson correlation coefficient, n = 449).
between WT and the abundance of cladocerans thus primarily owe to seasonality

(high abundance in summer, low in winter). Both parameters corresponded well
with each other (P = 0.61, p-value<0.001, Fig. 11.9d1). Rotifers, the most abundant
zooplankton group, showed a maximum peak in spring (April to May in Fig. 11.9c3),
their seasonal depression during the peak season being less pronounced (Fig. 11.9c3)
than those of calanoids and cladocerans. The relationship between zooplankton
warming trends was, however, not uniform throughout the year. The rotifers seemed
to be benefit from mild winters and their abundance was evidently suppressed in
autumn (Fig. 11.9c2). The relationship between long-term temperature and rotifer
abundance was almost as strong as for cladocerans (Fig. 11.9c1, P = 0.58,
p-value <0.001).
The long-term seasonal patterns of the four main zooplankton groups all show
spring peak abundances in the beginning to mid-May (calanoids, rotifers and cla-
docerans) or from the middle to the end of May (cyclopoids). In contrast, the degree
of summer suppression varied significantly between groups; thus, an early second
peak was observed in July for cyclopoids and rotifers and in September for cala-
noids and cladocerans. Moreover, the intra-annual variations in abundance of all
four groups were directly linked with WT in winter or spring, i.e. abundance was
higher in years with a mild winter and vice versa. According to long-term averages,
the two zooplankton groups exhibiting a pronounced midsummer decline, i.e. cala-
noids and cladocerans, were moderately inversely related to WT in summer. In con-
trast, the two zooplankton groups showing a short-term summer depression after the
spring peak, i.e. cyclopoids and rotifers, tended to be directly related to summer
WT. Furthermore, cyclopoids, which show the strongest coherence with the sea-
sonal cycle of WT in the long-term averages (Fig. 11.9a3) and which are positively
related to WT throughout the seasons (Fig. 11.9a2), were the zooplankton group
demonstrating the strongest relationship between WT and abundance (r = 0.83 in
Fig. 11.9a1) in Alte Donau. In turn, the long-term abundance of calanoids, which
exhibit a pronounced spring peak, a strong and long-lasting summer depression
(Fig. 11.9b3) and a positive relationship to WT in spring and a negative relationship
in summer (Fig. 11.9b2), corresponds to the lowest statistical relationship between
WT and abundance.
Fig. 11.9 (continued) a2: Correlogram displaying the correlation coefficient (R, Spearman rank)
for the relationship between WT and the abundance of cyclopoids every second week throughout
a year (n = 19 for each bar, covering the 19-year period from 1994–2012, for details see Method
Sect. 11.2.3). Lines indicate the statistical significance of R (p-value<0.05). a3: Seasonal distribu-
tion pattern of WT (bars) and the abundance of cyclopoids (line) displayed as long-term means at
biweekly intervals from January to December (j–d, averages over the19-year study period). Graphs
for rotifers (c1–c3), calanoid copepods (b1–b3), and cladocerans (d1–d3) are similar to that of
cyclopoids (a1–a3)
11.4 Discussion
Despite the large spatial heterogeneity of the banks and basins of the former river
branch (Table 3.1 and Fig. 3.1 in Chap. 3; Chaps. 18 and 19), the total biomass of
zooplankton does not differ statistically significantly when comparing the two
impoundments. The only exception during the 19-year study is the year 1994 (as
demonstrated by statistical results in this study and plankton description for 1994 in
Mayer et al. 1997). Also, the biomasses of all three zooplankton groups, rotifers,
cladocerans and copepods, differed significantly between the two basins in 1994
(large between-site differences were also found for the abundance of ciliates, as
shown in Table 12.1 in Chap. 12). The reason for the differences in zooplankton
biomass in this particular year might be the partial water exchange with Neue Donau
(inflow at the south end of the south basin, at ‘Seestern’, see Chap. 5), undertaken
with the aim to temporarily improve the water quality in Alte Donau (Mayer et al.
1997). The year 1994 thus exemplifies that disturbance by water management
impacts the zooplankton since the abundance in the south basin was obviously
lower than in the north basin. It is, however, beyond the scope of this study to anal-
yse the impacts of short-term water exchanges (washout effect to enhance water
transparency, Chap. 5) and of water drawdown in spring (stimulating the vernal
growth of macrophytes, details in Fig. 8.7 in Chap. 8) on zooplankton structure.
The species composition of the zooplankton community found in Alte Donau is
similar to that found in other shallow lakes. About 78% of the rotifer species, 64%
of the cladocerans and 43% of the copepod species found in Alte Donau also occur
in, for example, large and shallow Lake Võrtsjärv, Estonia (surface area 270 km2)
(Haberman and Virro 2004). Likewise, the main species of rotifers, cladocerans and
copepods observed in Alte Donau were found in lakes with a similar surface area,
for instance riverine lake Großer Müggelsee (Eckert and Walz 1998). All zooplank-
ton species listed by Baranyi et al. (2002) for the hydrologically dynamic waters of
the Danube River floodplain (Regelsbrunn near Vienna) were also common in Alte
Donau. The mean zooplankton abundance (March to October) of these floodplain
waters, characterised by a short water age ranging from hours to 39 days, is 1495
individuals L−1, which is, however, much higher than the number in the oxbow lake
Alte Donau (mean abundance from March to October over the 19-year study
period = 632 individuals L−1, the long-term annual mean being even lower, namely
494 individuals L−1 including 352 rotifers L−1, 83 copepods L−1 and 59 cladocerans
L−1). About 30% of the rotifer species described by Lair (2005) for a river ecosys-
tem, the Middle Loire, were also observed in our study in Alte Donau. It underpins
that some planktonic rotifers can adapt well to a habitat of an elevated current
despite their small size (hydrology see Chap. 4). The annual peak abundance of the
rotifers agrees well with those of the long-term observations in Middle Loire despite
the lower abundance in our study, which is ranging between 1000 to 2000 individu-
als per L during the 18 years of ongoing restoration in Alte Donau (1995–2012, cf.
decline of TP and phytoplankton in Chaps. 1, 6, 10 and 20).
11.4.1 R
esponse of Zooplankton to Lake Restoration: What
Are the Specifics of the Oxbow Lake Alte Donau?
11.4.1.1 M
ain Picture of Altered Crustacean Composition
Following the Trophic Change
In particular, large-bodied cladocerans are found to be efficient grazers on phyto-

plankton when considering that the size of food particles increases with animal size
(e.g., Brooks and Dodson 1965; Burns 1968, 1969; Arndt et al. 1993; Shapiro and
Wright 1984; Lampert et al. 1986; Gulati and Van Donk 2002; Gudimova et al.
2011). In Alte Donau, cladocerans such as Daphnia cucullata and Chydorus
sphaericus were found to be the dominant grazers when filamentous cyanobacteria
(Cylindrospermopsis raciborskii, Limnothrix redekei) bloomed in spring and sum-
mer in the year 1994 (Dokulil and Mayer 1996; Mayer et al. 1997; Dokulil 2015)
and macrophyte biomass was low (2 tons dry weight of macrophytes for the whole
water basin, see Chap. 8). The cladocerans were the only zooplankton group attain-
ing highest abundance before the chemical treatment (Fig. 11.3). When comparing
the relative contribution in the eutrophication period with later restoration periods,
zooplankton biomass was composed of high portions of cladocerans (Fig. 11.4-1).
In terms of biomass, this period hosted a cladoceran-dominated zooplankton assem-
blage (cladoceran-rotifer-rich), and in terms of abundance it was a rotifer-cladoceran-
rich community (Fig. 11.4-1). After restoration, the zooplankton biomass shifted
towards a copepod-rotifer-rich community and in terms of abundance to a rotifer-
copepod- rich community. The crustacean zooplankton therefore shifted from
mainly filter-feeding herbivorous cladocerans in the eutrophic algal-turbid state to
mainly selective-feeding omnivorous and herbivorous copepods in the mesotrophic
transparent-water (trophic states are illustrated Fig. 20.3 in Chap. 20). Such species
shifts from cladocerans to copepods are commonly described along trophic gradi-
ents from high to low total phosphorus in lakes (Pace 1986; Jeppesen et al. 2000;
Kasprzak and Koschel 2001; Köhler et al. 2005). In Alte Donau, the majority of the
copepod species consisted of cyclopoids (6 species in Table 11.1), calanoids only
being represented by one species (Eudiaptomus gracilis), as also found in other
studies (Anneville et al. 2007a). The dominance of the omnivorous cyclopoid cope-
pods indicates moderate food availability under eutrophic to mesotrophic condi-
tions (Kasprzak and Koschel 2000; Anneville et al. 2007b; Mehner et al. 2008). The
herbivorous calanoid copepods, constituting 19% of the copepods in a long-term
perspective, typically indicate a nutrient state lower than mesotrophic, i.e. it is gen-
erally assumed that the calanoid copepods are adjusted to lower food thresholds
than cyclopoid copepods (Adrian 1997; Kasprzak and Koschel 2000; Mehner et al.
2008). The suggestion that calanoids have a low demand for algal food agrees with
our observations in Alte Donau where an intermediate increase in calanoid abun-
dance occurred during the two Riplox-years (restoration periods 2a and 2b), i.e.
peak abundances of calanoids coincided with extremely low phytoplankton bio-
masses after phosphate precipitation (Fig. 11.3). According to Sommer and Stibor
(2002), large-bodied calanoid copepods and cladocerans might not be mutually
exclusive as calanoids feed on large-sized algae, while the remaining small-sized

algae might be still suitable as food for filter-feeding cladocerans. This finding is
supported by the results of our study showing coherent occurrence pattern of indi-
vidual species during the Riplox-year; thus, the peak abundance of the calanoid
Eudiaptomus gracilis coincided with the peak abundance of Daphnia cucullata in
Alte Donau (Fig. 11.2). Other aspects regarding the food quality for zooplankton
relative to the trophic state are described in Geller and Müller (1981), Capblancq
(1990), Jeppesen et al. (2000), Kurmayer et al. (2003), Kainz et al. (2004), Ferrão
Filho et al. (2005) and Brett et al. (2009), and these are all relevant for the zooplank-
ton shift in Alte Donau, which will be described in more detail in the following Sect.
11.4.1.2 for the years in which a drastic P reduction occurred.
11.4.1.2 S
hort-Term Zooplankton Response to Strong Phosphorus
Reduction During Riplox-Years
Benndorf (1987) stated that top-down control by zooplankton only efficiently con-
tributes to enhancing water transparency if the internal load of phosphorus has been
reduced a priori, thereby ensuring limited growth of algae through bottom-up
control. The Riplox-treatment (Ripl 1976), i.e. the chemical phosphate precipitation
and stabilisation of the oxidized sediment in April 1995 and April 1996, aimed at
creating a prompt phosphorus reduction in the water column and a long-term reduc-
tion of the internal phosphorus load at the sediment-water interphase (treatment
described in Chap. 5 and in Donabaum et al. (1999); for a review of the retention of
internal phosphorus load see, for instance, Søndergaard et al. 2001). Phosphorus is
the main limiting nutrient element for phytoplankton growth in freshwaters (Hecky
and Kilham 1988). The relationship between total phosphorus and phytoplankton
yield (estimated chlorophyll-a or phytoplankton biovolume) is depicted in Fig. 9.10
(Chap. 9) and Fig. 20.3 (Chap. 20), showing the strong phosphorus reduction lead-
ing to lower phytoplankton biomass in 1995 and 1996 (see also time series in
Fig. 11.5).
The total phosphorus concentration of 2.37 μmol L−1 before the restoration
decreased to 0.89 μmol L−1 in 1995 and 0.6 μmol L−1 in 1996 after the Riplox treat-
ment in Alte Donau (median values derived from Teubner et al. 2003). Triggered by
phosphorus precipitation, the planktonic community adjusted to efficiently utilising
the remaining phosphorus sources. Planktonic biota acted here as a sink for phos-
phorus under P-limitation which was seen in two ways: (a) with the tighter coupling
between food and consumer organisms, a larger portion of phosphorus was bound
by biota at the expense of the dissolved P-fractions, and (b) with the compositional
shifts from N-rich cyanobacteria to P-rich eukaryotic algae, the increase in the rela-
tive importance of P-rich bacteria and the increase of the zooplankton portion rela-
tive to phytoplankton, the stoichiometry of biota shifted towards a P-rich plankton
community (Teubner et al. 2003). The zooplankton played an important role in
accomplishing the higher phosphorus utilisation efficiency of the plankton
community. A tight coupling at both the producer-consumer and the nutrient-
producer interface was achieved mirroring higher phosphorus turnover rates under
phosphorus limiting growth conditions (e.g., Capblancq 1990; Teubner et al. 2003).
The carbon ratio between zooplankton and phytoplankton (Czoo:Cphyto) increased
clearly after the Riplox treatment and is indicating the coupling of zooplankton and
food but from a standing crop perspective (Fig. 11.6; see also Teubner et al. 2003).
This increase was mainly due to the low biovolume of phytoplankton under P-limited
conditions as many short-lived primary producers often respond promptly to drastic
nutrient reduction (e.g., Jeppesen 2000; Ibelings et al. 2007). In Alte Donau, her-
bivorous zooplankton was suppressed by the absence of large-sized phytoplankton
blooms. Instead, the small zooplankton mainly feeding on bacteria and small algae
such as rotifers (e.g., Fig. 11.3) and ciliates (Chap. 12) became increasingly abun-
dant in the Riplox-years when total phosphorus levels were low (Teubner et al.
2003). The existence of two alternate zooplankton assemblages dominated by either
small rotifers or large cladocerans is commonly found and discussed in the study of
fish stocks and top-down control on zooplankton in lakes (e.g., Gilbert 1988;
Järvinen and Salonen 1998; Jeppesen et al. 2000; Lehtovaara et al. 2014), as fish
primarily feed on large-bodied zooplankton, allowing growth of small-sized zoo-
plankton species. Lower abundances of Eudiaptomus gracilis and Daphnia cucul-
lata were observed in the small basin ‘Kaiserwasser’. This decrease was related to
the fish stocking experiment in 1998 (see Sect. 11.4.1.4). In contrast, the zooplank-
ton community in the two main basins was not affected by the stocking in 1995/96
as the fish community did not change remarkably. In addition, macrophyte canopies
were still missing during this early restoration period and thus could not serve as
spawning and feeding grounds for predatory fish species. Therefore, we assume that
the high abundance of rotifers and ciliates at low cladoceran levels, occurring in
particular in the first Riplox-year, was rather mainly due to the bottom-up control of
the large herbivorous zooplankton than to the top-down control by fish.
The food quality for zooplankton during both Riplox-years has rapidly changed.
While the phytoplankton biovolume significantly decreased year for year with
P-precipitation, bacterial biomass remained almost the same (Teubner et al. 2003).
With a significantly higher bacterial enzyme activity mobilising the P source from
the dissolved organic phosphorus fraction in Alte Donau, the planktonic bacteria
further gained relative importance over phytoplankton for P acquisition under
reduced phosphorus availability. The loss rates of bacterioplankton (not the stand-
ing crop) were in the same order of magnitude as those of phytoplankton. With the
reduction of total phosphorus, they even became slightly higher than those of phy-
toplankton (Teubner et al. 2003). These increased relative loss rates have been
regarded as indirect evidence of enhanced grazing on bacteria (e.g. by rotifers and
ciliates) which has been observed in other ecosystems (e.g., Berman 1990).
According to other studies (e.g., Jones and Cannon 1986; Capblancq 1990; Elser
and Goldman 1991; Loreau 1995; Gismervik et al. 1996; Queimaliños et al. 1998;
Vadstein 2000; Tadonléké et al. 2009), the planktonic bacteria in Alte Donau, the
eukaryotic phytoplankton and the small-bodied zooplankton (rotifers and ciliates),
seemed to be the most important factors for accelerating the circulation of matter
under reduced phosphorus availability in both Riplox-years 1995/96. After the
Riplox-years, total phosphorus concentrations slightly increased again (Fig. 11.5)

indicating a resilience phase which has been observed together with hysteresis pat-
terns discussed for trophic changes in other lakes including shallow to deep water
bodies (e.g. Carpenter et al. 1985; Beisner et al. 2003; Dokulil and Teubner 2005;
Jeppesen et al. 2005; Morabito et al. 2005; Ibelings et al. 2007; Anneville et al.
2007a; Schindler 2012, see also hysteresis for re-establishment for macrophytes in
Alte Donau in Fig. 20.4. in Chap. 20).
11.4.1.3 Impact of Macrophyte Re-establishment on Habitat Change
The time series of the annual mean body size in Alte Donau mirrored alternate shifts
between micro- and mesozooplankton, i.e. between small rotifers and large crusta-
ceans. During the 19-year development period, an increase in annual mean zoo-
plankton body size was observed two times, which could be both linked to an
increase of underwater vegetation cover. In 1998/1999, when the first intermediate
increase of mean body size was detected (see time series in Fig. 11.5c), seven of the
12 main crustacean species, but only 1 of the 12 main rotifer species, reached peak
abundances (see restoration period 2b in Fig. 11.2). The seven crustaceans were the
cladocerans Diaphanosoma brachyurum, Chydorus sphaericus, and Daphnia
cucullata and the copepods Eudiaptomus gracilis, Thermocyclops crassus, Cyclops
vicinus, and Mesocyclops leuckarti, and the rotifer Kellicottia longispina. The
increase of mean zooplankton size coincided with the spontaneous growth of mac-
rophytes in response to the chemical treatment in 1998, after macrophytes had
almost disappeared in the first year of the Riplox treatment (0.5 tons of dry weight
for macrophytes only in the whole lake). In 1998/1999, the submerged macrophytes
already yielded 12–14 tons of total dry weight (all macrophyte biomass from Fig.
8.7 in Chap. 8). The most dominant submerged vegetation species was Myriophyllum
spicatum. Nitellopsis obtusa together with five other charophytes as well as Najas
marina further gained remarkable yields (Table 8.2, Fig. 8.7 and 8.8 in Chap. 8).
Other studies confirm that even small increases of underwater vegetation can have a
large impact on the zooplankton community (Schriver et al. 1995; Burks et al.
2002). Although Secchi depth increased further and the standing crop of macro-
phytes resulted in a 30 times higher biomass yield (Chap. 8), the mean size of zoo-
plankton decreased again (Fig. 11.5c, d). A second increase of mean zooplankton
body size, observed in 2004, was in the beginning of re-planting of macrophytes
(period of ‘macrophyte re-establishment’). This coincided with a remarkable mac-
rophyte development (about 360 tons dry weight for the whole lake, details in Chap.
8). During this period, in particular Myriophyllum spicatum had high biomass. This
water plant typically builds up a canopy formation in the top water layer by profuse
branching of shoots near the surface. Other underwater vegetation, such as
Nitellopsis and Najas already mentioned before, were found again in 2004 but
reached only relatively low biomass yield (cf. Fig. 20.5 in Chap. 20). The small- to
medium-sized canopies of these ‘stoneworts’ and other ‘slow growing species’
(Chap. 20) are rather far from the surface. Different from the zooplankton situation
in 1998/1999, extremely large-sized cladocerans, such as Simocephalus vetulus and

Sida crystallina, commonly occurred in 2004 but contributed only low abundances.
Exceptionally large-sized crustaceans are well known from other macrophyte-
dominated habitats (e.g., Timms and Moss 1984; Jeppesen et al. 2000). Even though
these two cladoceran species were sporadically observed until 2012 in Alte Donau,
the mean size of zooplankton species declined again. The further increase of mac-
rophyte vegetation during our study was thus not mirrored by uniform dominance
of large zooplankton species. Nevertheless, the increased abundance of macro-
phytes coincided with the dynamics of the carbon ratio between zooplankton and
phytoplankton, which remained high until the end of the zooplankton investigation
period. Such an increase in the biomass ratio between zooplankton and phytoplank-
ton is commonly found at declining nutrient levels (e.g. Padisák 1993; Teubner et al.
2003; Jeppesen et al. 2000, 2005; He et al. 2017).
In shallow lakes, the canopy of underwater macrophyte serves as a daytime ref-
uge for zooplankton against visual predators such as planktivorous fish (e.g., Burks
et al. 2002). Macrophyte beds, however, may provide further benefits for zooplank-
ton than just acting as a spatial refuge. Depending on the macrophyte canopy struc-
ture (Carpenter and Lodge 1986, see ‘open’ and ‘closed’ canopy type in Sand-Jensen
and Pedersen 1999 for stream habitats) and on the strength of macrophyte allelopa-
thy to inhibit epiphyton and phytoplankton growth (e.g. Van Donk and van de Bund
2002; Berger and Schagerl 2004; Hilt and Gross 2008), submerged macrophytes
can serve as a valuable habitat for zooplankton development. The macrophyte
stands offer an environment of moderate flow velocity and abundant food supply for
zooplankton by stimulating the growth of epiphytic and planktonic algae. Even
though it is commonly noted that the allelopathy of submerged macrophytes con-
tributes to stabilising the clear-water state in shallow lakes (e.g., Hilt and Gross
2008), submerged macrophyte stands in lakes and rivers are not ‘algal free zone’ but
inhabit epiphytic algae and cyanobacteria (e.g., Carpenter and Lodge 1986;
Hofmann 1993; Schaumburg et al. 2004). As underwater vegetation utilises dis-
solved nutrients and as their life span (several months) exceeds the short life period
of algae and cyanobacteria (1 day to a few weeks), submerged macrophytes might
contribute to stabilising both the ecosystems state (Scheffer and van Nes 2007) and
the zooplankton habitat structure.
In the case of Alte Donau, Myriophyllum spicatum serves as the most common
refuge simply because it is the dominant macrophyte. The canopy formation of M.
spicatum at the water surface, however, might be less beneficial as a daytime refuge
for zooplankton compared with medium-high Charophyte-species, which have
evenly distributed branches along the water column. Underwater mowing in Alte
Donau removes the dense branches of the Myriophyllum spicatum canopy near the
surface. This mowing frequently applied in recent years on the one hand and the still
low presence of medium-sized macrophytes with abundant branching shoots (e.g.,
Charophyte-species) on the other hand seem to hamper the refuge effect for zoo-
plankton in Alte Donau (see also Sect. 15.2.3 in Chap. 15 on losses of young fish by
mowing of macrophytes). A more patchy underwater vegetation might further
enhance the refuge heterogeneity for zooplankton. The heterogeneity of ambient
habitat structure for zooplankton is difficult to describe with regular sampling (e.g.,
Van Donk and Van de Bund 2002) and is not analysed here because appropriate
high-resolution measurements of the patchy zooplankton habitat structure were
beyond the scope of this study.
11.4.1.4 Top-Down Control by Fish
Top-down control of zooplankton by fish can be of decisive importance in a lake. It

is commonly observed that large-bodied zooplankton (mainly Daphnia) becomes
predominant in shallow lakes when the number of planktivorous fish decreases and
vice versa (e.g., Brooks and Dodson 1965; Carpenter et al. 1985; Gilbert 1988;
Järvinen and Salonen 1998; Jeppesen et al. 2000; Benndorf et al. 2001; Mehner et al.
2008; He et al. 2017). In accordance with this, an increase of large zooplankton such
as Daphnia pulex has been observed to occur in years after natural fish mortality
under ice in winter (e.g., Rücker et al. 2003). In contrast, small-sized zooplankton
species have been recorded to benefit from fish stocking in high-altitude lakes, natu-
rally fishless mountainous freshwater basins (Schabetsberger et al. 2009).
Fish stock experiments with top predators indicate that a lower grazing pressure
due to loss of piscivorous fish triggers the establishment of large-bodied crustaceans
(Shapiro and Wright 1984; Lazzaro 1987; Ronneberger et al. 1993; Pace et al.
1999). In Alte Donau, such biomanipulation experiments were conducted in the
impoundment ‘Kaiserwasser’ in 1998. In the presence of piscivorous fish, the bio-
mass of crustaceans such as Eudiaptomus gracilis and Daphnia cucullata increased
(Fig. 15.15 in Chap. 15), confirming the findings mentioned before. The cladocer-
ans seem to be easier to capture than copepods by planktivorous fish (Drenner et al.
1978; Lazzaro 1987; Schriver et al. 1995; Mehner et al. 2008). During the 19-year
development period in Alte Donau, the zooplankton biomass exhibited a significant
shift from a cladoceran-rich towards a copepod-rich community (Fig. 11.4).
Therefore, our results suggest that the grazing pressure of fish on zooplankton still
remained high at the end of the 19-year period.
The species composition of the fish assemblages in Alte Donau did not change
significantly during the study period (fish surveys in Table 15.2, Chap. 15), which is
different from common internal-lake biomanipulation treatment aimed at removing
planktivorous and benthivorous fish (Søndergaard et al. 2007). Fish assemblage in
Alte Donau is still dominated by cyprinids, from the 1980s over the whole zoo-
plankton study period. Alte Donau is traditionally popular for angling, where the
most common stocking fish was the common carp over decades (Chap. 15, see also
about Löffler 1988 in the introduction in this chapter). With the reduction of total
phosphorus and the associated drastic reduction of phytoplankton (Fig. 11.5 in this
Chapter, further Chap. 9 and Fig. 20.3 in Chapter 20), fish catches declined in Alte
Donau (time series in Fig. 15.1, Chap. 15) as commonly described for other lakes
that undergo re-oligotrophication (e.g., Jeppesen et al. 2000, 2002; Gerdeaux et al.
2006). Before restoration, the mean chlorophyll-concentration was 35 μg L−1 and
the mean fish catch was 6157 kg biomass (period 1). At ‘stable conditions’ (period 4),
both declined to 4 μg L−1 and 4314 kg, respectively. A significant linear response

between lowering chlorophyll-a and decreasing fish catches is found for 1998 to
2012 (Fig. 20.7 in Chap. 20) and covers the last 15 years of observation. Furthermore,
fish catches were inversely associated with water transparency measured as Secchi
depth over the last 21 years (from 1992 to 2012, Spearman rank correlation,
r = −0.73, p < 0.001, not shown in graphs). This relationship indicates an over-
whelming control by fish on the plankton community. It seems that the grazing
pressure by fish on zooplankton in Alte Donau could not be dampened by a refuge
effect of re-established macrophyte stands as described from other studies (Schriver
et al. 1995; Tátrai et al. 2009), with the exception of a short-term increase in the
abundance of large zooplankton species in 2004 as discussed in Sect. 11.4.1.3. It
can be therefore concluded that fish populations have consistently controlled the
food web structure in Alte Donau for the whole study period and thus cannot explain
the large shift in zooplankton composition. The main change in zooplankton assem-
blages thus responded primarily to a bottom up control, i.e. to the strong reduction
of the phytoplankton food supply by one order of magnitude and associated effects
of water transparency increase as discussed in Sects. 11.4.1.1 and 11.4.1.2.
11.4.2 R
esponse of Zooplankton to Global Warming: What Are
the Specifics of Urban Shallow Alte Donau?
11.4.2.1 Climate Signal and Warming of Surface Water
The oxbow lake Alte Donau is located in the Vienna Basin belonging to the
Pannonian Plain. The Vienna Basin is one of the most vulnerable areas of climate
impact in Austria (Dokulil and Herzig 2009; Dokulil et al. 2010b; Olesen et al.
2011) alongside the Austrian Alps (e.g., Thies et al. 2007; Nickus et al. 2010;
Füreder et al. 2012; Tolotti et al. 2012). Alte Donau is located in the city of Vienna,
which has about 1.8 mill. residents and, thus, warming might be further strength-
ened locally by the effect of urban heat islands (UHI, IPCC 2014). The area of
Vienna is 41,487 km2 comprising 35.6% urban residential areas, 14.3% traffic areas,
45.5% parks and green space, and 4.6% covered by surface waters (MA23 2015).
According to the proximity to the urban area of Vienna, it could be assumed that
UHIs (e.g., Peterson 2003) have a potential impact on the microclimate of Alte
Donau. Böhm (1998) compared time series of air temperature and urban heat excess
temperatures of urban and suburban sites with those of rural reference sites in
Vienna for the study period 1991 to 1995. While heat excess temperatures were
pronounced in the urban areas of Vienna during the study period (mainly during
winter for the measuring site ‘city centre’ distant from parks), low values of heat
excess were measured at the suburban site ‘Donauinsel’, which is in a close distance
to Alte Donau. Böhm (1998) concluded that urban effects on air temperature are
more influenced by local surroundings than by the city Vienna itself. The significant
warming trend detected for the 22-year period in our investigation might thus be
less influenced locally by UHI, i.e. the city of Vienna; rather large-scale effects such
as climate warming may be of strong importance. Evidence for the response to
global warming by Alte Donau was derived from the significant correlation between
the climate signal NAO and SWT. The NAO index is commonly used as climate
signal in the Northern Hemisphere and Europe to detect year-to-year variability
driven by global climate change mediated by ‘local’ weather and subsequent filtered
by lake processes (Straile et al. 2003; Blenckner 2005; Adrian et al. 2009) through
physical conditions affecting lake biology (e.g., George and Hewitt 1999,
Weyhenmeyer et al. 1999, Gerten and Adrian 2000, Winder and Schindler 2004a, b,
Shatwell et al. 2008). In Vienna, NAO positive years are typically associated with
high total incoming shortwave radiation (long sunshine duration), low cloud cover
and low precipitation and thus relate to hot, dry years. In turn, NAO negative years
refer to cold wet years (unpublished results by KT; for other observations between
NAO and weather at specific sites in Europe; see George and Hewitt 1999, and
Blenckner 2005). Thus, NAO-associated phenomena mirror meteorological forcing
(examples given in Jennings et al. 2000, George et al. 2004). According to a long-
term study of a stratified alpine lake in Central Europe by Schmid and Köster (2016),
warming trends of the lake surface in spring and summer are caused by advection of
warm air (60%) and direct heating (40%) due to the incoming solar radiation.
In Alte Donau, the seasonally strongest correlation was found between the NAO
index of the winter period December to March (NAODJFM) and early year WT, from
late January to early March. This result is consistent with those of other studies,
which have reported a high impact of winter NAO on lake properties (Jennings et al.
2000; Gerten and Adrian 2000; Blenckner 2005; George and Hewitt 1999;
Weyhenmeyer et al. 1999; George et al. 2004; Blenckner et al. 2007; Adrian et al.
2009; Dokulil and Herzig 2009). Furthermore, the significant SWT increase with
1.52 °C per decade in late April together with a weakening of warming trends in
July in Alte Donau confirm the findings in other lakes studies of a pronounced
response to warming in early spring (e.g., Weyhenmeyer et al. 1999; Gerten and
Adrian 2000; Blenckner 2005; Dokulil et al. 2010b; Winder and Schindler 2004a;
Kainz et al. 2017). This result also holds true when comparing the response to the
climate signal during the seasonal hysteresis observed in the first and the second
half of the year in Alte Donau. While seasonal warming above 22 °C follows the
annual climate signal, seasonal cooling below 22 °C is not statistically significantly
related to annual NAO. In accordance with this, seasonal warming follows a pro-
nounced trend of an earlier onset of the threshold temperature of 22 °C. Seasonal
cooling below 22 °C is only slightly delayed. Such seasonal hysteresis of climate
response patterns, strengthening the time shifts in spring but weakening those later
in the year, as for instance the vernal onset and autumnal offset of thermal stratifica-
tion and associated development of phytoplankton described e.g., by Winder and
Schindler (2004a) and Deng et al. (2014), are also well-known from other pheno-
logical studies.
Besides the generally pronounced climate impact in winter and spring, Austria –
together with Switzerland and Poland – experiences the strongest warming anoma-
lies in summer in central Europe (O’Reilly et al. 2015; Woolway et al. 2016). From
a lake perspective and in addition to the persistence of climate signals passing lake
filters (Blenckner 2005) in winter and spring (e.g., Blenckner et al. 2007; Adrian
et al. 2009), the impact of climate warming in summer is also a topic of discussion
with particular focus on plankton dynamics (e.g., Wilhelm and Adrian 2008; Adrian
et al. 2009). A significant trend of summer warming has been verified for Alte
Donau where the period with SWT above 22 °C has increased by 10.5 days per
decade. This further confirms the importance of lake surface summer warming as
found also in other studies in this region (see Neusiedlersee in Sapna et al. 2015;
O’Reilly et al. 2015; Woolway et al. 2016), often promoting the proliferation of
cyanobacteria directly by temperature increase and also by associated early stratifi-
cation and/or reduced thermal turbulences in stratifying deep but also polymictic
shallow lakes (e.g., Joehnk et al. 2008; Pearl and Huisman 2008; Shatwell et al.
2008; Dupuis and Hann (2009); Wagner and Adrian 2009; Markensten et al. 2010;
Kosten et al. 2012; Posch et al. 2012; Dokulil and Teubner 2012; Dokulil 2014;
Rigosi et al. 2014).
11.4.2.2 W
arming and the Most Spectacular Zooplankton Species:
Freshwater Jellyfish
In Alte Donau, the most spectacular zooplankton species occurring in hot summer
periods is Craspedacusta sowerbii (Fritz et al. 2007). While many studies from the
1980s and onwards emphasise the increasing geographical spreading of this neobi-
otic species among freshwater habitats, including the Danube catchment in Germany
and Austria (e.g., Tittizer et al. 2000; Fritz et al. 2007), other studies focus on the
dates of the seasonal observations of the medusa stage from mid-June to July/
August or early September (Kronfelder 1989; Grohs 1998; Jankowski et al. 2005;
Peukert 2009). The seasonal development of the medusa stage of C. sowerbii
described in these studies matches the duration of the warm period above 21 °C in
Alte Donau (from late May to mid-September). This confirms the optimal tempera-
ture range for the development of the medusa stage suggested by Folino-Rorem
et al. (2016) from culture studies. More recent studies discuss the sporadic develop-
ment between years, as in the year after occurrence of the medusa stage it is absent
in the following year (e.g., Jankowski et al. 2005; Folino-Rorem et al. 2016). Folino-
Rorem et al. (2016) give their culture observations an eco-physiological perspective
by including the full life cycle of C. sowerbii. They conclude that warm tempera-
tures besides high food availability (feeding frequency and/or amount of food) are
of decisive importance for the optimal development of this freshwater jellyfish. The
significant trend recorded of prolonged summer periods with SWT exceeding 22 °C
supports the more frequent reports of the medusa stage of C. sowerbii in Alte Donau.
Our temperature trend analysis for the period 1993 to 2014 further illustrated that
the prolongation of the period with SWT above 21 °C is more pronounced than the
prolongation of the period with temperatures within the range of 19 to 25 °C, favour-
ing the medusa stage relative to the polyp stage given that the development is con-
trolled by temperature. Moreover, our estimate of 60 and 108 days with SWT > 21 °C
in Alte Donau exceeds the minimum life cycle (22 days) and life expectation of the
medusa form (21 to 77 days) proposed from culture work by Folino-Rorem et al.
(2016). However, the lack of data about the different life stages of C. sowerbii over
consecutive years (abundances or presence/absence data) in Alte Donau does not
allow us to draw any firm conclusions about whether the growth of this freshwater
jellyfish is primarily controlled by temperature or by food availability or by a com-
bination of both. According to mesocosm and enclosure experiments by Jankowski
et al. (2005), the medusa stage feeds in particular on small herbivorous crustaceans
(Bosmina longirostris and juvenile cyclopoid copepods) and may thus play an
important role as top invertebrate predator on zooplankton community.
11.4.2.3 P
henology of the Clear-Water Phase in Two Distinct Periods
with Regard to the P Decline
Lake phenology of the clear-water phase, i.e. the timing of short-term control of
phytoplankton by zooplankton grazing during the spring-summer transition, was of
early interest in the study of the impact of global warming on lake biology (Müller-
Navarra et al. 1997; Gerten and Adrian 2000; Jeppesen et al. 2003; Shatwell et al.
2008; De Senerpont Domis 2013; Berger et al. 2007; Huber et al. 2008; Berger et al.
2014). The relative robustness of the timing of the clear-water phase associated with
other physico-biological lake events led to a description of the general seasonal suc-
cession pattern found in many lakes in the temperate zone. The pattern mainly indi-
cates regime shifts between the two principal seasons of phytoplankton development
in a year, i.e. from winter/spring to summer/autumn (Teubner 2000, see also sea-
sonal pattern in Chap. 9) along the seasonal cycle (Lampert et al. 1986, Sommer
et al. 1986; De Senerpont Domis 2013). The timing of the clear-water phase between
early April to late May in our 21-year study is closer to the timing in other shallow
polymictic to dimictic lakes (early May to mid-June for shallow polymictic Großer
Müggelsee in Teubner et al. 1999; Straile and Adrian 2000; Huber et al. 2010; mid-
May to early June for stratifying lakes such as Plußsee in Müller-Navarra et al. 1997
and Lake Washington in Winder and Schindler 2004a, b) than in many other large
deep, mainly pre-alpine or alpine lakes in Central Europe (Straile and Adrian 2000;
Straile 2002). The trend towards an earlier clear-water phase in Alte Donau con-
firms general findings about an earlier onset of the spring clear-water phase in many
other lakes in recent decades (Müller-Navarra et al. 1997; Straile and Adrian 2000;
Straile 2002; Shatwell et al. 2008). The reason for the earlier timing of this spring to
summer event is partly due to the stronger warming at the beginning of the year as
discussed by seasonal hysteresis in Sect. 11.4.2.1. Another reason is the temperature
dependence of organism growth. Empirical data revealed that an increase in tem-
perature and an increase in food availability led to a shortening of the life span and
life cycles but to an increase of the reproduction rate for rotifers, cladocerans and
copepod species (e.g., Gophen 1976; Herzig 1983a, b; Orcutt and Porter 1983,
Sarma et al. 2002). Contemplating the climate change scenarios for lakes, it is
expected that the trophic structure might change from a simple but elongated food
web in a cold climate to a more complex (with a higher degree of omnivory) but
shortened food web in a warm climate (Jeppesen et al. 2010a; see also shifts in
dietary composition of (zoo)planktivorous fish which prey alternatively on (zoo)
benthic organisms in Alte Donau described in Chap. 15). Therefore, the effect of a
warmer climate relates to different and connected parts of the bottom-up and top-
down control in lakes, which includes also an enhanced probability of a clear-water
phase according to theoretical simulation models by Scheffer et al. (2001).
Early studies on the intra-annual variation of the clear-water phase without
including the climate aspect revealed that the probability of this short-term spring
event depends on the resource control of phytoplankton growth, on the one hand,
and the predation pressure on phytoplankton by zooplankton and fish, on the other
hand (e.g., Brett and Goldman 1997; Teubner et al. 1999). It is commonly known
that biomass yields are mainly highly dependent on the trophic status of a lake (e.g.,
Carlson 1977; Vollenweider 1968, not least shallow lakes e.g., Teubner et al. 1999;
Jeppesen et al. 2000; Teubner and Dokulil 2002), but it is assumed that the timing
of seasonal minima and maxima, including the timing of the clear water phase, is
less sensitive to trophic changes but responds to global warming (e.g., Straile and
Adrian 2000; Müller-Navarra et al. 1997; Winder and Schindler 2004a, b).
The long-term phenological shifts of the clear-water phase in Alte Donau pre-
sumably reflect the interference of two trends pointing in the same direction, the
trend of re-oligotrophication superimposed by the trend of global climate change.
The two periods with different trends for progressive earlier clear-water phase in
Alte Donau (earlier time shift of 33 days per decade for 1994–1998 and 7 days per
decade for 2000–2014) thus contradict the general robustness of the clear-water
phenology mentioned above. The pronounced earlier clear-water phase from 1994
to 1999 occurred concomitantly with the drastic reduction of the mean spring total
phosphorus pool from 1.4 to 0.7 μmol L−1 (spring mean value) and the associated
reduction of phytoplankton chlorophyll-a spring peak, decreasing from 29.2 μg L−1
to 10.7 μg L−1 (spring maximum value, mean values are 9.0 and 7.2 μg L−1). We did
not estimate the extents to which the earlier timing of the clear-water phase relates
to the re-oligotrophication and the superimposed climate forcing. A common
method to disentangle the two trends is the analysis of the residuals from the regres-
sion line of the detrended data (George and Hewitt 1999; Weyhenmeyer et al. 1999;
George et al. 2004; Seebens et al. 2007; Adrian et al. 2009). According to our expe-
rience, detrending of biological lake data responding to different nutrient levels
should be done with caution unless the ecosystem changes can be confirmed to be
only moderate. Alte Donau has undergone drastic changes in trophic state within a
short period. Detrending would here primarily satisfy statistics and thereby mathe-
matically eliminate the response to high nutrients levels in biological time series
rather than mirroring lake ecology responding to a complex regime shift passing the
transition from a nutrient-rich to a nutrient-poor ecosystem. Biological systems
respond to different levels of, for instance, phosphorus in different ways because the
absolute value of phosphorus (total pool size) is here of high importance (Teubner
et al. 2003). The drastic short-term reduction of the phosphorus pool with phosphate
precipitation enhances phosphorus utilisation efficiency of the planktonic community,
which was achieved by an increase of microbial activity and alterations in phyto-

plankton and zooplankton traits (Teubner et al. 2003, see also Sect. 11.4.1.2).
Therefore, we chose not to apply detrending.
An empirical analysis of a large number of Danish lakes by Jeppesen et al. (2003)
showed that phenological shifts of the clear-water phase toward an earlier date
occur when the spring phytoplankton peak declines from year to year in response to
reduced external loading. Alte Donau supports the findings by Jeppesen et al. (2003)
that bottom-up control affects the clear-water phase phenology. Van Donk et al.
(2003) also argued, based on a Danish lake survey, that shifts towards an earlier
clear-water phase could mainly be attributed to changing lake management prac-
tises aimed at improving water quality than to lake warming by climate change.
In Alte Donau, after the years with an initial strong reduction of phosphorus
availability (1994–1999), a moderate phenological shift of the clear-water phase
towards earlier onset in 2000–2014 coincided with the balanced intra-annual varia-
tions in spring chlorophyll-a concentration, which may be primarily attributed to a
climate change response as found in many of the above-mentioned studies.
11.4.2.4 Impact of Temperature Increase on Zooplankton Development
Seasonal differences in the relation between WT and individual micro- and meso-
zooplankton groups were assessed with correlograms, generated for biweekly inter-
vals in the 19-year period. The correlograms typically show a close temporal
coherence pattern between WT and all four zooplankton groups in winter and
spring. Extraordinarily high abundances of zooplankton early in the year corre-
spond with warm winters and springs, while the opposite is true for cold winters.
The closest relationship between year-to-year variations of temperature and zoo-
plankton development was found for cyclopoids and rotifers. In case of calanoids
and cladocerans, the positive relationship between abundances and WT is still pre-
dominant but much weaker. This general positive relationship between temperature
and zooplankton abundance early in the year confirms observations by George and
Hewitt (1999) for other lakes in the temperate zone, where small changes in the cold
seasons have significant effects on the development of zooplankton. During the
transition from the cold to the warm season, three aspects of temperature enhancing
the zooplankton development point in the same direction: (1) growth stimulation by
temperature dependent on the ontogenetic zooplankton development (see tempera-
ture dependent embryonic and postembryonic zooplankton development shortening
the life cycles and thus promoting faster growth rates at sufficient food availability
as discussed in Sect. 11.4.2.3), and (2) the vernal warming and (3) the climate driven
intra-annual temperature increase. The progressive growth of many organisms in
spring is linked to the excess of nutrient availability after winter stagnation, the
vernal increase of day length and water temperature, which is most beneficial for
fast-growing small primary producers. With the onset of the growing season for
phytoplankton and related plankton organisms, vernal warming thus stimulates the
increase of food availability for zooplankton (e.g., Adrian et al. 2006; Berger et al.
2014). In addition, the intra-annual temperature increase driven by climate warming

affects lake phenology mainly in winter and spring in Central Europe (Gerten and
Adrian 2002, Blenckner 2005, Huber et al. 2010; example for the subtropics see
e.g., Deng et al. 2014), as mentioned before in Sect. 11.4.2.1, and thus superimposes
ontogenetic and seasonal temperature effects. The coherent vernal temperature
response of all four zooplankton groups culminates in a narrow time window for
their peak abundance in Alte Donau.
The cyclopoids were most closely linked to water temperature (Fig. 11.9a1), i.e.
they mirrored a consistently positive response to temperature increase for all sea-
sons over the 19-year study period (Fig. 11.9a2) and did not built up a pronounced
early summer depression (Fig. 11.9a3) which was different than for the other crus-
taceans or the rotifers in our study. Their development thus seemed to be triggered
by different temperature effects pointing in the same direction. The temperature
increase, which is linked to both the seasonal warming (Fig. 11.9a3) and the year-
to-year warming trend by climate change (Fig. 11.9a2), led to an overall success of
these crustaceans in Alte Donau (Fig. 11.9a1). Mesocyclops leuckarti, Thermocyclops
crassus and T. oithonoides (Fig. 11.2, Table 11.1) are known for their tolerance of
high temperature during their embryonic development (e.g., Gophen 1976; Herzig
1983b; Wagner and Adrian 2011) and occurrence at maximum temperatures in sum-
mer (e.g., Tackx et al. 2004). Together, these three thermophilic species became
increasingly dominant after the phosphate precipitation in Alte Donau when com-
pared with the abundances of the remaining cyclopoids, which are known for a
rather temperature-indifferent development (Megacyclops viridis, Eucyclops ser-
rulatus) or for being adapted to relatively low temperatures (Cyclops vicinus).
Adrian and co-workers disentangled the seasonal temperature effects from global
warming by looking at the phenology response of individual cyclopoid species and
found evidence that the mentioned thermophilic ‘summer’ species benefit from
global warming trends. According to their results (Gerten and Adrian 2002; Adrian
et al. 2006), enhanced annual peak abundances of Mesocyclops leuckarti and
Thermocyclops oithonoides relate to the climate driven warm years. In case of the
latter species, a higher summer peak abundance was found to be accomplished by
an earlier start of growth in the beginning of the year (Gerten and Adrian 2002),
which corresponded to an earlier ice break up and an associated earlier spring peak
of phytoplankton in the studied shallow polymictic lake (Adrian et al. 2006). The
persistence of cyclopoids in our study could be accomplished by species alterations
among these crustaceans towards thermophilic species. The cyclopoid copepods as
a whole took advantage of seasonal and global warming on the one hand (Fig. 11.9a)
and were robust against trophic changes on the other hand (Fig. 11.3). The dynamic
of the calanoid copepods, which were less abundant and were represented by only
one species (Eudiaptomus gracilis), was different. They accomplished higher yields
for short periods during intermitted treatment years of extreme low phosphorus
availability (Fig. 11.3, see also discussion in Sect. 11.4.1.1) but benefited the least
from warming in general, which was indicated by seasonal decreases during par-
ticularly hot summer periods (Fig. 11.9b). The latter agrees with other studies show-
ing that E. gracilis is taking advantage from a relatively short embryonic development
at low temperatures, but fails at high temperatures (e.g., Herzig 1983b). In accor-
dance, unlike cyclopoid copepods, calanoid copepods can even reveal a negative
response to elevated annual mean temperatures as found by other long-term field
studies (Carter and Schindler 2012).
In Alte Donau, the second strongest response to the overall water temperature
increase was found for cladoceran crustaceans (Fig. 11.9d1). The suppression of
daphnids by climate warming, as discussed by Jeppesen et al. (2010b), Wagner et al.
(2013) and Tavşanoğlu et al. (2017), agrees with our finding that the pronounced
summer decline of cladocerans refers to the long-term increase of WT in summer
(Fig. 11.9d2, see also calanoids above). A further climate response most relevant for
all zooplankton from rotifers to crustaceans in Alte Donau was the predominant
positive response to intra-annual temperature variation in winter and spring in con-
cert with the common phenology of the close timing of their spring peak (Fig. 11.9,
cf. Adrian et al. 2006, Gerten and Adrian 2000, 2002). The responses of individual
zooplankton key species, which alternate between warm and cold year scenarios
and which also may alter the foodweb, are discussed in other climate studies (Adrian
1997; Benndorf et al. 2001; Winder and Schindler 2004b; Wagner and Benndorf
2007; Berger et al. 2014).
11.5 Conclusions
The main compositional shift from a cladoceran-rotifer-rich to a copepod-rotifer-

rich zooplankton assemblage seems to be driven by bottom-up control and thus
relates to drastic reductions of phytoplankton yield after initial phosphorus precipi-
tation and sustained low phosphorus availability during the subsequent years of
biomanipulation. With the suppression of filter-feeding herbivorous cladocerans
through reduced food supply, mainly selective-feeding omnivorous and herbivorous
copepods survived under mesotrophic transparent-water conditions. Rotifers and
calanoid copepods became particularly abundant during the two Riplox-years when
food availability drastically declined. Large-bodied crustaceans did not seem to
consistently benefit from predator avoidance by re-grown submerged vegetation as
assessed by the mean body-size of zooplankton community. The grazing pressure of
mainly cyprinid fish thus could not be dampened by the refuge effect of re-
established macrophyte stands in Alte Donau. During the long-term restoration
period, significant climate warming trends were detected such as an increase in
surface water temperatures in spring and a prolongation of the warm period in sum-
mer. Cyclopoid copepods seemed to benefit most from a warmed-up environment,
as it was the only zooplankton group with a robust positive long-term response to
climate warming throughout all seasons. In addition, the prolonged summer periods
linked to global warming may have supported the medusa stage of C. sowerbii in
Alte Donau.
Acknowledgments We thank David Livingstone and Susanne Wilhelm for valuable comments
on methods of climate research during European Union projects REFLECT (http://www.ife.ac.uk/
reflect/) and CLIME (http://clime.tkk.fi/) that were useful for analysing the climate impact in Alte
Donau. We are grateful to Thorsten Blenckner, Rita Adrian, Alois Herzig and two other colleagues
for critical comments improving the manuscript. We also thank Anne Mette Poulsen for improving
the language style. We would like to thank all of the numerous collaborators and the Municipal
Department for permission of publication. We further thank the ‘Wiener Fischereiausschuss’
(Austrian Fishery Association) for providing long-term fish catch records. The long-term lake
measurements were financially supported by Municipal Department – 45 (Water Management –
Vienna). ‘Österreichisches Komitee Donauforschung, Internationale Arbeitsgemeinschaft
Donauforschung’ partly funded data assessment (K.T.). Further data evaluation (I.T.) was partly
funded by the TU Wien Science award 2015 received by Wouter Dorigo (EOWAVE).
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Chapter 12
The Ciliate Assemblage in Alte Donau
Gerald Pfister
Abstract Within the framework of a comprehensive limnological survey of the

shallow urban lake Alte Donau in Vienna (Austria), the pelagic ciliate assemblage
has been continuously investigated throughout the years 1995–2000. The applica-
tion of a quantitative protargol stain (QPS) resulted in accurate data on ciliate spe-
cies composition, as well as reliable abundance and biomass numbers. The main
ciliate groups throughout the investigated period were identified as oligotrichs, pro-
stomatids and small hymenostomes, thus composing the so-called Oligotrichetea –
a ubiquitous and typical ciliate community found also in many other comparable
beta- to alpha-mesosaprobic pelagic environments. Significant seasonal variations
in ciliate abundance and biomass occurred, revealing lowest numbers during autumn
and winter. Despite the overall species composition being relatively unaltered dur-
ing the investigated years, absolute ciliate numbers and biomasses were high at the
beginning of the investigated period during the years of chemical RIPLOX-treatment
and decreased by more than 50% in the last 2 years of the study. This decrease of
ciliate numbers and biomass parallels thus the significant decrease of nutrients in
the water body as a result of phosphorus precipitation in 1995 and 1996, followed
by several restoration measures carried out during the years 1998–1999. Thus, low-
est ciliate abundance and biomass occurred in the last year of restoration and slightly
increased again with the re-colonization of macrophytes the year after.
Keywords Protozoa · Pelagic ciliates · Bioindicators · Oligotrichs · Prostomatids ·

Hymenostomes · Oxbow lake · Lake restoration · Riplox · Eutrophic · Mesotrophic
G. Pfister (*)
Qatar Biomedical Research Institute (QBRI), Hamad Bin Khalifa University (HBKU),
Doha, Qatar

https://doi.org/10.1007/978-3-319-93270-5_12
214 G. Pfister
12.1 I ntroduction: The Central Role of Ciliates in Pelagic

Freshwater Systems
The role of protozoa as major links in matter and energy fluxes within pelagic eco-
systems has been illuminated by various studies in the past and ciliates have been
attributed as a major component in all kinds of freshwater ecosystems (e.g., Arndt
1994; Simek et al. 1995; Pfister et al. 2002a, b; 2004). Ciliates are known to be an
important food source for metazoa (e.g., Weisse 1991; Sanders and Wickham 1993)
and effectively transfer picoplanktonic production to higher trophic levels (e.g.,
Fenchel 1987; Sherr and Sherr 1988; Dietrich and Arndt 2000). They can feed on
bacteria, auto- and heterotrophic pico- and nanoplankton (e.g., Porter et al. 1985;
Hall et al. 1993; Simek et al. 1995) and provide dissolved organic material (DOM)
as nutrients to bacteria (Posch and Arndt 1996). Different trophic levels can even be
influenced simultaneously by omnivorous feeding activities in certain protozoan
groups (Pfister and Arndt 1998).
Rapid growth, high turnover rates and short generation times allow protozoan
communities to respond quickly to changing environmental conditions. Hence,
many protozoan species can be highly valuable bioindicators in water quality analy-
ses (e.g., Sládecek 1973; Foissner 1988; Berger et al. 1997). Ciliates have been
reported as a significant component of the pelagic community in Alte Donau (Mayer
et al. 1997; Pfister 1997) and they have therefore been investigated as part of a com-
prehensive limnological survey in the small urban lake ‘Alte Donau’ in Vienna,
Austria. Alte Donau is a freshwater lake, which has artificially been derived by
disconnection from the river Danube some decades ago. This survey has started in
1993, and ciliates have been regularly investigated for 6 years from 1994–1996 to
1998–2000. In this chapter, the ciliate assemblages are described in more detail for
the 2 years of RIPLOX treatment (1995 and 1996), and 2 years later of the restora-
tion period (1998 and 1999, see general description of the restoration Chap. 5). The
last year of ciliate investigation outlined here, of the year 2000, refers to first year of
re-establishment of macrophytes. For ciliates, two sites in the southern (AD1) and
northern (AD4) basin have been sampled monthly (winter season) or twice a month
(summer season, morphometry see Table 3.1 in Chap. 3).
12.2 Methodology to Analyse Pelagic Ciliates
Sound eco-taxonomical studies require the exact determination of protozoan spe-

cies. Most investigations on the ecology of pelagic protozoan communities, how-
ever, do lack of a satisfactory taxonomic resolution, and surprisingly few of such
studies provide detailed taxonomic information (e.g., Lakey et al. 1993; Simek et al.
1995; Carrias et al. 1998; Montagnes et al. 1999; Auer and Arndt 2001). In Alte
Donau the pelagic ciliate community could be investigated over successive years
from 1994–1996 and 1998–2000 with respects to taxonomic and ecological
12 The Ciliate Assemblage in Alte Donau 215
demands and was accompanied by a comprehensive analysis of physical, chemical

and other biotic parameters (e.g., Mayer et al. 1997; Teubner et al. 2003).
Pelagic ciliates where obtained and integrated from three different depths (sur-
face, intermediate, ground). For that reason the obtained ciliate community is well
representative for the full water column. All samples were analysed by a quantita-
tive protargol stain (QPS). This method has extensively been evaluated and provides
both accurate quantitative and taxonomic information on most ciliate species
(Montagnes and Lynn 1987; Skibbe 1994; Pfister et al. 1999, 2004). For a detailed
method description and efficiency test see Pfister et al. (1999). In brief, ciliates are
fixed by Bouin solution, filtered on a membrane filter and stained by silver impreg-
nation. After the staining procedure, the filters are embedded on a cover slide and
can be analysed by bright field microscopy. A grid on the filter allows for enumera-
tion of ciliate numbers per defined sample volume.
The taxonomic identification of ciliates is a difficult task and requires both
sophisticated methodology and advanced skills of the investigator. The analysis of
the pelagic ciliate community in Alte Donau was carried out by the same investiga-
tor using the same methodology throughout the entire period. This makes the data
at least comparable and constant, if not highly valuable. Identification and taxo-
nomic nomenclature of all species is based on the most reliable and recent available
literature (Kahl 1930–1935; Maeda and Carey 1985; Maeda 1986; Song and Wilbert
1989; Krainer and Foissner 1990; Foissner 1994; Foissner et al. 1991, 1992, 1994,
1995, 1999; Foissner and Pfister 1997; Krainer 1988, 1995; Krainer and Müller
1995). Values for mean live biomass of most ciliate species were taken from Foissner
and Berger (1996). Specific biomass of species is not given in that work but was
calculated by approximations to simple geometric shapes and assuming a mean
specific weight of 1 for all species. Biomass, as a common denominator, was chosen
in order to facilitate quantitative comparisons. Shrinkage of cells due to the QPS
was considered and corrected by specific conversion factors for biomass calcula-
tions (cf. Pfister et al. 1999).
12.3.1 Ciliate Species Composition in Alte Donau
The main ciliate groups throughout the investigated period were oligotrichs and
prostomatids, as well as small hymenostomes. Many species from these groups are
amongst the most common pelagic ciliates in a wide range of freshwater habitats
such as lakes, rivers and even in many brackish waters (Pfister et al. 2002a, b, 2004;
Sonntag et al. 2006). Many species that were most abundant in Alte Donau through-
out all the investigated years have been recorded as major ciliates from numerous
freshwater lakes in Europe and America (e.g., Beaver and Crisman 1982, 1989;
Müller 1989; Laybourn-Parry and Rogerson 1993; Macek et al. 1996; Carrias et al.
1998). Additional records from Russia (Alekperov 1984; Oleksiv 1985) and China
216 G. Pfister
(Pai Kao-Tung 1962) demonstrate the world-wide distribution of these species.

Many of them have also been found in extreme habitats like caves (Griepenburg
1933; Gittleson and Hoover 1969) or the air-water interface of high mountain lakes
(Foissner 1979). Hence, these species are true ‘generalists’, obviously tolerating a
wide range of environmental conditions. Such a ciliate community (termed
‘Oligotrichetea’) is ubiquitous and typical for the pelagic ecosystems (Foissner
et al. 1995). Main species are the prostomatids Balanion planctonicum, Urotricha
spp., and the oligotrichs Halteria grandinella, Halteria chlorelligera, Stobilidium
humile, Strobilidium hyalinum, Tintinnopsis sp. (Figs. 12.1 and 12.2). According to
the saprobic assignation of the main species, this type of ciliate community is char-
acteristic for a beta- to alpha-mesosaprobic environment (Berger et al. 1997).
12.3.2 A
nnual Abundance and Biomass of Ciliates
in Alte Donau
Mean annual ciliate numbers showed high alterations in both basins of Alte Donau
during the investigated years (Table 12.1). Generally, no differences were seen
between the southern and the northern basin for ciliates confirming the results of
phyto- and zooplankton (Chaps. 9 and 11). The high abundance in year 1994 refers
to the good food availability (high phytoplankton biomass mainly composed by
cyanobacteria) before the chemical treatment started. The numbers ciliates in
1995/1996 were still high, i.e. about 40 cells per millilitre, but might refer rather to
a good food quality than an abundant food for ciliates in the environment at this
time. During these years of chemical precipitation of phosphorus (RIPLOX-
treatment) the phytoplankton declined drastically with low biomasses of cyanobac-
teria but high portions of eukaryotic algae and bacteria (Teubner et al. 2003,
phytoplankton composition see Chap. 9, bacterioplankton Chap. 13). Ciliate num-
bers significantly decreased in the following years, reaching minimal values of only
10–15 cells per millilitre in 1999. Most probably these low numbers were the result
of the restoration measures that were undertaken to improve water quality during
the years 1997 and 1998. However, at the end of the investigation period, ciliate
numbers increased again, reaching similar levels as in the year 1998. Obviously the
attempts to restore water quality in Alte Donau had only short-term effects with
regard to the pelagic ciliate community, which will be explained in a more detail in
the next paragraph about the ciliate biomass.
As observed for ciliate numbers, biomass changed significantly during the inves-
tigated years (Table 12.2). Despite the high cell numbers, the ciliates can reach only
a relatively low biomass in 1994, indicating the dominance of small species during
the eutrophied period. The mean ciliate biomass was by far highest in the years
1995/1996. At this time, total ciliate biomass was about 550 mg fresh weight per m3
water as the calculated annual mean. In 1995, the first year of phosphorus precipita-
tion by RIPLOX-treatment, a peak biomass of all the small-size zooplankton was
found, when compared with the nutrient-rich year before in 1994 and the second
Fig. 12.1 The ‘Oligotricheata’ – common pelagic ciliate community in freshwater lakes. All these
species have been identified in Alte Donau during the investigation period and the most common
species are encircled. (a) Coleps spetai, (b) Urotricha farcta, (c) Urotricha apsheronica, (d) Balanion
planctonicum, (e) Urotricha agilis, (f) Coleps hirtus, (g) Urotricha matthesi, (h) Monodinium balbi-
ani, (i) Lagynophrya acuminata, (j) Askenasia volvox, (k) Askenasia chlorelligera, (l) Mesodinium
pulex, (m) Didinium nasutum, (n) Cyrtolophosis mucicola, (o) Cyclidium glaucoma, (p) Uronema
nigricans, (q) Cinetochilum margaritaceum, (r) Strombidium viride, (s) Rimostrombidium lacustris,
(t) Tintinnidium fluviatile, (u) Halteria grandinella, (v) Codonella cratera, (w) Halteria chlorellig-
era, (x) Rimostrombidium hyalinum, (y) Rimostrombidium humile, (z) Pelagostrombidium fallax,
(aa) Rimostrombidium brachykinetum, (ab) Pelagostrombidium mirabile, (ac) Phascolodon vorti-
cella, (ad) Vorticella natans. Scale bar division = 20 μm. (Slightly modified from Pfister et al. 2004)
218 G. Pfister
Fig. 12.2 Micrographs from pelagic ciliates commonly found in Alte Donau from life. All pic-
tures modified from http://protist.i.hosei.ac.jp/PDB/Images/Ciliophora and size proportions
roughly adjusted. (a) Coleps spetai, (b) and (f) Mesodinium pulex, Urotricha farcta, (c) Urotricha
sp., (d) Balanion planctonicum, (e) Askenasia sp., (g) Halteria grandinella, (h) Uronema nigri-
cans, (i) Rimostrombidium sp., (j) Tintinnopsis sp., (k) Cyclidium glaucoma, (l) Cyrtolophosis
mucicola, (m) Codonella cratera, (n) Monodinium balbiani, (o) Vorticella sp.
Table 12.1 Mean annual ciliate numbers [cells ml−1] in Alte Donau during the years 1994–2000
Southern basin (AD1) Northern basin (AD4)
1994 1995/1996 1998 1999 2000 1994 1995/1996 1998 1999 2000
44.7 38.1 21.3 12.6 21.1 48.5 43.1 20.6 16.6 21.0
Table 12.2 Mean annual ciliate biomass [mg fresh weight m−3] in Alte Donau during the years
1994–2000
Southern basin (AD1) Northern basin (AD4)
1994 1995/1996 1998 1999 2000 1994 1995/1996 1998 1999 2000
291.6 548 275 160 238 474.1 562 198 216 245
year of RIPLOX-treatment, in 1996 (Teubner et al. 2003). The sudden increase of

biomass of ciliates and rotifers was thus a rapid but just a temporary response to the
large ecosystem change from phosphorus-rich to phosphorus poor conditions by
RIPLOX-treatment. The transient increase of these small planktonic animals coin-
cided with a short-term increase of carnivorous larger zooplankton, while herbivo-
rous zooplankton declined rapidly due to the significant decrease of phytoplankton
(Teubner et al. 2003, see also Chaps. 10 and 11). When cell numbers of ciliates
further declined during the late phase of the restoration period, also biomass
declined significantly in the years. Lowest annual mean ciliate biomass was observed
in the year 1999, the last year of the restoration period, when only 150–200 mg per
m3 where measured. With the re-colonization of macrophytes (see the period of re-
establishment of macrophytes in Chap. 8), the ciliates became again slightly more
abundant.
12.3.3 S
easonal Abundance and Biomass of Ciliates
in Alte Donau
Although mean annual values for ciliate numbers and biomass varied significantly
throughout the investigated period, the composition with regards to species stayed
surprisingly constant during the whole period. Therefore, data from the year 2000
are shown here as a representative example. As discussed above, from the years
1995 to 2000 main species belonged to the groups of the Prostomatida, Oligotrichida
and Hymenostomatida. In both basins of the lake prostomatid species (Urotricha
spp., Balanion planctonicum, Coleps spetai) were dominant in early spring, whereas
in summer and autumn the oligotrichs (Rimostrombidium spp., Strombidium spp.,
Tintinnopsis spp., Codonella cratera) gained highest importance. Hymenostomatdia
(Cyclidium glaucoma) just showed some significance in spring, whereas gymnos-
tomatids (Askenasia spp.) where usually found throughout the year, but in low
numbers (Fig. 12.3).
220 G. Pfister
a
50
45
40
35
30
25
20
15
10
5
0
16-Nov
16-Jun
16-Jul
16-May
16-Aug
16-Oct
16-Mar
16-Apr
16-Feb
16-Sep
b
50
45
40
35
30
25
20
15
10
5
0
16-Nov
16-Jun
16-Jul
16-May
16-Aug
16-Oct
16-Mar
16-Apr
16-Feb
16-Sep
Prostomatida Gymnostomatida Hymenostomatida Oligotrichida Others
Fig. 12.3 Abundance (cells ml−1) of pelagic ciliate groups in the southern (top) and northern basin
(bottom) of Alte Donau in the year 2000
Biomass of the different ciliate groups corresponds to the cell numbers through-
out the year. In all investigated years highest biomass values for prostomatid species
were found in spring. This peak was followed by a strong dominance of oligotrich
species throughout summer and autumn. As it was observed for cell numbers, bio-
mass of gymnostomatid species was usually low all year long. According to their
abundance, hymenostomatid species contributed to total ciliate biomass consider-
ably only in spring (Fig. 12.4). Oligotrichida and Prostomatida were also the two
predominant ciliate groups in the surface layer of oligotrophic alpine lake Traunsee
(Sonntag et al. 2006). Most of these epilimnetic ciliates referred to algivorous
species. Ciliates of the Hymenostomata were rather rare in the epilimnetic layer of
this deep alpine lake, which corresponds well to the minor importance of this group
among the ciliate assemblages of Alte Donau. The Hymenostomata, however, was
the dominant ciliate group in deeper strata from 60–160 m in Traunsee (Sonntag
a
600
500
400
300
200
100
0
16-Feb
16-May
16-Jun
16-Jul
16-Aug
16-Sep
16-Oct
16-Nov
16-Mar
16-Apr
b
600
500
400
300
200
100
0
16-Feb
16-May
16-Jun
16-Jul
16-Aug
16-Sep
16-Oct
16-Nov
16-Mar
16-Apr
Prostomatida Gymnostomatida Hymenostomatida Oligotrichida Others
Fig. 12.4 Biomass (mg fresh weight m−3) of pelagic ciliate groups in the southern (top) and north-
ern basin (bottom) of Alte Donau in the year 2000. The biomass time series for the year 1994 is
shown in Fig. 5 in Mayer et al. 1997
et al. 2006), where bacterioplankton becomes predominant (see depth profil of the
carbon ratio of phyto- to bacterioplankton in Teubner 2003). Most of these deep
living freshwater protozoans in Traunsee were known to be bacteriovorous species
(Sonntag et al. 2006) and refer thus to deep layer habitat conditions, which are not
relevant in shallow Alte Donau.
The ciliate abundance in Alte Donau in the year 2000 varied between 7 and 42
individuals per mL, which is shown in Fig. 12.3. According to a detailed study about
seasonal ciliate assemblages in Traunsee in 1997/1998, 2–12 individuals per mL−1
only were found in a nutrient-poor alpine lake (Sonntag et al. 2006). Comparing the
chlorophyll-a concentration in both lakes, which provides a rough estimation for
phytoplankton biomass, the annual mean in Alte Donau was 9 μg per L (7–11 μg L−1,
see Chaps. 6 and 9) and in Traunsee 1.04 μg per L−1 (range from 0.14 to 3.3 μg L−1).
As stated in the previous paragraph, most ciliate species living in the surface water
222 G. Pfister
layer of Alte Donau and Traunsee were algivorous. Phytoplankton is thus the main
food source for these protozoans. Comparing the both lakes, the ciliate abundance
in Alte Donau is only four times higher even if the chlorophyll is about even ten
times higher than in Traunsee. Ciliates in the mesotrophic Alte Donau thus seem to
play a less important role of rapid nutrient cycling than in an oligotrophic ecosystem
such as Traunsee.
Acknowledgments Many people in Mondsee and Vienna have contributed and helped during the
years of this survey. I want to address my special thanks to Martin T. Dokulil, who initiated this
project and brought me on-board during the initial years, and for his generous support throughout
the project. Thank you also to Arnold Nauwerck who was director of the Institute of Limnology in
Mondsee during the first years of the survey, for supporting me with material, advice and infra-
structure. Thomas Posch, Bettina Sonntag, Karl Donabaum and Monika Grossschartner are greatly
acknowledged for lots of help with coordination of sampling, sample preparation and protocol
optimization. Last, but not least, Katrin Teubner is to be acknowledged to have taken over the final
push and responsibility for publication as well as for valuable input and comments in finalizing this
manuscript.
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Chapter 13
Microbial and Viral Loop in Alte Donau:
A Case Study
Branko Velimirov, Ulrike R. Fischer, Alexander K. T. Kirschner,

and Claudia Wieltschnig
Abstract Attempting to describe the magnitude of the microbial carbon fluxes in

the water column of the Alte Donau we monitored basic parameters concerning
viruses, bacteria, and flagellates. Assuming a bacterial carbon conversion efficiency
of 31% and bacterial production of 2.56 μg C L−1 h−1 for the period from April to
September, the bacterial carbon demand would correspond to 8.26 μg C L−1 h−1,
which ought to be covered by primary production. For January to March and
October to December, we calculated a mean bacterial carbon demand of 3.06 μg C
L−1 h−1 and 3.48 μg C L−1 h−1 respectively. The balance calculations indicate that
heterotrophic bacteria would consume not all of the photosynthetically reduced car-
bon. Despite a positive C-balance in the water column its worthwhile mentioning
that benthic bacterial production needs to be considered before balance calculations
are forwarded for this ecosystem. The abundance of heterotrophic nanoflagellates
(HNF) and the high ratios of bacterial to HNF abundance (B:HNF range: 2 × 103 to
9 × 103, mean: 5.4 × 103) indicated a low bacterivorous impact by HNF. Calculated
ingestion rates were 2.2–26.5 bacteria HNF−1h−1 with a mean of 7.5 HNF−1h−1.
Grazing rates comprised between 0.3 and 20% of bacterial production, with a mean
of 5%. Virus-to-bacteria ratio (VBR) ranged from 4 in July to 39 in February, with
an average of 19. Viruses <60 nm capsid diameter dominated over all seasons
accounting for some 74% of the total, determined by transmission electron micro-
scope (TEM). To estimate the virus induced control of bacterial production the
number of lysed bacteria was obtained by dividing viral decay rates by the maxi-
mum burst size. On average 3.0 × 104 bacterial cells ml−1 h−1 had to be lysed to
maintain the monitored viral production implying that viruses controlled on average
56% of the bacterial production, ranging from 42 to 88%.
B. Velimirov (*) · U. R. Fischer · A. K. T. Kirschner · C. Wieltschnig

Institute of Medical Biology, Research Group General Microbiology, University of Vienna,
Vienna, Austria
e-mail: branko.velimirov@meduniwien.ac.at

https://doi.org/10.1007/978-3-319-93270-5_13
228 B. Velimirov et al.
Keywords Microbial carbon flux · Aquatic viruses · Heterotrophic bacteria ·

Heterotrophic nanoflagellates · Bacterial secondary production (BSP) · Viral loop ·
Microbial loop · Viral burst size
13.1 Introduction
The description of processes, which led to the formulation of the microbial loop
hypothesis (Pomeroy 1974) was a major breakthrough in the field of microbial ecol-
ogy and has certainly contributed to a better understanding of the functioning of
both marine and freshwater ecosystems. Major fluxes of organic matter, often
eclipsing the grazing food chain in quantity (Azam 1998) move as dissolved organic
matter into the bacterial compartment and the microbial loop (Azam et al. 1983;
Cho and Azam 1988; Münster 1991) It is now established knowledge that the
organic matter flux into the bacterial compartment is a major pathway in aquatic
systems and a number of field studies (Fuhrman et al. 1985; Cole et al. 1988) indi-
cate that up to one half of the oceanic primary production is channeled through the
bacterial compartment into the microbial loop. The amount of primary production
utilized by bacteria is highly variable between seasons and this variability has been
observed to influence the overall flux partitioning between the major pathways in
aquatic systems: the grazing food chain, the sinking fluxes of material, the microbial
loop and the ‘storage’ of dissolved organic matter in the bacterial biomass, and of
course carbon fixation itself.
Even though we still lack a conceptual framework to predict the variations in the
magnitude of the organic matter flux into the bacterial compartment, the microbial
loop hypothesis remains robust enough to explain the rapid remineralization and
recycling of inorganic nutrients under conditions of nitrogen or phosphorus limita-
tion in the water column above the thermocline.
The attractiveness of the microbial loop concept is seen in its dynamic behaviour,
which is the result of at least three major interacting ecological relationships such as
commensalism (mineral nutrient dependent production of ‘utilizable dissolved
organic matter’ by phytoplankton and subsequent utilization by bacteria), competi-
tion (for mineral nutrients by phytoplankton and bacteria) and predation (by micro-
flagellates on bacteria, influencing the outcome of the competition).
In the early 1990s, however, the recognition of the impact of virus induced lysis
on the bacterial production (e.g., Proctor and Fuhrman 1990; Bratbak et al. 1990;
Fuhrman and Suttle 1993; Børsheim 1993; Suttle 1994; Hennes and Simon 1995;
Fuhrman and Noble 1995) and its consequences (Proctor and Fuhrman 1990;
Bratbak et al. 1994; Mathias et al. 1995) led to the formulation of the viral loop
hypothesis, which had to be integrated into the microbial loop and which certainly
improved our understanding of the processes taking place within the microbial
compartment.
Also, one has to be aware of the fact that the concept of the microbial loop is as
dynamic as the functioning of the loop itself. As a consequence, the flow of informa-
tion emerging from recent studies on microbial ecology leads to a process of constant
13 Microbial and Viral Loop in Alte Donau: A Case Study 229
modification of the hypothesis, thereby contributing to the improvement of the pre-

dictive quality of models of the microbial compartment (Kirschner et al. 1998).
In this context the aim the study was not only to investigate the microbial loop in
order to model correct predictions on the carbon fluxes of the Alte Donau, but also
to contribute basic information for the ecological management of this eutrophic
aquatic system (scheme of restoration see Chapter 5). Even though all requested
parameters could not always be registered for similar time periods, a plausible
description on the magnitude of the carbon fluxes could be derived from the inves-
tigation. While the variability in bacterial numbers and bacterial production could
be followed up for 11 years in the stations Hofbauer (Obere Alte Donau = OAD,
north basin, morphometry of Alte Donau see Chap. 3) and Pirat (Untere Alte
Donau = UAD, south basin), viral numbers, production and the impact of viral
induced bacterial mortality on the microbial compartment was monitored from
January 1996–1997 (Fischer and Velimirov 2002). Similarly, abundances of hetero-
trophic nanoflagellates (HNF) were followed from 1995 to 1998 (Wieltschnig et al.
1999, 2001). Grazing rates of HNF on bacteria were estimated by the uptake of fluo-
rescently labelled bacteria (FLB) under in situ temperature conditions in 1995 and
1996 (Sherr et al. 1987; Wieltschnig et al. 1999). Additionally, growth parameters
of the HNF assemblage were determined in situ by selective filtration from 1996 to
1998 (Wieltschnig et al. 1999, 2001). For the sake of a better understanding of the
functioning of the microbial loop in the Alte Donau, the results of these studies were
integrated into the overall frame of the presentation. All bacterial parameters of the
water column were monitored on a biweekly bases for over 10 years, and for the
simplicity of the presentation it was decided to describe time courses of the various
parameters as yearly medians for the whole investigation period. As for the viruses
and HNF, the time courses over 12 months could be presented as mean monthly
values. When necessary, for comparative purposes time courses of bacterial num-
bers or production are mentioned for specific years where viruses or HNF were
investigated.
13.2.1 Heterotrophic Bacteria
The range of the median for bacterial numbers in the UAD and OAD remain within
2,5 × 109 cells L−1 and 4.8 × 109 L−1 from 1994 to 2003 (Fig. 13.1). For both sites
the lowest annual numbers were recorded for 1996, while all values from 1997 to
2003 were above 4 × 109 cells L−1. In contrast to abundance, the bacterial biomass
differences were less pronounced when comparing years 1994–96 (Fig. 13.2, see
also Teubner et al. 2003). These first 3 years of the 11-year study comprise the
eutrophied lake before the chemical treatment started and the two RIPLOX-years in
1995/96 (Chap. 5), when phytoplankton drastically decreased due to the reduction
OAD UAD
109 cells L-1
109 cells L-1

Fig. 13.1 Box-whisker plots of bacterial cell numbers in the water column of the Obere Alte
Donau (OAD) and Untere Alte Donau (UAD). Black bars: median, grey box: 50% confidence
interval (ci), whiskers: 75% ci, circles: extremes within 95% ci, asterisks: outliers outside 95% ci,
N = number of sampling events (Apr–Oct)
OAD UAD
µg carbon L-1
µg carbon L-1
Fig. 13.2 Box-whisker plots of bacterial biomass in the water column of the Obere Alte Donau
(OAD) and Untere Alte Donau (UAD). Black bars: median, grey box: 50% confidence interval (ci),
whiskers: 75% ci, circles: extremes within 95% ci, asterisks: outliers outside 95% ci, N = number
of sampling events (Apr–Oct)
of total phosphorus concentration from 2.37 μmol L−1 to 0.6 μmol L−1 (Teubner
et al. 2003, long-term time series of phytoplankton and total phosphorus in Fig. 6.4
in Chap. 6 and Fig. 9.1 in Chap. 9). In the long-term, bacterial biomass tended to
have the lowest values in 1996 (40 μC L−1) while values from 98 to 120 μC L−1 were
recorded between 1997 and 2003.
A different pattern was obtained for the time course of bacterial secondary pro-
duction (BSP), determined by following the incorporation of 3H-thymidine
12 12
11 11
OAD 10
UAD
10
9 9
mg carbon L-1 h-1
mg carbon L-1 h-1

8 8
7 7
6 6
5 5
4 4
3 3
2 2
1 1
0 0
N - 12 14 13 12 13 10 10 8 12 12 9 N - 11 14 13 12 13 10 10 12 9
1994 1996 1998 2000 2003 2005 1994 1996 1998 2000 2005
1995 1997 1999 2002 2004 1995 1997 1999 2003
Fig. 13.3 Box-whisker plots of bacterial secondary production in the water column of the Obere
Alte Donau (OAD) and Untere Alte Donau (UAD). Black bars: median, grey box: 50% confidence
interval (ci), whiskers: 75% ci, circles: extremes within 95% ci, asterisks: outliers outside 95% ci,
N = number of sampling events (Apr–Oct)
(Fuhrman and Azam 1982) and 3H-leucine (Simon and Azam 1989), where mini-
mum values of 1.5 μg C L−1 h−1 were obtained for OAD in the year 2000 and for
UAD in 1999 and 2000 (Fig. 13.3). Yet, the overall variability was not very pro-
nounced, ranging from 1.5 to 3.1 μg C L−1 h−1 at OAD and from 1.5 to 2.9 μg C l−1 h−1
in the UAD (BSP in comparison to primary production and other plankton param-
eters in the first 3 years of study see also Teubner et al. 2003, long-term time series
of primary production see Fig. 10.2 in Chap. 10). The values are well within the
range of bacterial production rates observed in other lakes such as Lake Mendota
(Pedros-Alio and Brock 1982), Lake Norrviken (Bell et al. 1983), Lake Mikolaskie
(Gajewski et al. 1993), the Kühwörter Wasser (Kirschner and Velimirov 1997) and,
with the exception of Australian billabongs (Boon 1991), maximal bacterial produc-
tion rates from other lakes were always in the magnitude of the values measured in
the Alte Donau.
Adopting a bacterial carbon conversion efficiency of 31% (Kristiansen et al.
1992), the carbon demand for the bacterial population of the water column was
estimated on the basis of the average monthly production rate from 1994 to 2005 for
the UAD and OAD. Even though we observed low variability of the bacterial pro-
duction rates, both within and between sites in the Alte Donau for the monthly aver-
ages, an overall mean was subsequently derived for those months where
phytoplanktonic primary production could be monitored (April to September; pri-
mary production see further Chap. 10). These bacterial production averages of 2.58
and 2.53 μg C L−1 h−1 for the UAD and OAD respectively, were higher than those
obtained from January to March (0.84 and 1.10 μg C L−1 h−1 respectively) and from
October to December (1.10 and 1.05 μg C L−1 h−1 respectively). Assuming that a
bacterial production of 2.56 μg C L−1 h−1 is representative of the water column from
Table 13.1 Heterotrophic nanoflagellate abundance (HNF-A), biomass (HNF-BM), growth rate
(HNF μ) and generation time (k) estimated in the Alte Donau in the years 1995 to 1998
HNF-A 106 cells L−1 HNF-BM μg C L−1 HNF μ d−1 HNF k h
1995 mean 0.87 7.9 nd nd
range 0.28–3.92 0.8–30.5
1996 mean 0.38 2.9 0.456 43
range 0.14–0.75 0.1–7.2 0.288–0.816 20–60
1997 mean 1.04 9.1 0.554 38
range 0.61–1.48 5.4–14.8 0.295–0.958 17–69
1998 mean 1.22 12.2 0.491 40
range 0.53–2.17 4.4–41.7 0.000–1.020 16–74
nd not determined
April to September, the corresponding bacterial carbon demand would correspond

to 8.26 μg C L−1 h−1, which ought to be covered by primary production. For the
periods from January to March and October to December, we calculated a mean
bacterial carbon demand of 3.06 μg C L−1 h−1 and 3.48 μg C L−1 h−1 respectively. To
estimate whether the water column’s trophic state is autotrophic or heterotrophic we
compared in a first step the average bacterial carbon demand from April to October
with the corresponding primary production. The outcome of the analysis was that
primary production could exceed all bacterial carbon demand including the winter
months from October to March (see also Fig. 10.2 in Chap. 10). The balance calcu-
lations indicate that heterotrophic bacteria in the water column would consume not
all of the photosynthetically produced carbon, but despite a positive balance it
should be pointed out that the benthic bacterial production should be considered
before a carbon budget can be forwarded for the ecosystem of the Alte Donau. First
data indicate that on average the production of the bacteria in the oxygenated sedi-
ment (i.e. 0–10 mm sediment layer) exceeds the bacterial production in the water
column by a factor of 800–1000. This would imply that the phytoplanktonic carbon
production alone cannot cover the total bacterial carbon demand in the Alte Donau.
More data on macrophyte production are needed in order to allow the development
of a representative carbon flow model for this ecosystem (standing crop of macro-
phytes for the period 1993–2005 see Fig. 8.7 in Chap. 8).
13.2.2 I mpact of Heterotrophic Nanoflagellates

on Heterotrophic Bacteria
The HNF assemblage was dominated by small cells of 2–5 μm cell length or cell
diameter, and a cell volume of <30 μm3, respectively. The abundance of HNF varied
from 0.14 × 106 to 3.92 × 106 cells L−1, and their biomass from 0.1 to 41.7 μg C L−1 h−1
(Table 13.1). The potential impact of HNF as bacterivores may be roughly estimated
when considering protozoan numbers, and the ratio of bacterial to HNF abundance
Fig. 13.4 Abundance of bacteria and heterotrophic nanoflagellates (HNF) measured in Obere Alte
Donau in year 1998
(B:HNF ratio). Yearly means of HNF abundance in the Alte Donau ranged from
0.38 × 106 L−1 to 1.22 × 106 cells L−1, which is on the lower end of the range
observed in other eutrophic and mesotrophic freshwater systems that often display
mean HNF densities larger than 1 × 106 cells L−1 with peak abundances of 107
cells L−1 (e.g., Nagata 1988; Sanders et al. 1992; Hansen and Christoffersen 1995).
This is a preliminary indication of a low bacterivorous impact by HNF, supported
by a lack of coupling of bacterial and HNF numbers during the course of a year
(time series for years 1998 see Fig. 13.4), and the observed high B:HNF ratio (range:
2 × 103 to 9 × 103, mean: 5.4 × 103). According to Sanders et al. (1992) this ratio
should not significantly exceed 1 × 103 if HNF were to exert a strong grazing impact
on heterotrophic bacteria.
Calculated ingestion rates were 2.2–26.5 bacteria per HNF per hour with a mean
of 7.5 bacteria HNF−1 h−1. The resulting grazing rates comprised between 0.3 and
20% of bacterial secondary production, with a mean of 5%. It is therefore evident,
that HNF were not able to control bacterial production and standing stock, indicat-
ing that factors other than consumption by HNF must be responsible for the regula-
tion of the heterotrophic bacterial compartment in this system (Amy and Morita
1983). It is worthwhile mentioning that HNF covered only a mean of 21% of their
carbon demand by the consumption of bacteria, despite the high abundance of bac-
terial carbon as a food source. This highlights that HNF use other carbon sources
(e.g., DOC or POM) in order to maintain production (Vaqué and Pace 1992).
Furthermore, it implies that the transfer of picoplanktonic carbon to higher trophic
levels via HNF was negligible, since we calculated that on average only 0.09 μg
bacterial C L−1 h−1 were consumed by HNF.
Fig. 13.5 Growth of HNF in unfiltered and filtrated samples shown by the number of cells during
the incubation time of 24 h. Filtering through a net of 10 μm mesh size was done to exclude
predators
Previous studies provide ample evidence for discrepancy between HNF grazing
and bacterial secondary production and an uncoupling of heterotrophic bacteria and
their protistan grazers (Sanders et al. 1989; Sherr et al. 1989; Pernthaler et al. 1996).
A number of investigations covering a broad range of system trophy has shown that
HNF densities do not keep pace with an increase in bacterial abundance with
increasing trophy because of growing predation pressure on HNF by ciliates and
metazooplankton (Šimek and Straškrabová 1992; Gasol and Vaqué 1993; Pace and
Vaqué 1994; Gasol et al. 1995).
In the Alte Donau, we have several indications for strong top-down control on
the HNF assemblage.
1. HNF growth rates ranged from 0 to 1.02 d−1, which corresponded to generation
times of 16–74 h (Table 13.1). The results correspond to values reported for other
freshwater systems (Nagata 1988; Carrick et al. 1992; Hansen and Christoffersen
1995; Weisse 1991, 1997). Yet, despite generation times being frequently less
than 1 day, HNF standing stock in the Alte Donau was low. Top-down control
was supported by filtration experiments conducted to compare net and gross
growth rates of HNF. While in unfiltered samples, HNF abundance remained
constant during a period of 24 h, it linearly increased in samples filtered through
a 10 μm mesh size gauze to exclude predators, inclusively ciliates (Fig. 13.5). So
we conclude that HNF production was completely consumed.
2. The B:HNF ratio also provides an indicator of the strength of top-down-control
on HNF (Gasol 1994). This ratio has been shown to be highly correlated with
metazoan zooplankton abundance and biomass in that increased abundance or
biomass of different zooplankton coincided with a greater number of bacteria per
flagellates. The observed B:HNF ratio indicated strong control of the HNF
assemblage during most of the year.
3. Finally, we have indirect indications for top-down control of the HNF assem-
blage from the taxonomic composition of the zooplankton in the Alte Donau.
Examining the relevant literature, we found that the dominating taxa are known
to ingest nanoplankton (Pace and Vaqué 1994; Cleven 1996; Jürgens et al. 1996).
13.2.3 Viral Impact on Heterotrophic Bacteria
A logical consequence of the above findings was to test the hypothesis that the low
grazing impact of HNF in the water column of the Alte Donau could be balanced by
a disproportionately high viral impact on bacterial production.
Variations in viral numbers over the investigation period of 12 months ranged
from 1.7 × 107 to 11.7 × 107 particles ml−1 and averaged 5.0 × 107 particles ml−1
(Fig. 13.6). The corresponding virus to bacteria ratio (VBR) ranged over an order of
magnitude from 4 in July to 39 in February, with an average of 19. Both, tailed and
non-tailed viruses, ranging in capsid diameter from 30 to 180 nm were observed
over the year. As a general pattern, viruses smaller than 60 nm dominated over all
seasons and accounted for some 74% of the total as determined by transmission
electron microscope (TEM, see Fig. 13.7). Bacteria were scored as virus-infected
when they contained at least 5 intracellular viruses, which were identified on the
basis of structure, size and intensity of staining. Distinction between two types of
burst size was made: minimum burst size, i.e. the average number of mature phages
in all visibly infected bacterial cells (VIC) and maximum burst size, i.e. average
number of viruses in cells that were completely filled with viruses. Of the 1800
14 14
Bacteria
12 12
Viruses
10 10
106 Bacteria ml -1
107 Viruses ml -1
8 8
6 6
4 4
2 2
0 0
J F M A M J J A S O N D J F
Months
Fig. 13.6 Seasonal variation in total viral and bacterial counts in the Obere Alte Donau in 1996/97.
Error bars represent standard deviations of 6 subsamples
100
<60nm 90-<150nm
90
60-<90nm >150nm
% of total viral population 80
70
60
50
40
30
20
10
0
J F M A M J J A S O N D J F
Months
Fig. 13.7 Seasonal variation of the viral capsid size classes. The capsid diameters were estimated
in four size classes in the Obere Alte Donau in 1996/97. A minimum of 70 capsids was measured
per subsample. Error bars represent standard deviations of 6 subsamples
virus-infected bacteria examined to determine the burst size at least 36% were com-
pletely filled with viruses. The average minimum burst size (average number of
phages in all visibly infected cells) was 22 viruses per cell, average maximum burst
size (average number of phages in cells that were completely filled with phages)
was 42 viruses per cell. The frequency of infected bacterial cells (FIC) can be esti-
mated by applying conversion factors presented by Proctor et al. (1993) to the fre-
quency of visibly infected cells (FVIC), which varied between 2.8 and 9% and
averaged 5%. Using the average conversion factor of 5.42 from Proctor et al. (1993)
it was shown that on average 28% of all bacterial cells were infected. Additionally,
the FIC was also estimated by taking into account the effect of grazing mortality on
the relationship between FVIC and FIC, assuming that infected and uninfected bac-
terial cells are grazed at equal specific rates (Binder 1999). Although this assump-
tion was not taken into consideration by Proctor et al. (1993) only a negligible
change was calculated, leading to 29% of infected cells in the bacterial population
instead of 28%.
In order to quantify viral production, viral decay experiments were performed
periodically, i.e. the decrease in viral concentration over time was recorded after
inhibiting the production of new viruses by the addition of potassium cyanide
(KCN), which arrested the bacterial metabolism over 24–30 h (Heldal and Bratbak
1991). For this approach, one has to assume that the viral abundance in a system
remains constant through time and that viruses from lysed bacterial cells replace
viruses lost by decay, thus leading to a measure for viral production Simultaneously
to the viral decay experiments, BSP was measured under the same environmental
conditions. All experiments were carried out at water temperatures ranging from
6
total <60 nm 60-90 nm
2
A
5 y (total) = -0.247 ln(x) + 2.393; R = 0.815; n=8
2
y (<60nm) = -0.213 ln(x) + 1.975; R = 0.763; n=8
2
(60-90nm) = -0.030 ln(x) + 0.288; R = 0.352; n=8
4
2
107 Viruses ml-1
06
total <60 nm 60-90 nm
2
y (total) = -0.065 ln(x) + 2.622; R = 0.312; n=8
B
5 2
y (<60nm) = -0.026 ln(x) + 1.707; R = 0.055; n=8
2
y (60-90nm) = -0.047 ln(x) + 0.809; R = 0.178; n=8
4
0
0 1 2 3 4 5 6 7 8 9 10
Time (h)
Fig. 13.8 Changes in total viral counts and of viral capsid size classes <60 nm and 60- < 90 nm as
a function of time after inhibiting production of new viruses in (a) Oct 96 (highest viral decay rate
of 1996/97) and (b) Dec 96 (lowest decay rate of 1996/97). Inhibition was induced by the addition
of KCN. Each data point represents the value of 1 triplicate sample
2.5 °C to 23 °C, being representative for the seasons. The viral decay can be described
as a negative exponential for all experiments during the first 9 h of the incubation
(Fig. 13.8), after which no obvious loss of viruses occurred. Thus, decay rates were
calculated with data from the initial 9 h. Two representative examples of a decrease
in viral numbers during decay experiments are shown in Fig. 13.8. Viral decay rates
varied significantly (Kruskal-Wallis one-way Anova, p < 0.05) between the seasons.
Between 50 and 84% of the initial viral population remained after 9 h in the experi-
mental jars, with an average of 68%. The number of lysed bacteria was obtained by
dividing viral decay rates by the maximum burst size (Table 13.2), which indicated
that between 1.5 × 104 and 5.5 × 104 cells ml−1 h−1 had to be lysed in order to main-
tain the viral production, averaging 3.0 × 104 bacterial cells ml−1 h−1. This implied
that viruses controlled on average 56% of the BSP, ranging from 42 to 88%.
Table 13.2 Viral abundance, viral decay rate, frequency of infected bacterial cells (FIC) calculated
after aProctor et al. (1993) using the average factor of 5.42, not considering grazing, and bBinder
(1999) considering grazing (assumption: specific grazing rate of infected bacterial cells equals
specific grazing rate of uninfected cells), virus-mediated bacterial mortality (calculated by dividing
the viral decay rate by the maximum burst size), and viral control of bacterial secondary production
(BSP)
Viral
Viral Viral decay Bacterial control
abundance rate Viral decay mortality of BSP
Date (107 ml−1) (106 ml−1 h−1) rate (h−1) FICa (%) FICb (%) (104 ml−1 h−1) (%)
7/96 2.3 (0.1) 1.0 (0.1) 0.055 (0.004) 41 (2) 41 (1) 2.2 (0.2) 42 (4)
8/96 3.1 (0.1) 1.3 (0.1) 0.050 (0.005) 48 (10) 43 (5) 3.1 (0.3) 57 (5)
9/96 5.3 (0.1) 1.2 (0.2) 0.025 (0.003) 17 (3) 19 (3) 2.4 (0.3) 51 (6)
10/96 3.7 (0.0) 2.1 (0.1) 0.077 (0.002) 15 (0.5) 18 (0.5) 5.5 (0.1) 88 (2)
12/96 3.0 (0.1) 0.5 (0.1) 0.019 (0.004) 19 (0.1) 22 (0.1) 1.5 (0.3) 43 (8)
Avg 3.5 (0.4) 1.2 (0.2) 0.045 (0.009) 28 (6) 29 (5) 3.0 (0.6) 56 (8)
Values in parentheses denote SEs of triplicate samples
13.3 Conclusion
Our data indicate that viruses play a major role in the control of the bacterial com-
munity in the Alte Donau. Viral lysis of BSP prevailed over grazing by HNF by a
factor of 11. Adding up the bacterial mortality due to viral lysis and the mean con-
tribution of HNF (5%), reveals that both factors together are on average responsible
for only 61% of the total bacterial mortality. So, some 40% of the bacterial produc-
tion may be directly available for the micro-and macrozooplankton. Yet, it should be
mentioned that viruses could occasionally remove far than 100% of the BSP, cor-
responding to the lysis of 1.6% h−1 of the bacterial standing stock. High bacterial
mortality due to viral lysis indicates that relative to the bacterial carbon production
an important amount of organic matter released into the water column became avail-
able for microheterotrophs.
However, it has to be emphasized that values for virus-induced recycled bacterial
carbon need to be corrected for the energetic costs needed for the replication pro-
cess of viral DNA and capsid protein synthesis within the infected bacterial cells.
Assuming a viral carbon content of 10−10 μg C (Gonzalez and Suttle 1993), it implies
that between 1.3 and 5 μg C L−1 d−1 is needed for the synthesis of DNA and capsid
proteins. Therefore the amount of carbon released via viral lysis of bacterial cells
was at most 5–39 μg C L−1 d−1 (mean: 15 μg C L−1 d−1) corresponding to 29–79% of
the BSP (mean: 46%, rather than the previously calculated 42–88%) whereby no
correction for the C-loss as CO2 during the above described process could be made
(Fischer and Velimirov 2002).
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Chapter 14
The Effect of Restoration Measures
on the Benthic Invertebrates of a Danube
Backwater (Alte Donau)
Berthold Janecek, Patrick Leitner, Otto Moog, and Katrin Teubner
Abstract Benthic invertebrates were used as bioindicators to document the effect

of restoration measures in the backwater Alte Donau in Vienna, a former side-arm
of the Danube. The study covers four periods of lake management: (1): the mesotro-
phic year before eutrophication (1987), (2): the 2 years of chemical iron chloride
treatment aimed at the phosphate precipitation in the water column and the oxidiza-
tion of nitrate-treated sediment surface layers (1995–1996), (3): further 3 years of
other lake management measures during the restoration period (1995–1999), and
(4): an early stage of the re-establishment of underwater vegetation (2000, 2003).
Over eight survey years from 1987 to 2003, about 330 benthic invertebrate taxa with
three most abundant systematic groups were identified: 37 species of oligochaetes,
23 species of molluscs (18 gastropods and 5 bivalves), and 190 species of the chi-
ronomids and other dipterans. The trophic classification index that refers to a habitat
quality score by chironomids indicates the year 1987 as mesotrophic (3.46, the
index range for mesotrophic conditions is 2.50–3.49). In this year the chironomids
and oligochaet species inhabited a variety of diverse habitats ranging from soft sedi-
ments (clay and mud), sand, gravel, pebbles and stones to dense stands of macro-
phytes. The biomonitoring record of 15 mollusc species was significantly higher in
this mesotrophic reference year than in any other following survey year. The trophic
classification index denotes 1995 as the most eutrophied year. With the restoration
and the re-establishment of macrophytes the values of this index decreased but
remained higher than during the mesotrophic condition. In the years 1995 and 1996,
when the chemical treatment with iron chloride, slaked lime and calcium nitrate was
applied and the macrophytes were rare, the biomass of chironomids and oligochaets
B. Janecek · P. Leitner · O. Moog

Department of Water, Atmosphere and Environment, Institute of Hydrobiology and Aquatic
Ecosystem Management, University of Natural Resources and Life Sciences, Vienna, Austria
e-mail: berthold.janecek@boku.ac.at; patrick.leitner@boku.ac.at; otto.moog@boku.ac.at
K. Teubner (*)

https://doi.org/10.1007/978-3-319-93270-5_14
244 B. Janecek et al.
was particularly low. Among the three important taxonomic groups, i.e. the chirono-
mids, bivalves, and oligochaets, only the first two groups achieved relative biomass
dominance. The relative importance of chironomids over oligochaets during these
both years of the Riplox-treatment might rather mirror the losses of oligochaets
being affected by nitrate exposure than the re-colonization by ‘new’ chironomid
species in the sediment. In addition, in particular active filter feeders such as bivalves
seemed to adjust well to the muddy sediment environment after chemical treatment.
In the following years of the restoration and the early stage of macrophyte re-
establishment, oligochaets and bivalves became the dominant groups mainly con-
tributing to the macrozoobenthic biomass, while the biomass of chironomids
remained relatively low. Different from the indication by enhanced water transpar-
ency and low phytoplankton biomass achieved by restoration measures in 2004, the
shift towards a mesotrophic environment seemed to be retarded when assessing Alte
Donau by the chironomid habitat quality score index. The main reason why the
chironomid assemblage did not follow the other indicators of mesotrophic condi-
tions in 2004 can be seen in the disruption of the sediment (e.g. by sedimentation of
precipitation chemicals) and associated loss of underwater habitat structure due to
still relatively low macrophyte biomass during the last invertebrate survey. The phy-
tophilic chironomid species still had a low abundance when only about 50% of the
macrophyte biomass was recorded if compared with the mesotrophic situation in
1987. Despite the wax and wane of benthic invertebrates described during the eight-
year survey, invertebrate biomass is relatively high in the long-term average and
thus characterises Alte Donau as a lowland environment along habitats from low- to
high altitudes.
Keywords Oxbow lake · Benthic fauna · Bio-indicator · Chironomids · Pupal

exuviae of chironomids · Bivalves · Oligochaets · Re-colonization of sediment ·
Lake restoration · Phosphate precipitation · Riplox
14.1 Introduction
Benthic invertebrates are a diverse group, containing a large number of species of

various systematic origins, widely used as bioindicators (Rosenberg and Resh 1993;
Woolsey et al. 2007; Moog et al. 2018). They are important consumers for accelerat-
ing the turnover of autochthonous and allochthonous organic matter in aquatic sys-
tems and serve as substantial food source for fish (Merritt and Cummins 1996;
Moog 2002; Jungwirth et al. 2003; Moog and Hartmann 2017). They are living on
the sediment surface, in the interstices and also colonize underwater vegetation
from littoral to deeper profundal zones in a lake. The survey of invertebrates in Alte
Donau was carried out sporadically in years of varying trophic states and restoration
14 The Effect of Restoration Measures on the Benthic Invertebrates of a Danube… 245
measures and thus differs logistically from those of planktic community of bacteria,
algae, ciliates and other animals described in Chaps. 9, 11, 12, and 13. Nevertheless,
the macrozoobenthos surveys in Alte Donau have the advantage of being consis-
tently studied by the same person (B. Janecek) when considering the species deter-
mination for 8 years over a whole period from 1997 to 2003.
The benthic invertebrates in Alte Donau have been described for year 1987
before the eutrophication and various periods of the restoration measures. The main
aim of this chapter is to characterise the benthic invertebrate community for these
different periods (the description of the four basic periods of restoration measures
are described in Chaps. 5 and 6). Beside the species description, a trophic index
(Fittkau 1992; Orendt 1993) is used to assess the water quality before eutrophica-
tion and along years of restoration. Other results derived from the pre-dominance
pattern of individual groups within the benthic community and from shifts in the
similarity of the community structure by multivariate ordination plots are used to
identify the main macrozoobenthic development. Among the benthic community a
strong focus is given on the chironomids because this taxonomic group represented
the highest species diversity throughout the 8-year benthic community survey in
Alte Donau. Chironomids are commonly used for assessing this type of freshwaters
(e.g., Wolfram 1996; Moog and Chovanec 2000).
14.2 Methods and Description of the Investigation Area
The oxbow lake Alte Donau in Vienna, a former side arm of Danube River was cut
off from the main river for more than 160 years (Dokulil et al. 2010). With the regu-
lation of the Danube River 1870–1875 (Chap. 2), a new river bed was built for navi-
gation (New Danube), while the former stretch of the Danube River was named Alte
Donau. Alte Donau thus became a groundwater-seepage lake, which was used for
boating, fishery, and poultry farming (goose husbandry). This oxbow lake has also
a long tradition as a popular recreational area (Chaps. 2 and 19).
Macrozoobenthos biomonitoring surveys were carried out about two times a year
from 1995 onward in Alte Donau. The peak season for sampling was autumn, fol-
lowed by summer contributing about 40% and 30% to the data set, respectively.
Taking into account the spatial heterogeneity along the elongated shape of Alte
Donau, in total more than 25 sampling sites were surveyed (Fig. 14.1), which
included four sites for the bottom samples and seven sampling sites for chironomid
pupal exuviae. Lake bottom samples were taken by scuba diving using a hand net
(625 cm2 area, 100 μm mesh size) or with a PVC-corer (27 cm2 area). Pupal exuviae
of chironomids were sampled by sweeping the surface with a pond-net. The sam-
ples were stored in plastic bottles and preserved with 4% formaldehyde.
Fig. 14.1 Sampling sites.

Transects of bottom
samples taken in 1987 are
marked by I–VI. Bottom
samples at sites A and D
were taken from 1995 to
2003, at B in 1995 and at
G in 1999, pupal exuviae
at Pe1 to Pe5 in 1996,
1997 to 2000 and 2003, at
Pe6 in 2000 and 2003, and
at Pe7 in August 1997 and
September 1997
14.2.1 S
pecies Identification and Determination
of the Biomass
The separation of benthic invertebrate specimens was done by hand-picking, for

some samples warm sucrose solution (30 °C, conc. c. 1.13 g per ml H2O) was used
to separate the invertebrates from the remaining sample (Kajak et al. 1968).
All larger animals with a length of up to 2–3 mm were collected and separately
labelled according to major taxonomic affiliations. A fraction of smaller animals
was obtained by sub-sampling (volumetric method) to reduce the effort required for
sorting and identification. For calculating the biomass of the benthic invertebrates,
the weight was measured using a precision balance (accuracy of 1 per 10.000 g).
The conversion of fresh weight into ash free dry weight and caloric equivalent is
described in Janecek (1985, 1995).
After taxonomic identification, the specimens have been preserved in 80% etha-
nol. Many chironomids and especially a lot of their pupal skins have been processed
Fig. 14.2 Chironomidae found in Alte Donau – (a) Larva of Polypedilum sp. (Chironominae –
Tribus Chironomini), (b–f) Heads of larvae, (b): Paratendipes cf. albimanus (Chironominae –
Tribus Chironomini), (c): Virgatanytarsus cf. arduennensis (Chironominae – Tribus Tanytarsini),
(d): Cricotopus sylvestris (Orthocladiinae), (e): Nanocladius rectinervis (Orthocladiinae), (f):
Prodiamesa olivacea (Prodiamesinae). (Photos a, c and f by Wolfram Graf; b, d and e by Sabine
Schiffels)
to permanent slides embedded in Euparal. For the identification of chironomids, a

microscope with a magnification of at least 400x (for some details 1000x with oil
immersion) was most appropriate. Some common macrozoobenthos species are
shown by photographs in Fig. 14.2.
14.2.2 Graphical Display and Statistical Methods
The snapshot sampling of macrozoobenthos from 1987 to 2003 did not rely on a
regular sampling frequency but took advantage of presenting a synoptic measure of
species distributions and environmental situations over a long-time perspective in
Alte Donau. Data were graphically and statistically treated by distribution free
methods, taking into account the sporadic sampling in space and time for the ben-
thic invertebrates described before (see non-parametric tests that are statistically
satisfying large data sets by snap shot sampling in ecology in Krienitz et al. 2016).
For the graphical display of the main biomass pattern among the dominant inver-
tebrate groups along the large range of biomass (1.13–2789 g fw m−2), we reduced
the information of metric invertebrate biomass to a binary set of presence and
absence data (Fig. 14.3). We separated the invertebrates in oligochaetes, bivalves
and chironomids, which refer to the most species-rich or most persistent inverte-
brate groups, and in the remaining invertebrates (see species and taxonomic affilia-
tions in Table 14.1). In addition, the predominant invertebrates among these groups
for each sample were identified (the predominance of an invertebrate group is
defined by contributing a higher biomass to the total biomass of macrozoobenthos
than individually other groups, compare with the method of Krienitz et al. 2016).
The original data averaged for each year are shown for chironomids and oligochaets
in Fig. 14.4.
The trophic chironomid index (Fig. 14.5, see further method description below)
was depicted as box-whisker plots using SigmaPlot 10 (SPSS Inc., Chicago, USA).
The line inside the box indicates the median, both ends of the box the 25th and 75th
percentile, and the error bars the 10th and 90th percentile.
Multivariate analysis was used to determine whether the sampling sites were
placed into specific groups that will minimize variance within groups and maxi-
mize variance among chironomid groups. We applied NMS (Non-metric Multi-
dimensional Scaling) as an ordination technique that seeks to explain the variation
in species community data using as few dimensions as possible (Kruskal 1964).
NMS was analysed based on log (x + 1) transformed benthic invertebrate data
using PC-ORD version 5. In this analysis sites with similar taxa composition are
plotted close to each other in the scatter plot while dissimilar sites are plotted far
from each other (Fig. 14.6). The following parameters were used in NMS ordina-
tions: distance measure = Sorenson Bray-Curtis; starting configuration = random;
runs with real data = 50; step-down in dimensionality from 6 dimensions to 1
dimension; initial step length = .2; maximum number of iterations per run = 400;
stability criterion = .00001; iterations to evaluate stability = 10; Monte Carlo (ran-
domized data) runs = 50.
a: 1995-1996
Others predom
presence
Bivalvia
Oligochaeta
Chironomidae
0.1 1 10 100 1000 10000
b: 1997-1999
Others
Bivalvia
Oligochaeta
Chironomidae
0.1 1 10 100 1000 10000
c: 2000, 2003
Others
Bivalvia
Oligochaeta
Chironomidae
0.1 1 10 100 1000 10000

-2
Total biomass (g fw m )
Fig. 14.3 Presence of invertebrates displayed along the total invertebrate biomass during three
periods of restoration: (a) – chemical phosphate precipitation by RIPLOX-method, (b) – further
restoration measures, (c) – re-establishment of macrophytes (n = 36). The composition of inverte-
brates is depicted by the three groups Chironomidae, Oligochaeta and Bivalvia that are mainly and
most persistently contributing to the invertebrate biomass in addition to remaining invertebrates
(others) (species see Table 14.1). An invertebrate group is displayed as present (dots) if the contri-
bution to the total invertebrate biomass exceeds 5%. Note that each invertebrate sample can be
represented by all four groups. A bar indicates the predominant invertebrate for each sample. Note
the log-scale of the total biomass of invertebrates. Further description see methods
Table 14.1 Macrozoobenthic taxa found in Alte Donau from 1987 to 2003
TAXONOMIC AFFILIATION TAXON
CNIDARIA: Hydra sp. WARINGER in LÖFFLER, ed.,
Pelmatohydra oligactis PALLAS; Craspedacusta sowerbii LANKESTER
TURBELLARIA: Dendrocoelum lacteum O.F. MÜLLER, Dugesia tigrina GIRARD
NEMATODA: Nematoda Gen. sp.
GASTROPODA: Acroloxus lacustris LINNAEUS, Bithynia tentaculata LINNAEUS, Potamopyrgus antipodarum GRAY, Galba truncatula O.F. MÜLLER,
Radix auricularia LINNAEUS, R. ovata DRAPARNAUD,
Physella acuta DRAPARNAUD, P. heterostropha SAY,
Gyraulus albus O.F. MÜLLER, G. crista L., G. sp., Planorbis planorbis L., Hippeutis complanatus LINNAEUS, Valvata cristata O.F. MÜLLER,
V. piscinalis O.F. MÜLLER, V. studeri BOETERS & FALKNER,
Viviparus acerosus Bourguignat, V. contectus Millet
BIVALVIA: Dreissena polymorpha PALLAS, Casertiana nitida JENYNS,
Anodonta anatina LINNAEUS, A. cygnea LINNAEUS,
Unio pictorum latirostris KÜSTER
OLIGOCHAETA: Aeolosoma sp., Criodrilus lacuum HOFFMEISTER,
Eiseniella tetraedra SAVIGNY, Stylodrilus heringianus CLAPAREDE, Amphichaeta leydigii TAUBER, Chaetogaster diaphanus GRUITHUISEN,
Dero digitata MÜLLER, D. obtusa D’UDEKEM, D. sp.,
Nais christinae KASPRZAK, N. communis PIGUET,
N. pseudobtusa PIGUET, N. elinguis MÜLLER, N. simplex PIGUET, N. sp., Ophidonais serpentina MÜLLER, Pristina longiseta EHRENBERG,
Slavina appendiculata D’UDEKEM, Specaria josinae VEJDOVSKY,
Stylaria lacustris LINNAEUS, Vejdovskyella comata VEJDOVSKY,
V. intermedia BRETSCHER, Propappus volki MICHAELSEN,
Aulodrilus japonicus YAMAGUCHI, Branchiura sowerbyi BEDDARD, Ilyodrilus templetoni SOUTHERN, Limnodrilus claparedeianus RATZEL, L.
hoffmeisteri CLAPAREDE, L. profundicola VERILL, L. sp.,
Potamothrix bavaricus ÖSCHMANN, P. hammoniensis MICHAELSEN,
P. moldaviensis VEJD. & MRÁZEK, Psammoryctides barbatus GRUBE, P. moravicus HRABE, P. sp.,
Tubifex cf. ignotus STOLC, Tubifex tubifex MÜLL
HIRUDINEA: Dina punctata JOHANNSON, Erpobdella octoculata LINNAEUS,
Alboglossiphonia hyalina MÜLLER, Helobdella stagnalis LINNAEUS
CRUSTACEA -MYSIDACEA: Limnomysis benedeni CZERNIAWSKI
CRUSTACEA -AMPHIPODA: Chelicorophium curvispinum SARS,
Dikerogammarus haemobaphes EICHWALD, D. sp.,
Echinogammarus ischnus BEHNING, Gammarus roeselii GERVAIS,
CRUSTACEA -ISOPODA: Asellus aquaticus LINNAEUS
ARACHNIDA-ACARINA: Hydrozetes lacustris MICHAEL, Trimalaconothrus novus SELL.,
Hydrodroma despiciens O.F. MÜLLER, Lebertia sp.,
Frontipoda musculus O.F. MÜLLER, Limnesia undulata O.F. MÜLLER,
Hygrobates longipalpis HERMANN, Unionicola aculeata KOENIKE,
U. crassipes O.F. MÜLLER, Neumania deltoides PIERSIG,
Piona discrepans KOENIKE, P. pusilla NEUMAN,
P. pusilla rotundoides THOR, Hydrochoreutes krameri PIERSIG,
Forelia curvipalpis VIETS, Axonopsis complanata O.F. MÜLLER, Brachypoda versicolor O.F. MÜLLER, Mideopsis orbicularis O.F. MÜLLER,
Arrenurus sp., Porolohmanella violacea KRAMER
EPHEMEROPTERA Caenis horaria L., C. lactea BURMEISTER,
C. luctuosa BURMEISTER, C. luctuosa/macrura,
C. rivulorum EATON, C. sp., Cloeon dipterum L., Ephemerella sp.,
Ephemera danica MÜLLER, E. vulgata LINNAEUS, E. sp.
ODONATA: Coenagrion puella LINNAEUS, Erythromma najas HANSEMANN, Ischnura elegans Van der LINDEN Platycnemis pennipes PALLAS, Orthetrum
sp.
PLECOPTERA: Nemoura sp.
HETEROPTERA: Gerris sp., Ilyocoris cimicoides LINNAEUS
PLANIPENNIA: Sisyra sp.
COLEOPTERA: Haliplus sp., Oulimnius tuberculatus MÜLLER, Donacia sp.
TRICHOPTERA: Ecnomus tenellus RAMBUR, Hydropsyche contubernalis McLACHLAN, Orthotrichia costalis CURTIS, Oxyethira flavicornis PITET, O. sp.,
Athripsodes cinereus CURTIS, Athripsodes juv.,
Ceraclea dissimilis STEPHENS, Leptocerus tineiformis CURTIS, Mystacides azurea LINNAEUS, Mystacides longicornis LINNAEUS,
M. sp., Oecetis lacustris PICTET, Oecetis ochracea CURTIS, O. sp., Psychomyia pusilla FABRICIUS, Tinodes waeneri LINNAEUS
(continued)
Table 14.1 (continued)
DIPTERA: Bezzia sp., Monohelea sp., Chaoborus flavicans MEIGEN, C. Sp.,
Ablabesmyia longistyla FITTKAU, A. monilis LINNAEUS,
A. cf. monilis LINNAEUS, Acricotopus lucens ZETTERSTEDT,
Brillia bifida KIEFFER, Bryophaenocladius cf. nidorum EDWARDS, B. sp., Chironomus acutiventris/obtusidens, C. cf. annularius nec DE GEER,
C. balatonicus DÉVAI W. & S., C. balatonicus/plumosus,
C. longipes STAEGER, C. longistylus GOETGHEBUER,
C. luridus STRENZKE, C. nudiventris RYSER S. & W., C. nuditarsis KEYL, C. plumosus LINNAEUS, C. plumosus, C. pseudothummi STRENZKE,
C. sp., Cladopelma bicarinata BRUNDIN, C. goetghebueri,
C. virescens MEIGEN, C. viridula LINNAEUS, C. sp.,
Cladotanytarsus atridorsum KIEFFER, C. lepidocalcar KRÜGER,
C. mancus WALKER, C. mancus, C. nigrovittatus GOETGHEBUER,
C. vanderwulpi EDWARDS, C. wexionensis BRUNDIN, C. sp., Corynoneura arctica KIEFFER, C. carriana EDWARDS,
C. coronata EDWARDS, C. gratias SCHLEE, C. scutellata WINNERTZ, C. sp., Cricotopus albiforceps KIEFFER, C. cf. arcuatus HIRVENOJA,
C. bicinctus MEIGEN, C. cylindraceus KIEFFER, C. festivellus KIEFFER,
C. flavocinctus KIEFFER, C. fuscus KIEFFER, C. intersectus STAEGER, C. cf. laricomalis, C. pilitarsis ZETTERSTEDT, C. reversus HIRVENOJA,
C. ricotopus FABRICIUS, C. sylvestris, C. trifasciatus MEIGEN,
C. sp., Cryptochironomus albofasciatus STAEGER, C. obreptans WALKER, C. psittacinus MEIGEN, C. supplicans MEIGEN, C. sp., Cryptotendipes
holsatus LENZ, C. ryptotendipes usmaensis PAGAST,
C. sp., Demicryptochironomus vulneratus ZETTERSTEDT,
Dicrotendipes lobiger KIEFFER, D. nervosus STÆGER,
D. notatus MEIGEN, D. pulsus WALKER, D. tritomus KIEFFER, D. sp., Einfeldia pagana MEIGEN, Endochironomus albipennis MEIGEN,
Glyptotendipes mancunianus EDWARDS, G. lyptotendipes glaucus/pallens, G. pallens MEIGEN, G. paripes EDWARDS, G. signatus KIEFFER,
G. viridis MACQUART, G. sp., Harnischia curtilamellata MALLOCH, H. arnischia fuscimana KIEFFER, H. sp.,
Heterotrissocladius marcidus WALKER, Hydrobaenus lugubris FRIES, Kiefferulus tendipediformis GOETGHEBUER,
Labrundinia longipalpis GOETGHEBUER,
Limnophyes minimus MEIGEN, L. sp., Microchironomus tener KIEFFER, Microtendipes britteni EDWARDS, M. britteni/confinis,
M. chloris MEIGEN, M. icrotendipes pedellus DE GEER,
Nanocladius balticus PALMÉN, N. anocladius bicolor ZETTERSTEDT, N. anocladius rectinervis KIEFFER, Nilotanypus dubius MEIGEN,
Orthocladius consobrinus HOLMGREN, O. rthocladius fuscimanus KIEFFER, Parachironomus arcuatus GOETGHEBUER, P. biannulatus
STAEGER, P. mauricii KRUSEMAN, P. tenuicaudatus MALLOCH,
P. varus GOETGHEBUER, P. vitiosus GOETGHEBUER,
P. conversus WALKER, P. sp., Parakiefferiella bathophila KIEFFER,
P. coronata EDWARDS, P. sp., Paralauterborniella nigrohalteralis MALLOCH, Paramerina cingulata WALKER,
Paratanytarsus bituberculatus EDWARDS, P. dimorphis REISS,
P. dissimilis JOHANNSEN, P. aratanytarsus inopertus WALKER,
P. laetipes ZETTERSTEDT, P. tenellulus GOETGHEBUER,
P. tenuis MEIGEN, P. sp., Paratendipes albimanus MEIGEN,
Paratrichocladius rufiventris MEIGEN, Phaenopsectra cf. flavipes MEIGEN, P. sp., Polypedilum bicrenatum KIEFFER, P. convictum WALKER,
P. cultellatum GOETGHEBUER, P. nubeculosum MEIGEN,
P. sordens V.D. WULP, P. sp., Procladius choreus MEIGEN,
P. lugens KIEFFER, P. rufovittatus V.D. WULP (Psilotanypus rufovittatus), P. sagittalis KIEFFER, P. signatus ZETTERSTEDT, P. sp., Prodiamesa
olivacea MEIGEN, Psectrocladius limbatellus HOLMGREN, P. sectrocladius cf. brehmi KIEFFER, Psectrocladius oxyura LANGTON,
Psectrocladius oxyura/sordidellus,
P. psilopterus KIEFFER, P. sordidellus ZETTERSTEDT,
P. schlienzi WÜLKER, P. sp., Pseudochironomus prasinatus STAEGER, P. sp., Schineriella schineri STROBL, Stempellina almi BRUNDIN,
S. subglabripennis BRUNDIN, S. minor EDWARDS,
Stenochironomus gibbus FABRICIUS, Stictochironomus sp., Synorthocladius semivirens KIEFFER, Tanypus kraatzi KIEFFER, T. punctipennis
MEIGEN, T. sp., Tanytarsus bathophilus KIEFFER, T. chinyensis GOETGHEBUER, T. cretensis REISS, T. ejuncidus WALKER, T. eminulus
WALKER, T. excavatus EDWARDS, T. glabrescens EDWARDS, T. gregarius KIEFFER, T. heusdensis GOETGHEBUER,
(continued)
T. inaequalis GOETGHEBUER, T. lactescens EDWARDS,
T. lestagei GOETGHEBUER, T. lestagei-Agg. Typ 1&2,
T. cf. longitarsis KIEFFER, T. mancospinosus EKREM & REISS,
T. medius REISS & FITTKAU, T. mendax KIEFFER, T. mendax/occultus, T. nigricollis GOETGHEBUER, T. nigricollis/usmaensis,
T. occultus BRUNDIN, T. pallidicornis WALKER, T. sensu Langton,
T. signatus V.D. WULP, T. smolandicus BRUNDIN,
T. sylvaticus V.D. WULP, T. cf. usmaensis PAGAST,
T. verralli GOETGHEBUER, T. spp., Thienemanniella sp., Virgatanytarsus sp. sensu Langton & Visser,
Xenochironomus xenolabis KIEFFER, Zavreliella marmorata V.D. WULP
Sites and sampling period see Fig. 14.1
3500
0
3000
0 Oligochaeta
Abundance (ind. m )
-2
Chironomidae
2500
0
2000
0
1500
0
1000
0
5000
0
1995 1996 1997 1998 1999 2000 2003
25
20
Biomass (g m )
-2
15
10
0
1995 1996 1997 1998 1999 2000 2003
RIPLOX restoration macrophytes
Years
Fig. 14.4 Abundance and biomass of oligochaetes and chironomids displayed as bar charts cover-
ing the main three periods of restoration as in Fig. 14.3
14.2.3 Trophic Classification Index
In addition to abundance and biomass data of invertebrates described in the method

before, also habitat quality scores were applied to track the long-term habitat
changes in Alte Donau. Such scores are retrieved from a meta-analysis of empirical
observations of biotic community structure or other phenomena discovered across
individual habitats (Karr 1991; Hofmann 1994; Brettum 1989; Moog and Chovanec
2000; Moog 2002; Henderson 2003; Blenckner et al. 2007; Moog and Hartmann
2017). Habitat scores are applied to infer an ecologically sound integrity of aquatic
habitats without referring to the individual data again. The trophic classification
index suggested by Fittkau (1992) and Orendt (1993) was applied because in Alte
Donau (1) the chironomids were a representative and species-rich group among the
invertebrates (Table 14.1), and (2) the habitat change of Alte Donau was mainly due
to nutrient enrichment and subsequent nutrient reduction passing restoration
4.5
Trophic index 4.0
3.5
3.0
87 95 96 97 98 00 03
Year
Fig. 14.5 Chironomid assemblages used as bioindicators: Trophic index calculated from the
abundance of chironomid species for individual years (1987–2003)
Fig. 14.6 Ordination scatter plot for data points that refer to the chironomid species communities
in benthic samples from individual years (1987–2003)
measures (Chaps. 5 and 6). This habitat quality score refers to those chironomid
species that are most frequently contributing to the chironomid community within a
narrow trophic range. Empirical regressions described in detail in, e.g., Orendt
(1993) provide evidence that the population density of many chironomid species
responds significantly to the total concentration of phosphorus, i.e. to the total pool
of the nutrient element which is primarily understood to control the bottom up

growth in freshwaters. The chironomid trophic index displayed in Fig. 14.5 covers
many species, and thus is a community metric mirroring the habitat conditions
along the phosphorus gradient and associated effects. The index consists of five
ranks and indicates the trophy as follows:
1.00–1.99 oligotrophic
2.00–2.49 oligo- to mesotrophic
2.50–3.49 mesotrophic
3.50–3.99 meso- to eutrophic (in some lists up to 4.49)
4.00–5.00 eutrophic
14.3 Results
14.3.1 B
enthic Macroinvertebrate Species Inventory of Alte
Donau
A total number of 330 benthic invertebrate taxa were identified during the extended
invertebrate surveys over 8 years from 1987 to 2003 (Table 14.1). The oligochaetes
(37 species), the molluscs (18 species of gastropods and 5 bivalves), and the chi-
ronomids together with other Diptera (190 species) were the three main groups of
benthic invertebrates (Tables 14.1 and 14.2). Specimens from these systematic
units have been found in all years of the habitat surveys in Alte Donau. Eighty spe-
cies of other taxonomic affiliations such as Cnidaria, Turbellaria, Nematoda,
Hirundinea, Crustacea (Ostracoda), Mysidacea, Amphipoda, Isopoda, Arachnida-
Acarina, Ephemeroptera, Odonata, Plecoptera, Heteroptera, Planipennia,
Coleoptera, and Trichoptera were further recorded during this study (Table 14.1).
Comparing the chironomid species inventories of 1987 with those of each follow-
ing year, 5–24 chironomid species were found that haven’t been observed during
the very detailed initial survey in the mesotrophic year 1987 (Table 14.2). The
settlement of this countable number of newly observed chironomid species illus-
trates the dynamic and also the persistence of chironomids throughout surveys dur-
ing restoration and macrophyte re-establishment in Alte Donau.
The observation of these invertebrates is attributed to four periods: (1) the meso-
trophic reference condition in 1987, before the oxbow lake has undergone the
strong eutrophication in the early nineties, (2) the 2 years of chemical phosphate
precipitation (iron chloride treatment, 1995, 1996), (3) a 3 year period of further
restoration treatments (1997–1999) and (4) 2 years of the re-establishment of mac-
rophytes (2000, 2003) (more details about the periods see Chap. 6). In addition to
a description of particular species and habitat change during these lake periods, the
main pattern of invertebrate development is graphically shown in Figs. 14.3, 14.5
and 14.6.
Table 14.2 Number of collected species of the seven main taxonomic affiliations (n.c. not
collected, * only bottom samples). The number of chironomid species, which were newly found in
Alte Donau when compared with the survey in 1987 are listed from 1995 onward (last row)
1987 1995 1996(97) 1997 (98) 1998 1999 2000 2003
Gastropoda 13 5 7 4 4 2 2 3
Bivalvia 2 3 3 2 1 4 1 2
Oligochaeta 7 13 10 16 16 21 8 14
Crustacea – Malacostraca n.c. 3 2 3 3 4 4 2
Ephemeroptera 3 3 3 2 2 3 3 3
Trichoptera 5 5 8 5 3 12 5 2
Chironomidae 68 39* 91 77 67 114 114 90
Chir. spp., new for AD 15 24 10 6 31 18 5
14.3.2 Year 1987 – Mesotrophic Conditions
The survey in year 1987 characterises the period before eutrophication of Alte
Donau with luxuriant stands of macrophytes, which were mainly built up by water-
milfoils (Myriophyllum spp.) and stoneworts (charophytes, most widespread spe-
cies was Nitellopsis obtusa) (macrophyte survey see Chap. 8). In the extended
surveys in spring (April, May) and early summer (June), the Naididae, which can
occur on both, the sediment and the macrophytes, dominated the Oligochaeta com-
munity. The motility of these species is well recognised as they can leave the sedi-
ment climbing up to higher parts of the plants when faced with oxygen deficiency
near the lake bottom. Stylaria lacustris and Pristina aequiseta were the most abun-
dant species of Naididae, followed by Nais communis, N. elinguis and N. pseudob-
tusa. In contrast to the years after the iron chloride treatment, Tubificidae (Tubifex
tubifex, Potamothrix hamoniensis) were rarely found in the sediments in 1987. An
explanation for the lack of these Oligochaeta species under mesotrophic conditions
may be the relatively coarse-grained to sandy sediments, the large fraction of coarse-
textured detritus of plant material (further details see e.g., Learner et al. 1978;
Schneider et al. 1988; Bauer and Waringer 1987; Hauer et al. 2018).
The record of 15 mollusc species (spp.) in 1987 was higher than in any of the
following survey years of this study (Table 14.2) (Haberlehner 1987). In compari-
son to the benthic fauna of some larger sized backwaters in the alluvial forests east
and west of Vienna (e.g. floodplain of Stopfenreuth: 34 spp., Lobau floodplains: 27
spp., Haberlehner 1986), however, Alte Donau serves as fragmented and modified
habitat for a smaller number of species and lower animal densities (history of river
cut-off Alte Donau see method and Chap. 2).
The Chironomidae were the third important benthic invertebrate group, with
respect to abundances (individual numbers and biomass) and a large variety of spe-
cies during the whole study period. In general, the chironomid communities in the
fine sediment exhibited low species numbers even though high densities of 4200
individuals m−2 were found in January (1357–12,995), 13,650 individuals m−2 in
March (3. March: 5913–31,490 ind. m−2), 3060 individuals m−2 in May (27. May:
1676–5579 ind. m−2) and 2810 individuals m−2 in June (15. June: 1410–5599 ind.
m−2). The biomass values ranged from 0.81 in May to 2.02 in June and thus were
moderate despite the peak abundances. The comparably low total biomass of chi-
ronomids was due to the small size of the abundant species, e.g., belonging to the
Tanytarsini (Tanytarsus glabrescens, T. spp., Paratanytarsus tenuis, P. spp.) and
Orthocladiinae (Parakiefferiella bathophila, Psectrocladius sordidellus). Merely
two Chironomus species (C. plumosus, C. balatonicus) were of large size and were
only common at single sites. Chironomini were in general not numerous, with the
exception of up to 480 individuals m−2 for Chironomus plumosus and up to 560
individuals m−2 for Parachironomus arcuatus in June. Tanypodinae mainly con-
sisted of Procladius (Holotanypus) spp. and Tanypus punctipennis. Only a few spe-
cies have been found exclusively on sediments, for example Cryptochironomus
spp., Stictochironomus spp. or Cladotanytarsus mancus. A larger number of species,
however, was found on both the macrophytes and the bottom sediments and was
thus able to use the diverse habitat structure during this mesotrophic period with
dense macrophyte stands. Members of this transition group were Ablabesmyia
monilis, Parakiefferiella bathophila, Psectrocladius sordidellus, Dicrotendipes tri-
tomus and some species of Paratanytarsus and Tanytarsus (e.g., T. glabrescens).
Polypedilum nubeculosum was found on the submerged plant species Ceratophyllum
demersum and Myriophyllum spicatum, and even Chironomus plumosus can occur
on macrophytes such as Chara tomentosa, Nitella obtusa and Myriophyllum
spicatum. The following species were collected exclusively from macrophytes
Chara tomentosa and Nitella obtusa or from periphyton: Nanocladius bicolor,
Cricotopus sp., Dicrotendipes lobiger, Endochironomus albipennis, Parachironomus
tenuicaudatus and Glyptotendipes viridis (mining in macrophytes Sagittaria and
Alisma, see Kalugina 1975). Glyptotendipes signatus refers to the same habitat
behaviour but is also known to settle on the bryozoans such as Plumatella fungosa
(Janecek 2005).
The trophic indices were calculated from the pupal exuviae collections and refer
three times to mesotrophic and another three times to meso-eutrophic conditions.
The range of variability in the trophic index is much larger in 1987 when compared
with later years (Fig. 14.5). In the following survey years mesotrophic conditions
could only be identified for certain sites and only twice: site Pe3 sampled on 14.8.
1998 (value for the trophic index = 3.49) and site Pe1 on 19.8. 2000 (value for the
trophic index = 3.47) (sites see Fig. 14.1).
The NMS ordination plot of scores illustrating the similarity or dissimilarity of
chironomid assemblages is shown in Fig. 14.6. One symbol represents an investiga-
tion site. The benthic communities of the scores from 1987 (depicted as black tri-
angles) are not overlapping with the scatterplots of the other years and thus confirm
the unique status of chironomid community structure. They are separated most dis-
tant from those of the other years along the first NMS axis mirroring substantial
differences in the community structure (Fig. 14.6). These main changes in the chi-
ronomid species composition can be well verified when going back to the original
survey information. In the years after 1987 and with chemical treatment of phos-
phate precipitation, some of the most abundant species such as of the Tanytarsini
(Tanytarsus glabrescens) completely disappeared.
14.3.3 Y
ears 1995 and 1996 – Chemical Treatment
for Phosphate Precipitation
In the period of strong eutrophication of Alte Donau in 1992 and 1993, the formerly
clear-water oxbow lake turned into a turbid water body with large biomass blooms
of phytoplankton (mainly Cylindrospermopsis raciborskii and Limnothrix redekei,
see Chap. 9). The submerged water plants almost disappeared under low underwater
light availability (Chaps. 5 and 8), the annual mean Secchi depth was 0.85 and
0.89 m (shown in Fig. 11.5 in Chap. 11). The benthic invertebrate surveys for 1995
and 1996 were conducted weeks to months after the chemical precipitation treat-
ment with iron chloride (April 95 and April 96). With the deposition of sedimentary
layers rich in organic carbon (biodegraded material formed during eutrophication
together with precipitate containing phosphate, slaked lime and calcium nitrate by
chemical treatment, Ripl 1976), large areas of the sediment surface were then cov-
ered by sapropel while sandy river sediments of a grain diameter 100–250 μm
became rare (Janecek 2000). In accordance with the sparse underwater vegetation in
1995 (Fig. 8.1 in Chap. 8, 1.6 m annual mean Secchi depth in Fig. 11.5 Chap. 11),
in particular, the phytal macrozoobenthos species of the Chironomidae and
Trichoptera were rarely found. The abundance of the phytal species, however,
immediately increased again when the submerged vegetation started to recover due
to elevated water transparency in 1996 (Chap. 8, 2.1 m annual Secchi depth in
Chap. 11). Particular abundant macrozoobenthic species observed in 1987, such as
the Chironomidae Tanytarsus glabrescens, Paratanytarsus laetipes, P. tenuis, none-
theless, could not be detected in 1995 and 1996 (Janecek 2000). In turn, many new
species of oligochaets and Chironomidae were recorded in these 2 years, when
compared with the macrozoobenthos survey in year 1987 (Table 14.2). 14 species
of Chironomidae were identified for the first time in Alte Donau and three of them
became quite frequent in the sediments: Procladius rufovittatus (Psilotanypus rufo-
vittatus), Stempellina subglabripennis and Stempellinella minor. An even higher
number of newly found species were recorded for the second year of chemical treat-
ment (Table 14.2). Some of the newly found species in Alte Donau prefer sediments
(e.g. Cryptotendipes sp., Glyptotendipes paripes, Microtendipes cf. britteni,
Polypedilum sordens and Tanytarsus medius) or even live on water plants or stones
(e.g. Glyptotendipes pallens, G. viridis and Polypedilum sordens). Procladius rufo-
vittatus (Psilotanypus rufovittatus) is known for migration and was found in larger
densities. Some species newly found in these 2 years might be called pelophilic and
are more or less tolerant to oxygen deficiency. They are typical species of eutrophic
waters. Large species such as Chironomus plumosus, Ch. balatonicus, Tanypus
punctipennis and T. kraatzi could reach peak abundances of up to 520 individuals
per m2 and thus became dominant in terms of individual species biomass. According
to the large distance of data point for 1987 and 1995 along the first NMS axis
(Fig. 14.6) the species composition of the chironomids shifted largely from meso-
trophic conditions to the first year of chemical treatment. Despite the high number
of species observed, the abundance and biomass of chironomids and also of
Oligochaets were in general low in 1995 and 1996 (Figs. 14.3a and 14.4), which can
be seen as an initial negative effect of the chemical phosphate precipitation treat-
ment. When comparing the biomass of these two macrozoobenthos groups, the chi-
ronomids became predominant against the oligochaets (Fig. 14.3a). Beside the
chironomids, which were often pre-dominant when low total biomass of macrozoo-
benthos was found, bivalves became also often the most dominant group during
both years of the chemical treatment (Fig. 14.3a) but stand for sampling dates and
sites of high total biomass of macrozoobenthos. It thus seems that bivalves, as active
and efficient filtrating collectors, can cope well with the chemical phosphate floc-
culation. In this view the two iron chloride treatments had some positive effects on
Bivalvia and Chironomidae as the large number of species at low abundance
increased the diversity in the macrozoobenthic community in particular for the lat-
ter. When looking at the trophic index, which is calculated from the abundance of
chironomid species, the community indicated eutrophic conditions in 1995 and a
meso-eutrophic environment in 1996 (Fig. 14.5).
A clearer picture of the compositional shifts in the macrozoobenthos community
is recognised when assessing the functional feeding groups. In accordance with
large sediment surface areas that are covered by fine particulate organic matter,
detritivorous macrozoobenthos became the most important feeding group in 1995
and 1996. Filtrating collectors that are able to accumulate sinking organic particles
were quite abundant too. We further found macrozoobenthic predators especially
among the Chironomidae, namely Tanypodinae and Cryptochironomus spp. that are
known to be at least partially carnivorous.
The compositional changes in the chironomids toward species that are primarily
living in gravel and sandy sediment slightly altered the values of the trophic index
(Fig. 14.5) and the overall species composition (Fig. 14.6) toward a chironomid
community structure that was described under mesotrophic conditions
14.3.4 Years 1997–1999– Further Restoration Measures
In the 3 years of the “restoration period” after chemical phosphate precipitation, the
annual Secchi depth reached values from 1.7 to 2 m (Fig. 11.5 in Chap. 11). The
macrophyte cover did not increase substantially and remained at moderate biomass
levels comparable with the year 1997. The abundance and biomass of both the oli-
gochaets and chironomids increased when compared with the 2 years of chemical
phosphate precipitation (Fig. 14.4). In view of the biomass contribution of species
groups allocated to large taxonomic affiliations, the macrozoobenthos community
changed from the predominance of chironomids and bivalves during the chemical
treatment to a community dominated by oligochaets and bivalves in subsequent
years (Fig. 14.3b). From 1997 to 1999 the muddy sediment in particular at the
deeper parts of the lake seemed to build up a stable environment that provides an
ideal habitat for large mussels such as Anodonta cygnea and Unio pictorum. Among
the chironomids a number of species such as Parakiefferiella bathophila and
Paratanytarsus spp. could reach higher densities on less muddy sediments, i.e.
gravel and sand. In case of oligochaets the Naididae became dominant again as
commonly described for mesotrophic conditions in 1987. In 1999, the number of
species and biomass of chironomids and oligochaets was highest when compared
with those of the previous and the following years (Fig. 14.4). The peak abundance
and biomass for four main taxonomic affiliations for individual sampling sites were
36,900 individuals per m2 with a corresponding biomass of 42.20 g fw per m2 for the
oligochaets, 81,790 individuals per m2 with a corresponding biomass of 19.42 g fw
per m2 for the chironomids, 4200 individuals per m2 with a corresponding biomass
of 1.53 g fw per m2 for the Ephemeroptera, and 900 individuals per m2 with a cor-
responding biomass of 1.75 g fw per m2 for the Trichoptera. In eutrophic lakes
annual averages of macrozoobenthos biomass can reach up to 650 g fw per m2 in the
profundal zone (Brinkhurst 1974). In such ecosystems the biomass of Chironomidae
usually contributes from 30 to more than 100 g fw per m2 (Janecek 1985, 1987).
Beside active filterers also predators were found such as the free-living flatworm
Dugesia tigrina and the leech Helobdella stagnalis. Among the Chironomidae a
number of carnivorous (or at least partially carnivorous) species have been observed:
Ablabesmyia longistyla, A. monilis, Procladius spp., Cryptochironomus spp.,
Tanypus kraatzi and T. punctipennis. Some species that are commonly known for
running waters have been observed in 1999 in Alte Donau, such as Propappus volki
belonging to the Oligochaeta (this oligochaet species occurred also in the Danube
River, see Moog 2002; Moog and Hartmann 2017), Hydropsyche contubernalis of
the Trichoptera and Polypedilum convictum of the Chironomidae. Most species of
the Chironomidae, but also of the Oligochaeta and Trichoptera, were living on the
open bottom areas and thus became quantitatively important whereas phytophilic
species (such as for example Ophidonais serpentina belonging to the Naididae) did
not contribute significantly to abundances and biomass within the macrozoobenthic
community. In case of the chironomids this result is also supported by the percent-
ages of pupal exuviae: Here the most phytophilic species did not even reach 1% of
the total drift. The dominance of macrozoobenthos species that are living in the
sediment in association with rarely found phytolitic species indicates that the sedi-
ment surface had recovered as valuable habitat for the benthic fauna while macro-
phytes were still missing during this period in Alte Donau.
The compositional changes in the chironomids shifted toward species that were
primarily living in gravel and sandy sediment. These changes correspond to a slight
decrease of the trophic index indicating eutrophic-mesotrophic conditions
(Fig. 14.5). The overall species composition displayed in the NMS-ordination plot
shows that the chironomid community structure tends to be less different from
mesotrophic conditions than in year 1995 (Fig. 14.6, along the first NMS-axis points
for 1997–1999 are less distant to points of the mesotrophic year 1987 than com-
pared with the distance of points of mesotrophic year 1987 and eutrophic year
1995).
14.3.5 Y
ears 2000 and 2003 – Period of the Re-Establishment
of Macrophytes
The 2 years of macrozoobenthos surveys within the period of re-establishment of

macrophytes (2000–2006, Fig. 5.4 in Chap. 5) are in the beginning and middle of
the 7-year period. In 2000 and 2003 the annual mean Secchi depth was 1.9 and
2.3 m, respectively. The water transparency in these years was thus similar to that in
1999 (Fig. 11.5 in Chap. 11). While the macrophyte biomass surveyed for the whole
basin of Alte Donau had increased consecutively from 1996 to 1999 by 10 to almost
15 tons dry weight (Figs. 8.1 and 8.7 in Chap. 8), the biomass did not exceed 15 tons
in 2000. The second year of macrozoobenthic survey, year 2003, however, is the
first “macrophyte transition” year with a significant increase to 125 tons dry weight
biomass of underwater vegetation. From 2004 onward the annual mean Secchi
depth increased significantly from 2.8 to 3.9 m and macrophytes reached remark-
able stands of 300–500 tons dry weight in Alte Donau (under mesotrophic condi-
tions in 1987 macrophytes reached 721 tons dry weight, Chap. 8). Macrozoobenthos
surveys, however, are only described for 2000 and 2003 and, thus, relate to an envi-
ronment of still low water transparency and a moderately enhanced macrophyte
biomass in 2003. The abundance was much lower than in the years before. In 2000
and 2003 about a half of the oligochaets and a third of the chironomids of the peak
abundance in 1999 could be reached (Fig. 14.4). Nevertheless, the biomass of oligo-
chaets was still high while the biomass of chironomids was considerably low. The
oligochaets dominated the benthic community most frequently when comparing the
biomass of the main taxonomic affiliation groups (Fig. 14.3c) even though the total
macrozoobenthos biomass was relatively low when compared with previous periods
of restoration (Fig. 14.3a, b). More subtle differences in compositional shifts were
found when considering single macrozoobenthic species. Stempellinella minor was
a common chironomid species in the years of chemical phosphate precipitation
inhabiting the littoral sediment areas without submerged vegetation. Their abun-
dance declined slowly in years of stepwise re-establishment of macrophytes. Some
chironomids that are commonly found at less nutrient rich habitats, such as
Cladotanytarsus nigrovittatus (trophic index 3.6), for example, became more abun-
dant in Alte Donau while C. mancus (trophic index 4.0) declined. Large profundal
living Chironomus species such as C. balatonicus and C. plumosus that are com-
monly found under eutrophic conditions and that are expected to survive even in
anoxic periods became less often observed. They reached lower abundances in 2000
and 2003 than compared with the restoration period in Alte Donau before. The pro-
fundal chironomid fauna typically associated with sustained mesotrophic condi-
tions (e.g., Tanytarsus bathophilus, T. gregarius and T. inaequalis) were,
nevertheless, quantitatively still of minor importance. When assessing the whole
chironomid community, the median values of the trophic index for 2000 and 2003
tended to decrease slightly (Fig. 14.5) indicating a slow development toward
eutrophic-mesotrophic conditions. In addition, the chironomid species composition
did not differ considerably from those in 1997–1999 (see NMS-ordination plot in
Fig. 14.6) and thus indicates again that the chironomid community structure is still
very different from observations in mesotrophic 1987.
In patchy sediment areas where the bottom structure clearly indicated the
approved iron chloride treatment to reduce phosphate availability by precipitation,
a community of active filter feeders was reaching high biomasses in the two survey
years. Dreissena polymorpha that was in particular found in 2000 and Pisidium niti-
dum and Pea mussels that were commonly observed in 2003 thus seem to be the
most persisting active filter feeders which can still occur under more stable condi-
tions after the restoration period has been passed.
14.3.6 C
omparison of Macrozoobenthos Biomass
across Habitats
The abundance and biomass of benthic fauna varied among large taxonomic affilia-
tions corresponding to the environmental disturbance by restoration measures and
different trophic states in Alte Donau. It must be stated here that the relatively low
trophic index of 1987 (3.46) representing mesotrophic conditions (2.50–3.49,
Fittkau 1992) was never reached again during the following years of the investiga-
tion. The habitat biomass of macrozoobenthic communities, however, can also vary
across altitudes (Jungwirth et al. 1980), (Fig. 14.7). The biomass of macrozooben-
thos increases exponentially when following the aquatic habitats from the source at
high altitude to the river mouth at low altitudes in Austria. Among Austrian water
bodies, Alte Donau represents an ecosystem at particular low altitude that is located
in the lowland close to the Danube river flood plain in Vienna. The enhanced bio-
mass for Alte Donau (137 g per m2) thus corresponds well to the exponential increase
of macrozoobenthic biomass at low altitude habitats (see also the correspondence of
the data point for Alte Donau in the log-log plot in the inset of Fig. 14.7).
14.4 Discussion
14.4.1 Species Inventory of Alte Donau
The knowledge on the benthic invertebrate fauna of the Austrian Danube section has
shown a remarkable input in the last 15 years of the twentieth century. Nine years
after Russev (1985) who reported 167 species, Humpesch and Moog (1994)
observed 897 species of the macrozoobenthos for the River Danube in Austria. Of
these, 306 species were restricted to the free-flowing main river channel, 354 spe-
cies to the backwaters of impoundments and 683 to the backwaters of the riverside
forests. A couple of years before the European Water Framework Directive was put
140 2.5
Biomass (LOG gm-2)

2.0
120 1.5
1.0
100
Biomass (g m-2)
0.5
0.0
80 -0.5
2.0 2.2 2.4 2.6 2.8 3.0 3.2 3.4
60 Altitude (LOG m)
40
20
0
0 500 1000 1500 2000 2500
Altitude (metres above sea level)
Fig. 14.7 Biomass of invertebrates sampled from benthic habitats across altitudes. Inset displays
the same data but on double logarithmic scale. The black dot refers to the median biomass of Alte
Donau (1995–2003). The white dots are from Dückelmann 2001 in Jungwirth et al. 1980, 2003
into practice, the upcoming European environmental water policy stimulated a

number of Danube and large river studies with a focus on environmental status
assessment and discussions about mitigation measures of flood protection, naviga-
tion and hydropower use effects. As a result, Moog et al. (1995) published a list with
1142 invertebrates species in the Austrian Danube followed by a total of 1289 spe-
cies 5 years later (Moog et al. 2000).
The study on the benthic fauna of Alte Donau contributes valuably to the knowl-
edge of the biodiversity of the Austrian Danube system. A total of 330 benthic
invertebrate taxa were recorded by Löffler et al. (1988) and during the current inver-
tebrate surveys from 1995 to 2003. This is a comparably high number if one takes
into consideration, that the entire water body of Alte Donau is located in the urban
area close to the centre of Vienna. The importance of Alte Donau as an urban diver-
sity hot spot is also confirmed by the fact that from the whole floodplain area around
Vienna (downstream Greifenstein, river-kilometre 1940, to Vienna-Freudenau,
river-kilometre 1920) a total of 511 species and 625 species have been reported in
1995 and 2000 respectively (Moog et al. 1995, 2000; Moog and Hartmann 2017).
This means that the comparatively small water body Alte Donau offers a habitat for
65% and 53% respectively of the benthic invertebrate species that are known from
the floodplains around Vienna.
14.4.2 I mpact of Habitat Disturbance on Benthic Community

in Alte Donau
Benthic invertebrates are the most preferred biomonitoring organisms studying

stressors and drivers in freshwaters from temperate to subtropical and tropical zone
(e.g., McLachlan 1974; Wolfram et al. 1999; Stendera et al. 2012; Mengistou 2016;
Cai et al. 2017; Moog et al. 2018). Despite their high value as bioindicator in natural
habitats (e.g., for chironomids in Fittkau 1992 and Orendt 1993) and their metric-
based differences in benthic invertebrate communities between restored and non-
restored sites (Sundermann et al. 2011), several studies emphasise the difficulties of
tracking the success of restoration by retarded re-colonization of benthic inverte-
brates (Langford et al. 2009; Stendera et al. 2012). It is rather argued that the link-
age between habitat change and structural change in the invertebrate community is
less pronounced than assumed by restoration practitioners (Jähnig et al. 2010;
Sundermann et al. 2011). Our results of benthic invertebrate biomonitoring surveys
are in accordance with these studies as the improvement of water quality indicated
by these organisms is much slower than expected by other restoration indicators. In
case of Alte Donau, however, the delay in the recovery of the benthic community
might be mainly due to coating of biota and sediment surface by chemical precipi-
tants, on the one hand, and a too abrupt habitat shift by drastic nutrient reduction on
the other. In the year 2004, when periods of chemical phosphate precipitation
(1995–1996) and the moderate recovery of submerged macrophyte stands have
been passed, the chironomid trophy index indicated only a slight recovery of the
species composition compared with the mesotrophic conditions in 1987 in Alte
Donau. Other organisms than benthic invertebrates, such as phytoplankton, that are
commonly used to assess water quality, indicate the “success” of a restoration more
rapidly. Already from year 2001 onward, the chlorophyll-a concentration revealed
low values indicating a decrease of phytoplankton biomass under mesotrophic con-
ditions in Alte Donau (Fig. 5.5 in Chap. 5, see period 3 in Fig. 9.10 and 9.11 in
Chap. 9). An assessment by phytoplankton relies on species that have a short gen-
eration time ranging from hours to few days. Only phytoplankton taxa living in a
more persistent environment in deep chlorophyll layers in deep lakes (Teubner et al.
2003b) are known to have a longer life span of up to 5–7 days (e.g., Gervais 1998,
Zotina et al. 2003). The rapid decline of phosphorus in the water column due to
phosphorus flocculation in Alte Donau thus immediately suppressed the growth of
short-lived phytoplankton and was associated with significant compositional shifts
from the dominance of cyanobacteria to mainly eukaryotic algae (Teubner et al.
2003a). Different from these planktonic primary producers, all other planktonic
organisms such as heterotrophic bacteria, protozoans and metazoans, responded dif-
ferently and also not necessarily coherently to the shortage of the total phosphorus
pool (Teubner et al. 2003a). Unlike these planktonic assemblages the benthic inver-
tebrates are taxonomically and ecologically a very heterogeneous group. On the one
side, they are comprised of species that may live for only a few weeks to months as
part of their life as larvae (e.g. chironomids larvae in warm bodies). On the other
side, this group also includes species with a life span of five to ten or even much
more years (for tubifex see e.g., Jónasson and Thorhauge 1972, for Bivalvia
Cummings and Graf 2009). In view of an ecological perspective the habitat struc-
ture of benthic species is much more complex than of planktonic organisms and,
thus, benthic invertebrates were affected in different ways by the restoration mea-
sures in Alte Donau. Taking into account this ecologically heterogeneity, the re-
settlement of benthic invertebrates after the Riplox-treatment toward species that
are associated with mesotrophic conditions might have been driven by three aspects:
–– Suppression by the initially abrupt and further lasting shortage of planktonic
food that was primarily achieved by the growth control of phytoplankton under
reduced phosphorus availability.
–– Vulnerability to the short-time habitat destruction and toxicity from the chem-
ical sediment treatment and the settlement of the phosphorous precipitate on
sediment surface aimed at reducing internal phosphorus load and thus to reduce
the total phosphorus pool in the water column and.
–– Colonization of the new habitats by the long-term re-settlement of macro-
phytes concomitantly with the increasing water transparency.
The impact of these three aspects of habitat disturbance by Riplox-treatment
1995/96 and other restoration measures on benthic invertebrates acted concurrently
and cannot be detangled by analysing snap shot surveys during 7 years of observa-
tions in Alte Donau. As submerged macrophyte vegetation was sparse, most inver-
tebrates were found in the sediment in Alte Donau. The majority of these species
observed during the eutrophication and Riplox-treatment were not able to live
attached to submerged macrophytes as an alternative. The bottom sediment was
thus the main habitat for species of the benthic community in Alte Donau before
macrophytes were successfully re-establishment in Alte Donau in year 2003
onwards (Chap. 8).
It is generally agreed that anthropogenically generated fine sediment deposition
by terrestrial degradation, eutrophication or restoration (e.g., Naden et al. 2016) can
rapidly change the sediment’s habitat structure and thus impact the benthic inverte-
brates in manifold ways (Wood and Armitage 1997; Gundacker 2000; Jones et al.
2012; Schröder et al. 2013; Leitner et al. 2015; Murphy et al. 2015; Graf et al. 2016;
Hauer et al. 2018). Such rapid fine sediment settlement alters not only the sediment
architecture due to the small grain size (clogging of sediment and embeddedness of
larger grains) but also the associated sediment properties such as hydrological
exchange at the water-sediment interface, the oxygen content and related redox-
chemical characteristics. In Alte Donau, the quality of fine sediments has been
changed. During the eutrophication period, the fine sediment was built from subse-
quent sequestering of huge biomass of planktonic cyanobacteria, algae and other
organisms and contained chemical precipitates due to the Riplox-treatment (Ripl
1976) in 1995/1996. The total biomass of benthic invertebrates and also the biomass
and abundances of individual groups such as the chironomids and the oligochaets
were particularly low during the chemical treatment with iron chloride, slaked lime
and calcium nitrate in Alte Donau. The rapid and sustained oxidization of the sedi-
ment by nitrate that aimed at phosphate precipitation in the water column and a
suppressed P-release from sediment, seem to enhance microbial activity (e.g.,

Wauer et al. 2005) on the one hand but is also described to be toxic for benthic
invertebrates on the other hand. According to mesocosm experiments by Sueitt et al.
(2015) that were designed for assessing the ecological risk of the calcium nitrate
exposure to benthic invertebrates, the abundance of benthic community mainly
composed by oligochaetes and chironomids was significantly lowered after 25 days
of incubation. During their experiments the abundance of 900 oligochaets was
reduced by 87%, those of about 50 chironomids by 20%. The relative importance of
chironomids over oligochaets at generally low biomass of benthic invertebrates dur-
ing both years of the Riplox-treatment in Alte Donau thus might rather mirror the
losses of oligochaets by nitrate exposure than the re-colonization by ‘new’ chirono-
mid species in the sediment. It is worth mentioning that oligochaetes are known to
be particularly hypoxia resistant (Tubifex species are surviving 10–11 weeks in the
hypoxic sediment under ice in winter and 2 weeks in hypoxic sediments at around
16 °C during stratification in summer, see review by Grieshaber et al. 1993), and
thus seem to be especially vulnerable to superficial over-saturation of oxygen by
access of nitrate in the sediment. Remediation experiments by Janke et al. (2011)
and Sueitt et al. (2015) that were designed to tackle the time schedule of re-coloni-
zation by benthic invertebrates after sediment treatment with nitrate confirm labora-
tory experiments by Ripl (1976) ‘that the restored sediment can support chironomids,
tubificids and other animals’. After the decline of the population densities of benthic
invertebrates due to the disturbance of benthic sediments by chemical or mechanical
treatment (e.g., dredging declines 82% of benthic biomass, Moog et al., 2015), chi-
ronomids and oligochaets are commonly the primary or early secondary colonizers.
The re-settlement of invertebrate animals after restoration is strongly supported by
invertebrates from the surrounding environment (Langford et al. 2009; Sundermann
et al. 2011). According to Graf et al. (2015) the Diptera with 174 taxa and the
oligochaets with 53 taxa are the most heterogeneous groups within the benthic
invertebrate community in the Danube River (see also Moog et al. 1995, 2000 and
Sect. 14.4.1). Although it can be assumed that chironomid species that inhabit
mesotrophic freshwaters in the nearby Danube River floodplain would have a major
contribution to the recolonization, the species composition of chironomids did not
significantly alter from 1995 to 2003 in Alte Donau. A concurrent increase of the
abundance of phytophilic chironomids and the re-settlement of submerged macro-
phytes, however, was observed during this period and supports the importance of
macrophytes as habitat for benthic invertebrates (Cyr and Downing 1988; Cheruvelil
et al. 2002; Kirby and Ringler 2015). The invertebrate survey ended in 2003 before
a more stabilized macrophyte biomass has been developed from 2004 onward in
Alte Donau. While the macrophyte biomass was still relatively low in 2003 (17% of
the biomass in 1987) about 50% macrophyte biomass of the mesotrophic reference
year 1987 has been observed from 2004 onward (Figs. 8.7 and 8.11 in Chap. 8).
The invertebrate activity by feeding and bioturbation mediates detritus process-
ing and subsequently affects the properties of the sediment habitat (McCall and
Fischer 1980; Mermillod-Blondin et al. 2002; Nogaro et al. 2006, 2009, Hunting
et al. 2012). The shift from the predominance of chironomids during both years of
the Riplox-treatment (1995/96) to a predominance of oligochaets (1997–1999 and

2000, 2003) might have altered the sediment structure as the mode of bioturbation
by these two invertebrate groups affects the water-sediment exchange in aquatic
ecosystems differently. While the tubificids are described as upward conveyors
feeding on deeper sediment but ejecting the faecal pellets at the sediment-water
interface, the chironomids are rather known as biodiffusors living in U-shaped tubes
at the near surface layer of the sediment (e.g., McCall and Fischer 1980; Mermillod-
Blondin et al. 2002; Nogaro et al. 2006, 2009). The tubificid worms are thus seen as
the more efficient benthic animals for reducing interstitial clogging when compared
with chironomids.
In Alte Donau, mussels occurred during all three phases from the chemical treat-
ment onward, which might relate to an alternate environmental adjustment of single
species. According to Gundacker (2000), fine sediment habitats are preferred by
Anodonta sp., while Unio pictorum is rather found in patchy areas of low fine sedi-
ment deposition in the Danube River. Both bivalve species were also observed dur-
ing the invertebrate surveys in Alte Donau. These efficient filtrating collectors
seemed to cope well with both fine sediment types, i.e. the particles sequestered
from planktonic biota and those deposited by chemical treatment.
Despite the variation of benthic invertebrates between years of different trophic
state and restoration measure, the relatively high biomass and the large number of
species discussed above, characterise Alte Donau as lowland habitat. The exponen-
tial biomass increase of macrozoobenthos from high to low attitude habitats might
be a function of a number of indirect local effects (e.g., nutrient enrichment in the
lowland ecosystems, lower flow velocity might favour large-bodied benthic animals
and stable potamal habitat conditions and thus might also protect the animals against
washing out during floods) superimposed by global impacts (e.g. decreasing atmo-
spheric pressure and less exposure to UV radiation across high to low attitude habitats).
The different effects modifying macrozoobenthic biomass and community structure
thus need to be disentangled when further assessing Alte Donau compared with
other habitats across altitudes. An increase of both, the invertebrate biomass and the
number of benthic invertebrate orders and families from high to low altitude habi-
tats, however, are commonly observed in stream ecology and are mainly discussed
as direct impact of temperature and associated habitat characteristics (e.g. heavy
metal concentration, ‘stream hydraulics’) (e.g., Clements and Kiffney 1995;
Jacobsen et al. 1997; Jungwirth et al. 2003; Schmutz and Moog 2018).
14.5 Conclusion
The macrozoobenthos biomonitoring surveys covered the year 1987 before eutro-
phication and years of restoration (including the 2 years of chemical treatment of
phosphate precipitation) and an early stage of the period of re-establishing the sub-
merged vegetation in the oxbow lake Alte Donau. Despite the rapid decrease of
phytoplankton biomass associated with an increase of water transparency and a
subsequent growth of macrophytes along restoration measures, the chironomid

assemblage did not clearly show a species shift towards a mesotrophic habitat. A
main reason can be seen in the still relatively low macrophyte biomass during the
last recent invertebrate survey in 2004, when only about 50% of the macrophyte
biomass was recorded compared to the mesotrophic situation in 1987. Thus in 2004
many invertebrates living in the sediment have been found while phytophilic spe-
cies, which typically live attached to the submerged vegetation, were still less com-
mon. It can be expected, however, that with an increase of submerged water plants
in Alte Donau also more clean-water colonizers as found in 1987 will further reset-
tle in Alte Donau. Despite the wax and wane of benthic invertebrates described
during the 8 years of observation, the relatively high long-term average of inverte-
brate biomass characterises Alte Donau well as lowland environment across low- to
high altitude habitats.
Acknowledgments During the 8 years of research many colleagues have contributed to the work.
We want to thank Hasko Nesemann (Turbellaria, Hirudinea, Mollusca and Crustacea), Dr. Michal
Horšak (some Mollusca), Dr. Elisabeth Haberlehner (Mollusca: 1987), Dr. Ferdinand Sporka
(Oligochaeta), Dr. Christa Thurnherr (Hydracarina: 1987) and Dr. Wolfram Graf (Trichoptera) for
their contribution to determining the species. The following colleagues who have not only sup-
ported in taxonomy but also contributed evaluating the results of the survey 1987 we also want to
thank: Dr. Elisabeth Bauer (benthal fauna generally), Dr. B. Grasl (benthal), Dr. Alexander
Schneider (phytal faunistics), Univ.-Prof. Dr. Johann Waringer (benthal- and phytal faunistics) and
Dr. Harald Wintersberger (phytal faunistics). We further thank for field work Dipl. Ing. Reinhard
Wimmer and Mag. Gabriele Weinzettl. Many thanks also to the colleagues contributing in the lab
work: Dipl.-Ing. Dr. Gabriele Bannhofer, Dipl.-Ing. Karin Egger, Mag. Peter Hecher, cand. rer. nat.
Verena Hirzinger, Dipl.-Ing. Mik Hochfellner, Dipl.-Ing. Catherine Hörl, Isabella Leeb, Univ.-
Doz. Dr. Seval Muminović, Dipl.-Ing. Andreas Römer, cand. phil. Martin Seebacher and Dipl.-Ing.
Susanne Steiner. Finally, we express our thanks for collecting pupal exuviae and identifying
Ephemeroptera: Agnes und Lisa Beier, Dipl.-Ing. Rosemarie Forster, Mag. Thomas Geretschläger,
Richard Petrik and Mirjam Suchomel; Dipl.-Ing. Dr. Astrid Schmidt-Kloiber.
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48(10):1859–1872
Chapter 15
Fish Assemblages of the ‘Alte Donau’
System: Communities Under Various
Pressures
Herwig Waidbacher and Silke-Silvia Drexler
Abstract ‘Alte Donau’ was formerly a major arm of the braided Danube in Vienna
and served as habitat for a cold-water river fish assemblage. After river regulation,
the habitat types for fish changed towards a warm-water fish environment with high
temperature, high nutrient content and reasonable macrophyte stands. Accordingly,
the fish species and biomasses changed in the direction of backwater fish composi-
tions. Currently, 22 fish species are recorded. In the early 1990ies, the macrophyte
stands collapsed and heavy blooms of algae and cyanobacteria produced very insta-
ble habitat conditions, particularly in summer, close to oxygen depletion, and fish
kills. A main nutrient source for algal growth, phosphorus, was removed from the
water column by an Iron-III-Chloride treatment, and the fish community responded
by consuming food particles from benthic and surface areas instead of the missing
plankton. Despite these ecosystem changes, no increased mortality in the fish com-
munity was observed during this period. With the intention of keeping high trans-
parency in the water column, a bio-manipulation experiment was introduced by
increasing predators via stocking to reduce plankton feeding Bleak. The species
Asp was the most effective predator (consuming up to 5.8 individuals of bleak/day),
followed by Pike and Pike-Perch. The water transparency – expressed as Secchi
depth – increased by up to 15 centimeters in the experimental area ‘Kaiserwasser’,
compared to the control sections.
Most of the fish species occurring in the Alte Donau are connected to macro-
phyte stands, at least in some stages of their life span. Denser water plant communi-
ties (expressed in CKI – Cumulative Kohler Index) are not only preferred as habitats
by young of the year-class fishes, fish larvae and foragers, but are also used as
spawning grounds. There is a clear correlation between the density of plants and fish
H. Waidbacher (*) S.-S. Drexler

Department of Water, Atmosphere and Environment, Institute of Hydrobiology
and Aquatic Ecosystem Management, University of Natural Resources
and Life Sciences, Vienna, Vienna, Austria
e-mail: herwig.waidbacher@boku.ac.at; silke.drexler@boku.ac.at

https://doi.org/10.1007/978-3-319-93270-5_15
276 H. Waidbacher and S.-S. Drexler
in the horizontal distribution across the water body of the Alte Donau. Furthermore,
the vertical distribution of fish was also associated with plant stock density.
Management of macrophytes is done regularly through mowing activities and plant
harvesting. It is important to note that young fish should have the opportunity to
escape from a dense macrophyte canopy before the plant biomass is removed from
the water column. Small boats with mowing bars and separated macrophyte har-
vesting devices exert clearly less pressure on accidentally removed individuals out
of the living habitat (31.3 kg of harvested macrophytes with 1 removed fish – imme-
diate convey belt harvesting after mowing remove 1 fish every 2.1 kg).
Keywords Biomanipulation · Fish survey · Fish stocking · Cyprinids · Enclosure-

experiment · Stomach and intestinal content · Feed analyses · Food chain ·
Iron-III-Chloride treatment · Ecological measures
15.1 Introduction
The fish assemblages of the Alte Donau system have changed dramatically within
the last 150 years. In addition to human activities such as game fishing, (over)
exploitation of stocks, species supplying plus a wide variety of recreational activi-
ties in general, the overall characteristics of the approx. 150 ha habitat have changed
completely since April 1875, when the River Danube was directed into a newly
excavated and straightened river bed. The river regulation (history see Chap. 2)
provoked an irrevocable change in the fish assemblages and abundances in the Alte
Donau environment. Pristine Danube is characterised by running water conditions,
fit for habitation by individuals belonging to the rheophilic fish guild (Schiemer and
Waidbacher 1992). Highly productive backwaters with rare correspondence to flow-
ing river arms have historically been an exceptional case in the Austrian Danube
stretch (Hohensinner et al. 2011). The habitat preference for cool temperate running
waters is well documented for characteristic native Danube species assemblages.
Haidvogl (2010), for example, reports approximately 60 native fish species occur-
ring in the pristine Austrian section, 96% inhabit breakthrough sections as well as
anabranched areas.
The modified situation in the Alte Donau favoured fish species out of the indif-
ferent- and stagnophile guild, such as Roach, Bleak and Rudd. Macrophytes became
a dominant structuring element used by warm water species as spawning- and feed-
ing grounds. Organic based turbidity correlated with nutrient availability (Chap. 6),
while at the same time fish stocking for game fishing became a yearly repeated
event. A connection to the flowing river with corresponding migration activities of
the fish fauna has been blocked since the days of separation even at highest water
discharge (flow rates see Chap. 4). Fish have since penetrated the Alte Donau sys-
tem through human activities such as stocking and releasing or via water fowl where
fish eggs can stick on legs and feathers and spread to water bodies even in remote
distances.
15 Fish Assemblages of the ‘Alte Donau’ System: Communities Under Various Pressures 277
Fig. 15.1 Total fish stocking [kg] versus total fish catch [kg]
Figure 15.1 gives an overview on long-term data of the angling society

(Österreichische Fischereigesellschaft – ÖFG 1880) for fish stocking and catching
amounts at the Alte Donau from 1984 to 2012. Stocking has always been dominated
by one species: the common carp. Approximately 4–6 tons of catchable fish have
been stocked every year. The catch was 250% higher than stocking in the late
1980ies and decreased to approximately the same amount of stocked fish in recent
times. This trend could be explained by a parallel decrease of the trophic level of the
water body (compare Sect. 11.4.1.4 in Chap. 11, Fig. 20.7 in Chap. 20). Latest
developments in stocking activities (from 2013 to today – not included in the above
shown data) implement a more ecological based fisheries management by forcing
an increased stocking of predatory fish like Asp, Pike and Pike-Perch following
scientifically based management measures (see also Chap. 15.3) to foster today’s
high water quality of the Alte Donau.
A major danger for the already far from pristine developed fish community
occurred in the early nineties of the last century. Macrophyte stocks collapsed;
eutrophication ensued, and serious oxygen problems in the water body threatened
fish populations dramatically.
As a swift and lasting remedy, of an Iron-III-Chloride treatment to remove phos-
phor out of the water column was proposed and discussed very intensely. The reac-
tion of the fish community to such a treatment had not been confirmed on a big data
set in those days.
15.2 T
he Impact of the Iron Chloride Treatment on Fish
Abundances
During restoration by applying the Riplox-treatment (Chap. 5), Iron-III-Chloride

was added to the Alte Donau in 1995 and 1996 to counteract the hypertrophic level
of the aquatic system. The treatment implements a chemical reaction on phosphor
as well as a mechanical response. The insertion of iron particles effects a mechani-
cal precipitation in the way that phosphor is linked to the particles and sinks to the
ground of the water column. The water column clears up; phytoplankton growth is
limited by phosphorus and thus unable to develop large biomass (Chaps. 9 and 10),
and zooplankton is limited by a reduced food supply (Chap. 11).
An earlier investigation (Löffler 1988) classified the Alte Donau as a Cyprinid-
dominated water body with high numbers of planktivorous fish and low quantities
of carnivorous fish – a classic predator-prey-community. In estimating the effects of
the Iron–III-Chloride treatment, two main questions were analysed from the fish
ecological point of view:
1. Is there a change in the species assemblage due to the Iron-III-Chloride

treatment?
2. Is there a shifting of the food particle spectrum for the three dominant fish spe-
cies, i.e. Bleak (Alburnus alburnus L., Fig. 15.2), Roach (Rutilus rutilus L.) and
White Bream (Blicca bjoerkna L.)?
15.2.1 Methods
Between mid of March and mid of September 1995 five main fish surveys were
conducted at three different places in the Alte Donau. Gill nets with mesh sizes of
12, 15, 28 and 38 mm were used. Those nets with a mesh size of 12 and 15 mm were
used for capturing Bleaks and juvenile fish of any other species. Nets with meshes
Fig. 15.2 Bleak (Alburnus

alburnus L.) is a common
fish species found during
investigation in 1987 and
1995–2007 in Alte Donau
and which is majorly
feeding on plankton
of 28 and 38 mm size were exposed for capture of adult fish. Nineteen stretches
(each 30 m of length) were seined with a trawl net of 2 mm mesh size. The investi-
gation areas were sampled semi-quantitatively to 1 m depth with a seining width of
about 3 m. This method was used especially for proof of juvenile fish stocks
recruited out of successful reproduction of single species. In addition to the main
fishing events, gill nets with a mesh size of 80 mm were exposed at the deepest
places in the Alte Donau for capture of any adult individuals.
All captured individuals were determined to species level, and length was mea-
sured. Additionally, weights of 1300 fish were recorded. For most fish, sexual matu-
rity could be identified.
Intestinal and stomach content analysis was undertaken for 578 individuals dur-
ing the investigation period in 1995. Stomach content was only analysed of preda-
tory fish like Pike-Perch (Sander lucioperca L.) and Pike (Esox lucius L.). Food
particle analysis was performed on deep-frozen fish, respectively, in the case of
smaller individuals, on material fixed with 4% formol. The ingestion rate was deter-
mined via the filling degree of the open intestinal tract using the following catego-
ries (Table 15.1).
Detailed analyses were performed – depending on the filling degree and the het-
erogeneity of the food particles (classified in 10–100% of the intestinal content) –
by using a 50-fold magnification of a binocular. The samples were numerically
investigated. A valuation of the surface ratio in comparison to the size of the sam-
pling dish was estimated for the following food components:
• Zooplankton (cladocera, copepoda, rotifera)
• Plant material (macrophytes, crop and maize)
• Filamentous algae
• Animal traces (terrestrial insects, etc.)
• Herbal traces (willow seeds, leaves of the riparian vegetation, etc.)
• Benthos (diverse aquatic insect larvae and pupae, especially chironomidae,
hydracarinae, ostracoda, etc.)
• Mollusca (Dreissena polymorpha, aquatic gastropoda)
• Sediment (mainly small stones)
• Detritus (dead organic material)
• Pisces (fish)
• Decapoda (crustaceans)
Table 15.1 Categories for Category Filling degree

estimation of intestine filling
0 No content
degree
1 0–25%
2 26–50%
3 51–75%
4 76–100%
15.2.2 Results and Discussion
A total number of 17 species could be detected during the investigation period in

1995 (the first year of the Iron-III-Chloride treatment) with the above mentioned
methods. Two species (Grass Carp and European Eel) have been added from angling
reports. Therefore, 19 species have been listed (Table 15.2). With this result no
Table 15.2 Species assemblage of the Alte Donau

Investigation
Species period
1987 1995 1996–2013
ACIPENSERIDAE
Hybrid acipenser Acipenser sp. x sp. *
ESOCIDAE
Northern pike Esox lucius (L.) * * *
CYPRINIDAE
Freshwater bream Abramis brama (L.) * * *
Bleak Alburnus alburnus (L.) * * *
Asp Leuciscus aspius (L.) * *
White bream Blicca bjoerkna (L.) * * *
Crucian carp Carassius carassius (L.) * *
Prussian carp Carassius gibelio (B.) *
Grass carp Ctenopharyngodon idella (Val.) A A A
Common carp Cyprinus carpio (L.) * * *
Chub Squalius cephalus (L.) A * *
Roach Rutilus rutilus (L.) * * *
Rudd Scardinius erythrophthalmus (L.) * * *
Bitterling Rhodeus amarus (B.) *
Tench Tinca tinca (L.) A * *
SILURIDAE
Wels catfish Silurus glanis (L.) * * *
ANGUILLIDAE
European eel Anguilla anguilla (L.) * A *
CENTRARCHIDAE
Pumpkinseed Lepomis gibbosus (L.) * * *
PERCIDAE
Ruffe Gymnocephalus cernua (L.) E E
European perch Perca fluviatilis (L.) * * *
Pike-perch Sander lucioperca (L.) * * *
GOBIIDAE
Western tubenose goby Proterorhinus semilunaris (H.) * * *
Abundance 16 19 22
(A) proofed by sport fisheries
(E) single specimen
(*) proofed by investigation
outstanding changes in comparison to the examination in 1987 are detectable. Three

more species (Crucian Carp, Ruffe and Asp) were identified in 1995. This can be
explained by the more intensive study design as well as stocking activities of the
Asp. European Eel and Grass Carp could not be proofed again, because of the cho-
sen sampling methods, which are not appropriate for capture of these species.
In the time range between 1995 and the yearly investigations up to 2013, an
overall species list of 22 fish species can be presented.
Cyprinid species like Bleak, White Bream, Roach and Carp, are known as poly-
phagous generalists, who ingest a wide spectrum of food particles (Table 15.3). The
Bleak takes up mainly zooplankton, like Copepoda and Cladocera and sporadically
also rotifers (Keratella sp. and Kellicottia sp.). White Bream and Roach consume
from the plankton solely crustaceans. For example, Leptodora kindtii could only be
detected from White Bream stomachs. Carp pick from zooplankton beside single
copepoda basically one species of the cladocera – namely Eubosmina coregoni (for
abundance of zooplankton species see Figs. 11.1 and 11.2 in Chap. 11). In a few
Carps, Chaoborus sp. from the order ‘Diptera’ could be detected.
Table 15.3 Proofed prey clusters and species, investigation in 1995

Alburnus Blicca Rutilus Cyprinus Sander Esox
Food components alburnus bjoerkna rutilus carpio lucioperca lucius
PLANKTON x x x x
Cladocera x x x x
Eubosmina coregoni x x x x
Daphnia cucullata x x x
Leptodora kindtii x x
Alona sp. x x x
Chydorus sphaericus x x x
Copepoda x x x x
Keratella sp. x
Kellicottia sp. x
Chaoborus sp. x
BENTHOS x x x x
Nematoda x
Oligochaeta x x x x
Chironomidae L x x x x
Ephemeroptera L x
Trichoptera L x
Ceratopogonidae L x
Diptera pupae x x x x
Coleoptera adult x
Hydracarina x x x
Aquatic Heteroptera x
(continued)
Alburnus Blicca Rutilus Cyprinus Sander Esox
Food components alburnus bjoerkna rutilus carpio lucioperca lucius
Amphipoda x
Asellus aquaticus x x
Harpacticoida x x
Ostracoda x x x
MOLLUSCA x x x
Dreissena sp. x x x
Glochidia x x
Aquatic Gastropoda x x x
Ancylus fluviatilis x
ANIMAL REMAIN. x x
Insecta indet. x x
Trichoptera adult x
Diptera adult x x
Formicidae x
Terrestrian Araneida x
HERBAL REMAININGS x x
ARTIFICIAL FEEDS x x x x
Maize/crop x x x
FILAMENT. ALGAE x
SEDIMENT x x x
DETRITUS x x x x
+ sediment x x x
+ herbal material x x x
Stato-/Flotoblast x x x
Cocoon x x
PISCES x x x x
Indet. x x
Alburnus alburnus x x x x
Rutlius rutilus x x x
Perca fluviatilis x x
Sander lucioperca x
Lepomis gibbosus x
DECAPODA x
The food component benthos was proofed in all four analysed fish species,
whereas the White Bream ingests the largest spectrum. Highest amounts of
Chironomidae, Diptera Pupae and Oligochaeta were detected in stomachs of Bleak,
White Bream, Roach and Carp. Carps sporadically feed additionally on Asellus
aquaticus. Hydracarina and Ostracoda are eaten by Bleak, White Bream and
Roach. Representatives of Harpacticoida were proofed rarely in the intestine of
Bleak and White Bream. Water beetles were only detected in Bleak’s stomachs,
while larvae of Ceratopogonidae were solely ingested by Roach. All other represen-
tatives of this food group could only be found in the intestines of the White Bream.
Individuals of the food cluster mollusca are especially present in intestines of
White Bream, Roach and Carp, while the Bleak does not feed on them. Larvae of
mussels (Glochidiae) are only ingested by White Bream and Roach.
Animal remaining is mainly food bases for Bleak, except adult Diptera and all
other not specified species can be detected in Roach exclusively. The food group of
herbal remainings (like seeds of the willow) is only found in Bleak and White
Bream. Plant material from submerged macrophytes as well as pieces of maize
and crop are food for all four groups of investigated cyprinids. Filamentous algae
(mainly green benthic algae; the biomass of planktonic filamentous cyanobacteria
was already low in 1995; further details about phytoplankton composition see
Chap. 9) are proofed in Roach; sediment (stones and fine sandy material) can be
detected in the intestine of White Bream, Roach and Carp. The group of detritus,
which includes dead organic material, with fine sediment and herbal material, is
proofed in the intestines of all four cyprinids.
A number of piscivorous species like Pike-Perch and Pike, which ingest mainly
pisces, occur in the research area. In detail, the prey fish are Bleak (Fig. 15.1),
Roach, Perch, Pike-Perch and Pumpkinseed. Amazingly, even White Bream and
Roach were found to have some fish prey in their intestines. Pieces of Decapoda
(thoracal bones) were only proofed once in the stomach of a Northern Pike.
Analysis and comparison of all net samplings did not show any lasting negative
effects on fish population due to the Iron-III-Chloride treatment. Small captures
during the first fishing period as shown in Fig. 15.3 result from the low water tem-
Fig. 15.3 Sampling and treatment design with captured individuals

peratures at this time of the year. In all fishing events, the Bleak is the species with
the highest abundance in reference to the other fish species, with an outstandingly
high occurrence in the catches after the end of the main Iron-III-Chloride treatment.
This might be due to a higher migration and foraging activity caused by changes in
their natural habitats. A short-term effect is visible for adult Bleaks, but increased
mortalities could not be detected during the investigation period.
The ingestion rate of fish normally increases with increasing water temperatures
due to their higher physiological and locomotional activity. Therefore, an increase
of the intestine filling degree can be expected. This particular aspect is only visible
in the results for White Bream within the first and the third fishing event. Bleak and
Roach show a clear decrease from the first to the second respectively to the third
fishing event (Figs. 15.4, 15.5 and 15.6). Such reduced food intake values appear
under normal conditions during spawning time. In the present investigation, the
lower ingestion rates can also be explained by the Iron-III-Chloride treatment,
which had already started. A separation of these effects is not possible from the
available data.
Feed analyses of predatory fish like Pike-Perch and Northern Pike show similar
results. While the filling degrees of the Pike-Perch increased again, those of Pike
could not be analysed due to a reduced sample size.
0 1 2 3 4
5. fish survey
4. fish survey
3. fish survey
2. fish survey
1. fish survey
0% 20% 40% 60% 80% 100%
Fig. 15.4 Intestine filling degree (according to Table 15.1) for Bleak (Alburnus alburnus)
0 1 2 3 4
5. fish survey
4. fish survey
3. fish survey
2. fish survey
1. fish survey
0% 20% 40% 60% 80% 100%
Fig. 15.5 Intestine filling degree (according to Table 15.1) for White Bream (Blicca bjoerkna)
0 1 2 3 4
5. fish survey
4. fish survey
3. fish survey
2. fish survey
1. fish survey
0% 20% 40% 60% 80% 100%
Fig. 15.6 Intestine filling degree (according to Table 15.1) for Roach (Rutilus rutilus)
Concerning the dietary composition, cyprinid species ingest a wide range of prey
and are therefore ranked among the polyphagous generalists. The investigated spe-
cies are highly flexible in food intake. During the first fishing period the Bleak feeds
mainly on plankton, although this changes with end of April where plant material
and animal- plus herbal trace become more important, as well as benthic inverte-
brates (at least for a short time span). End of May, the Bleak resumes grazing on
zooplankton species with a peak at the end of September (Figs. 15.7, 15.8 and 15.9).
Reduced plankton intake from Bleak in the early summer might be interpreted to be
a reaction to the Iron–III-Chloride treatment. However, in an earlier investigation at
the Alte Donau (Löffler 1988) the same pattern became visible. Samples for the
study by Löffler were taken in the middle of June in 1987 and the food particles
consumed by Bleak were found to consist of 43% animal- and 50% herbal remain-
ings; plankton was mainly absent. Changes in food intake of the Bleak are therefore
not due to the Iron–III-Chloride treatment but are better explained by increasing
energy demand caused by increasing water temperature and by the relatively low
plankton rates at this time of the year. In general, results from Löffler (1988) con-
firm the wide spread nutrition spectrum (plankton, detritus, benthos and herbal
material) of the Bleak as well as the main food components for the Roach (32%
molluscs, 20% herbal material, 26% detritus and 11% filamentous algae).
100%
80%
60%
Volume
40%
20%
0%
1. survey 2. survey 3. survey 4. survey 5. survey
Zooplankton Benthos
Plant material Mollusca
Filamentous algae Sediment
Animal remainings Detritus
Herbal remainings Pisces
Fig. 15.7 Food spectrum of Alburnus alburnus

100%
80%
60%
Volume
40%
20%
0%
Zooplankton Benthos
Fig. 15.8 Food spectrum of Blicca bjoerkna
100%
80%
60%
Volume
40%
20%
0%
Zooplankton Benthos
Fig. 15.9 Food spectrum of Rutilus rutilus

The main nutrition for Carp belongs to the food clusters benthos, detritus and
herbal material, where 100% of the latter is represented by maize and artificial fish
feeds. Maize and crop are typical baits for Carp sport-fishing. Their high detection
rate in the intestines proves the well-accepted feeding places as nutrition source,
especially for Carps. The investigated predatory fish (Pike-Perch and Pike) feed on
fish exclusively. Mainly Bleak and Roach serve as food for predatory fish.
15.3 Bio-manipulation to Improve the Water Quality
Due to the situation in 1995/1996 and the on-going measures in the following years,
the Alte Donau was classified as a eutrophic water body with a high abundance of
cyprinid fish species and low rates of predators. For a more balanced system with
corresponding lower algae biomass, bio-manipulation can be a successful input.
There are two manipulative types of intervention described for lentic water bodies
like the Alte Donau: Regulation can be effected via the basis of the food chain
(‘bottom-up’) or via the effort of the predators (‘top-down’). The ‘bottom-up-
hypothesis’ indicates that biomasses from all trophic levels are positively correlated
and depend on limiting factors of the habitats (Lampert and Sommer 1993). This
means that an increasing nutrient content would raise the algae biomass, which
would then lead to higher abundances of zooplankton. A higher number of plank-
tivorous fish can feed on these resources and, in turn, provide the food base for a
higher number of predatory fish. The ‘top-down-hypothesis’ has also been referred
to as the ‘cascade effect’ (Schwoerbel 1993). The ‘top consumer’ (predatory fish)
decreases pressure on zooplankton by reducing planktivorous fish and, indirectly,
favours the development of herbivore zooplankton. In terms of a ‘top-down con-
trolled’ water body the result would be a decreasing density of phytoplankton,
which effects a higher Secchi depth in the water column. The clearing up of the
water column becomes visible.
In the Alte Donau, the nutrient level was reduced by the Iron-III-Chloride treat-
ment and could not be influenced further in the sense of starting a ‘bottom-up’ strat-
egy to increase water transparency. Dokulil et al. (1995) reported that size and
composition of the zooplankton populations show high predation pressure by fish,
indicating that fish management in accordance with the ‘top-down-hypothesis’
could improve the transparency of the water column of the Alte Donau. The 19-year
development of zooplankton (1994–2012, Chap. 11) shows that a temporary increase
in the zooplankton size was associated with the initial increase of macrophyte bio-
mass (Chap. 8), which underpins the importance of spatial refuge against visual
predators for zooplankton. Larger-sized herbivorous zooplankton, which forms the
most effective filtering component of phytoplankton, is majorly reduced by plank-
tivorous fish, especially by the Bleak (Alburnus alburnus L.). According to Lampert
and Sommer (1993) and Dawidowicz (1990), for example, the smaller zooplankton
is known to be not able to reduce efficiently the phytoplankton by grazing. Decreased
biomass of filtering zooplankton, dominated by small-bodied individuals, benefits
the reproduction of algae and, therefore, causes turbid water. This phenomenon
results in worse light conditions for the growth of macrophytes. By increasing the
abundance of predatory fish, planktivorous fish, such as Bleak, which mainly ingest
large cladocera, are reduced and the filtering activity of the zooplankton increases
(Begon et al. 1991; Benndorf et al. 1994 in Bohle 1995). Predatory fish are, there-
fore, also called ‘keystone predators’, as they influence more trophic levels, which
leads to long-term effects in the whole water body (Lampert and Sommer 1993).
The ‘top-down-hypothesis’ was tested in the Alte Donau in 1998 to prove the
assumption that the value of the Secchi-depth would increase. The effect of bio-
manipulation could not be examined in the complete Alte Donau system, since
150 ha are simply too voluminous for massive stocking with predators. Therefore,
the experiment was conducted in the better to handle ‘Kaiserwasser’ with an area of
approx. 6 ha (a nearly separated part of the ‘Alte Donau’), where mainly Pike-Perch
(Sander lucioperca L.) was stocked as a top predatory fish. In those days under
moderate turbidity the Pike-Perch was the predestined predator for the system,
which was characterised by: moderate productivity in the water column, high prey
quantity, high water temperatures, low depth with missing stratification in summers
and with species-preferred turbidity of the water for efficient foraging strategies
(Spindler 1997; Willemsen 1978; Nagięć 1977; Hokanson 1977; Popova and Sytina
1977; Hansen 1987).
The Ranking of Top Predator Fish
Following the theory of ‘top-down’ bio-manipulation the question arose which fish
would be the most effective in reducing small planktivorous cyprinids and specifi-
cally Bleak (Alburnus alburnus L.). Five fish species were discussed for stocking;
three species were tested in an enclosure-experiment. The five ventilated species
are:
Pike-Perch (Sander lucioperca L.)
Pike (Esox lucius L.)
Asp (Leuciscus aspius L.)
Wels – Catfish (Silurus glanis L.)
Perch (Perca fluviatilis L.)
Silurus glanis is a bottom preferring species and not specialised in hunting small
cyprinids close to the water surface in high quantities. Perca fluviatilis preys on
small cyprinids most prominently only in an adult stage and can show a volatile
behaviour of switching to planktivory when high numbers of zooplankton are avail-
able as seen in Lake Constance in the 1970ies (Hartmann 1975). Such behaviour
can be an unpredictable parameter when trying to support the bigger stages of the
filtering plankton in an experiment.
The species Pike-Perch, Pike and Asp all fit in the natural species composition of
the described water body. All three species were tested in enclosures as shown in
Fig. 15.10 and were fed with Bleak ad libitum while consumption behaviour was
controlled.
Fig. 15.10 Cages for the enclosure-experiment
15.3.1 Methods
(1) Enclosure-Experiment
For the experimental station, metal cages with dimensions of 150 cm × 150 cm
base area and a height of 180 cm were constructed. For higher safety of the fish, the
inner part of the cages was lined with textile mesh. The ten cages were arranged
around a float and, for lowering in the water, were handled with an electric winch,
which was fixed on a flexible crane. Each cage was assembled with a flap. Via the
open flap, the fish inside were controlled, and the cages could be checked for dam-
ages. One of the cages was used as a storage reservoir for prey fish. The experiments
took place between 1998 and 2000. Nine Pike-Perch, eight Asps and four Pikes
were tested for a period of at least 1 month. The mean starting weight of Pike was
626.5 g, for Pike-Perch 373 g and for Asp 1134 g. Fish were fed ad libitum with
Bleak – attention was paid that at least two living Bleaks were always swimming in
an enclosure.
Tested Pike-Perch were taken from the Alte Donau ecosystem, while Asps were
caught in a Danube side arm eastward of Vienna and Pikes originated from ponds in
Austrian Waldviertel region. Prey fish were taken from the Kaiserwasser with seine nets.
The length of prey fish was measured; weight was calculated from regressions.
While running the experiments, the following steps had to be performed daily:
• Fishing for Bleaks
• Checking vitality, health and any lesions of the individual predators per
enclosures
• Checking the nets of the cages for damages

• Taking out and measuring the length of dead and not-ingested prey-fish
• Appraisal of number and length of resting Bleaks in each cage
• Determining the length of ingested Bleaks with the protocol
• Stocking each cage with fresh and measured prey fish
• Recording all relevant data
After finishing the experiment period and removing all fish from the enclosures,
the weight of each predator was taken.
(2) Kaiserwasser bio-manipulation
As mentioned above, the Kaiserwasser is a 6 ha nearly separated side water body
of the Alte Donau. It is connected by a small channel with a maximum width of 3 m
and a bridged gap of approximately 5 m. During the experiment, a dense net fence
closed both connections. The channel was closed permanently; the bridged gap was
closed with a special type of lock where rowing boats could trespass.
Gill nets with a mesh size of 12 mm were used in April and September 1998 to
estimate the comparative density of Bleak populations with the measure of catch per
unit effort (one-hour unit). In addition, a seine net of 30 m length and 1.5 m depth
with the mesh size of 3 mm was exposed by boats at a distance of 30 m parallel to
the shoreline. Seining was done by pulling ropes of the net simultaneously in direc-
tion of the shore with at least four people on each side. As a third method, three
people additionally operated a small shore seining net, which seined an area of
100 m2, and compared catches per unit.
The stocking density with Pike-Perch was 375 kg for the whole Kaiserwasser.
That means an introduction of prey fish of 62.5 kg per hectare was the starting con-
centration at the beginning of the experiment. A slight reduction of individuals hap-
pened during the investigation period by poaching. Pike-Perch was the only species,
which was available in required quantities (Fig. 15.11).
Fig. 15.11 The stocked

predator Pike-Perch
(Sander lucioperca L.)
Production of Asp as a stocking fish has to be planned a long time before bio-
manipulation experiments start – this was not possible in the described case study.
15.3.2 Results
15.3.2.1 Enclosure Experiments
The box-plot of Fig. 15.12 shows the daily intake of Bleak by the three tested pisciv-
orous fish species in the Alte Donau, Pike, Pike-Perch and Asp. The comparison
between these three predators is displayed by the percentage of weight increase due
to the intake of Bleak and was about 3–4% (median) for all the three species during
the enclosure experiment. The largest variation was found for Asp where the intake
can reach double values compared with the other two fish species.
Analysing the medium weight of the ingested Bleak (Fig. 15.13), it becomes
significantly visible that Pike-Perch prefers the smallest sizes over the whole period.
The results are highly significant between Pike and Pike-Perch as well as between
Pike-Perch and Asp. Asp and Pike prefer bigger prey individuals; the highest
weights are consumed by Asp.
According to the results that Asps fed the highest weight-quantities of Bleaks per
day, Fig. 15.14 shows numbers for the individual intakes of Bleaks in 24 h. Larger
Asps fed more individuals and higher quantities. It was shown that they fed up to
5.82 individuals when the predator weighs 2247 g. Large fish are very effective in
reducing prey as shown by the presence of Bleak in the enclosures.
Ranking the three species, Asp (Leuciscus aspius) was shown to be most effec-
tive, followed by Pike and Pike-Perch with similarly low values.
10
8
% of starting weight
0
N= 4 9 8
Pike Pike perch Asp
Fig. 15.12 Daily intake of Bleak as the percentage of weight increase for Pike, Pike-Perch and
Asp
16
14
Medium weight of bleak [g]
12
10
5
8
6
N= 4 9 8
Pike Pike-perch Asp
Fig. 15.13 Medium weight [g] of Bleak intake for Pike, Pike-Perch and Asp
100
g/24h
Ind/24h
80
60
Gram
40
20
0
g
1g
6g
0g
g
98
90
39
47
30
37
74
85
17
22
22
Fig. 15.14 Intake of Bleak (as mean values in gram) per 24 h and of individuals of Bleak per 24 h
15.3.2.2 Bio-manipulation Stocking in ‘Kaiserwasser’
The results from seining the Kaiserwasser are shown in Fig. 15.15. The mean value
in spring was 97 individuals per catch, and this stock was reduced to 58 individuals
per catch in autumn. The effect of the bio-manipulation through stocking of Pike-
Perch is clearly visible. Bleak was reduced by approximately 30% of the starting
density. An even more dramatic reduction is shown by the comparison of the 100 m2
shore line seining. There, a reduction from 11.5 individuals per 100 m2 in spring to
4.1 individuals per 100 m2 in autumn becomes visible.
400
Number of caught individuals/effort

22
300
200
100
N= 16 15
Spring Autumn
Fig. 15.15 Number of Bleaks expressed in catch per unit effort from seine net catches in the
experimental area Kaiserwasser
350
Reference
300 Kaiserwasser
250
200
-1
µg L
150
100
50
0
98
98
98
98
98
98
98
98
9
9
.1
.1
.1
.1
.1
.1
.1
.1
5
2
.0
.0
.0
.0
.0
.1
.1
.1
01
01
01
01
01
01
01
01
Fig. 15.16 Total biomass [μg L−1] of Eudiaptomus gracilis and Daphnia cucullata in the experi-
mental area Kaiserwasser compared with the reference water body
In controlling the principle of the ‘top-down-biomanipulation cascade’ Fig. 15.16

shows the total biomass of larger filtering plankton particles (∑ Eudiaptomus graci-
lis and Daphnia cucullata) over the time period. Densities in Kaiserwasser com-
pared with densities in the major Alte Donau water column show a slightly higher
biomass during July, August and September in Kaiserwasser. The reduced Bleak
stock forces the development of larger plankton particles, which are able to filter
more effectively. Fig. 15.17 points out quantities of chlorophyll-a content in the
water column during the term of the experiment.
20
18
16
14
-1
chl-a µg L
12
10
8 Reference
Kaiserwasser
6
98
98
98
98
98
98
98
19
19
19
19
19
19
19
5.
6.
7.
8.
9.
0.
1.
.0
.0
.0
.0
.0
.1
.1
01
01
01
01
01
01
01
Fig. 15.17 Same as Fig. 15.16 but for chlorophyll-a content [μg L−1]
0,0
Reference
Kaiserwasser
-0,5
Secchi depth [m]
-1,0
-1,5
-2,0
-2,5
98
98
98
98
99
99
99
19
19
19
19
.1
.1
.1
5.
6.
7.
1.
8
0
.0
.0
.0
.0
.0
.1
.1
01
01
01
01
01
01
01
Fig. 15.18 Same as Fig. 15.16 but for Secci depth [m]
A third indicator for the efficiency of bio-manipulation in Kaiserwasser is given

through the values of measured Secchi depth (Fig. 15.18) in the compared water
columns. The difference shown in summer 1998 is not drastic, but a higher Secchi
depth of 10–15 cm can be detected in Kaiserwasser due to reduced phytoplankton
biomass, which is based on more effective filtering zooplankton.
15.3.3 Discussion
In 1998 – 2 years after Iron-III-Chloride was added to the Alte Donau to counteract
the hypertrophic level – specific bio-manipulation in accordance with a ‘top-down-
hypothesis’ was tested. Effects of focused fish stocking, which introduced and
intensified trophic interactions within the food chain, were investigated. The
intended practical application of the experiments was to achieve a significant clear-
ing-up of the water column for a certain time period at least.
(1) Enclosure experiment in artificial cage habitats
Nine Pike-Perch (Sander lucioperca), eight Asps (Leuciscus aspius), and four
Pikes (Esox lucius) were kept for at least 1 month in a cage system and fed ad libi-
tum. Regular food intake of protocolled Bleak (Alburnus alburnus) was recorded by
a daily inspection. Abiotic parameters were equal to the surrounding Alte Donau
environment. Appropriate cage-keeping in 4 m3 enclosures proved a suitable meth-
odology for studying the pressure of predatory fish species on Bleak populations.
The condition factors of Pike-Perch – taken at the beginning and at the end of the
experiment – confirmed normal feeding. There was no significant difference of the
nutritional status of the fish at the beginning and at the end of the experiments. Also,
a comparison of condition factors of fish kept in cages with those living in the wild
did not show any significant discrepancy.
In the enclosure experiment, Asps generally had the highest predatory perfor-
mance, pinpointing this species as a favourite for stocking. Results show that an Asp
with 528 g starting-weight ingested the highest amount of fish (650 g) in compari-
son to the heaviest Pike-Perch with a similar starting-weight of 614 g, which took
only 567 g of Bleak within a comparable period. The Asp with 528 g starting weight
also had a significantly higher gain through the daily consumption of 2.93 weight
percent of its bodyweight than a Pike-Perch of approximately the same weight.
Smaller individuals of Pike-Perch ingested lower amounts of prey fish, but – in
comparison to their body weight – significantly more than the largest fish. Stocking
younger individuals, generally more readily available for purchase, was thus con-
firmed as a promising management action. Sander lucioperca with starting weights
of 203 g and 258 g respectively had daily intake rates of 3.63–4.06 weight percent
of their body weight. Heavier specimens, with starting weights of 547–614 g, had
daily intake-rates of only 1.87–2.21% body weight.
Statistical analysis displays significant differences in weight percent of the daily
ingested amount of Bleaks for different starting weight classes. Additionally, a
strong negative correlation between weight of Pike-Perch and weight of daily
ingested Bleak, expressed in percent of the starting weight of the Pike-Perch, was
observed.
Literature confirms that two of the experimental fish species (Sander lucioperca
and Leuciscus aspius) have highest growth rates during the first 4 years of their life
(Lehtonen et al. 1996; Salonen et al. 1996; Kangur and Kangur 1996; Martyniak
and Heese 1994; Gaygalas 1977; Fickling 1981). This backs the chosen strategy of
stocking younger individuals and highlights the fact that offering constructed
spawning grounds and supporting self-sustaining growth of predator populations is
preferable to a simple stocking scheme of adding large fish.
(2) Biomanipulation in a natural habitat – the experimental area Kaiserwasser
The study focused mainly on the effects of predating fish on Bleak stocks
(Alburnus alburnus) to strengthen the populations of bigger-sized planktonic filter
feeders.
Seine-fishing data show influences of bio-manipulation measures with stocked
Pike-Perch. In spring, before stocking predators in high quantities (62.5 kg ha−1),
more individuals of Bleak were recorded in the Kaiserwasser. Depending on the
fishing method used, the quantified influence on the targeted Bleak population was
visible in a clear decreasing trend. Significant results were achieved with the method
of shore seine netting (100 m2). In autumn significantly smaller amounts of Bleak
(65%) were recorded, which was confirmed via statistical tests. A lower reduction
is shown by numbers generated by sampling with the ‘large trawl net’. The statisti-
cally insignificant result was caused due to a relatively low number of repeats.
Nevertheless, a trend is clearly visible in the comparison of the data of the fishing
efforts with the trawl net. The mean value of the estimated Bleaks was 97 per fishing
event before stocking and decreased to 58 per run in autumn. Beside the investi-
gated Bleak, additional fish species like Roach, Rudd, Ruffe and European Perch
were sources of food for predators. Especially the massive appearance of small
European Perch (Perca fluviatilis) as prey item was remarkable but fits perfectly in
the predator’s search-pattern.
Pike-Perch additionally chase from the bottom of the water body and therefore
Ruffe (Gymnocephalus cernua) and European Perch, which graze for benthic inver-
tebrates in shallower areas, are often consumed. Predatory pressure of Pike-Perch is
therefore observed for several prey species. Nevertheless, the impact of bio-
manipulation remains unaffected by this bottom feeding pattern, as most of the
small fish in the Alte Donau feed on zooplankton in the pelagic zone. Results of the
plankton investigations showed a distinctly positive effect on zooplankton abun-
dancy from decreasing zoo-planktivorous fish populations.
A suitably modified food chain in the ecosystem of the Alte Donau requires a
self-sustaining, high level of predatory fish. Therefore, the aim must be to achieve
reproduction of predators under natural conditions. Following Lampert and Sommer
(1993), the system should stabilize through habitat-driven improvements in popula-
tion structures, thus avoiding external inputs such as stocking.
In this study, short-term effects by bio-manipulation were generated through
massive stocking with predators. However, the level of specific predators needs to
be supported by long-term management strategies to achieve an effective and sus-
tainable restoration of the water body.
15.4 Macrophytes as Habitats for Fish
Macrophytes are noted to be very important habitats for fish (Savino and Stein
1982). The species spectrum and the biomass of submerged water plants in Alte
Donau are described in Chap. 8. Many fish need submerged structures for spawning.
Most of the cyprinids fix their sticky eggs on macrophytes and even on terrestrial
plants in case of flooding periods. Some fish build their nests between water plants
for spawning. Drastic drawdowns of the water level can bring serious damages for
eggs fixed on plants directly below the water surface, as well as the removing of
submerged vegetation during spawning time (Petr 2000).
For juvenile fish, macrophytes are also intensively populated habitats. Mixed
submerged plant assemblies are very complex in structure and therefore very impor-
tant as growing areas for juveniles. Dense stocks support the fish society and maxi-
mize amount, biomass and diversity (Conrow et al. 1990).
Presence of predatory fish evokes special habits in food intake. Roach normally
prefer prey of the free water column, for example zooplankton. But the occurrence
of predators leads to changes in food preference. The Roach, which uses the macro-
phytes as hiding place, starts feeding on less nutrient-rich food, like detritus and
algae (Persson 1993).
Defense strategies of Roach and Perch are also influenced by the presence of
predatory fish. Roach flee to dense stocks of submerged plants and Perch, which are
an easy prey in open water, hide in vegetated structures and even use gaps in the
ground. Those plant structures reduce the chasing speed for predators due to their
size and function as obstacles, but small prey can easily flee and hide in the plants,
and the visual contact between predator and prey is decreased too (Christensen and
Persson 1993; Berg et al. 1997).
Macrophytes can serve as direct or indirect food basis. Most fish species are
opportunists, who change their nutrition rates according to abundance and avail-
ability. Habitats for feed search even change along with fish age and season
(Opuszynski 1992; Opuszynski and Shireman 1995). The before-mentioned authors
described herbivorous fish as those fish whose nutrition consists – at least in most
life stages – of more than 50% of macrophytes. Food analysis findings in some
individuals in the Alte Donau samples concur with this definition. Roach (Rutilus
rutilus) and Rudd (Scardinius erythrophthalmus) can consume high amounts of
macrophytes. Bream (Abramis brama), White Bream (Blicca bjoerkna), Crucian
Carp (Carassius carassius), Tench (Tinca tinca) and Carp (Cyprinus carpio) ingest
low amounts of macrophytes and Perch (Perca fluviatilis) and juvenile Pike (Esox
lucius) only sporadically feed on herbal components (Prejs 1984). The main con-
sumers of macrophytes, namely Roach and Rudd, use almost all species of sub-
merged plants as nutrition, preferring soft parts (Werner et al. 1981).
Macrophytes and their epiphytes are important resources for fishes. Submerged
vegetation also contains high densities of small animals, like invertebrates (Crowder
and Cooper 1982; Heck and Crowder 1991; Diehl 1993; Persson 1993; Diehl and
Kornijów 1998). Herbivorous species feed directly on water plants while other spe-
cies sometimes benefit from high densities of epiphytes and invertebrates.
Cowx and Welcomme (1998) summarised the most relevant parameters of plants
for fish:
• Water purification – a. direct: for example, by oxygenation and conversion of
toxic ammonia to usable nitrates; − b. indirect: provision of huge surface area for
microbes to do the same tasks
• Nutrient recycling – removal during growth season, return during senescence
• Physical link between water and air for many invertebrates, e.g. larvae and
nymphs of caddis flies, mayflies, chironomids, which are food for fish and have
aquatic larval stages and aerial adults
• Refugia for zooplankton which graze phytoplankton and keep water clear
• Cover for huge range of invertebrates, many of which are food for fish
• Cover for fish – value and type vary with age/species of fish as well as type of
vegetation
• Spawning areas and sites of oviposition for many cyprinid and percid fish, e.g.
Tench, Roach, Rudd, Perch and also for Pike (all fish of the temperate waters of
Western Eurasia)
• Food source (living and dying) – direct source of vegetarian fish, e.g. Grass Carp;
indirect source via other animals
• Affect flow patterns – accrete sediments and deflect flow thus providing quieter
waters and faster shallows
• Create discrete habitat, which is a functional as physical structure.
The aim of this study was to investigate interactions between macrophytes and fish
biocoenoses in two different areas, which were chosen due to the aggregation of
different densities of plant composition and biomass. Major attention was paid to
the occurrence of any variation in fish length, number of individuals or species
diversity between the two types of habitats.
Area (1)
As mentioned above, the Kaiserwasser is a 6 ha nearly separated side water body
of the Alte Donau. It is connected by a small channel with a maximum width of 3 m
and a bridged gap of approximately 5 m.
The experimental area was situated in the small channel and was quite homoge-
nously covered with macrophytes of different densities. The dominant submerged
vegetation was built by Myriophyllum spicatum, which was accompanied by spo-
radic occurrence of Ranunculus circinatus. Additionally, branches of Salix were
touching and/or penetrating the water, underwashed roots and root-beards served as
fish-shelters and habitats along the shore-lines. The mean depth varied between 130
to 160 cm; fine sediments dominated the location; a small area was covered with
gravel.
Area (2)
This experimental site was located 10 to 15 meters in front of a reed belt covering
the shore line of the western part of the ‘Gänsehäufel’ island. The average water
depth of 200 cm was comparatively deeper than in Area (1). Eurasian Watermilfoil
(Myriophyllum spicatum), the dominant submerged macrophyte, occurred in a gra-

dient from sporadic- but mainly very dense stocks. Muddy to silty sediments built
the bottom area.
15.4.1 Methods
The method for classifying the occurring density of the macrophytes in the sampled
areas was the ‘Kohler-Index’ after Kohler (1978) and Kohler and Janauer (1995).
The amounts of plants were estimated by species via the following classes:
1 = very rare
2 = rare
3 = common
4 = abundant
5 = very abundant, in masses
The ‘Cumulative-Kohler-Index’, also called CKI (Pall 1996), was used for the
characterisation of mixed macrophyte stocks, according to the classes mentioned
above.
For the evaluation of fish densities in submerged plant stocks the modified ‘air-
lift method’ (Löffler 1988; Adelmann 1999) was used very effectively. The air-lift
has a quadratic frame of 2 × 2 m, made of plastic tubes with a diameter of 110 mm.
On the upper part of the tubes four ball taps, serving as air outlets, are fixed (see
Fig. 15.19). On the bottom side of the tubes, small holes are installed to enable the
entrance and elusion of water. Additionally, an air pipe with a back-pressure valve
is fixed on the main tube and connected via a hosepipe to a boat located compressed
air bottle, which supplies the system with compressed air. On the quadratic frame
construction, a fine net is fixed with a height of 2 m and a sinker rope at the lower
end (see Fig. 15.20).
Before starting the procedure, submerged plants were determined on species
level as well as the average height, and the frequency was estimated after Pall
(1996). The net was put on top of the tube frame, the valves were opened and the
air-lift-apparatus sunk at the chosen site. As soon as the air-lift is placed on the bot-
tom of the water body a scuba diver controls the exposition, puts the net under the
tubes and closes the four ball-taps (Fig. 15.21). After approx. 30 min, during which
time the slightly disturbed environment can swing back to normal and scared fish
will turn back to their preferred area, compressed air is pumped into the system and
the air-lift starts to swim up while the net unfolds itself (Fig. 15.20). Then the diver
goes down again and starts folding the net, beginning from the ground up to the
water surface (Fig. 15.22). At the end of the procedure, assisting persons on the boat
can take out the caught fish with a dip net. Fish are determined to species level,
measured and returned to the water.
Fig. 15.19 The airlift
Fig. 15.20 Schematic diagram of the airlift. (Source: Moser 2001; Reiter 2001)
Fig. 15.21 The diver closes the valves. (Source: Moser 2001; Reiter 2001)
Fig. 15.22 Removing the fish from the airlift net. (Source: Moser 2001; Reiter 2001)
15.4.2 Results
The investigations of different densities of macrophyte stocks display a clear picture

when airlift-net-samples are analysed: sparse standing plant densities host the low-
est numbers of fish in their surroundings. Areas with medium densities show slightly
increased values in fish abundances, but dense stocks, mainly built of Myriophyllum,
attract significantly the highest values of fish specimens (see Fig. 15.23).
Fig. 15.23 Number of fish 120

individuals in sparse-, 18
medium- and dense
macrophyte stocks from 100
Number of individuals
investigations with the
airlift in 2006 80
60
17
40
20
0
sparse middle dense
In a detailed analysis of the species density in correlation with the ‘Cumulative-

Kohler-Index’, results show 0.71 individuals of Perca fluviatilis, 0.28 of Scardinius
erythrophthalmus and 0.32 of Rutilus rutilus in areas with lower plant-densities
(CKI class 3) expressed as mean values per ‘air-lift’ catch. 7.75 individuals of Perca
fluviatilis and 0.4 of Gymnocephalus cernua could be proofed in macrophyte stocks
of a density of 3.5. In the following class ‘four’, 5.27 Perca fluviatilis and 0.9
Gymnocephalus cernua as well as 0.9 Alburnus alburnus were detected. At a den-
sity class of 4.5, the mean comparable catch per unit displays 5.6 Perca fluviatilis,
0.5 Rutilus rutilus, 0.13 Proterorhinus semilunaris and the same value for
Gymnocephalus cernua.
Most individuals were proofed in macrophyte structures with a density of ‘five’.
8.34 Perca fluviatilis, 2.0 Rutilus rutilus, 0.23 Proterorhinus semilunaris, 0.9
Lepomis gibbosus and 0.1 Blicca bjoerkna were caught as mean value per air-lift
sampling. The T-test confirms a decrease of abundances with lower densities.
Not only the number of individuals, but also the fish species diversity is higher in
denser macrophyte structures. Fig. 15.24 gives a pooled overview on the amount of
fish species in CKI 3–3.5 and 4–5. It shows the dominance of Perca fluviatilis very
clearly. The species White Bream, Pumpkinseed and Western Tubenose Goby were
captured only in macrophyte habitats with densities of 5. In areas with densities of
4 and 4.5, the Ruffe was caught with 1.0 and 0.4 individuals per trial. Perch was the
species which dominated in most samplings. In plant-structured areas with CKIs of
4.5 and 5, mean abundances of 6.38 individuals per air-lift were captured. The
Roach is a ubiquitous species; it could be proofed in all densities.
An additional result shows differences in the body-length of fish habituating the
different plant–density structures: In areas with a density of 4.5 a small range of
lengths between 55 and 67 mm is detected for Perch (median is 60 mm). At a CKI-
density of 5, the median is 62 mm; but the length spectrum ranges from 25 to
95 mm. In macrophyte-structures with CKI-class 4, mainly Perch with almost the
7
CKI classes 3 - 3.5
6 CKI classes 4 - 5
5
Individuals (CPUE)
is
ha is
us
na
us
us
us
nu
til
ar
rn
os
lm
til
rk
ia
hr ilun
er
ru
bu
oe
bb
v
c
flu
s
al
bj
em
gi
ht
us
ilu
a
op
us
ca
is
al
ut
s
rc
m
rn
ic
ph
R
s
Pe
po
ni inu
Bl
bu
yt
ce
er
Le
ar orh
Al
no
us
ym
Sc ter
o
G
di
Pr
Fig. 15.24 Pooled data on the amount of fish species in CKI 3–3.5 and 4–5
same length was found. The T-test confirms a significant difference between class
4.5 and 5.
Roach showed the highest length variety in areas with a CKI density of 5. The
median is situated at 50.5 mm and the length values vary from 45 to 59 mm.
Although the Roach inhabited a larger variety of types of macrophyte stands, no
statistically significant preference for a particular macrophyte-habitat structure
could be derived from Kruskal-Wallis-Test.
These findings of different habitat structures favoured by individual fish species
along the water basin of Alte Donau, illustrated the horizontal distribution of fish
colonisation. Additionally, differences of the plant biomasses become visible in a
vertical view of the same stocks. Close to the surface, more plant biomass is estab-
lished – specifically in Myriophyllum accumulations – than in the bottom area.
A comparison between these two vertical ‘densities’ shows the same general pat-
tern as seen in the already described areal situation (CKI, horizontal density).
As shown in Fig. 15.25, the species Roach, Pumpkinseed, Perch, Ruffe and
Tench occurred in varying numbers and densities in the two different vertical strata
of Myriophyllum stocks. The results are generated by airlift-net-sampling. The hori-
zontal differentiation between dense and loose strata is given by a specific depth –
the mowing depth – used in macrophyte management (Fig. 15.25). When the upper
part was removed by a mowing boat, the remaining loose plant parts could be sam-
pled again by airlifting. Perch was still proven in quantities of two thirds in compari-
80
70
60 n = 152
50
40
30
20
dense
10
0
us
ilis
ca
u
u
til
os
tin
rn
at
ru
Mowing layer
vi
ce
bb
a
flu
s
nc
gi
us
ilu
Ti
a
is
al
ut
rc
m
ph
R
Pe
po
ce
Le
no
ym
G
80
70
60 n = 36
50
sparse 40
30
20
10
0
us
us
ilis
ua
ca
til
os
tin
rn
at
ru
vi
ce
bb
a
flu
s
nc
gi
s
ilu
Ti
a
is
al
ut
rc
m
ph
R
Pe
po
ce
Le
no
ym
G
Fig. 15.25 Number of fish specimens in dense and sparse layers of macrophyte stocks from airlift-
investigations in 2003
son to the ‘before mowing situation’, but the other species were reduced much more
dramatically (e.g. Pumpkinseed reduction by 90%).
Mowing macrophytes is used as a tool to accommodate the users’ needs of this
150 ha water body as a ‘downtown’ recreation area. Within the last decades several
types of mowing boat were used. Different techniques for removing the macro-
phytes from the water body resulted in different threats for the fish assemblages,
which use the water plants as shelter, nursery, spawning ground or foraging area.
The biggest boat – operated by the Water Authority of Vienna – does not have a
mowing bar, but an elevator, which tears off the upper part of the plants and delivers
them into a storage hold on the boat. A mid-sized boat and a small boat use a mow-
ing bar, plant parts swim up to the surface and need to be collected/removed after-
wards. The principle of the mowing activities is shown in Fig. 15.26.
For the fish, the mowing activity is a disturbance, which forces them either to flee
deeper into the biomass bundles or to stay close to the cut plants, thus running into
danger of being trapped by collecting activities.
Fig. 15.26 The principle of the mowing boats
35
Large mowing boat
30 Medium mowing boat
Weight of macrophytes [kg]
Small mowing boat

25
20
15
10
0
2004 2005 2006 2007
Fig. 15.27 Weight of macrophytes [kg] per fish individual and boat for 2004–2007
Figure 15.27 gives an overview of different years sampled with different sizes of
mowing boats and the mean weighted plant biomasses examined, to find an indi-
vidual fish. In this figure, no details of the species and size are given, but the catch
is definitely dominated by young of the year class fish.
As a result, it becomes clearly visible that the small sized boats are much more
protective of the fish. The method of the big boat, to collect the torn macrophytes
immediately by a conveyer belt and elevate them to the boat surface, gives fish only
a very short time for a decision to escape. Considering yearly mowing quantities in
tons of macrophytes, it is quite ruinous for a fish population to lose one young of the
year individual with every 2.1 kg macrophytes harvested (result for 2006). At a
count of one ‘young of the year’ as by-catch with 31.3 kg of macrophytes, the small
boats with mowing bars and separately harvested biomasses give the impression of
being 10 times safer than the scenery described earlier. These results motivated the
responsible authority to develop mowing concepts with small boats only. The activi-
ties are more intensive and costly but more protective for the fish.
Weight [g] Number of eggs
0
1
2
3
4
5
6
14.05.2009
0
500
1000
1500
2000
15.05.2009 14.05.2009
16.05.2009 15.05.2009
17.05.2009 16.05.2009
18.05.2009 17.05.2009
19.05.2009 18.05.2009
20.05.2009 19.05.2009
21.05.2009 20.05.2009
22.05.2009 21.05.2009
23.05.2009 22.05.2009
24.05.2009 23.05.2009
25.05.2009 24.05.2009
26.05.2009 25.05.2009
27.05.2009 26.05.2009
28.05.2009 27.05.2009
29.05.2009 28.05.2009
30.05.2009 29.05.2009
31.05.2009 30.05.2009
01.06.2009 31.05.2009
02.06.2009 01.06.2009
Fig. 15.29 Weight of eggs [g] found per day in 2009

03.06.2009 02.06.2009
04.06.2009 03.06.2009
05.06.2009 04.06.2009
06.06.2009 05.06.2009
06.06.2009
07.06.2009 07.06.2009
08.06.2009 08.06.2009
09.06.2009 09.06.2009
10.06.2009 10.06.2009
11.06.2009 11.06.2009
12.06.2009 12.06.2009
13.06.2009 13.06.2009
14.06.2009 14.06.2009
15.06.2009 15.06.2009
16.06.2009 16.06.2009
17.06.2009 17.06.2009
dently low values (Fig. 15.29) correspond to the numbers of eggs.
18.06.2009 18.06.2009
19.06.2009 19.06.2009
20.06.2009 20.06.2009
Fig. 15.28 Number of fish eggs collected from macrophytes within 6 h (per day) in 2009
outstanding damage to fish eggs by harvesting in May. These fish eggs were sticking
numbers and biomasses during the spawning season. The results clearly show an
Numbers of collected eggs are shown in Fig. 15.28, the biomasses which show evi-
to the plants after spawning where the major family of fish, the Cyprinids, r eproduce.
In 2009, the major investigation of harvested macrophytes focused on fish egg
As a result of these investigations in 2009, the critical time of spawning activities

in May must be observed more carefully and mowing activities have to start later in
the year.
15.4.3 Discussion
It has been the most central target of the monitoring activities at the Alte Donau
since the 1980ies to investigate the ecological stability and to restore it to the high-
est extent possible. The reestablishment of depleted macrophyte stocks after their
collapse in the early 1990ies was a highly relevant aspect. Submerged plants fix
nutrients of the open water column, produce oxygen and provide heterogeneous
habitats for fish assemblages. Submerged vegetation is not only important as spawn-
ing area but also a relevant refugium for juveniles and foragers (Conrow et al. 1990).
The applied Cumulative Kohler Index (CKI) is a very robust index for expressing
the densities of macrophyte communities. The comparison of CKI 3–3.5 and 4–5
with the abundance of various fish species showed clearly that densely stocked areas
were more frequented than loose ones. Investigations of Randall et al. (1996) con-
firm these results. They detected that vegetated habitats establish higher fish densi-
ties, smaller individuals and higher diversity compared to less vegetated areas. In
the present investigation, seven species (Perch, Roach, Western Tubenose Goby,
Ruffe, Pumpkinseed, Bleak and White Bream) were captured in dense sectors,
whereas only four species (Perch, Roach, Rudd and Ruffe) were found in loose
plant stocks. Rudd occurred only in loose densities of submerged plants. Similar
results are confirmed by Bean and Winfield (1995) by detecting Scardinius eryth-
rophthalmus in the open water zone or in less dense macrophyte stocks.
A finding similar to the ‘CKI - fish density correlation’ seems to exist in the ver-
tical distribution of plant biomasses. After removing the uppermost part by a mow-
ing boat, the remaining loose plant parts could be sampled again by airlifting. Only
Perch was able to stay in the remaining loose plant structure, but occurrence was
reduced by one third. The other species were reduced even more dramatically.
Future investigations could develop an index analogous to the CKI, which would be
useful for applied macrophyte harvesting management.
The local post-mowing redistribution of the dissipated fish, and any further
effects, such as overcrowding and foodweb – and/or foraging changes, could not be
ascertained sufficiently in the present study. It is not clear, whether adjoining can-
opy habitats soak up the dispersed populations, or if some fish simply move deeper
to the looser plant cover.
In the present investigation, the number of individuals of Perca fluviatilis is dom-
inating in dense macrophyte areas. The number of individuals is more than double
in comparison to the next species. Investigations on ‘defend the structure’ actions of
Perch due to presence of predators in different habitats showed that they prefer more
vegetated areas to open waters (Christensen and Persson 1993; Berg et al. 1997).
In the Alte Donau system, the predating species Pike, Pike-Perch, Asp and Wels are
additionally managed by an angler organisation and are both naturally and artifi-
cially kept in high stocks. A pressure on Perch from the above predators is assumed.
Therefore, the findings of Christensen and Persson (1993) can be interpreted as
valid for the Alte Donau.
In a comparison of three mowing boats for macrophyte management, it became
visible that the smaller boats with mowing bars and separate plant collections are
more fish friendly than big units working with conveyer belt harvest, tearing plants
instead of cutting. Giving fish time to escape is one of the essential points in a gen-
eral ecological macrophyte management. Another easy to manage issue is the tim-
ing of mowing activities by considering the high spawning season of the Cyprinids
in May. Postponing management activities by a fortnight at least will give young of
the year fish class a better start. The new methods used by the Viennese Water
Authority are based on the results of the fish monitoring from the past years and thus
represent the state of the art standards for future activities.
15.5 Conclusions
All of the fish ecological measures tested in the Alte Donau were successful in sta-
bilising the hydro-biological system in general, as well as enhancing fish communi-
ties. It is vital to achieve a balanced composition of predatory and prey fish to lower
the feeding pressure on filtering zooplankton particles. This contributes to a clearer
water column as a basic parameter of a well-functioning Alte Donau system – serv-
ing the many purposes of an urban environment. In a ranking of predatory efficiency
based on enclosure experiments, the Asp proved most successful in reducing the
Bleak population, followed by Pikeperch and Perch. A stocking density of Pikeperch
at 65 kg ha−1 resulted in an improved visibility by approx. 15 cm in “Kaiserwasser”
(compare with long-term measurements of Secchi depth assessing the water trans-
parency in Chap. 6). Furthermore, stocking with species for sport fishing has to be
in line with a general, fish ecologically balanced, ecosystem management.
Constructing spawning grounds will be a further valuable contribution.
In addition, technical interventions, such as mowing activities, need to be man-
aged sustainably. Spawning seasons must be taken into account and prudent mow-
ing methods chosen. Specifically, it is recommended to operate small boats with
mowing bars and separate plant collection units, where ten times less young-of-the-
year fish are harvested in the macrophyte clippings, compared to convey belt
harvesters.
The Alte Donau as an ecological system is exposed to several pressures, as it is
one of the major urban recreation areas in Vienna. Keeping the current status of
water quality and all its biotic communities will require constant management to
support a sustainable system for future days. Fish are a key factor for success.
45 (Water Management – Vienna). We would like to thank all of the numerous collaborators,
master students, PhD students and the Municipal Department for permission of publication.
Additionally, we are grateful for the long lasting generous support of the ‘Österreichische
Fischereigesellschaft gegr. 1880” and the support with data. We also want to thank Ms. Erika
Thaler for her support in improving our English writing and the “Wiener Fischereiausschuss” for
providing fish catch and stocking long-term data of the Alte Donau.
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Chapter 16
The Stocks of Waterbirds on the Alte
Donau in Vienna
Rainer Raab and Ulrike Goldschmid
Abstract The whole area of the River Danube in Vienna, with its three more or less
parallel river stretches Donau, Neue Donau and Alte Donau is an important hiber-
nating area for waterbirds. The “Alte Donau” is an intensely used recreation area.
Here especially, people take an interest in birds and feed them in winter. For the
inhabitants of a city, this close contact with birds has a highly social aspect, notably
for children and elderly people. Nevertheless, the high number of birds has a nega-
tive impact on water quality. In the years 1997/1998 to 2000/2001 a total of 391.313
individuals were observed in the Viennese Danube area, consisting of 44 species of
water birds, 6 gulls and 8 hybrid species. Most frequent during the whole investiga-
tion was the Black-Headed Gull (Larus ridibundus) with a maximum of 15.579
individuals on a single day; Mallard (Anas platyrhynchos) and Eurasian Coot
(Fulica atra) dominate among the waterbirds (not counting gulls). Out of these 44
species 23 species of waterbirds, 3 gulls and one hybrid species and a total of 71.913
individuals were observed on the waterbody of the Alte Donau by regular countings
during the winters from 1997/1998 to 2000/2001. The three most frequent species
on the Alte Donau were the Black-headed Gull with a maximum of 5.965 individu-
als on a single day, followed by the Mallard and the Mute Swan.
Keywords Waterbird censuses · Hibernating waterfowl · Black-headed Gull ·

Mallard · Mute swan · Ecological relevance · Neozoa
R. Raab (*)
Technisches Büro für Biologie, Deutsch-Wagram, Austria
e-mail: rainer.raab@tbraab.at
U. Goldschmid
MA 45 – Wiener Gewässer, Wien, Austria

https://doi.org/10.1007/978-3-319-93270-5_16
314 R. Raab and U. Goldschmid
16.1 Introduction
A great deal of information exists on Anatidae in the Eurasian region, not least
because of the International Waterfowl Census, which has provided site-based
data for many species since 1967 (Scott and Rose 1996; Boere et al. 2006). The
Alte Donau in Vienna is a site that supports nationally important numbers of
waterbird species in winter (Aubrecht and Böck 1985; Aubrecht and Winkler
1997; Raab 2003a, b).
The aim of this study was to show the impact of various management measures,
the construction of the flood protection system for the Neue Donau, the restoration
of the Alte Donau and the construction of the hydroelectric power plant Freudenau,
on the stocks of water birds on the Alte Donau in Vienna.
16.2 Methods
The investigated stretch along the Alte Donau has a length of 5.6 km and a sur-
rounding area of approximately 158 ha. The Alte Donau is divided in the Untere
Alte Donau (Lower Old Danube, the south basin) and the Obere Alte Donau (Upper
Old Danube, the north basin) including the so-called “Wasserpark” (4,12 ha). The
wintering stocks of waterbirds were investigated on the Alte Donau (Old Danube),
Neue Donau (New Danube) and Donau (River Danube) in Vienna. The investigation
area was 1.118 ha in total (Table 16.1).
From winter 1997/1998 to winter 2000/2001 the number of waterbirds on the
Alte Donau, on the Donau and on the Neue Donau was counted in the course of 16
field trips per year from 23rd September to 22nd April. Additional data exist from
the winters of 1994/1995 to 1996/1997 (Raab 2001, 2003a, b).
The colony of the Grey Heron (Ardea cinerea) in the “Wasserpark” was investi-
gated on 10–17 days per year from February till July 1998 to 2003 (Raab 2003a, b).
Table 16.1 Investigation stretches (IS) used for the analysis along the Alte Donau, Neue Donau
and Donau with their length in km and area in ha
Alte Donau (158 ha) Neue Donau (368 ha) Donau (582 ha)
IS Name km ha IS From To km ha IS From To km ha
1 0 1.82 1.82 26 1 1916.4 1918.3 1.9 55
2 1.82 4.5 2.68 43 2 1918.3 1921.1 2.75 75
3 4.5 7.5 3 52 3 1921.1 1924 2.95 80
4 7.5 9.59 2.09 39 4 1924 1926 2 52
5 Lower Alte Donau 2.6 97 5 9.59 12.5 2.91 47 5 1926 1929 3 80
6 Upper Alte Donau 3 61 6 12.5 14.5 2 40 6 1929 1931 2 50
7 14.5 17.5 3 48 7 1931 1934 3 86
8 17.5 21.33 3.83 67 8 1934 1938 4 105
9 21.33 21.67 0.34 6 9 1938 1938.3 0.3 9
16 The Stocks of Waterbirds on the Alte Donau in Vienna 315
In the years 1997/1998 to 2000/2001 a total of 391,313 hibernating individuals were

observed in the Viennese Danube area, consisting of 44 species of water birds, 6
gulls and 8 hybrid species. Most frequent during the whole investigation was the
Black-Headed Gull (Larus ridibundus) with a maximum of 15,579 individuals on a
single day, Mallard (Anas platyrhynchos) and Coot (Fulica atra) dominate among
the waterbirds (without gulls; Tables 16.2 and 16.3).
Out of these 44 species 23 species of waterbirds, 3 gulls and one hybrid species
(71,913 individuals in total) were observed on the water body of Alte Donau during
the regular counting from winter 1997/1998 to winter 2000/2001. Most frequent on
the Alte Donau was the Black-Headed Gull (Larus ridibundus) with a maximum of
5.965 individuals on a single day, the Mallard (Anas platyrhynchos) and Mute Swan
(Cygnus olor, Fig. 16.1).
The Black-Headed Gull reached the highest numbers at the Alte Donau in the
early 1990s with a maximum of about 15,000 individuals on 1 day during winter
(Steiner 1985). The birds used the Alte Donau as a resting and sleeping place, espe-
cially in winter when it was covered by ice. In the daytime most of the birds fed at
the rubbish tip and came back to the sleeping place in the afternoon.
The first change took place when the flood protection system of the Neue Donau
was finished in 1988 (for more details about the history of Alte Donau see Chap. 2).
The 21 km long new water body runs parallel to the Alte Donau and divided the
population of the Black-Headed Gulls in two parts. The new water body was imme-
diately used as resting and sleeping place as well.
After the completion of the hydroelectric power plant Freudenau and after filling
the impoundment in November 1997, the population again split. Since then nearly
the same amount of birds (in total 15.579 individuals on 15th January 1999) rest and
sleep on the three more or less parallel waterbodies. So, the population on the Alte
Donau was reduced to 6,000 individuals at most.
Since 1967/1968 a lot of results of the Waterbird Census exist for the Alte Donau,
but also for other water bodies in the Danube region of Vienna (Table 16.4).
Before the Neue Donau (New Danube) was built the number of water birds in the
Viennese area was about 3,500 individuals in January (cf. 1984/1985), in years with
a large amount of birds hibernating in this area. Only a few years after the Neue
Donau was built (winter 1990/1991) an increase of water birds up to 4,600 individu-
als were observed (Laber et al. 1991) at Donau and Neue Donau. This number is
consistent with the observations of the years 1998/1999 till 2000/2001 (Fig. 16.2).
The total number of waterbirds in the Danube area of Vienna can be estimated at
about 5,500 individuals in the winter of 1990/1991, if we assume a number of about
1.000 individuals at the Alte Donau. Because of the construction of the Neue Donau,
the number of waterbirds in the area rose by about 2,000 individuals during midwin-
ter in years with high numbers of hibernating waterbirds.
An interesting aspect is the comparison of ecological relevance of the 9 investi-
gation stretches based on the maximum of individuals per hectare and day as well
Table 16.2 Observed waterbird species (excl. gulls) in the 4 winter half years 1997/1998 to
2000/2001, separated for Alte Donau, Neue Donau and Donau; maximum number of individuals
observed 1 day (max.), sum of individuals (tot, the number for the relative frequency larger than
1% is indicated in brackets)
Alte Donau Neue Donau Donau
Species English name Max. Tot (%) Max. Tot (%) Max. Tot (%)
Gavia immer Great Northern 1 3
Loon
Gavia arctica Black-throated 3 21 1 1
Loon
Gavia stellata Red-throated Loon 1 4
Tachybaptus ruficollis Little Grebe 2 6 67 1063 (1) 26 258 (1)
Podiceps cristatus Great Crested 3 8 20 245 22 64
Grebe
Podiceps grisegena Red-necked Grebe 1 5
Podiceps auritus Horned Grebe 1 5
Podiceps nigricollis Black-necked 1 2
Grebe
Phalacrocorax carbo Great Cormorant 16 52 142 1690 (1) 180 3054
(15)
Phalacrocorax Pygmy Cormorant 2 4 5 14 5 14
pygmeus
Ardea cinerea Grey Heron 20 400 (1) 9 75 26 342
Ardea alba Great Egret 1 1
Cygnus olor Mute Swan 249 4065 721 4821 (4) 53 312 (2)
(14)
Cygnus atratus Black Swana 10 96 1 7
Anser fabalis Taiga Bean Goose 7 7
Anser albifrons Greater White- 1 18 1 3
fronted Goose
Anser anser Greylag Goose 2 55 7 17 2 5
Branta canadensis Canada Goosea 14 687 (2)
Tadorna ferruginea Ruddy Shelduck 1 2 1 1
Anser sp. hybrid Goose hybrida 1 35
Cairina moschata Muscovy Ducka 4 65
Nettapus auritus African Pygmy 5 61
hybrid Goose hybrida
Callonetta leucophrys Ringed Teala 1 11
Aix sponsa Wood Ducka 2 14
Aix galericulata Mandarin Ducka 21 370 (1) 10 15
Anas penelope Eurasian Wigeon 24 201 6 14
Anas strepera Gadwall 13 77 11 41
Anas crecca Eurasian Teal 11 32 35 64
Anas platyrhynchos Mallardb 882 19457 1336 19358 719 10084
(66) (15) (51)
Anas acuta Northern Pintail 1 1 1 1
(continued)
Anas querquedula Garganey 6 18
Anas clypeata Northern Shoveler 4 17 1 5
Netta rufina Red-crested 1 6 38 349 6 10
Pochard
Netta rufina x Aythya Red-crested 1 1
fuligula Pochard x Tufted
Duck
Aythya ferina Common Pochard 22 154 770 7144 (6) 500 1012
(5)
Aythya fuligula Tufted Duck 132 810 (3) 1394 31977 504 1961
(25) (10)
Aythya fuligula x Anas Tufted Duck x 1 1
platyrhynchos Mallard
Aythya marila Greater Scaup 12 82 1 1
Aythya marila x A. Common Pochard 1 1
ferina x Greater Scaup
Somateria mollissima Common Eider 1 1
Clangula hyemalis Long-tailed Duck 1 3 1 5
Melanitta fusca Velvet Scoter 2 7 7 7
Bucephala clangula Common 2 2 49 543 20 152
Goldeneye
Mergellus albellus Smew 68 481 4 10
Mergellus albellus x Smew hybrid 1 2
M. serrator
Mergus serrator Red-breasted 3 13 17 27
Merganser
Mergus serrator x M. Red-breasted 1 1
albellus Merganser hybrid
Mergus merganser Common 53 395 27 176 (1)
Merganser
Gallinula chloropus Common Moorhen 19 292 (1) 14 158 5 25
Fulica atra Eurasian Coot 125 2708 2,742 60660 594 2287
(3) (47) (12)
Sterna caspia Caspian Tern 6 6
Total 1476 29,378 7186 129,520 2634 19,940
a
Members of the “Wasserpark” population
b
Incl. wrong coloured crossbreeds; bird names after Gill and Wright (2006)
as the total number of species (without gulls) per stretch before and after the full
damming up of the Danube (winter 1994/1995 to 1996/1997 and winter 1997/1998
to 2000/2001 respectively).
When comparing the 9 investigated stretches of the Donau and Neue Donau to the
2 at the Alte Donau distinct differences in numbers of individuals as well as in num-
bers of species can be detected before and after the filling of the hydroelectric basin.
Table 16.3 As Table 16.2 but for the observed gulls

Larus ridibundus Common Black- 5965 42,292 5855 73,843 7828 88,625
headed Gull (99) (98) (94)
Larus canus Mew Gull 60 218 391 1277 (1) 1104 4493 (5)
Larus argentatus European Herring 0 0 0 0 5 9
Gull
Larus cachinnans Caspian Gull 7 25 35 205 126 1480 (1)
Larus fuscus Lesser Black-backed 0 0 1 1 2 3
Gull
Larus hyperboreus x Glaucous Gull x 0 0 0 0 1 1
L. argentatus European Herring
Gull
Larus marinus Great Black-backed 0 0 0 0 1 3
Gull
Total 5988 42,535 6252 75,326 9059 94,614
Fig. 16.1 Waterbirds observed in Alte Donau: (a) Canada Goose (Branta canadensis), (b) Mute
Swan (Cygnus olor), (Photo credit: K. Teubner)
The numbers also varied between the stretches themselves. At the Donau, before the
reservoir was filled, more than ten species were only found in stretches 5, 7 and 8;
after filling the reservoir, this number of species was reached at the whole Donau
(Fig. 16.3). In the winter of 1995/1996, 20 times more waterbirds (excl. gulls) per
km were counted in the area downstream of the hydroelectric power plant Freudenau
(stretch 1 and 2) than in the reservoir of the power plant (stretch 3, 4 and 5).
In the winters of 1996/1997, 1997/1998 and 1998/1999 about three times more
waterbirds were found downstream from the power plant and in 1999/2000 only
half as many, as in the impoundment. Three years after the construction was final-
ised, the ratio between the impoundment and downstream area changed for the first
time. This change was confirmed in the 4th year. Both periods show that the number
of individuals in the upper part of the Alte Donau (Obere Alte Donau) is seven times
higher than in the lower stretch of the oxbow (Untere Alte Donau) (Fig. 16.3). The
explanation for this phenomenon is the intense regular feeding of the animals in the
Table 16.4 Results of the waterbird censuses in the Danube region of Vienna between 1967/1968
and 2000/2001
Year R Period n Tot SN n Tot SN n Tot SN
67/68 1 Jan 1* 6053 5
68/69 1 Jan 1 1912 5
69/70 1 Jan 1 1267 3 1 486 15
70/71 1 Jan 1 791 2 1 936 9
71/72 1 Jan 1 947 3 1 1480 12
72/73 1 Jan 1 1174 9
73/74 1 Jan 1 1738 4 1 214 8
75/76 1 Jan K 1 172 7
76/77 1 Jan K 1 839 9
77/78 1 Jan 1 2331 3 1 244 7
78/79 1 Jan 1 1069 3 1 452 9
80/81 1 Jan 1 1223 2 1 219 6
81/82 1 Jan 1 1116 3 1 136 5
82/83 1 Jan 1 1790 4
83/84 1 Jan 1 852 3
84/85 2 Jan 15 29,683 17
(3031)
84/85 1 Jan 1 2090 5 1 1124 14
85/86 1 Jan 1 1184 3
86/87 1 Jan 1 1534 3 1 667 11
87/88 1 Jan 1 1729 4
89/90 1 Jan 1 774 7 1 1430 10
89/90 3 20.10.– 23 21,439 15
30.03. (1986)
90/91 1 Jan 1 1589 15
90/91 4 06.10.– 27 25,692 25
17.03. (2307)
91/92 1 Jan 1 1683 14
92/93 1 15.11.– 4 1248 (505) 10
14.02.
93/94 1 14.11.– 5 6035 (2144) 18 5 940 (301) 10
13.03.
94/95 1 13.11.– 4 1861 (606) 14 5 11,235 22 5 1599 (523) 14
12.03. (2778)
94/95 5 11.10.– K 6 12,375 18 5 464 (146) 10
17.03. (3153)
95/96 5 26.09.– 9 6515 (1082) 14 16 32,329 26 16 3663 16
23.04. (4451) (1029)
96/97 5 23.09.– 15 7426 (1000) 16 15 20,783 25 15 2431 (749) 12
10.04. (2118)
(continued)
Year R Period n Tot SN n Tot SN n Tot SN
97/98 5 23.09.– 16 9361 (1085) 16 16 19,946 26 16 2928 (725) 15
22.04. (2271)
98/99 5 23.09.– 16 6768 (763) 17 16 41,109 33 16 3751 (680) 17
21.04. (5089)
99/00 5 23.09.– 16 6979 (898) 17 16 32,512 23 16 7204 15
22.04. (4635) (2083)
00/01 5 24.09.– 16 6270 (672) 17 16 35,953 28 16 6057 26
22.04. (4760) (1313)
R = references with 1 = archive BirdLife Austria (no data available for winter 1974/1975,
1979/1980, 1988/1989), 2 = Steiner (1985), 3 = Zuna-Kratky (1990), 4 = Laber et al. (1991) und
5 = Raab (2003a, b), n = number of field trips, tot = sum of the number of individuals (in case of
more than one field survey the maximum of observed individuals is given in brackets), SN = total
species number
* = value for Alte Donau is doubted, because the number of mallards (3.500) seems to be too high.
Since winter 1964/65 the upper stretch of the Donau (river-km 1938 to 1925) is part of the interna-
tional waterfowl census therefore continuous data (with some exceptions) is available. Data form
1964/65 to 1968/69 (Festetics & Leisler, 1971, Böck & Scherzinger, 1975) could not be taken into
consideration in this table as the data was aggregated for a larger region. The publication of Laber
(1991) could not be considered as well
4.12 ha area of the “Wasserpark” at the upper end of the Alte Donau, where about
50 individuals of domestic water birds also lived. In this area abnormal densities up
to 235 individuals per ha could be found.
Compared to the Donau, more species and individuals were found per hectare in
the Neue Donau. In stretches 1, 7 and 8 the differences at both investigation periods
were distinct. Noticeable is the strong increase in the number of individuals in
stretches 5 and 6 (Fig. 16.3) after the impoundment was completely filled. At the
time, there was a strong increase in macrophytes in the Neue Donau, which influ-
enced the troop-sizes of Eurasian Coot (Fulica atra) and attracted them from other
areas (Raab 2003a, b) During mid-winter the Alte Donau is of national importance
for only two waterbird species, the Mallard and the Mute Swan (Aubrecht and
Winkler 1997; Raab 2003a, b). The Mallard is the most common of all waterfowl-
species in Austria (Dvorak et al. 1994; Aubrecht and Winkler 1997). This is also the
case at the Alte Donau. The mid-winter population is about 1,000 individuals
(Fig. 16.2, Table 16.5).
In wintertime the population of the Mute Swan (Cygnus olor) on the Alte Donau
increased strongly because of migrating birds from Eastern Europe. Most of the indi-
viduals came from breeding areas in Poland and Czech Republic, but also from
Germany (especially the area of the former Eastern Germany) and Hungary
(Zuna -Kratky and Ranner 1993; Zuna-Kratky 2003). In 1984/1985 the average num-
ber of Mute Swans was 210 (Steiner 1985). In 1996/1997 the number was reduced to
95 and in the winter of 1998/1999, it further decreased to 53 individuals. The reason
for this reduction was less feeding by visitors. Within the period from January 1998
to February 2008 a total 171 individuals of the Mute Swan died in the area of
7000
Urban Danube area 39 other species
6000 Mute Swan
Common Pochard
5000 Tufted Duck
Eurasian Coot
4000 Mallard
3000
2000
Number of individuals of waterbirds
1000
2000
Alte Donau
1000
0
5000
Neue Donau
4000
3000
2000
1000
2000
Donau
1000
0
23.09.
07.10.
21.10.
04.11.
18.11.
03.12.
17.12.
31.12.
14.01.
28.01.
11.02.
25.02.
11.03.
25.03.
08.04.
22.04.
Fig. 16.2 Number of individuals of water birds by species (without excl. gulls) observed in the
Danube region of Vienna during the four winter half year from 1997/1998 to 2000/2001 (left to
right) for separated days of survey; arrows = more than 85% of the water surface of the Neue
Donau were frozen
“Wasserpark”. A lot of these birds died through collisions with the overhead contact
line of the newly built railway or collisions with trains. Between January 2007 and
February 2008 one of the dead individuals had been marked with a ring. This bird
died on 15.1.2008 (N Museum Praha, LB 4157) through collision with a train. It was
found on the track near “Wasserpark”. In the same period 2 other marked birds died
on the Donau near “Wasserpark”. One bird died on 27.2.2007 (N Museum Praha LB
4280) and one died on 12.9.2007 (Vogelwarte Radolfzell, RV 3832).
1994/95 to 1996/97
Number of individuals / ha
30 73 48
Number of species
25 40
20 32
15 24
10 16
5 8
0 0
9 8 7 6 5 4 3 2 1
AD ND 1
9 2
8 4 3
7 6 5
Danube
HPP F
1997/98 to 2000/01
Number of individuals / ha
30 48
Number of species
25 40
20 32
15 24
10 16
5 8
0 0
9 8 7 6 5 4 3 2 1
number of species AD number of species ND number of species Do

number of species ND individual/ha AD individual/ha Do
Fig. 16.3 Comparison of ecological relevance of the nine investigation stretches on the basis of
maximum daily number of individuals per hectare as well as the total species number (excl. gulls)
per stretch before (winter 1994/1995 to 1996/1997, top) and after the full damming up (winter
1997/1998 to 2000/2001, bottom) of the three sites studied, AD = Alte Donau, ND = Neue Donau,
Do = Danube; Further abbr. in the map: HPP F = hydroelectric power plant Freudenau
When the water body is frozen over, the waterbirds gather at the “Wasserpark”
(4.12 ha) which is most attractive for feeding (Fig. 16.4). The high density of birds
keeps the surface of the “Wasserpark” from freezing, because it is always in motion.
In this part of the Alte Donau system the dense population of water birds, up to 235
individuals per ha (Raab 2003a, b), has a negative impact on water quality.
In the period from 1969/1970 to 1987/1988 the amount of all water bird species
(excl. Mute Swan, Cygnus olor) introduced utility breed and ducks with abnormal
pigmentation was approximately 1,400 individuals on average per day in
mid-winter.
Table 16.5 Comparison of 9 waterbird species on the basis of maximum numbers in January of the three study areas and for Austria separated for the three
time periods 1970–1983 (cp. Aubrecht and Böck 1985), 1985–1994 (cp. Aubrecht and Winkler 1997) and 1995–2001 (cp. Raab 2003a, b)
Austria Alte Donau Neue Donau Donau in Wien
1970– 1990– 1970– 1985– 1995– 1985– 1995– 1970– 1985– 1995–
Species 1983 1994 1983 1994 2001 1994 2001 1983 1994 2001
Great Cormorant Phalacrocorax 520 4326 0 0 16 34 156 0 205 170
carbo
Mute Swan Cygnus olor 1195 2963 0 127 252 51 335 19 40 7
Mallard Anas platyrhynchos 40,581 42,154 920 1109 717 301 510 747 1243 719
Common Pochard Aythya ferina 6169 9497 67 0 14 54 890 305 252 500
Tufted Duck Aythya fuligula 11,175 24,414 64 31 60 345 1204 329 716 415
Common Goldeneye Bucephala 4961 3027 0 0 0 0 46 95 114 31
clangula
16 The Stocks of Waterbirds on the Alte Donau in Vienna
Smew Mergellus albellus 54 95 0 0 0 6 51 2 7 3

Common Merganser Mergus 1398 807 0 0 0 11 21 24 51 27
merganser
Eurasian Coot Fulica atra 29,980 31,328 1347 1219 197 651 2673 464 658 337
The values for Austria between 1990 and 1994 correspond to the sum of the five annual average values and are therefore no maximal values; the values for
Neue Donau from 1985 to 1994 include only data from the Untere Neue Donau. No data were available for Neue Donau for the period 1970–1983
323
Fig. 16.4 Alte Donau is popular for feeding water birds. The photo is taken in autumn 1996
From winter 1995/1996 to winter 2000/2001 the population declined to about

580 individuals per day, which is about 40% of the original population. The reduc-
tion of the waterfowl originated from the dramatic loss of macrophytes (see Chaps.
8 and 18), which were the main food source. In 1986 and 1987 the macrophytes of
the Alte Donau played a main role in the ecosystem of the water body (Löffler
1988). They disappeared almost completely until 1995 (Janauer et al. 1997).
The loss of macrophytes at the Alte Donau affected mainly the Eurasian Coot
(Fulica atra). In 1966/1967 (Festetics and Leisler 1971) and 1984/1985 (Steiner
1985) more than 1,700 individuals were counted. In 1997/1998 till 2000/2001 a
maximum of only 125 individuals was observed.
The non-indigenous Black Swan (Cygnus atratus), originating from Australia,
was introduced in about 1955 as attraction in the public park “Wasserpark”. By
1984 the population had grown to 150 individuals, which spread along the water
body. They had a breeding colony on a small island in the area of the “Wasserpark”
at the upper end of the Alte Donau. Because of the intense feeding by the visitors of
the park, the birds were breeding successfully all year round, with a peak in winter.
In 1985 some pairs bred successfully at temperatures of –15 °C. For example: In
1983 a total of 54 nestlings successfully hatched (Steiner 1985). To reduce the pos-
sibility of further expansion to other water systems in the area and a new awareness
of the problem of neozoa a strict reduction program was started. A person was hired
to shake the eggs in the breeding colony. Partly the shaken eggs put back, partly they
were exchanged for dummies to fix the parents as long as possible to the breeding
activity of the useless eggs. The dummies were readily accepted and incubated for
up to 342 days (Brugger and Taborsky 1994). In winter 1997 the population was
down to ten individuals, in 1998 the whole population was extinct.
70
60 breeding pairs
Number of individuals fledged offspring
50
40
30
20
10
0
1997 1998 1999 2000 2001 2002 2003
Years
Fig. 16.5 Development of the Grey Heron colony in the “Wasserpark, Wien-Floridsdorf”, in the
period from 1997 to 2003
These reduction measures were accompanied by intense information of the visi-

tors about the negative impact of feeding waterfowl on the ecosystem of the water
body. When the Black Swan vanished, the feeding activity of visitors and animal
welfare activists declined strongly. Only in cold winters, when other water bodies
freeze and all the waterbirds of the area gather at the “Wasserpark” the discussion
of feeding waterbirds comes up again.
Only a couple of years ago the Grey Heron started to breed in parks of large cit-
ies. Since 1998 Vienna also has a colony of Grey Heron like London and Amsterdam
(Berg, personal correspondence). The colony started in 1998 with the first 4 nests of
Grey Heron in the “Wasserpark”. In 1999, 5 nests could be found and in the breed-
ing season of 2000 11 nests of Grey Heron could be observed. In the 15 years prior
to this development there was no colony in the Viennese district.
On February 25, 1998 the first nest was discovered on a large poplar on a small
island in the “Wasserpark”. In all 4 new nests breeding was successful. The 4 breed-
ing pairs of 1998 had 15 (3 × 4 and 1 × 3) fully-fledged offspring. In 1999 the 5
breeding pairs had 18 (3 × 4 and 2 × 3) fully fledged offspring, in 2000 9 breeding
pairs had 24 (6 × 3 and 3 × 2) and in 2001 at least 62 offspring (1 × 6, 7 × 4, 6 × 3
and 5 × 2) fledged out of 19 nests. In 2002 24 broods took place in 20 nests of which
19 were successful. At least 62 offspring (2 × 5, 5 × 4, 8 × 3 und 4 × 2) fledged in
2002. The 18 breeding pairs of 2003 had at least 31 fledged offspring (2 × 4, 5 × 3,
3 × 2 und 2 × 1), so only half as many as in the previous 2 years (Fig. 16.5). The
main reason for this decline was a strong hail in Mai, which killed at least 17 off-
spring (Raab 2003a, b).
16.4 Conclusions
Investigations on water quality often neglect water birds. Here we have shown that
over-wintering birds, particularly large numbers, may affect water quality at least in
certain areas. Effects come from bird faeces and the remains from feeding introduc-
ing nutrients and pathogens to the water (see Chap. 17). New constructions in the
vicinity, restoration measures and recreation activities all influence the behaviour of
the bird flocks. In addition, management practices can influence and regulate spe-
cies composition and abundance.
45 (Water Management - Vienna). We would like to thank all of the numerous collaborators and
the Municipal Department for permission of publication. We further thank Lisa Greis for critical
reading and improving the linguistic style of an early draft of the manuscript.
References
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Reihe des Bundesministeriums für Gesundheit und Umweltschutz, Band 3, Wien, 270 pp
Aubrecht G, Winkler H (1997) Analyse der Internationalen Wasservogelzählungen (IWC) in
Österreich 1970–1995 – Trends und Bestände. Biosystematics and Ecology Series, Band 13,
Österreichische Akademie der Wissenschaften, 175 pp
Böck F, Scherzinger W (1975) Ergebnisse der Wasservogelzählungen in Niederösterreich und
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swan. Cygnus atratus Ethol 96:138–146
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Alten Donau in Wien in den Winterhalbjahren 1994/1995 bis 2000/2001. Unveröff. Studie im
Auftrag der MA 45 – Wasserbau, 22 pp.
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überwinternden Wasservogelbestände im Wiener Donauraum. Denisia 10:159–179
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22 – Naturschutz, 43 pp.
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1989/90. Vogelkundliche Nachrichten aus Ostösterreich 1:5–7
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Part IV
Recreation, Landscape Planning and
Synthesis
Chapter 17
Hygienic Aspects of a Recreational
Water-Body
Wolfgang Zoufal
Abstract The recreational use of surface waters always has been and still is a mat-
ter of grand concern for Viennese people. As bathing is most popular in Alte Donau,
there is a focus on the hygienic aspect of monitoring water quality. In this chapter
general, legal and operational facts are given and the hygienic status in the year
1986 is compared with a recent period. As hygienic water quality in general is
excellent in Alte Donau some rare singular events are depicted.
Keywords Bathing · Water hygiene · Indicator organisms · Water quality

management
17.1 Introduction
Bathing in the Danube has been important for the Viennese population for centuries.
In the eighteenth century the municipality felt even impelled to prohibit bathing due
to moral and safety reasons. But the Viennese ignored this law. Therefore the first
baths were built on the banks of the river Danube to legalize bathing and to be able
to protect the majority of non-swimmers by ropes or cages.
In the years 1870–1875 the regulation of the river Danube resulted in the creation
of Alte Donau as a stagnant waterbody that has been used for bathing since then. On
the island of Gänsehäufel a private open air bath was established by Florian Berndl
with a focus on nudism. Again, due to moral reasons this bath was closed in 1905,
but the municipality started a public open-air bath in the same place with a separated
beach for nudism that still exists today.
Offering 1.6 km2 of surface and approximately 10 km of bathing beach Alte Donau
is the largest stagnant water body in Vienna and is a prominent place of recreation for
1.8 million inhabitants. Since Neue Donau was constructed in the years 1975–1987,
W. Zoufal (*)
Vienna Municipal Department 39, IFUM – Laboratories of Environmental Medicine (head:
T. Nadiotis-Tsaka), Vienna, Austria
e-mail: wolfgang.zoufal@wien.gv.at

https://doi.org/10.1007/978-3-319-93270-5_17
332 W. Zoufal
Alte Donau may be in second place of importance for bathing in Vienna but it is still
the most traditional site. There are three municipal open-air baths, the biggest one
being Gänsehäufel, which has a capacity for 30,000 guests. In addition there are sev-
eral private baths and also widespread areas of the shoreline which are used as a
public beach. For many Viennese, bathing in natural waters is an important aspect of
the quality of life and therefore expectations are high for the quality of their water.
17.2 G
eneral Conditions for the Hygienic Aspect of Water
Quality of Alte Donau
In Austria it is illegal to introduce sewage into stagnant water bodies. Recently the
whole surrounding area of Alte Donau was connected to the municipal sewer sys-
tem and therefore there is not any risk left of leaky cesspools affecting Alte Donau
via ground water.
Between Alte Donau and Neue Donau there is an abandoned site that was used
to deposit domestic and construction waste after World War II. Because of the
hydraulic gradient from Neue Donau to Alte Donau it is necessary to operate several
wells near the shoreline of Alte Donau to produce a sink that prevents an inflow of
contaminated ground water, which could afflict the water body with chemicals and
nutrients in particular.
A very special problem of all water bodies in large cities are the adverse effects
of big populations of water birds (Chap. 16). The feeding of birds often serves as a
substitute for the experience of wildlife and the drawbacks of unnatural high popu-
lation densities are ignored. The feeding of birds is forbidden by law in the whole
area surrounding Alte Donau during the bathing season, but it proves challenging to
enforce this regulation especially in areas of extensive public use.
Bathing itself is a prominent risk for the quality of bathing waters. Due to the
strictly delimited water body of Alte Donau and the high degree of utilisation the
input of bacteria by bathers is capable of producing negative effects during periods
of extremely fair weather. For private and municipal baths, sanitary equipment (e.g.
toilets and showers) is regulated by law. On public beaches the administration pro-
vides fixed and portable toilets in increasing numbers.
Summing up, from the hygienic point of view, the risks for degraded water qual-
ity in terms of natural surface waters are low in Alte Donau.
17.3 Modalities of Hygienic Monitoring of Alte Donau
For decades the hygienic water quality of Alte Donau has been investigated. The
sampling points always focused on municipal open-air baths. These are Angelibad,
Strandbad Alte Donau and the three beaches of Gänsehäufel. Public beaches such as
17 Hygienic Aspects of a Recreational Water-Body 333
Kaiserwasser and the eastern shoreline of the southern part of Alte Donau were
added to the monitoring program in recent years. At the beginning of the bathing
season of 1997, these seven sampling points were reported to the European Union
as being bathing sites. Since then the sampling points are part of the monitoring
program according to council directives of 8 December 1975 concerning the quality
of bathing water (76/160/EEC) and 15 February 2006 concerning the management
of bathing water quality (2006/7/EC), respectively. Bathing season 2013 was the
last one within the transitional period. Both directives have been put into national
legislation by Austrian acts (BGBl. Nr. 254/1976, BGBl. Nr. 42/2012) and decrees
(BGBl. Teil II, Nr. 321/2012, BGBl. Teil II Nr. 349/2009).
The set of parameters investigated and almost any method performed have been
altered over the years of investigation. For example, Escherichia coli and faecal
coliforms, resp., were detected using a lactose-peptone-broth before 1998, from
1998 until 2000 a MUG-laurylsulfate-broth was used, beginning in 2001 the tests
were performed using Colilert 18 Quanti-Tray/2000 kits and since 2010 Bio-Rad
microplates are in use according to EN ISO 9308-3. The test for Salmonella sp. was
removed from the monitoring program at the beginning of the EC-program, because
of the low risk of contamination in Alte Donau, whereas total coliforms were
included according to the directive 76/160/EEC and dropped again according to
directive 2006/7/EC. Overall data on E. coli/faecal coliforms and faecal strepto-
cocci, both commonly used indicators for faecal pollution, are now available for a
long period. Before 1997 monthly sampling intervals were usually performed, but
beginning in 1997 the frequency was adapted to fortnightly ones according to the
EC-directive. The bathing season is determined by law as being the months June,
July and August, but samples are also taken from the end of April until September,
because during this period of bathing Viennese demand information about “their”
Alte Donau.
In the following specific hygienic aspects will be described in some detail.
17.3.1 Hygienic Status 1986
Looking at the data of the year 1986 two outstanding results can be seen. The first
one is the positive record of Salmonella typhimurium on April 14th at Angelibad. At
that time the upper part of Alte Donau where Angelibad is situated was afflicted by
the so-called Wasserpark. This is the northern end of Alte Donau, which was adapted
to be an urban park in the year 1927. Feeding of water birds was, and still is, a com-
mon practise in this area. This resulted in extremely high densities of water birds,
which had adverse effects on the water quality. To solve this problem a dam was
built between Wasserpark and the main part of Alte Donau in the 1980ies, but it had
not been efficiently constructed in those early days. In the years 1991–1996 this
dam was reinforced in the course of constructing an underground trail and since
then it is capable of efficiently separating the two water bodies concerned.
334 W. Zoufal
200
Strandbad Alte Donau
180
160
E. coli [MPN 100 ml ]
-1
140
120
100
Guide Value
80
LTA
60
40
20
0
Gänsehäufel Ost
160
140
E. coli [MPN 100 ml ]
-1
120
Guide Value
100
80
60
LTA
40
20
0
1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014
Fig. 17.1 Most probable numbers of Escherichia coli as MPN per 100 ml of surface water at two
sites in Alte Donau (Strandbad Alte Donau, upper basin; Gäsehäufel Ost, lower basin). The guide
value and the long-term average (LTA) is indicated
The second peculiar result was the detection of 5000 E. coli at the eastern beach
of Gänsehäufel on April 14th 1986. High numbers of indicators for faecal pollution
are rare. As long as there cannot be found any causality they have to be accepted as
a singular phenomenon within the range of water quality of a natural surface water.
17.3.2 Hygienic Status in the Years 1993–2014
Monitoring data from Strandbad Alte Donau as an example for water quality of the
upper basin and data from the eastern beach of Gänsehäufel representing the lower
basin indicate very good hygienic water quality throughout the years (Fig. 17.1).
Pollution from Escherichia coli as an indicator for the hygienic status of the system
17 Hygienic Aspects of a Recreational Water-Body 335
is well below the guidance value given in the directives concerning the quality of
bathing water in most of the years at both locations. Long-term average most prob-
able numbers (MPN) of E.coli are 64 and 40 for the two sites (Fig. 17.1). Sporadic
higher values are within the range of fluctuations expected in natural surface waters.
They usually occur in the middle of summer at times of high input of bacteria from
bathers and it has to be accepted due to the intense use of recreational water bodies.
In fact the data gained from the hygienic monitoring gave proof for the risk assess-
ment for Alte Donau as mentioned previously. An interesting aspect is the fairly low
numbers of indicators for faecal pollution especially in the years 1993–1996,
although there were problems with blooms of cyanobacteria and attempts of resto-
ration by RIPLOX-treatment were successfully carried out. It can be assumed that a
high biological activity in a water body is capable of depressing allochthonous bac-
teria and therefore supports the self-purification processes after faecal pollution.
17.3.3 Singular Events with Relevance to Hygiene
In the years 1993–2014 the only remarkable event was a ban of all beaches of
Gänsehäufel for 1 week in the year 1998, as a consequence of violation of total
coliforms’ statutory values. Looking back these non-conformities can be explained
by the method used for detection. In this year the MPN-MUG-laurylsulfate method
was used in Vienna on general official agreement in Austria. This method merely
gives numbers of 4600 and 11,000 MPN per 100 ml respectively, for a high concen-
tration thus being slightly below or slightly above the imperative value of the direc-
tive. In consequence of this event the method was improved by increasing from
three to five the number of vials used per each step of dilution. But this remarkably
increased the handling times. Therefore the Colilert kit was introduced in 2001.
This test gave data from 1 to 2419 MPN per 100 ml and 10 to 24,190 MPN per
100 ml, resp., using a dilution of 1:10 in a fine graduation throughout the range
achieved with an acceptable amount of handling time. According to Directive
2006/7/EC the test for total coliforms was finally removed from the monitoring
program in 2010.
Beginning with the bathing season 2000, swimmers in Alte Donau complained
about swimmers itch. Since then this phenomenon can be observed every year in the
middle of summer when water temperatures are high. The larvae of Trichobilharzia
szidatii have to be accepted as a part of biocoenosis of natural surface waters. There
are almost no possibilities to eliminate Trichobilharzia szidatii. The use of
molluscicides or the collecting of all snails has to remain a theoretical approach
under the general framework of Austria’s laws and environmental interests. The
treatment of wild water birds with drugs against the parasite is also not in accor-
dance with Austria’s law (BGBl. Nr. 16/1975, GZ 39.262/0-VII/B/10/03). The only
acceptable management measure that can be taken is the minimisation of feeding of
water birds in order to reduce the population densities of the host. Information to the
336 W. Zoufal
public therefore is the only way to handle this problem, which concerns administra-
tion as well as providers of baths.
17.4 Conclusions
When the general framework already fulfils ambitious legal demands it is not easy
to further improve the hygienic quality of a water body. Low risk impels sparse
chances for advancement. A limnologically stable state of the water body is impor-
tant for the usability as bathing water, but does not guarantee perfect water quality
in a hygienic meaning. Improvement can merely be achieved by giving the users
information that enables them to enhance their behaviour. The more information
they get the more they will feel responsible for their water and the more they will
take care of it. Providing the public with data about water quality combined with the
necessary background information about themes of actual interest (e.g. via internet)
is the way of choice to enable the users of a recreational water body in a large city
to achieve or maintain excellent water quality of their favourite bathing water.
References
Änderung des Bäderhygienegesetzes, BGBl. Nr. 42/2012

Bäderhygienegesetz, BGBl. Nr. 254/1976
Badegewässerverordnung, BGBl. Teil II Nr. 349/2009
Bäderhygieneverordnung, BGBl. Teil II, Nr. 321/2012
Bundesministerium für Soziale Sicherheit und Generationen 2003: Arzneimittelanwendung bei
freilebenden Wildtieren (GZ 39.262/0-VII/B/10/03)
Council directive of 8 December 1975 concerning the quality of bathing water (76/160/EEC)
Directive of the European Parliament and of the Council of 15 February 2006 concerning the man-
agement of bathing water quality and repealing Directive 76/160/EEC (2006/7/EC)
Tierärztegesetz, BGBl. Nr. 16/1975
Chapter 18
Restoration of the Littoral Zone
Karin Pall and Ulrike Goldschmid
Abstract In the context of the restoration of the Alte Donau, the planting of reed
and marsh plants (helophytes) was carried out in the early 1990s. The three planting
methods applied (planting young reed or reed cuttings, using a rhizome/soil mixture
and sowing), are described in detail. The development of the introduced helophyte
stands is further analysed. To allow helophyte plantings, some parts of the lake
shore had to be modified. The measure generated habitats in this urban oxbow lake
not only for the introduced species but also for various other species, which settled
spontaneously in the littoral zone.
Keywords Helophytes · Marsh plants · Shore zone · Lake restoration · Reed

planting · Reed belt · Phragmites australis
18.1 Introduction
Since the regulation of the Danube River in the late nineteenth century, the Alte
Donau has become an oxbow without connection to the main river (details about the
history and morphometry in Chaps. 2 and 3). Since that time, it has been used inten-
sively for recreation (Chaps. 17 and 19). For this purpose, the littoral zones were
partly fixed with concrete walls, especially at the outer bank of the oxbow. The
adjacent shore areas are mainly in private property. In the north-west basin small
gardens with weekend houses mostly reach the shoreline. The remaining space is
predominantly occupied by public swimming or recreation areas of different clubs.
In this part of the Alte Donau only small areas could be used for restoration mea-
sures. Around the south-east basin longer stretches of the shoreline, belonging to the
city of Vienna, could be used for the restoration of the littoral zone.
K. Pall (*)
Systema GmbH, Vienna, Austria
U. Goldschmid
Vienna Water Management, Vienna, Austria

https://doi.org/10.1007/978-3-319-93270-5_18
338 K. Pall and U. Goldschmid
The water quality suffered a great deal due to a long and intensive use for recre-
ation (boating, bathing, swimming, rowing, sailing, fishing), an enormous amount
of water bird habitation (Chap. 16), the lack of an appropriate sewage system and
the absence of an ecological water body management scheme. In 1987, an extensive
study of the ecology and the water quality was carried out (Löffler 1988), which
among other results pointed out the deficits in the littoral zone in particular. In addi-
tion to a variety of suggestions for improvements, this study marked the starting
point of a series of management measures carried out by the City of Vienna, Water
Management (Municipal Department 45).
On the one hand, the main aim of lake restoration management was to reduce the
input of nutrients by building a proper wastewater channel and including lake-
internal restoration measures (Chaps. 5 and 20). On the other hand, the natural
potential of self-purification of the water body itself should be improved by the
initiation of reed zones. Thus, the re-introduction of helophyte species was an
important step of sustainable management of lake restoration in Alte Donau. In the
following, the methods used for reed planting and the further development of the
species introduces are described.
18.2 Methods and Material
18.2.1 Reed Planting Method
The reed plantings in the Alte Donau were carried out from 1989 to 1994. To rein-
troduce reeds three different methods were used. The selection of the method
depended on the morphology of the shoreline.
18.2.1.1 Gentle Sloped Shoreline, Former Point Bar (Method 1)
In shallow riparian zones, reeds could be directly planted into the lake bottom. The
planting area reached from the shoreline to a water depth of about 0.5 m. Young reed
(Phragmites australis) was planted into small ditches. The young plants were fixed
with wooden sticks and coconut fibre cords (Fig. 18.1). In order to prevent swim-
mers and boats entering the newly planted reed area, a swimming fencing, built of
tree trunks, was installed (Fig. 18.2). Figures 18.3 and 18.4 show the planted area
one and five years later.
Usually, gentle sloped areas like these can be planted with reed cuttings that are
fixed to the ground with ropes. Therefore, it is necessary to cut the reed before
blooming. Using this technique, the plant material has to be submerged for about
18 Restoration of the Littoral Zone 339
Fig. 18.1 Planted reed
Fig. 18.2 Swimming fencing

Fig. 18.3 Planted area one year later
Fig. 18.4 Planted area five years later

15–20 cm. It takes new reed plants only a few weeks to grow out of the nodes. In the
case of the Alte Donau, however, it was not possible to use this method, as the young
plants would have been pulled out immediately by the enormous number of ducks
and swans (Chap. 16).
18.2.1.2 Steep Sloped Shoreline, Former Cut Side (Method 2)
In front of steep shorelines, mostly fixed by concrete walls, no simple reed planting
was possible. Therefore, initial structural preparations in the form of timber con-
structions had to be made. Along the shoreline, the timber poles, firmly connected
with wooden boards, were hammered into the ground at different distances to the
shore.
A gentle sloped shore area was created by filling the space between the shoreline
and the wooden construction with gravel (Fig. 18.5). In some places the concrete
behind the timber construction was removed, in others it was simply covered by
gravel. A mixture of rhizomes of reed (Phragmites australis) and soil was put on top
of the gravel in a 30–50 cm layer, just at the water level for avoiding a permanent
flooding (Fig. 18.6, further details about preparing this rhizome/soil mixture see
Sect. 18.2.2).
18.2.1.3 Additional Planting and Sowing (Method 3)
Additionally, typical marsh plants such as Alisma plantago-aquatica, Butomus

umbellatus, different Carex- and Juncus-species, as well as different species of
Mentha, Schoenoplectus and Typha were planted and/or sowed in some areas
(Figs. 18.7 and 18.8). Figures 18.9 and 18.10 show the planted area three and ten
years later.
18.2.2 Plant Material
For all methods, the plant material was collected from natural or semi natural stands.
For method 1, whole reed plants were dug up and carefully transported to the Alte
Donau in spring. For method 2, the reeds were harvested in retention basins along
the Wienfluss, a river flowing through Vienna. The plants were cut about 10–20 cm
Fig. 18.5 Timber construction, filled up with gravel
Fig. 18.6 Readily prepared rhizome/soil mixture

Fig. 18.7 Planting of additional marsh species, here mainly Juncus ssp.
Fig. 18.8 Planting of additional marsh species, here mainly Carex ssp.
Fig. 18.9 Planted area three years later
Fig. 18.10 Planted area ten years later

Fig. 18.11 Collection of seeds of different marsh plants
above the ground, just before a Caterpillar machine took out the roots together with
the soil. The mixture of roots and soil was transported to the Alte Donau and imme-
diately brought onto the prepared shoreline to a layer of about 30–50 cm. The best
time for this work is in early springtime after frost, but before the reed starts to grow.
The rhizome/soil layer should be kept wet, but not covered by water, because in this
case the roots would start to rot.
In some areas additional marsh plants were sowed in between the planted reed
respectively on the soil/rhizome layer. In order to get authentic, genetically correct
plant material, seeds were collected from natural locations, as seeds from Alisma
plantago-aquatica, Carex acuta, C. elata, C. riparia, Eupatorium cannabinum,
Glyceria fluitans, G. maxima, Juncus articulatus, J. compressus, J. effusus, J.
inflexus, Lythrum salicaria, Mentha aquatica, M. longifolia, Schoenoplectus lacus-
tris, Scirpoides holoschoenus, Typha angustifolia and T. latifolia (Fig. 18.11).
Rare or protected plants as Acorus calamus, Butomus umbellatus, Iris pseud-
acorus, Lysimachia thyrsiflora, Sagittaria sagittifolia, Scirpoides holoschoenus,
Typha angustifolia and T. latifola were grown artificially in special marsh plant
nurseries maintained by the City of Vienna, Water Management (Municipal
Department 45) (Fig. 18.12).
The following Table (Table 18.1) provides a list of all species used for planting
including later on abbreviations of the species names.
Fig. 18.12 Marsh plant nursery of the Municipal Department 45, Vienna
Table 18.1 Plant material Species Abbr.

(species spectrum) and list of
Acorus calamus L. Aco cal
abbreviations
Alisma plantago-aquatica L. Ali pla
Butomus umbellatus L. But umb
Carex acuta L. Car ssp.
Carex elata All. Car ssp.
Carex riparia Curtis Car ssp.
Eupatorium cannabinum L. Eup can
Glyceria fluitans (L.) R. Br. Gly flu
Glyceria maxima (Hartm.) Holmb. Gly max
Iris pseudacorus L. Iri pse
Juncus articulatus L. Jun art
Juncus compressus Jacq. Jun com
Juncus effusus L. Jun eff
Juncus inflexus L. Jun inf
Lysimachia thyrsiflora L. Lys thy
Lythrum salicaria L. Lyt sal
Menta aquatica L. Men aqu
Mentha longifolia (L.) L. Men lon
Phalaris arundinacea L. Pha aru
Phragmites australis (Cav.) Streud. Phr aus
Sagittaria sagittifolia L. Sag sag
Schoenoplectus lacustris (L.) Palla Sch lac
Scirpoides holoschoenus (L.) Soják Sci hol
Sparganium erectum L. em. Rchb. Spa ere
Typha angustifolia L. Typ ang
Typha latifolia L. Typ lat
Table 18.2 Plant Mass Index PMI Interpretation

(PMI) and interpretation
1 Very rare
2 Rare
3 Common
4 Abundant
5 Very abundant, in masses
18.2.3 Monitoring
18.2.3.1 Fieldwork
In 1994, just after the completed work of planting, periodical investigation of the
imported reed and marsh plants started. The monitoring was carried out annually in
July and/or August by driving along the entire shoreline with a boat. Every 100 m
the species spectrum and the individual abundances of the reed and marsh plants
were noted as a plant mass index (PMI; Pall and Moser 2009). According to Kohler
(1978) the five levels of the PMI can be interpreted as follows (Table 18.2):
18.2.3.2 Data Evaluation
In order to derive the plant mass development of all reed and marsh plants in the
Alte Donau the Cumulative Plant-Mass Index (CMI; Pall and Moser 2009) was
calculated. This term describes the overall plant mass of the plant group concerned,
regardless of the species composition. The CMI is calculated for each shoreline sec-
tion separately. The calculation algorithm is given in Formula 18.1.
Formula 18.1 Calculation of Cumulative Plant-Mass Index (CMI) after Pall and
Moser (2009)
 n 
CMI  min  3 PMI i3 , 5 
 i 1 
CMI = Cumulative Plant Mass Index (rounded down)

PMI = Plant Mass Index of different species
i = Current index of different species
In this paper, the five-level scale was reduced to three levels for practical pur-
poses. The used symbols and their interpretation are depicted in Table 18.3.
Table 18.3 CMI (Pall and Symbol CMI Interpretation

Moser 2009) as a measure for
1 + 2 Only single plants or single plant stands
vegetation density
3 Moderate dense plant stands
4 + 5 Dense to very dense plant stands
The Relative Plant Mass (RPM) after Pall and Janauer (1995) is used to analyse
the quantitative composition of the vegetation. The RPM-value of a species gives
the percentage of the plant mass of this species of the entire plant mass of all occur-
ring species. The calculation is to be done according to following formula:
Formula 18.2 Calculation of the Relative Plant Mass (RPM) after Pall and Janauer
(1995)
 M 
3
i •Li •100
RPM %  i 1
k  n 
 M 
•Li 
3
ji
j 1 i 1 
RPM = Relative Plant Mass of a species

Mi = for a section i estimated plant mass of this species,
Li = Length of section i
j = Current index of different species
18.3 Results
Only single stands of four different helophyte species (Iris pseudacorus, Phragmites
australis, Schoenoplectus lacustris and Typha sp.) were recorded from the Alte
Donau in 1987 (Löffler 1988; Fig. 18.13, picture left). Based on these results a pro-
posal for potential helophyte planting areas has been worked out (Löffler 1988;
Fig. 18.13, picture right).
During 1989–1994, large areas along the shoreline were reorganized and planted
with reed and marsh plants. The proposal of Löffler (1988) was followed to a great
extent in the selection of the planting areas. All in all, in about 20% of the shoreline
respective measures were set. Figure 18.14 (picture left) shows the distribution and the
vegetation density of helophyte stands after finishing the planting measures in 1994.
Until 2005 the reed and marsh plants grew up to dense vegetation over wide
areas. Figure 18.14 (picture right) shows the status of the helophyte vegetation in
2005. It is obvious that the reed and marsh plants from 1994 to 2005, not only have
increased their vegetation density, but also could spread out to further areas in the
Alte Donau, where no planting measures had been carried out.
As already shown in Table 18.1, overall 26 different species of reed and marsh
plants have been introduced in the Alte Donau. The species were brought into the
water body as entire plants, rhizome/soil mixture or as seeds.
Fig. 18.13 Helophytes in the Alte Donau. Left side: Stands of helophytes in the Alte Donau in
1987. Right side: Proposed areas for helophyte plantings. Both pictures from Löffler (1988)
In the years after the planting measures, not only the introduced species could be
found, but also some additional species established more or less dense plant stands
in the Alte Donau. These species were mainly found in the rebuilt shore sections but
also in some other regions of the lake shore.
Table 18.4 provides a list of the additional helophyte species, which settled spon-
taneously in the Alte Donau.
Table 18.5 shows the development of all – planted and spontaneously settled –
helophyte species in the Alte Donau in the years 1987–2005. As it can be taken from
this table, nearly all of the introduced species could establish more or less dense
plant stands in the Alte Donau. In general, there was a difference between planted
and sowed species. While the planted species could be found from the beginning of
the monitoring program in 1994, the sowed species mostly occurred only later. In
the end, however, both methods lead to satisfying results. Particularly the sedges
(Carex ssp.) and the reed (Phragmites australis) spread rather quickly from the
planting areas to many other regions in the Alte Donau. Some species, such as Iris
pseudacorus, Juncus articulatus, J. inflexus, Lythrum salicaria, Mentha aquatica,
Phalaris arundinacea and Typha angustifolia, after initial problems, finally reached
a rather widespread distribution. Other species, such as Acorus calamus, Alisma
Fig. 18.14 Helophyte vegetation in the Alte Donau. After finishing the planting measures in the
years 1994 (left side) and 2005, eleven years later (right side)
plantago-aquatica, Butomus umbellatus, Juncus compressus, J. effusus,

Schoenoplectus lacustris, Scirpoides holoschoenus, Sparganium erectum and Typha
latifolia, lost a part of their initial locations but could survive with at least some
plant stands. Only a few species, such as Glyceria maxima and Sagittaria sagittifo-
lia, disappeared altogether or could not be found again in the years following plant-
ing, such as Eupatorium cannabinum, Glyceria fluitans, Lysimachia thyrsiflora and
Mentha longifolia.
It is here worth emphasizing that even additional helophyte species found habi-
tats in the newly constructed shore areas. Some of them, such as Cyperus fuscus,
Equisetum palustre, Juncus filiformis, Ranunculus repens, Rumex hydrolapathum,
Schoenoplectus mucronatus, Solanum dulcamara, Sparganium emersum and Typha
laxmannii, could be found only in single years, while others, such as Alisma lanceo-
latum, Eleocharis palustris, Lysimachia nummularia, Myosotis scorpioides,
Persicaria dubia P. hydrolapathum and Scirpus sylvaticus, reached a relatively
wide-spread distribution in subsequent years.
To get an impression of the quantitative composition of the vegetation, the
Relative Plant Mass (RPM; Pall and Janauer 1995) was calculated (Fig. 18.15).
Immediately after planting the reed (Phragmites australis) clearly dominated the
Table 18.4 Additional Species Abbr.

helophyte species, found in
Alisma lanceolatum With. Ali lan
the years 1994 to 2005
Cyperus fuscus L. Cyp fus
Eleocharis palustris (L.) Roem. et Ele pal
Schult.
Equisetum palustre L. Equ pal
Juncus filiformis L. Jun fil
Lycopus europaeus L. Lyc eur
Lysimachia nummularia L. Lys num
Myosotis scorpioides L. Myo sco
Persicaria dubia (A. Braun) Fourr. Per dub
Persicaria hydropiper (L.) Delarbre Per hyd
Ranunculus repens L. Ran rep
Rumex hydrolapathum Huds. Rum hyd
Schoenoplectus mucronatus (L.) Palla Sch muc
Scirpus sylvaticus L. Sci syl
Solanum dulcamara L. Sol dul
Sparganium emersum Rehmann Spa eme
Typha laxmannii Lepech. Typ lax
Typha minima Funck in Hoppe Typ min
vegetation with a proportion of about 75% of the total plant mass. Until 2005, the
RPM-value of the reed was reduced to about 50% due to the spreading and mass
increasing of the other species. When considering the whole period, the sedge
(Carex ssp.) and Typha angustifolia represented the second and the third rank in the
RPM-scale. The RPM-values of all other species constantly remained below 5%.
These results demonstrate a quite high stability within the mainly artificially intro-
duced helophyte vegetation.
18.4 Conclusion
The results of this study demonstrate a successful re-introduction of reeds in an

urban lake. By restructuring the former mainly artificially modified shoreline, habi-
tats for many different helophyte species could be re-established. Most of the spe-
cies introduced between 1989 and 1994 showed an excellent growth in the following
years, independently of the planting method (planting, sowing or using a rhizome/
soil-mixture). Over the years additional helophyte species, which were not
introduced by planting measures, enriched the species spectrum. The current distri-
bution of helophytes along the shoreline represents to a large extent the aim estab-
lished in the 1980s and contributes to the general restoration of the Alte Donau (see
also Chap. 20).
Table 18.5 Dispersal of the different species in the Alte Donau in the years 1994 to 2005:
Occurrence in one 100 m section = 1, occurrence in 2–5 = 2, 6–25 = 3, 26–50 = 4 or more than
100 m sections = 5. Technique: P = planting, S = sowing, Rh = using a rhizome/soil mixture.
Abbreviations of species names see Tables 18.1 and 18.4
Relative Plant Mass of Helophyte Species

80
1994
70 1995
2005
60
50
RPM [%]
40
30
20
10
0
us
ax
al
ol
al
r
f
la
ar
in
eu
ss
an
ps
aq
ar
ts
ih
du
co
ra
m
p
a
c
ar
Ly
Sc
Ty
Iri
si
Ju
p
en
Ju
ly
n
Ph
Ly
Ph
Ty
re
C
Ju
G
Fig. 18.15 Relative Plant Mass of the different helophyte species after planting (1994/1995) and
about ten years later (2005). Abbreviations of species names see Tables 18.1 and 18.4
Acknowledgments This study was funded by the City of Vienna, Municipal Department 45
(Water Management). We would like to thank for permission of publication.
References
Kohler A (1978) Methoden der Kartierung von Flora und Vegetation von Süßwasserbiotopen.
Landschaft + Stadt 10/2:73–85
Löffler H (1988) Alte Donau. Limnologische Projektstudie – Ökosystem Alte Donau. Endbericht,
p 272
Pall K, Janauer G (1995) Die Makrophytenvegetation von Flußstauen am Beispiel der Donau
zwischen Fluß-km 2552,0 und 2511,8 in der Bundesrepublik Deutschland. Archiv für.
Hydrobiologie Suppl 101, Large Rivers 9/2:91–109
Pall K, Moser V (2009) Austrian index Macrophytes (AIM-module 1) for lakes: a water framework
directive compliant assessment system for lakes using aquatic macrophytes. Hydrobiologia
663:83–104
Chapter 19
New Planning Activities in Vienna’s
Water-Management
Brigitte Hozang
Abstract The oxbow lake ‘Alte Donau’ and its surrounding green spaces are one
of the most popular recreational areas in Vienna. Since the 1990s Vienna is steadily
growing, this has induced raising land use impacts and intensifying conflicts
between competing interests. As a consequence Vienna’s water-authority (MA 45
Wiener Gewässer) assigned the ‘Masterplan Alte Donau’ – an innovative planning
process integrating ecological and free-space planning into watershed management.
The main objective of this masterplan is thus to improve the usability of green
spaces and recreational areas in accordance to the sensitivity of the ecosystem and
nature protection. The Masterplan Alte Donau comprises an action plan for the lake
and the vicinity and for the water-related recreational facilities, including water
management and nature protection measures. Furthermore it outlines a freespace –
concept for the waterfront and green areas taking into account the balance between
the needs of all groups and the standards of green space planning. The Masterplan
Alte Donau thus strives to meet all requirements for the preservation of the sensitive
urban habitat of the oxbow lake on the one hand, and to keep the residential area
attractive by providing green spaces and water access for recreational activities on
the other hand.
Keywords Masterplan Alte Donau · Freespace- and water management ·

Upgrading urban waterfronts · Vienna’s recreational paradise
B. Hozang (*)
Technisches Büro für Landschaftsplanung, Zweiersdorf, Austria
University of Vienna, University of Natural Resources and Life Sciences, Vienna, Austria
e-mail: office@hozang.at

https://doi.org/10.1007/978-3-319-93270-5_19
356 B. Hozang
19.1 I ntroduction – The Alte Donau – A Sensitive Lakeside

and Highly Frequented Recreational Paradise
Cities are more than just a conglomeration of buildings: parks, playgrounds, rivers
and lakes are also important focal points of a city (Vassilakou 2014). Lakeside Alte
Donau, a former river arm, which became a very popular urban lake-site and
recreation-area, is a good example. The charming ambience of the famous natural
beaches (Strandbäder), the water-front restaurants and the boat jetties attract visitors
from all parts of the City of Vienna. The lakeside of Alte Donau is of vital interest
for the people and their identification with the city of Vienna. At the same time the
lakeside Alte Donau is characterized by its very specific nature and habitats (see
protection zones in Fig. 19.1). Several of the rare species and habitats occurring
here, are listed in the Council Directive on the conservation of natural habitats and
of wild fauna and flora (Richtlinie 92/43/EWG 1992).
19.1.1 A
Brief Historical Portrait of Vienna’s
‘Danube – landscape’
Vienna’s Danube – landscape covers the northern part of the city and was separated
from the historic city by a large area of wet meadows (history of Alte Donau see
Chap. 2). This 6 km wide area was hardly useful for permanent settlement since it
was prone to flooding and massive relocation of sediments. The interaction of the
river dynamics and settlement development results in a specific pattern of the struc-
ture of the town and its land-use (Alexander et al. 1977).
In the nineteenth century, regulation of the river Danube was forced by
industrialisation and an increasing demand for trade routes, particularly rail-
roads road bridges and a navigable river channel. While severe damages from
flooding made a regulation of the Danube indispensable, technical progress
Fig. 19.1 Protection zones at the Alte Donau. Strictly protected species with high priority
occurring at the Old Danube are Little bittern (Ixobrychus minutus), Common toad (Bufo bufo),
Edible frog (Pelophylax esculentus), Green toad (Bufo viridis), Common pipistrelle (Pipistrellus
pipistrellus), Beaver (Castor fiber)
19 New Planning Activities in Vienna’s Water-Management 357
Fig. 19.2 Map of the Danube before its regulation in 1868, the northern channel equates to recent
Old Danube. (Ladinig not dated)
made it possible. A straight artificial riverbed of 6.638 m length and 284.5 m

width was constructed which left the original main river stretch behind a dam.
This newly created freshwater are was called the ‘Alte Donau’ meaning the
‘Old Danube’. Additionally, an Area of 260 hectare land was reclaimed by fill-
ing up and levelling off some minor branches of the river (Fig. 19.2). New
trades were established such as ice factories and leather works (Hinkel 1994).
Urban expansion reached Alte Donau in the 1870s. In the period of promoterism
(Gründerzeit), town planning focused on the infrastructure for transport and traffic.
Five bridges were built across the new riverbed in the course of the regulation. They
connected the new suburbs with the city center.
Garden settlements were created during this period, which were utterly contrary
regarding to their social structure. On the right bank there were the so-called
‘Brettldörfer’, which were rather wild accumulations of huts scattered along the
edges of dump sites near the outbound bridge (Reichsbrücke). On the other side of
the river well-cultivated settlements and summer cottages were built simply for plea-
sure. Currently these former garden allotments are being transformed into prestigious
real estates with exorbitant property prices (Fig. 19.3). Alte Donau serves as a wel-
come promotion factor comparing the area to the famous beauties of the alpine lakes.
Because the city center of Vienna is sideways from the right river bank, Vienna’s
urban development planning focused since the 1960s on the north-eastern outskirts,
following the motto ‘get Vienna to the Danube’. (Stadtentwicklungsplan 1985, 2005;
Heiss 1964a, b). The Lakeside Alte Donau has gained increasing importance to pro-
vide recreational areas within reach of the town expansion areas. Former wilderness
358 B. Hozang
Fig. 19.3 Prospect of Kaisermühlen in 1929 (Eichert, in Lading, not dated)). Residential areas
were laid out alongside the Alte Donau, but settlement was kept at a distance – promoterist plan-
ning kept its symbolic distance to nature. Within two decades, a great number of boat clubs and
hires and bathing areas were established
areas along the riverbanks were cultivated and promoted as green corridors as part of
Vienna’s green belt.
This development altered the lakeside Alte Donau massively. Former waste
dump sites were cleared, sewers and motorways built. Skyscrapers spotted the
waterfront and three new subways connected this outstanding water landscape with
the city center (Fig. 19.4).
To cope with the significant increase of the population by 45,000 inhabitants
within 25 years in the vicinity, a so-called ‘Second City’ has been created.
19.2 M
asterplan Alte Donau – Novel Methods in Urban
Water- Management
The objectives of Masterplan Alte Donau are to improve the usability of green
spaces and recreational areas in accordance with the sensitivity of the ecosystem
and with nature protection. The challenge was to adapt innovative methods using
social survey and spatial planning tools (Hozang 2013). Therefore a crossover con-
ception was designed that comprises social objectives and relates them to hydro-
logical and ecosystem data.
Fig. 19.4 Town expansion reached the Alte Donau in the 1970s after the subway was built. (Photo
was taken in 2012)
The following thesis for innovative strategies and measures were considered:
• Quality in urban planning must come from a site- and group specific approach,
which perceives different needs of users. Innovative planning approaches and
methods must capture the different needs of target groups and clarify gender-,
age- and group-specific interests (Damyanovic et al. 2013; Kose and Licka 1995).
• Cooperative planning structures are essential to enforce a long-term process,
requiring a long – term partnership with relevant people, parties and participants
especially on a local level.
• The tense situations in upgrading urban areas require a forward-facing public dis-
cussion and political decisions how common land should be available and used.
• Involving the people and comprising their interests and needs gives way for inte-
grative solutions also on a spatial scale.
• Multifunctionality of green spaces is a valuable planning principle with benefits
according to reduced spatial and financial resources.
• Near nature, multiple use and availability are key words for innovative measures.
19.3 Results – Planning Activities
Urban development planning at Alte Donau has the advantage that the lake is sub-
ject of long term monitoring program.
360 B. Hozang
The investigations of hydrobiology and habitat-functions started after the dra-

matic nutrient increase and heavy blooms of cyanobacteria in the years 1992–1994
(Chaps. 6, 9, and 10). The situation of the Alte Donau is unique in Europe, for its
inappropriate relation of the shallow lake with approximately 3.6 millions of cubic
meters volume to its extensive utilization. In hot summers, more than a million
swimming visitors are counted. No other European lake can come up with these
many visitors in comparison to its size. Unlike other urban lake sites like Lake
Wannsee in Berlin (Germany), Lake Baldeney in Essen (Germany) or Lake Zurich
in Zurich (Switzerland), with rivers passing through, the Alte Donau has no flow-
through and its groundwater discharge is strongly reduced because of protection
measures against leakage from the old dump site.
19.3.1 Action Plan to Secure Water Quality
The Masterplan Alte Donau sums up the measures of the monitoring project and
extends them to collateral fields (Donabaum et al. 2005; DWS 2007; Pall 2007) The
main actions, which are pointed out in this plan, are the following:
• To secure the water quality by a permanent input of 50 l of water per second to
compensate the lack of groundwater-discharge (Taschke 2005)
• To safeguard compensation zones in nature near lakesides with less visitors
attendance
• To conceptualise and construct water bodies in forthcoming urban development
areas
• To campaign on water-quality-safeguarding and problematic nutrient upload
such as suntans and organic particle and to line out self-responsibilities and per-
sonal contributions
19.3.2 Public Green-Spaces at the Lakeside
I nnovative Methods of User Group Oriented Approach

The Masterplan Alte Donau is based on the Logical Framework-Approach, which
structures cause-effect-relations and helps to visualize relations and responsibilities.
The matrix-system is a reliable tool for cyclic project management and control of
results. In its present version special attention is being paid on the integration of stake-
holders and the adaptation of target oriented initiatives. Stakeholders from government
departments, industry, science and local people claimed their aims and interests.
The project design scheduled a three-step investigation to capture different needs
of target groups and relate them to behavioral patterns and spatial categories
(Hozang and Diran 2008; Lacina 1998).
• A typology of water – related free spaces and recreational facilities was gener-
ated including:
–– lakeside meadows, parkland, wilderness areas, nature protection areas, stages

and platforms, high banks, shallow bays and others
–– recreational facilities, playgrounds, landing stages, barriers, boat-hires, mari-
nas, clubs, boats and boat-types, restaurants and others
• The following survey of activities and utilization was based on these free-space
categories thus showing different habits and frequency of users according to the
different spatial types
–– Survey and recording of activities according to gender, age and groups.
–– Records of the number of visitors joining bathing areas ‘Strandbäder’ and
public lake sites.
• Especially local bound groups were approached for interviews and
questionnaires.
–– Interviews at schools and kindergarten of the Alte Donau vicinity
–– Interviews and questionnaires with tradesmen at the Alte Donau
–– Interviews with experts of administration and science
The results supported a more differentiated assessment of utilization, the various
interests of schools and child-care institutions and also the local traders.
ction Plan for Follow Up Activities on Public Green-Space and Recreational

A
Facilities
The main goals of the Masterplan Alte Donau are to outline a waterfront and green
space concept taking into consideration the needs of all user-groups in accordance
with standards of green-space planning and to adjust them to nature protection goals
(Hozang 2015).
The Masterplan Alte Donau points out the following main actions:
• To safeguard a public waterfront and free access to the lake. Free access is endan-
gered by private or club owned land use. To keep and to extend the public areas
at the water front is a main goal and a great challenge for town planning in
upgrading areas. The Masterplan Alte Donau provides accurate arguments
according to sustainability and nature protection.
• This includes a balanced and coordinated strategy to safeguard public space at the
water front. Vienna’s City development plan 2005 (STEP Stadtentwicklungsplan
2005) serves as a strong backing for a greener city. As a result of the analysis,
specifications and locations for further recreational facilities were pointed out.
Further private clubs or marinas with selective rented access were declined.
• Increasing public use of green spaces and recreational facilities especially on
the promenades, in the swimming pools and on the lakeside causes problems
and conflicts. Masterplan Alte Donau lines out where and how segregating
solutions are appropriate e.g. dog zones and where to apply multifunctional
solutions.
362 B. Hozang
• Standards and gender and group specific goals according to safety aspects on
paths and promenades and gender sensitive playgrounds are implemented, e.g. a
safe network of paths and promenades for pedestrians and cyclists. Seventy five
percent of the circuit paths and promenades achieve quality and security stan-
dards. Several sections are, however, neither attractive nor barrier-free. Especially
old bridges and landing stages should be adapted for people with special needs
according to the Austrian law (Bundesbehindertengleichstellungsgesetz 2005;
Groiss 2000; ÖNORM B 1600 2005).
• The survey showed clearly that almost 100% of the nature near and sensitive habi-
tats are located on public ground. ‘Near to nature appearance’ is also highly
favoured by a great number of visitors as can be seen in the interviews.
Multifunctional green-spaces match with nature near zones. Masterplan Alte
Donau lines out new design concepts, which value the uniqueness of morphology
and plantation of the Lakeside Alte Donau instead of substituting them by ordi-
nary parkland conceptions.
• Integrative measures in cooperation with owners and tenants follow the ambition
to convey the traditional mix of recreational facilities such as boat hire and sport-
ing facilities especially in the urban development areas and to adjust them to
ecological and hydrological demands. The measures comprise the redesigning of
a formerly enclosed tribune and open it to the public as well as bank-side planta-
tion works in cooperation with private tenants.
• Alongside rivers numerous competences are found according to historical status
causing inefficient maintaining. Reorganizing competences following the motto
‘to avoid parallelism’ bears a considerable potential.
19.3.3 B
athing Areas (‘Strandbäder’), Boat-Clubs
and Restaurants
raditional Mix But Ongoing Privatisation

T
Alte Donau bathing areas and public lakesides are well known and popular with the
Viennese. They have a long tradition. 1907 saw the opening of the municipal pool
Gänsehäufel – well known for the beauty of its old poplar groves, its layout and
also for the outstanding achievement of making it affordable for everyone. It is
generally considered a political exhibit of the former socialist’s ‘Red Vienna’s
housing policies’.
Gradually the Alte Donau developed into a highly appreciated recreational and
bathing paradise offering pools, sporting facilities, boat hires and landing stages,
clubs and marinas, restaurants and bathing huts.
Alike the bathing activities also the boat hires and boat sports have a long tradi-
tion. In 1887 the first boat hire was founded, followed by sailing- and rowing-clubs
(Fig. 19.5).
Fig. 19.5 Public landing-stages (left), marinas, cabins for rent (middle) and public lake-sides
(right, the photos were taken in 2008)
BesucherInnenfrequenz Alte Donau Strandbäder 1919 - 2017

Angelibad Strandbad ADO Gänsehäufel Bundesbad Summe ADO
1,300,000
1,200,000
1,100,000
1,000,000
900,000
800,000
700,000
600,000
500,000
400,000
300,000
200,000
100,000
0
1941
1951
1987
1997
1919
1921
1923
1925
1927
1929
1931
1033
1935
1937
1939
1943
1945
1947
1949
1953
1955
1957
1959
1961
1963
1965
1967
1969
1971
1973
1975
1977
1979
1981
1983
1985
1989
1991
1993
1995
1999
2001
2003
2005
2007
2009
2011
2013
2015
2017
Fig. 19.6 Recording of annual attendances of visitors attending the three bathing places (‘Strand-
bäder’) at ‘Angelibad’, ‘Strandbad ADO’ and ‘Gänsehäufel’, and the total number from 1919 to
2007. (Source: Hozang and Diran 2008)
The survey of recreational facilities, landing stages, boats, boat-types and vari-
ous water-bound activities and recordings of visitors distinguished spatial differ-
ences and conflict zones (Fig. 19.6). It also shows a significant increase of
attendances especially within in the last two centuries. The seven highest annual
visitor attendances were found in the last decade. The average annual attendance
was 731,000, the maximum annual attendance was 1 218,745 people. Nevertheless
more than 90% of the visitors attended the ‘Strandbäder’, where hygienic infra-
structure and additional pools relieve the tension of anthropogenic inputs into the
sensitive lake Alte Donau.
In question of boating activities, one can choose from various boats for rent, such
as electro-, pedal-, sailing and rowing-boats or memberships in one of the sailing
and rowing-clubs. During the last two decades the number of boats on the Alte
Donau has risen to a total number of 1477 boats (according to recordings and
interviews).
364 B. Hozang
The investigations stated a trend from public hires to private ownership. The most
expansive sectors are boat hiringand private yachts.. This tendency causes negative
effects such as reduction of free access and conflicts in highly frequented zones. In
addition the growing square measures of restaurants and clubsentail the enlargement
of bank protection measures, landing stages, pontoons, and the uprooting of riparian
vegetation. This development of increased commercial use of the lakeside leads to a
loss of ambiance as well as to a depreciation of biological values.
Other expanding lakeside utilizations are the marinas, often equipped with land-
ing stages for private yachts. New cabin sites are built on former brown-land or by
conversion of boat hires.
ction Plan for Recreational Facilities

A
Main goals of the Masterplan Alte Donau are to preserve the traditional mix of pub-
lic free-spaces, bathing areas, boat sports and restaurants and to make free access
for the people available. It points out main actions, such as:
• To secure access to the banks of the Alte Donau and to public boating facilities,
no further conversion of boat-hires or construction of marinas etc. In 2015/16 the
City of Vienna successfully took over two former club areas alongside the Alte
Donau, revitalized the banks and vegetation and permitted public access. To limit
boat frequency in the central zones of the lake, as boat traffic is dense regularly
causing conflicts and collisions.
• To enforce a multifunctional waterfront zoning form ‘public to private’ as shown
in Fig. 19.7. Demand will presumably increase in two development areas. This
comes into conflict with lakeside and water protection. The riparian vegetation
and habitats are of an outstanding ecological value and require protection mea-
sures. As new boating facilities and restaurants have to keep in distance from
protections zones, free access to the lakeside and natural riparian structures are
to be secured. The existing landing stages and locations instead should be used
more efficiently.
• To balance public and private land-use by assessment-plans and compensation
actions described by the water authority. Protection zones should be imple-
mented in land-assignment plans.
• To enforce partnership models aiming multifunctional usage. This can include
access to ramps for boat-clubs and licensees or the construction of a multifunc-
tional regatta tribune on former club owned and enclosed banks as shown in
Fig. 19.7.
• To provide incentives to private and commercial tenants to take part in regenerat-
ing the riparian ecosystem.
Fig. 19.7 ‘Green Regatta Tribune’ was constructed on former club rented land. The composition
put into practice public access to the water, a green tribune and typical tree and weed plantings.
(Design: Hozang and Meier 2011)
19.3.4 Riparian Structures and Vegetation
apid Transformation of Riparian Habitat and Vegetation

R
Riparian structures and vegetation are important factors of aquatic ecosystems, and
habitats, and for the specific scenery of lakesides. Despite urban development and
river regulation some characteristics of lowland riparian forests are still to be found
at Alte Donau, such as the typical diversity of steep and shallow banks and a zoned
vegetation with macrophytes, weeds, willow belts to poplar groves and other ripar-
ian forests (Fig. 19.8).
A detailed survey of morphology and vegetation was operated, including a map-
ping of weed, willow and poplar zones and completed by bioengineering parame-
ters, such as age, vitality, stability and aesthetic values.
Along the Alte Donau typical natural riparian structures and vegetation (Fig. 19.9)
are found mostly on the southern banks. Only 29% of the riparian structures and
vegetation of the Alte Donau are in a near to nature condition, based on classifica-
tions of morphology, connectivity and typical vegetation as parameters (Muhar
et al. 1996).
More than 53% of the lakeside are in a heavily modified condition with quays
and walls as bank protection bare of vegetation. Eleven percent are in an intermedi-
ate status with riprap bank protection and narrow vegetation zones.
366 B. Hozang
Eco morphological classification versus percentage

50
45.0
45
40
35
29,0
30
25
20
15
11.0
10 7.0 8.0
5
0
Nature near banks bank stabilisation with riprap bank protection side wall bathing beach
reed zones
Fig. 19.8 Eco morphological classification of the banks of Alte Donau versus percentage, general
goal is the extension of nature near banks
Fig. 19.9 Reed belt (mainly built by Carex riparia and Phragmites australis) and the fringes of a
bathing area. (Photo taken in 2013 by N. Novak)
The typical Danube lowland forests with belts of willows (Salicion albae) and
poplars (Populus alba, Populus nigra) are still found along the lakeside (Margl
1972). Their impressive height and colours distinguish the Alte Donau from other
areas of Vienna’s green belt. Sixty seven tree species were ascertained, with 35%
willows and poplars, 30% other typical Danube lowland species and an alarming
proportion of invasive neophytes, mainly Robinia pseudacacia growing along the
lake-side.
The forecast stated that 60% of the old poplars will have to be cut down for secu-
rity reasons within the next decade and that Robinia pseudoacaia, an invasive neo-
phyte, will be the dominant tree species.
Furthermore the effects of building activities lead to a considerable loss of ripar-
ian vegetation, due to design- and gardening styles which neglect the natural vegeta-
tion and especially replace trees by hedges.
ction Plan for the Regeneration and Protection of Riparian Habitats
A
and Vegetation
Lakes and watercourses are exceptionally rich in species and widely accepted as
invaluable habitats. Protection, restoration and management of aquatic and riparian
habitats are implemented in the guidelines of water rights, of construction measures
and maintenance of the water authorities.
Main goals of the Masterplan Alte Donau are regenerate the typical reed-zones,
willow and poplar groves on municipal land, to set up protections zones and to
cooperate with private tenants and traders in revitalisation measures (Fig. 19.10)
(Hozang 2015).
Masterplan Alte Donau points out main actions, which are as follows:
• To implement judicial protection, such as an obligate authorisation according to
the Austrian water rights with reference to Austrian Water Act 1959. It commits
users to restore riparian structures and vegetation when building or uprooting.
• To revitalize the natural riparian habitats and vegetation up to 70% of the total
length of the lakeside and at the same time to reduce the derogated sections from
55% to 46% and to enhance the well structured lakesides sections from 11% to
30% (Hozang and Novak 2013).
• To establish 11 protection and restoration zones according to the Water Act 1959
and Vienna’s Environment Protection Act (AVL and TBK 2006; Bluewaters and
Grimm 2002)
• To regenerate the riparian woodland and to enlarge the weed zones according to
the type-like species of EU – FFH guidelines. A General Draft for tree planting
for the recreational parklands was worked out in 2013 (Hozang and Diran 2013)
to arrange measures between the three municipal authorities responsible.
• To plant typical species such as willows (Salix alba, S. rubens, S. purpurea, S.
viminalis, S. incana, S. triandra) and poplars (Populus alba and P. nigra) in
riparian habitats and also in the surrounding parklands.
368 B. Hozang
Fig. 19.10 Lakeside zonation – natural lakeside and multifunctional green spaces by preservation
and expansion of the willow belt and the reed zones, plantation and protection of pondweed; A –
willow belt, B – reed zone, C – Charales and further submerged macrophytes
• To integrate all lakeside tenants and owners to cooperate with the restoration
measures. So far almost all of the measures were conducted on municipal prop-
erty by the water authority.
• To continue the macrophyte management and monitoring and to accomplish
temporary charophyte restoration zones in accordance with the demands of boat-
ing and swimming.
• To realize the aesthetic relations of the Danube landscape with its large dimensions.
• To work out and implement new design concepts allowing ‘nature to express
herself’ instead of antagonizing her’.
19.4 Résumé and Conclusion
Lakeside Alte Donau and its surrounding green spaces and recreational facilities are
very popular and highly frequented by a great number of people. The main objective
of Masterplan Alte Donau is to improve the usability of green spaces and recre-
ational areas in accordance with the sensitivity of the ecosystem, and with nature
protection.
Therefore, a crossover conception was designed that comprises social objectives
and relates them to hydrological and ecosystem data. A monitoring of hydro-
chemical, macrophyte and hydrobiological data from over 10 years of investiga-
tions, served as the main input for the analysis, combined with representative
statistics of census and the City development plan STEP 2005 (Stadtentwicklungsplan
1985, 2005). In addition, a three-step investigation was designed to capture different
needs of target-groups, behavioral patterns were related to spatial categories and
utilisation was surveyed and recorded according to gender, age and groups.
Masterplan Alte Donau comprises goals, measures for the lake and the surround-
ing green-spaces, and for the water-related recreational facilities, as well as coordi-
nated water-management and nature-protection measures.
A main goal and a great challenge for town planning in upgrading areas, is to
keep the water front open for the public. Masterplan Alte Donau provides arguments
according to sustainability and nature protection. It includes a balanced strategy to
safeguard free access to the waterfront, and to protect nature near sections.
Guidelines for forthcoming design and development follow the principles of multi-
functional waterfront zoning form “public to private”, and nature near plantation
and appearance.
Partnership models were launched aiming multifunctional usage, such as access
to ramps for boat clubs or the construction of a regatta tribune on former club owned
banks. Additionally incentives for nature near measures were given to owners or
tenants who cooperated in regenerating the banks of the lake.
Strategies and proposals were worked out to preserve the traditional middle scale
mix of public free space, swimming areas, restaurants and boat hires and to limit
further conversion to private marinas. A safe and barrier free network of paths and
promenades for pedestrians and cyclists was worked out.
Design principles for the green spaces were generated following the criteria user
oriented, multifunctional and nature near. It is of great aesthetic and social value to
restage the uniqueness of morphology and plantation of the lakeside Alte Donau.
The typical Danube lowland forests with belts of willows and poplars are still found,
but they have a share of only 32% of the lake-side’s vegetation. The investigations
also provided insight into an alarming proportion of invasive neophytes, mainly
Robinia pseudacacia.
A mission statement proposes the revitalization of natural riparian habitats and
vegetation up to 70% of the total length of the lakeside. The corresponding mea-
sures include plantations of the typical reed zones, willow and poplar groves on
municipal land. Eleven protection and restoration zones, according to the Water
rights and Vienna’s Environment Protection Act, were proposed.
The interview results asserted an approval of visitors with the ‘Near to nature
appearance’. This coincides well with the nature protection measures, and it assists
the ambitions to decrease maintaining costs. Alongside rivers you can find numer-
ous competences, according to historical status, causing inefficient maintaining.
Reorganising competences bears a considerable potential by obstructing differing
responsibilities.
Masterplan Alte Donau is a long-term, differentiated strategy and action plan. It
relies on a considerable number of people to permute to practice and reality. Its most
needed forces are the key persons on different levels.
Half term review of 2014 shows that:
• Water quality safeguarding is successful despite increasing visitor’s attendances.
• Public relations and social media offer information and questionnaires, on local
scale group specific guided tours and workshops are well attended.
370 B. Hozang
• Public Lakeside has been amplified by designing a Green tribune on former club
owned land.
• Bathing stages were redesigned according to aesthetical and functional criteria.
• Over 300 poplars and willows were planted to restore the typical vegetation,
attended by neophyte removal.
• Eleven private tenants have cooperated in restoration and plantation measures.
For the second half-time coordinated measures are scheduled within the
EU-LIFE+ Urban Lake Alte Donau focussing on further revitalization of banks and
vegetation, the enlarging of weed zones and new fish habitats, barrier free access to
the lakeside and intensified dissemination activities.
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Floridsdorf Hrsg.: Magistratsabteilung 22 der Stadt Wien, 28–44pp
Bauordnung für Wien 2015, Wiener Stadtentwicklungs-, Stadtplanungs- und Baugesetzbuch
(2013/2014) Gestaltungskonzept für gärtnerisch auszugestaltende Flächen, § 63, §77–§86, §79
Bluewaters ZT, Grimm DI (2002) Wiener Arten- und Lebensraumschutzprogramm, Leitlinien
Donaustadt: Hrsg. Magistratsabteilung 22 der Stadt Wien, 38–51pp
Bundesbehindertengleichstellungsgesetz 2005, BGStg, BMVIT, 4pp
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Donabaum K, Nagel G, Riedler P (2005) Die Alte Donau – Beispiel für ein erfolgreiches
Gewässermanagement der Stadt Wien, in: Perspektiven: Gewässerschutz in Wien. Stadt, Wien, 3pp
DWS, Hydro – Ökologie (2007) Limnologische Untersuchung – Alte Donau, Zustandsanalyse –
Hydrochemie und Plankton, i. A. der Stadt Wien, MA 45 – Wasserbau, 3pp
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Arbeitsgemeinschaft für Rehabilitation, Wien, 3–6pp
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aufbau, 3, 113pp
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report, 36pp
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Stadtgestaltung, Bd. 56, Wien, 22–28pp
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Wien, 34pp
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Naturgeschichte Wiens, Bd. 2, Wien, 499–654pp
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-Design principles, ON – Österreichisches Normungsinstitut, 4pp
Pall K (2007): Limnologische Untersuchung – Alte Donau, Zustandsanalyse – Makrophyten,
Magistratsabteilung 45 der Stadt Wien, 49-52pp
Richtlinie 92/43/EWG (1992) Flora – Fauna – Habitat – Richtlinie zur Erhaltung der natürli-
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BMLFUW, § 30 and § 31, 134pp
Chapter 20
Eutrophication, Management and
Sustainable Development of Urban Lakes:
General Considerations and Specific Solutions
for Alte Donau – A Synthesis
Martin T. Dokulil, Karl Donabaum, Karin Pall, Georg A. Janauer,

and Katrin Teubner
Abstract Intensively used urban water bodies are vulnerable to eutrophication.

The shallow lake Alte Donau (Vienna) can be seen as an example for the extent of
anthropogenic influence. Human impacts paired with changes in environmental
conditions gave way to eutrophication processes in Alte Donau. Due to the great
public interest restoration concepts and subsequently management programs were
established. This chapter provides a synthesis of the key aspects to evolve and
implement a successful water management plan. An attempt is made to generalise
our specific solutions to serve as a basis for the development of similar strategies for
other urban lakes.
Keywords Restoration · Groundwater · Seepage · Management · Improvement ·

Shallow lake
M. T. Dokulil (*) · K. Donabaum

e-mail: martin.dokulil@univie.ac.at; karl.donabaum@dws-hydro-oekologie.at
K. Pall
Systema GmbH, Wien, Austria
G. A. Janauer · K. Teubner
e-mail: georg.janauer@univie.ac.at; katrin.teubner@univie.ac.at

https://doi.org/10.1007/978-3-319-93270-5_20
374 M. T. Dokulil et al.
20.1 Introduction
Urban lakes are different types of natural or man-made water bodies within densely
populated areas. For the purposes of management, these lakes can be defined by
several operational criteria. They tend to be small and shallow, with surface areas of
less than 2.5 km2 and a mean depth of 6 m or less. The ratio of the watershed area
to lake area can be variable but is often 10:1 or higher, meaning that their water-
sheds exert a strong influence on the lake. As an overall index of development, the
urban lake watershed must contain at least 5% impermeable cover. The water bud-
get often depends on ground-water and precipitation or is entirely artificially con-
trolled. Most of these urban lakes must be managed for recreation, water supply,
flood control or some other direct human use regardless whether natural or man-
made (Birch and Mc Caskie 1999).
Curiously, the unique problems and conditions of urban lakes have received little
attention in scientific literature. This is particularly surprising given that many of the
management efforts are devoted to lakes and reservoirs that are distinctly urban in
character. While the watershed management literature is filled with phosphorus
budgets and watershed models, it is unusual to find overviews about the influence of
watershed development on lake quality and it is exceptionally rare to find studies
that have tracked changes in lake quality as a function of watershed development
over time (Schueler and Simpson 2001).
However, restoration and recovery of already eutrophicated systems can only be
achieved if external measures like watershed management are combined with inter-
nal measures.
20.2 Eutrophication
Small and shallow aquatic ecosystems generally have lower resilience than large,
deep lakes. Urban lakes are therefore more sensitive to water pollution and eutro-
phication suffering from natural or anthropogenic impacts. If man-made, these envi-
ronments tend to have rather regular shapes and higher shoreline development when
recreationally used resulting in negative impacts on their functioning (Naselli-
Flores 2008).
Metropolitan runoff flowing over impermeable surfaces collects large amounts
of nutrients resulting in higher unit area phosphorus load from storm-water than
other watersheds. Many urban watersheds receive additional loads from storm-
water overflow, failing septic systems or pollutant seepage. Urban lakes also have
unique internal nutrients sources such as water bird droppings, boat sewage and
sediment release (Traut and Hostetler 2004). As a consequence, phytoplankton
blooms and uncontrolled macrophyte growth may severely impair water quality and
cause sanitary risks. Massive growth of submerged plants may form an obstacle to
several forms of recreational use. Moreover, highly developed shorelines, including
20 Eutrophication, Management and Sustainable Development of Urban Lakes… 375
tourism exploitation (Dokulil 2014a), housing and development sites (Cappiella and
Schueler 2001) may significantly contribute to eutrophication and pollution. Urban
lakes may potentially be contaminated by various compounds particularly their
sediments causing long-term environmental problems and human health risks.
Because of these many facets impacts have on urban lakes, management efforts
must be integrative and sustainable (e.g. Sorensen 1996) especially when impacts
are amplified by climate warming (Dokulil 2014b).
20.3 Restoration and Management
Urban lakes are complex systems strongly influenced by disturbances within their
watershed. Therefore, thorough management of urban lakes must be tightly coupled
with the management of the watershed. The entire catchment must be supervised
and necessary strategies developed to optimise land use, erosion, housing, wastewa-
ter treatment, public transport, recreation, tourism or any other factor which might
be important in the watershed (NALMS 1988).
Internal restoration techniques are abundantly described particularly for shallow
water bodies (Carvalho 1994; Xu et al. 1999; Morscheid and Maehlmann 2005).
Methods have been summarized and classified in Fig. 20.1 after Singh (1982).
Among the many methods available, chemical stabilisation of phosphorus (Welch
and Schrieve 1994) and removal of lake sediment (Van der Does and Frinking 1993;
Björk 1978, 1994) are most popular. In many food web oriented lake rehabilitation
LAKE RESTORATION TECHNIQUES
PREVENTIVE (direct) AMELIORATIVE (indirect)
1. Drainage basin alterations

2. Diversion of wastes
3. Legal control measures
IN-LAKE OUTSIDE-LAKE
PHYSICAL CHEMICAL BIOLOGICAL PHYSICAL CHEMICAL BIOLOGICAL
1. Mechanical harvesting 1. Phosphorus Species 1. Mechanical 1. Clarification Biological

of biomass precipitation manipulation treatment 2. Active carbon treatment
2. Lake deepening or 2. Use of algicides 2. Reduction in adsorption
dredging and herbicides sediment input 3. Ammonia
3. Admission of stripping
unpolluted waters 4. Nutrient and
4. Aeration of water ion removal
5. Hypolimnetic drainage 5. Disinfection
6. Lake drawdown
7. Lake bottom sealing
Fig. 20.1 Classification of lake restoration techniques. (Modified from Singh 1982)
activities, manipulation of the fish community is a prime focus (Hansson and Butler
1994; Berg et al. 1997; Hansel-Welch et al. 2003).
As aquatic plants may easily become a nuisance in man-made waters, manage-
ment of aquatic vegetation is a major requisite usually by controlling abundant
growth of macrophytes, e.g. by extensive harvesting (Pieterse and Murphy 1990;
Bowmer et al. 1984; Madsen et al. 1988). More recently, macrophytes are recog-
nised as an essential component in suppressing algal dominance during rehabilita-
tion of eutrophic water bodies (Körner 2002). Regrowth of macrophytes was
reported in a majority of lake restoration studies (Hansson and Butler 1994; Perrow
et al. 1997). The re-establishment of aquatic vegetation in these studies occurred
mainly by natural propagation and without any active management (Hansel-Welch
et al. 2003). Under such conditions, the recovery of the water body towards the
macrophyte dominated state may take long and the ecosystem is at risk to shift back
to the algal dominated state.
Investigations on the essential environmental conditions promoting macrophyte
growth are numerous (e.g Riis and Hawes 2002; Crisman et al. 2005; Janse et al.
1998). The complexity of ecological pressures in lake environments, ranging from
sediment characteristics to herbivory, often impairs the proliferation of the aquatic
plants (Lau and Lane 2002; Irfanullah and Moss 2004).
The submersed aquatic vegetation is the essential factor determining water qual-
ity by preventing the growth of phytoplankton in shallow lakes (Bailey et al. 2002;
Gopal and Goel 1993; Wium-Andersen 1987; Berger and Schagerl 2003).
Several experimental approaches were developed for the transplantation of sub-
mersed plants for pools (Irfanullah and Moss 2004), lake littorals (Wychera and
Humpesch 2002; IGB 2005; Hilt 2005; Gross and Hilt 2005; Morscheid and
Maehlmann 2005) and for oxbows (Janauer 1995; Janauer and Pall 1998).
20.4 Sustainable Development
Water is so essential to life and the life processes of all living beings, that manage-
ment of water resources requires a new paradigm, the concept of sustainability
(Heintz 2004; Taylor and Goldstein 2010). This concept describes the dynamic con-
ditions and the resiliency or robustness of complex systems to adapt and thrive in
the face of change. Sustainable development shall meet the needs of the present
without compromising vital ecosystems as well as the ability of future generations
to meet their own needs. Application of the concept to water resources management
involves integrative components. Once sustainable goals have been defined and
adopted, sustainable infrastructure principles can operate. These principles must be
based on the best integrated technology and institutional capacity. The outcome
must then be evaluated and adapted if necessary in an iterative process. All activities
in sustainable development of water resources shall rely on basin wide
perspectives.
20.5 Specific Solutions for Alte Donau
Restoration and management measures for sustainable development of the urban

lake Alte Donau were extensively described and discussed in this volume. The key
measures and their results shall be briefly summarised here.
Excessive nutrient concentrations were strongly reduced by phosphorus floccu-
lation with ferric chloride and by sediment oxidation with calcium nitrate (Chap.
5 – Riplox treatment, Chap. 6). The efficacy of this treatment was predicted to be
10 years. Algal blooms disappeared and across the years phytoplankton composi-
tion shifted from blue-greens to green algae, diatoms and later on to chrysophytes
(Chaps. 9 and 10). Concomitantly the metazoan zooplankton altered from mainly
filter-feeding herbivorous cladocerans under eutrophic algal-turbid state to mainly
selective-feeding omnivorous and herbivorous copepods under mesotrophic
transparent-water state (Chap. 11). Assemblages of microzoans (rotifers, Chap. 11)
and protozoans (ciliates in Chap. 12, heterotrophic nanoflagellates in Chap.13) dif-
fered between treatment periods by lowered food supply (see also bacteria in Chap.
13 and discussion Chap. 11) or increasing grazing pressure.
With the reduction of the phosphorus concentration in the water column and
associated suppression of phytoplankton development, the transparency increased
significantly (Chap. 6) promoting an initial recovery of macrophyte-stands
(Fig. 20.2). But in the following years the macrophytes showed no significant
increase in biomass, even though regrowth was assisted by plantings. Figure 20.3
shows that the hysteresis of the year by year trajectory indicates considerable
Fig. 20.2 Annual changes of macrophyte biomass (BM) in t dry mass (bars) and phytoplankton
biovolume (B) as mg fresh weight per litre (line) from 1987 to 2011. Arrows with continuous line:
Riplox-treatment, arrow with dashed line: start of periodical water level lowering
Fig. 20.3 Trajectory of total phosphorus (TP) versus chlorophyll-a (Chl-a) in Alte Donau during
eutrophication and restoration (1987–2011). Years with macrophyte or algal domination are indi-
cated by white and grey background respectively separated by a line equivalent to Chl-a = 0.5TP. TP
and Chl-a as annual averages. Trophic delineations are indicated by dashed lines
resilience after perturbation. The starting point for intensive regrowth of macro-
phyte was given by an intervention into the hydrological regime of Alte Donau
(Fig. 20.2). From 2002 onwards every spring the water level was lowered in the
range of 15–30 cm to improve the light availability at the lake bottom at the begin-
ning of the growing season and to imitate partly the former natural hydrological
dynamic. The watermilfoil started to grow as well from planted areas as spontane-
ously in other lake areas. Other species that have been planted showed initially good
growth but were more and more suppressed by shading due to the dense stands of
Myriophyllum spicatum.
Seven years after the Riplox treatment the system switched back to the macro-
phyte dominated clear-water state. The response of macrophyte biomass to the
increase and subsequent decrease in TP concentrations is shown in Fig. 20.4. The
trajectory indicates large hysteresis in the loss and re-colonization of under-water
vegetation. Management plans to stabilise rehabilitated ecosystems need to take
hysteretic behaviour and return time into consideration.
Water quality in terms of total phosphorus and chlorophyll-a is now, 22 years
after the forward shift, better than it was back in 1987 (Fig. 20.3, details in Chap. 6),
but the quantity and composition of submerged vegetation is still quite different.
Fig. 20.4 Hysteresis in the decline and recovery trajectories of macrophyte biomass versus total
phosphorus concentration (TP) in the open water during the eutrophication and rehabilitation
phases between 1987 and 2011. Macrophyte biomass as in Fig. 20.2, TP as in Fig. 20.4
Macrophyte beds are dominated up to more than 90% by Myriophyllum spicatum,

whereas the ground-covering charophytes are under-represented.
The relative contribution of different plant groups to total macrophyte biomass is
documented for the period before eutrophication with macrophyte domination for
the year 1987 and for the years 1993, 2002 and 2015 following treatment (Fig. 20.5,
see also Fig. 20.2).
Myriophyllum spicatum is characterised by a very successful growth strategy
but the dominance was also triggered by additional effects. At the beginning of the
1990th the plant stands regularly died back in the winter months. By the end of the
1990th the Myriophyllum stands started to overwinter as green plants, possibly due
to a series of mild winters in the last decade that might be an effect of global warm-
ing. The impact of global change on Alte Donau was verified by the correspon-
dence between the climate signal (North Atlantic Oscillation Index) and water
temperature (WT) in winter and early spring, the increase of 1.52 °C per decade for
surface water temperature in April and the prolongation of warm period in summer
(Chap. 11).
Two years after full recovery of macrophytes and due to intensive photosynthesis
of macrophytes and also of phytoplankton (algal primary production see Chap. 10)
and low ground water influx especially into the main basin of Obere Alte Donau,
pH-values tended to rise up to 10 in the summer months. Water from the i mpoundment
100%
Relative Plant Mass
90%
80%
70% Myriophyllum spicatum
60%
50% Other low growing
species
40%
Myriophyllum/
30% Potamogeton
20% Stoneworts
10%
0%
1987 1993 2002 2015
Fig. 20.5 Relative shares of different plant groups in the years 1987 (last documented good status
before the eutrophication phase), 1993 (nearly total loss of macrophyte vegetation), 2002 (7 years
after the Riplox treatment at the starting point of the periodical water level lowering in spring) and
now (2015)
Neue Donau with a higher buffer capacity was used to exchange with water from
Alte Donau to overcome this problem. This measure was done regularly from 2006
up to now in summer or autumn, exchanging quantities of 1.5–4.5 Mio m3 within
several weeks. The buffer capacity in Alte Donau could thus be raised sufficiently
and the pH-values settled between 8 and 9.
A special soil filter, which is in use since 2016, was constructed in the northwest-
ern part of Alte Donau (Wasserpark) to find a sustainable solution to stock up the
main basin of Obere Alte Donau with calcium and to raise alkalinity. This soil filter
can be fed with water from Neue Donau. Phosphorus, suspended materials and
chlorophyll-a are reduced when passing the soil filter. The water from Neue Donau
is marked by higher calcium concentrations and by higher alkalinity than the water
of Alte Donau. But the special feature of the soil filter is, that the Ca content can be
stocked additionally by passing a reservoir that is filled with calcium carbonate
before entering the system of Alte Donau. The filter allows a constant discharge of
2500 m3 per day, which should be sufficient to raise buffer capacity of Alte Donau
permanently. This prototype of a soil filter was planned and built within an EU-Life
project (EU-Life 12 ENV/AT/000128).
Because of the overall dominance of the high-growing species Myriophyllum
spicatum intensive mowing is necessary to ensure bathing and other activities
(Chaps. 8 and 19). Mowing the macrophytes also enhances the availability of light
in deeper zones stimulating the growth of the low-growing vegetation. To support
the proliferation of stoneworts intensive plantings have been done (Chap. 8). In
contrast to helophyte planting, which turned out to be feasible with reasonable effort
(Chap. 18), planting and re-establishing dense stonewort stands was a great
Fig. 20.6 Macrophyte cover in the Alte Donau in 1987 after Löffler (1988) – light grey: charo-
phytes, dark grey: Myriophyllum ssp. and Potamogeton spp. – serving as objective of the macro-
phyte management
c hallenge. As well-documented by an automatic underwater video-trap (Chap. 8)

fish effectively hindered the growth of charophytes by grazing (amur and rudd) or
by digging out the new planted stoneworts (bream and carp, fish assemblages in
Chap. 15). In order to allow a successful re-establishment of charophytes in the Alte
Donau fish management has to be optimised. Charophytes can preserve a good
water quality equally well as the currently dominating high-growing vegetation
(Van den Berg et al. 1998), therefore the stoneworts are an ideal aquatic weed group
for water quality management of intensively used urban lakes. The conditions in the
Alte Donau in the 1980ies (Löffler 1988) can serve as a benchmark for a sufficient
cover of charophytes in a mesotrophic urban lake (Fig. 20.6).
10
Catch = 1.490 + 0.624 Chl-a
r2 = 0.815, p < 0.0001
2000 1998
8
Total fish catch [t]
6 2006
2002
2004 2008
4 2010
2012
0
0 5 10 15
-3
Chl-a [mg m ]
Fig. 20.7 Dependence of total fish catch (tons) on phytoplankton chlorophyll-a (mg m−3) for the
post-restoration years (1998ff.). Chl-a data from DWS-Hydro-Ecology, total fish catch data from
Austrian Fishery Association
In the time frame of the restoration process of Alte Donau a lot of efforts have
been made to trigger changes in the fish community. Structure of fish assemblages
was modified by biomanipulation in 1998 in ‘Kaiserwasser’ (Chap. 15) and changes
in fisheries management and practice. Predators like pike-perch, asp and later on
pike were intensively stocked to reduce the planctivorous and non-predatory fish
population (Chap. 15). Since macrophyte stands recovered pike is the main predator
in Alte Donau. Restructuring of the fish community promoted also changes in the
zooplankton assemblage (Chap. 11), stimulating the upgrowth of mainly copepods
and some large species of macophyte-habitat associated cladocerans.
As a result of a consequently performed fish management, total fish catches have
declined considerably and became significantly dependent on chlorophyll-a con-
centrations (r2 = 0.81, p < 0.001, n = 15, see also Chap. 15) since 1998, the post-
restoration period (Fig. 20.7.). A recent study of the fish biocenosis according to the
European Water Framework Directive methodology (Gassner et al. 2013) attested
the Alte Donau a “good ecological status” (Gassner et al. 2014). Since 2007 phyto-
plankton is also assessed according to the WFRD and the results also indicate the
“good ecological status”.
20.6 Conclusions
It can be concluded that suitable concepts and methods exist to establish and stabi-
lise a macrophyte-dominated state in shallow lakes. Successful re-introduction of
the aquatic macrophyte vegetation must be accompanied by adjusting the overall
nutrient balance and the protection of the young aquatic plants against the negative
influence of fish (herbivory, enhanced turbidity, nutrients) and waterfowl (herbiv-
ory, nutrients). Public awareness for the importance of ‘aquatic weeds’ has to be
raised because of the potential of macrophytes to compete successfully with phyto-
plankton. Using macrophytes to keep water bodies transparent may include macro-
phyte management and short term restrictions to water sports.
According to Heintz (2004) and Taylor and Goldstein (2010) the management of
water resources requires a new paradigm, the concept of sustainability. Restoration
of the eutrophied urban lake Alte Donau was achieved by the implementation of an
integrated management plan. Seven years after chemical treatment and after intro-
duction of biomanipulation and other ecotechnical measures a switch back to a
macrophyte dominated clear water stable state was observed. Routine monitoring of
water quality and hydrology was essential for fine tuning and performing of a con-
tinuous management. Macrophyte domination and associated with this – a good
water quality – could be stabilised for more than 20 years, demonstrating the quality
of our management. In case of the urban lake Alte Donau the concept of sustain-
ability is suitable and successful. Actually, in the frame work of the EU-Life proj-
ect – Urban Lake Alte Donau (Life 12 ENV/AT/000128), two plans are compiled
for maintaining and ensuring the “good ecological status”. The Integrated Lake
Management Plan will include all management measures and a plan for monitoring.
The Risk Management Plan will give advices how to reduce the vulnerability of the
ecosystem to effects like climate change and other anthropogenic pressures. This
plan will be made accessible to authorities in other cities and can serve as a model
for management of urban lakes in Europe.
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Subject Index
A Austrian standards, 109, 113, 125, 126

Accelerated circulation of matter, 164 Austrian Trophic State Index, 109
Accumulation, 50, 74, 159, 304, 357 Autumn, 63, 116, 117, 119, 132, 174, 185,
Aeration, 48, 52 187, 196, 198, 219, 220, 245, 293, 297,
Air-lift method, 300 324, 380
Algae, 4, 8, 46, 50, 51, 60, 61, 66, 109, 114,
131, 136, 137, 159, 165, 190, 191, 193,
216, 245, 266, 267, 279, 282, 283, 286, B
288, 289, 298, 377 Bacillariophytes, 117, 119, 122–124, 128, 136
Algal blooms, 7, 56, 150, 377 See also Diatoms
See also Bloom Backwater, 28, 43, 47, 53, 56, 62, 64, 66,
Algal classes, 71, 79–81 243–270, 276
Algivorous, 8, 172, 174, 220, 222 Bacteria, 50, 60, 78, 138, 190, 191, 214, 216,
Alkalinity, 58, 380 228–238, 245, 266, 332, 335, 377
Alternative stable states, 1–9, 53 Bacterial biomass, 191, 228–230
Ammersee, 113 Bacterial carbon, 231–233, 238
Ammonia, 57, 60, 77, 78, 129, 299 Bacterial cell numbers, 230
Anaerobic denitrification, 49 Bacterial production, 70, 79, 228, 229,
Angling, 194, 277, 280 231–233, 235, 238
Annual, 5, 6, 20, 29–33, 39, 43, 46, 60, 63–66, Bacterial secondary production (BSP), 156,
97, 111, 113, 117, 125, 126, 128, 130, 230, 231, 233, 234, 236–238
138, 150, 155–161, 164, 168, 169, 171, See also Bacterial production
179, 180, 184, 185, 187, 188, 192, 196, Bacterioplankton, 191, 216, 221
199–202, 216–219, 221, 229, 260, 262, Bacterivorous, 172, 174, 233
263, 323, 363, 377, 378 Baiting, 63, 64, 66
Annual cycle, 185 Bathers, 22, 63, 332, 335
Anthropogenic, 4, 6, 8, 11, 165, 363, 374, 383 Bathing, 66, 129, 331–333, 335, 336, 338,
Apatite, 75, 76, 82 358, 361–364, 366, 370, 380
Aquatic gastropoda, 279, 282 Benthic, 47, 73, 79–81, 199, 232, 243–270,
Aquatic vegetation, 133, 150, 376 283, 286, 297
Aquatic viruses, 228, 229 Benthic fauna, 258, 262, 264, 265
Aquifer, 4, 11, 17, 38 Benthic invertebrates, 73, 79, 243–270, 297
Asp, 277, 280, 281, 289, 290, 292, 293, 296, Benthos, 279, 281, 282, 286, 288
309, 382 See also Benthic invertebrates

https://doi.org/10.1007/978-3-319-93270-5
388 Subject Index
Bioindicators, 244, 259, 266 Carbon demand, 231–233

Biomanipulation, 11, 50, 52, 135, 164, 165, Carbon flux, 229
194, 202, 297, 382, 383 Carbon phytoplankton, 158, 159, 167
Biomass-macrophytes, 92–97, 104, 159, 181, Carbon ratio, 167, 180–181, 191, 193, 221
189, 192, 263, 269, 270, 288, 377–379 Carbon uptake, 150–153, 155, 157
Biovolume:TP relationship, 135, 137 Carbon zooplankton, 167
Bird faeces, 326 Carnivorous, 172, 174, 219, 261, 262, 278
Birds, 8, 50, 64, 66, 315, 317, 320–322, 324, Carp, 8, 25, 46, 52, 104, 108, 194, 277,
326, 332, 338, 374 280–283, 288, 298, 299, 381
Bistable theory, 158 Catchment, 11, 197, 375
Bivalves, 248, 257, 261 Catfish, 280, 289
Black-headed Gull, 315, 318 Chl-a, see Chlorophyll-a
Black Swan, 316, 324, 325 Cell diameter, 232
Bleak, 46, 50, 276, 278, 280–284, 286, Cell length, 232
288–294, 296, 297, 308, 309 Cell volume, 232
Blooms, 4, 7, 42, 48, 56, 60, 62, 66, 118, 119, Charophytes, 46, 52, 57, 92, 97, 101–105, 117,
129–132, 136, 150, 152, 165, 167, 181, 122, 124, 131, 135, 192, 193, 256, 368,
189, 191, 260, 335, 360, 374, 377 379, 381
Blooms of algae, 56, 150, 275, 377 Chemical phosphate precipitation, 93, 109,
of cyanobacteria, 46, 48, 107, 129–132, 110, 125, 166, 180, 181, 190, 249, 257,
136, 152, 165, 189, 260, 275, 335, 360 261, 263, 266
of phytoplankton 7, 60, 62, 66, 118, 167, Chemical treatment, 57–60, 62, 66, 79, 119,
181, 191, 374 164, 166, 167, 171, 175, 176, 180, 189,
Blue-greens, 46, 60, 66, 81, 377 192, 216, 229, 259–262, 268–270, 383
See also Cyanobacteria Chironomids, 114, 244, 245, 248, 256–264,
Boats, 22, 24, 82, 90, 291, 300, 304–306, 266–270, 279, 281, 282, 299
308, 309, 338, 347, 356, 358, 361–364, Chl-a:TP relationship, 135
369, 374 Chloride, 58, 59, 62–65, 78, 257, 258, 260,
Body-length, 174, 177, 303 261, 264, 267, 278–288, 296, 377
Bottom feeding, 297 Chloride balance, 62
Bottom-up control, 109, 137, 164, 190, 191, Chlorophyll-a, 5, 6, 9–11, 46, 50, 52, 60–62,
195, 200, 202 66, 70, 71, 79, 80, 108, 110, 111, 113,
Boundary conditions, 36, 38, 43 117–119, 122–127, 130, 133–138,
Breeding pairs, 325 150–158, 160, 161, 165, 190, 195, 200,
Brettum index, 113–114 221, 222, 267, 294, 295, 378, 380, 382
Bryozoans, 259 content, 68, 80, 122–126, 135, 136
BSP, see Bacterial secondary production Chlorophyll:TP, 111, 113
Budget Chlorophytes, 81, 117, 119, 122–124, 126,
nutrient, 4 136
water, 43, 374 Chroococcales, 112, 122, 131
Buffer capacity, 36, 57, 58, 380 Ciliates, 188, 191, 214–222, 234, 245, 377
abundance, 215, 216, 219, 221, 222
biomass, 128, 216, 219, 220
C community 214–217 (see also (Species
Ca(NO3)2, 49, 58, 60, 70, 76, 81, 83, 166 spectrum))
Calanoida, 172 fresh weight 216, 219, 221, 246, 377
Calanoids, 173, 175, 176, 185, 187, 189, 190, Cladocera, 172, 279, 281, 289
200–202 Cladoceran-rich community (cladoceran-rich
See also Calanoida assemblage), 180, 189
Calcium, 21, 51, 58, 59, 65, 70, 73, 74, 76, 77, Cladocerans, 46, 50, 138, 170–177, 179, 185,
81, 82, 260, 268, 377, 380 187–194, 198, 200, 202, 377, 382
biogenic, 51, 58 See also Cladocera; Copepoda
Calibration data set, 113, 114 Clear water condition, 93
Subject Index 389
Clear water phase, 119, 133, 167–169, 315, 317, 319, 321, 322, 331, 337,
183–185, 198–200 356–358, 367–369
Climate Daytime refuge for zooplankton, 193
change, 133, 165, 196, 198–201, 383 (see Decapoda, 279, 282, 283
also (Global warming)) Decoupling, 136
research, 132 Deep lake, 7, 10, 35, 131, 132, 135, 266, 374
signal, 164, 168, 183, 184, 195–197, 379 Deterioration, 90, 165
warming, 27, 163–202, 375 Detritus, 256, 269, 279, 282, 283, 286, 288,
Compositional changes, 123, 262, 263 298
Compositional changes of chironomid Diatoms, 49, 50, 60, 62, 66, 79, 109, 112, 116,
assemblage, 259, 270 117, 128, 131, 137, 377
Compositional shifts, 109, 118, 119, 126, 132, Dietary composition, 199, 286
134, 136, 137, 190, 192, 261, 264, 267 Dissolved organic carbon (DOC), 62, 71,
Concepts, 3–11, 164, 228, 306, 358, 361, 362, 79, 233
368, 376, 383 Dry weight, 51, 70, 72, 74, 76, 81, 83, 90, 92,
Conductivity, 58, 65 93, 128, 138, 158, 189, 192, 246, 263
Consumer, 134, 190, 244, 288, 298 Dump site, 21, 43, 46, 48, 357, 358, 360
Contamination, 21, 333
Continentality, 30
Copepoda, 172, 279, 281 E
Copepod-rich community (copepod-rich Echo-sounding, 90, 93
assemblage), 180, 194 Ecological measures, 309
Copepods, 138, 170–177, 179, 180, 185–189, Ecological relevance (of bird habitats,
192, 194, 198, 201, 202, 377, 382 stretches), 131, 315, 322
Council directive, 333, 356 Ecosystem degradation, 165
Crustaceans, 46, 166, 174, 177–179, 189–190, Enclosure-experiment, 198, 289–293,
192–194, 198, 201, 202, 279, 281 296–297, 309
Cryptophytes, 52, 112, 117, 120, 122, 123, Energy efficiency, 150
127, 128, 131, 136, 137 Enzyme activity, 131, 191
Cumulative Plant-Mass Index (CMI), 347, 348 Ephemeroptera, 251, 257, 262, 281
Cyanobacteria, 6, 48–52, 109, 112, 114–117, Epiphytes, 298
119, 122–124, 126, 127, 129–132, 136, Equilibrium, 7–10
137, 152, 165, 189, 190, 193, 197, 216, Euglenophytes, 120, 128
267, 268, 283, 335, 360 Euphotic zone, 57, 150, 152, 155, 161
bloom, 4, 129, 136, 165 (see also Eurasian Coot, 317, 320, 323, 324
Blooms) European Eel, 280, 281
Cyclopoida, 172 Eutrophic, 46, 75, 113, 126, 127, 134, 152,
Cyclopoids, 164, 173, 175, 185–187, 189, 198, 154, 155, 174, 175, 179, 189, 229, 233,
200–202 248, 261–264, 288, 376, 377
See also Cyclopoida See also Eutrophied; Eutrophied period
Cylindrospermopsis raciborskii, 46, 116, 117, Eutrophication, 4, 35, 46–48, 53, 55–57, 60,
122, 127, 129–132, 152, 165, 189, 260 62, 66, 75, 77, 108–110, 129, 133, 150,
Cyprinid(s), 46, 194, 202, 281, 283, 286, 288, 165, 166, 171, 179, 189, 245, 257, 258,
289, 298, 299, 307, 309 260, 267, 269, 277, 373–383
Cyprinid-dominated, 165, 278 Eutrophied, 108, 113, 119, 122, 123, 125, 130,
171, 216, 229, 383
lake, 113, 130, 229
D period, 122, 123, 125, 216
Daily intake, 292, 296 Evaporation, 32, 33, 36–39
Danube River/Donau/Danube region, 5, 9, 10, Experiments
17–22, 24, 28, 32, 33, 36, 37, 40, 43, lake, 90, 91
56, 59, 89, 110, 128, 133, 165, 171, macrophyte, 94–96
188, 197, 214, 243–270, 276, 290, 314, Extremely hot summer (period), 184
390 Subject Index
F Freshwater jellyfish, 183, 184, 197–198

Faecal pollution, 333–335 Freshwater medusa, 164, 184
Fast-growing, 200 Functional phytoplankton groups, 109,
Feed analyses, 284 111, 117
See also Dietary composition; Food,
components; Food, intake; Food,
spectrum; Nutrition spectrum G
Feeding, 50, 191, 197, 214, 261, 269, 276, Generation time, 214, 232, 234, 267
278, 288, 296–298, 309, 318, 320, 322, Geology, 17–18, 55
324–326, 332, 333, 335, 377 Gill net, 278, 279, 291
Ferric chloride (FeCl3), 49, 58, 59, 70, 74, 75, Global warming, 129, 165, 166, 195–202, 379
81, 82, 166, 377 Grass Carp, 8, 52, 280, 281, 299
Filament(s), 112, 129–131 Gravel, 17, 20, 21, 262, 263, 299, 341, 342
Filamentous, 50, 60, 66, 112, 122, 129–132, Grazing, 8, 46–48, 104, 191, 194, 195, 198,
165, 189, 286 202, 228, 229, 233–236, 238, 286, 288,
See also Filament(s) 377, 381
Filamentous algae, 279, 283, 286 Grazing pressure, 46, 48, 194, 195, 202, 377
Filamentous cyanobacteria, 50, 122, 129–132, Green algae, 46, 50, 51, 60, 66, 109, 131,
189, 283 137, 377
Filter-feeding, 138, 189, 190, 202, 262, 377 Grey Heron, 314, 316, 325
Fine sediment, 258, 267, 269, 283, 299 Groundwater
Fish flow, 4, 20, 21, 37, 38, 40–43, 130
age, 298 input, 4, 28, 38
assemblages, 165, 194, 275–309, 381, 382 seepage, 3, 28, 33, 36, 130, 133, 150, 245
catch(es), 108, 138, 194, 195, 277, 382 seepage lake, 3, 130, 150, 245
community, 46, 191, 277, 309, 376, 382 Growth, 4, 8, 40, 42, 50–52, 57, 58, 60, 65, 66,
eggs, 276, 307 79, 90, 91, 93, 94, 96, 103, 104, 108,
exclosure, 52 109, 111, 119, 122, 128–137, 150, 165,
re-introduction, 383 167, 169, 188, 190–193, 198–201, 214,
stocking, 108, 138, 191, 194, 276, 277, 296 229, 234, 248, 267, 270, 278, 289, 296,
survey, 194, 278 297, 299, 351, 374, 376, 378–382
Fish species, 8, 191, 276, 278, 281, 282, 284, Growth rate, 50, 96, 111, 117–119, 130, 137,
288, 289, 292, 296–298, 303, 304, 308 167, 180–181, 200, 232, 234, 296
See also Species spectrum Gulls, 315–318, 321, 322
Flocculation, 11, 49, 52, 70, 74, 82, 109, 134,
261, 266, 377
Flood control, 374 H
Flood plain, 9, 10, 17, 18, 42, 43, 188, 256, Habitat change, 192–194, 248, 254, 266
265, 268 Habitat quality score, 244, 248
Flood protection, 18–20, 265, 314, 315 Habitats across altitudes, 265, 270
Flood water, 5, 32, 36–38, 43 Helophytes, 90, 338, 348–351, 353, 380
Flow-velocity, 193, 269 Heterocytes, 129
Foliage, 63, 64, 66 Heterotrophic bacteria, 50, 229–238, 267
Food Heterotrophic nanoflagellates (HNF), 229,
availability, 189, 197, 198, 200, 202, 216 232–235, 238, 377
chain, 108, 228, 288, 296, 297 Hibernating waterfowl, 315
components, 279, 281, 282, 286 High-growing macrophytes, 135
intake, 284, 286, 296, 298 Historic development, 17–26
particles, 189, 278, 279, 281, 286 History, 46–48, 55, 56, 133, 165, 166, 258,
quality, 190, 191, 216 276, 315, 337, 356
source, 214, 222, 233, 244, 299, 324 Hydracarinae, 279
spectrum, 286, 287 Hydraulic barrier, 39
Freespace-and water management, 360, 361, Hydroelectric power plant, 20–21, 314, 315,
364, 369 318, 322
Subject Index 391
Hydrological conditions, 36, 37 phenology, 119, 132, 133, 198, 201

Hydrological data, 36, 37 quality, 374
Hydrology, 9, 36, 55, 110, 166, 188, 383 recover, 134
Hygienic monitoring, 332–336 restoration, 6, 108, 109, 120, 129, 138,
Hymenostomes, 215 164, 165, 169, 171, 177–179, 189–195,
Hypertrophic, 10, 107–138, 278, 296 338, 375, 376
Hysteresis, 10, 134, 192, 196, 198, 377–379 shallow, 5, 7, 9, 10, 56, 60, 65, 129, 130,
150, 188, 193, 194, 197, 199, 360,
376, 383
I shore, 349
Illumination of littoral sediments, 263 urban, 3, 8, 11, 17–26, 33, 36, 89, 129,
Improvement, 19, 36, 52, 74, 156, 159, 229, 134, 138, 150, 160, 161, 164, 165, 214,
266, 297, 336, 338 351, 360, 370, 373–383
In-and outflow, 3, 28, 36, 39, 40, 42 Large-celled, 136
Increased public access, 361 Leaves of the riparian vegetation, 279
Indicator organism, 333, 334 Leisure time activities, 22–26
Infected bacterial cells, 238 Lengthening of the warm period, 183
Ingestion rates, 233, 279, 284 Life-span, 193, 198, 267
Input, 39–43, 63–66, 75, 133, 264, 288, 297, Light, 9, 47, 50–52, 79, 81, 91, 93, 94, 96,
332, 335, 338, 360, 363, 368 102, 110, 128, 130, 131, 134–137,
Insect larvae and pupae, 279 150–152, 154–157, 160, 161, 260, 289,
Internal loading, 11, 50, 70, 74 378, 380, 381
Interstitial water, 71, 74–79 Light harvesting, 136, 155, 160
Intestinal content, 279 Limitation, 9, 47, 61, 129, 228
Intra-annual, 164, 180, 187, 199–202 Littoral, 174, 175, 244, 376
Invertebrate abundance, 255 Littoral sediment, 264
Invertebrate biomass, 244, 249, 269, 270 Littoral zone, 47, 50, 337–353
IPCC, 165, 195 Load, 5, 63, 64, 66, 134, 190, 267, 374
Iron content, 70, 72–74 Loading
Iron-III-Chloride treatment, 257, 258, 261, external, 200
264, 278–288 internal, 11, 50, 70, 134, 190
See also Phosphate flocculation; Phosphate Lobau, 43, 62, 64, 66, 256
precipitation; Riplox treatment Loire, 188
Long-term changes, 55–66
Long-term development, 29, 119–122, 154,
J 165, 169, 180–187
Juvenile fish, 278, 279, 298 Long-term re-settlement of sediment, 267, 268
See also Re-colonization of sediment
Long-term trend, 39, 58, 62, 65, 168
K Loss on ignition, 70, 72, 73, 76, 77, 81
Kaiserwasser (KW), 29, 52, 70, 80, 191, Lowering water level, 26, 96, 97, 103, 104,
194, 289–291, 293–295, 297, 299, 309, 377, 380
333, 382 Lowland habitat, 269
Kohler-Index, 300
Kohler method, 300
M
Macroinvertebrates, 53, 257–258
L Macrophyte
Lake biomass, 90, 96, 101, 135, 304, 306, 308
assessment, 114, 132 decrease, 47, 48
biomanipulation, 135 domination, 8, 9, 52, 55–57, 59–61, 66,
groundwater, 3, 4, 48, 150 379, 383
management, 3, 7, 45–53, 92, 96, 109, experiments, 94–96
122, 133, 137, 138, 166, 174, 180, 181, growth, 40, 46, 51, 52, 57, 58, 65, 90, 93,
200, 383 374, 376
392 Subject Index
Macrophyte (cont.) Mowing, 51, 52, 135, 193, 276, 304–306, 308,
planting, 52 309, 380
recovery, 62 See also Underwater weed cutting
re-establishment, 110, 123, 124, 166, 169, Mowing boat, 304–306, 308, 309
171, 174, 175, 180, 181, 192, 244, 254 Mud, 49
underwater vegetation/plants, 128, 138, Multifunctional and nature-near waterfront,
164, 165, 181, 192, 193, 244, 260, 263 369
Macrozoobenthos, 109, 245, 247, 260–265, Mussels, 262, 264, 269, 283
269, 270 Mute Swan, 315, 316, 318, 320–323
abundance, 260, 261, 263
biomass, 244, 263–265, 270
Magnesia, 58 N
Main periods of (the ecosystem-based) lake N2, 129
management, 166 See also Nitrogen
Maize, 279, 282, 283, 288 Nanoplankton, 214, 235
Mallard, 315–317, 320, 321, 323 NAO, see North Atlantic Oscillation
Management, 4, 5, 7, 11, 20, 26, 40, 43, Neozoa, 324
45–53, 62, 90, 92, 93, 96, 102, 105, Nests, 298, 325
108, 109, 122, 129, 133–135, 137, 138, Net growth rate, 111, 117–119, 137, 167, 178,
161, 165, 166, 169, 174, 175, 179–181, 180
188, 200, 203, 229, 276, 277, 288, 296, Net growth rate of zooplankton, 167, 180
297, 304, 308–310, 314, 326, 333, 335, Neue Donau, 5, 10, 20, 21, 23, 24, 32, 36–38,
338, 345, 353, 355–370, 373–383 40, 42, 43, 47, 49–52, 58–60, 62–66,
Marsh plant nursery, 346 75, 110, 133, 171, 184, 188, 314–323,
Marsh plants, 52, 90, 341, 345–348 331, 332, 380
Masterplan Alte Donau, 358–362, 364, Nitrate, 11, 49, 60, 66, 78, 129, 134, 244, 260,
367–369 268, 299, 377
Mean Mass Index (MMT), 92, 101, 102 Nitrate oxidation, 49, 134, 166
Mean zooplankton body size, 180, 192 Nitrite, 60, 78
Meso-eutrophic, 156, 259, 261 Nitrogen, 4, 49, 60, 66, 78, 81, 112, 129, 133,
Mesotrophic, 51, 60, 66, 78, 92, 107–138, 152, 137, 228
156, 165, 189, 202, 222, 233, 244, North Atlantic Oscillation (NAO), 160, 168,
257–259, 261–264, 266–268, 270, 169, 181, 184, 196, 379
377, 381 North basin, 110, 119, 120, 128, 166,
Mesozooplankton, 192, 200 169–171, 188, 229, 314
Metalimnetic, 131 See also Obere Alte Donau (OAD)
Meteorological forcing, 196 Northern Pike, 280, 283, 284
Microbial carbon flux, 229 Nostocales, 112, 122, 129
Microbial loop, 228, 229 N/P ratio, 50
Microflagellates, 228 N-rich, 134, 190
Microzooplankton, 172 Numeric model, 38, 40
Midsummer decline, 185, 187 Nutrient(s), 3–11, 21, 36, 37, 46, 48–50, 58,
Model, 4, 6, 7, 10, 37–43, 63, 90, 150, 157, 60–65, 74, 77, 81, 103, 108, 109, 122,
199, 229, 232, 364, 369, 374, 383 125, 128–131, 133–135, 150, 164, 165,
ecological, 7, 36, 56, 90, 113, 114, 131, 168, 171, 180, 189–191, 193, 199, 200,
132, 214, 228, 229, 255, 267, 268, 214, 216, 221, 222, 228, 264, 266, 269,
276–278, 308, 309, 315, 322, 338, 362, 276, 288, 298, 299, 308, 326, 332, 338,
364, 376, 382, 383 360, 374, 375, 377, 383
hydrological, 10, 20, 27, 28, 35–43, 47, 59, availability, 8, 130, 200, 276
62, 110, 130, 188, 268, 358, 362, balance, 36, 62, 383
368, 378 concentrations, 37, 48, 50, 103, 133, 377
Mollusca, 270, 279, 282, 283, 286, 287 element, 109, 135, 164, 190, 255
Morphometry, 96, 214, 229, 337 enrichment, 108, 150, 165, 255, 269
Subject Index 393
input, 4, 21, 42, 48, 133 Phosphorus, 4–6, 9–11, 43, 46, 49, 50, 57,
loading, 3–11, 46 60–66, 70, 71, 74–76, 81, 82, 92,
spectrum, 286 108–111, 114, 119, 122, 123, 128, 130,
Nutrient-rich, 4, 98, 130, 164, 199, 216 131, 133–135, 137, 138, 152, 169, 174,
See also Nutrient, enrichment 178, 180, 181, 189–192, 194, 199–202,
216, 219, 228, 230, 248, 256, 257, 266,
267, 277, 278, 374, 375, 377–380
O acquisition, 131
Obere Alte Donau (OAD), 57, 58, 62–66, 70, balance, 62, 63, 66
72–74, 77–80, 110, 229–231, 233, 235, (or nutrient) pool size, 50, 164, 199
236, 314, 318, 379, 380 release, 49, 70, 82 (see also Nutrient
Ochrophytes, 117, 119, 122, 123, 127, 128, element)
131, 132, 136 Photoacclimation, 136, 137
Oligochaets, 244, 248–250, 258, 260–263, Photosynthetically produced carbon, 232
267–269, 281, 282 Photosynthetic pigment, 135, 136
Oligotrichs, 215, 216, 219 pH-values, 21, 51, 52, 57, 58, 70, 76, 379, 380
Oligotrophic, 5, 10, 113, 125–127, 194, 199, Phycobilines, 136
222, 248 Physico-chemical coniditions, 55–66
Omnivorous, 138, 172, 174, 189, 202, Phytobenthic biomass, 79–81
214, 377 Phytoplankton, 4, 6, 8–11, 46, 47, 49–51, 56,
ÖNORM, 109, 113, 126, 362 57, 59–62, 66, 74, 90, 92, 107–138,
Open-air bath, 331, 332 149–161, 164–169, 179–181, 184, 185,
Optimal growth, 130, 169, 184 188–191, 193–196, 198–202, 216, 219,
Ostracoda, 254, 279, 282 221, 222, 228–232, 244, 260, 266, 267,
Output, 37, 38, 40, 62–66 269, 278, 283, 288, 295, 299, 374, 376,
Oxbow, 21, 43, 89, 108, 110, 129, 130, 165, 377, 379, 382, 383
166, 188, 189, 195, 245, 257, 260, 270, assemblage metric, 108, 113, 114, 126,
318, 337, 376 127, 132
Oxbow lake, 21, 32, 43, 108, 110, 129–132, biomass, 9, 51, 113, 124–126, 135, 136,
165, 166, 188, 189, 195, 245, 257, 164, 165, 169, 180, 181, 189, 190, 216,
260, 269 221, 244, 267, 270, 295
Oxygen content, 57, 267 bloom, 60, 62, 66, 118, 181, 191, 374
Oxygen deficit, 50, 75 development, 108, 109, 117–122, 126, 128,
132, 184, 198, 377
net growth, 167, 180 (see also (Net growth
P rate))
Paradigm, 376, 383 structure, 120
Particulate phosphorus, 60 yield, 124, 130, 133–138, 169, 185,
P/E curves, 150, 151 190, 202
Pelagic ciliates, 214, 215, 218 Phytoplankton biovolume (PB), 6, 11, 108,
Pennate diatoms, 112, 117, 128, 131 110–114, 116–131, 133–137, 152, 166,
Periods of lake management, 57–61, 122, 167, 190, 191, 377
174, 181 Phytoplankton:chl-a relationship, 108, 111,
Phosphate flocculation, 261 123–126, 135, 137, 138
Phosphate precipitation (chemical), 93, 109, Picoplankton, 115, 214, 233
110, 125, 166, 169, 179, 181, 189, 190, Pike, 25, 46, 50, 277, 279, 280, 283, 284,
199, 201, 244, 249, 257, 260–261, 263, 288–293, 296–299, 309, 382
266, 267, 269 Pike-Perch, 45, 277, 279, 280, 283, 284,
See also Iron-III-Chloride treatment; 288–293, 296, 297, 309, 382
Phosphate flocculation Piscivorous fish, 194, 292
Phosphor binding capacity, 73, 77, 82 Planktivorous fish, 193, 194, 199, 278, 288,
Phosphor release, 49, 70, 82 289, 297
394 Subject Index
Plankton (planktonic), 4, 8, 50, 82, 109, 110, Profundal, 244, 262, 263
114, 119, 128, 129, 132, 134, 137, 138, Prolongation of the period with an optimum
159, 165–167, 175, 188, 190, 191, 193, temperature, 184
195, 197, 199, 200, 219, 231, 267–269, Prostomatids, 215, 216, 219–221
278, 281, 283, 286, 289, 294, 297 Protection zones, 356, 364
Plant, 4, 7, 8, 11, 19, 20, 52, 63, 90–97, Protozoa, 214, 221, 222, 232, 267, 377
100–103, 110, 133, 135, 150, 183, 192, Pumpkinseed, 280, 283, 303–305, 308
258–260, 270, 276, 279, 283, 286, Pupal exuviae of chironomids, 244, 245
298–300, 302–309, 314, 315, 318, 322,
338, 341, 345–351, 353, 367, 374, 376,
379, 380, 383 Q
group, 97, 101–103, 347, 379, 380 Quantitative protargol stain (QPS), 215
height, 91, 93–96
mass, 91, 92, 101–103, 347, 348, 350, 351,
353, 380 R
Planting measures, 103, 348–351 Radiation, 33, 150–152, 154–156, 160, 161,
Plant mass index (PMI), 91, 92, 347 196, 269
P-limitation, 190 Rates
Polymictic, 29, 33, 56, 65, 110, 130, 133, 138, chlorophyll specific, 150, 152
150, 160, 166, 197, 198, 201 photosynthetic, 150–152, 154–157,
Polyp stage, 184, 197 160, 161
Pre-alpine, 113, 198 Recolonisation, 57, 58, 60, 66
Precipitation, 20, 28, 30–33, 36–39, 51, 58, Re-colonization of sediment, 245
62, 63, 65, 66, 70, 74, 76, 93, 109, 110, Recovery, 8, 11, 23, 50, 52, 56, 57, 59, 61, 62,
119, 122, 123, 125, 134, 137, 166, 169, 70, 122, 133–135, 152, 161, 177, 266,
174, 180, 181, 189–191, 196, 199, 201, 374, 376, 377, 379
202, 216, 244, 249, 257, 260–261, 263, Recreation, 20, 21, 23, 25, 36, 46, 89, 96, 103,
264, 266, 267, 278, 374, 375 108, 135, 164, 165, 245, 276, 305, 309,
Predation, 199, 228, 234, 288 326, 331–338, 356–358, 360–364,
Predator, 50, 193, 194, 198, 202, 234, 261, 367–369, 374, 375
262, 278, 288–292, 297, 298, 308, Recreational activities, 36, 96, 103, 276
309, 382 Recreational water, 331–336
Predatory fish, 104, 108, 109, 138, 191, 277, Reduced ressources, 359
279, 284, 288, 289, 296–298, 382 Reed belt, 50, 134, 299, 366
Predatory pressure, 297 Reed planting, 338, 341
P reduction, 190 Reed species spectrum, 348
Presence of invertebrates, 249 Refugia for zooplankton, 299
P-rich, 134, 190 Regime shift, 6, 8, 10, 151, 160, 161, 198, 199
Primary producer, 8, 108, 134, 137, 191, Rehabilitation, 10, 52, 154, 160, 375, 376, 379
200, 267 Relative Plant Mass (RPM), 91, 92, 97, 101,
Primary production, 57, 72, 74, 82, 150, 348, 350, 351, 353, 380
155–161, 228, 231, 232, 379 Resilience, 7, 10, 35, 50, 52, 53, 103, 192,
Pristine fish community, 277 374, 378
Production Resilience phase, 50, 192
annual, 155 Resting and sleeping place, 315
carbon, 158–160, 232, 238 Restoration, 3–11, 45–53, 55–57, 59–62, 66,
lake, 150, 159, 160 70, 73, 81, 108–111, 119–122, 125,
macrophyte, 158, 159, 232 126, 129–135, 138, 150–152, 154–157,
primary, 57, 72, 74, 82, 150, 155–161, 228, 159–161, 163–197, 214, 216, 219, 229,
231, 232, 379 243–270, 314, 326, 335, 337–353,
submersed, 159 367–370, 374–378, 382, 383
water column, 150, 153, 160 Restoration plans, 51
Productivity, 150, 152, 153, 161, 289 Rhizomes of reed, 341
Subject Index 395
Riplox (treatment), 49, 50, 52, 60, 66, 70, Shoreline, 29, 35, 89–91, 93, 291, 332,
73–77, 79, 81–83, 92, 93, 109, 110, 333, 337, 338, 341, 345, 347, 348,
120–123, 126, 127, 133, 134, 138, 157, 351, 374
170, 174–176, 179, 190–192, 202, 214, Shore zone, 52, 90
216, 219, 229, 244, 249, 255, 267–269, Short-term development, 60
278, 335, 377, 378, 380 Short-time habitat destruction, 267
River, 9, 17–27, 29, 32, 33, 36, 42, 43, 56, 59, Silica, 62, 137
89, 110, 128, 132, 133, 164, 165, 188, Small-size, 216
193, 214, 215, 245, 258, 260, 262, Snapshot sampling, 248
264–266, 269, 276, 314, 320, 331, 337, Soluble reactive phosphorus, 61, 74
341, 356, 357, 360, 362, 365, 369 Soluble unreactive phosphorus, 60
Riverine, 21, 109, 130, 133, 188 South basin, 110, 119, 120, 128, 166, 170,
River regulation, 276, 365 171, 188, 229, 314
Roach, 46, 276, 278, 280–286, 288, 297–299, See also Untere Alte Donau (UAD)
303, 304, 308 Spawning area, 299, 308
Robustness, 198, 199, 376 Species composition (of phytoplankton), 46,
Rotifers, 164, 166, 170–174, 176, 178, 185, 56, 103, 104, 108, 114, 127, 128, 136,
187, 188, 191, 192, 198, 200–202, 219, 137, 155, 165, 172, 188, 194, 215, 259,
279, 281, 377 261–263, 266, 268, 289, 326, 347
Rudd, 46, 104, 276, 280, 297–299, 308, 381 Species spectrum, 90, 96–98, 103, 104, 298,
Ruffe, 280, 281, 297, 303, 304, 308 345–347, 351
Sperrbrunnen, 42, 62
Spontaneous growth, 134, 192
S Spring, 5, 26, 40, 49, 51, 52, 63, 75, 108,
Sand, 193, 262 116–119, 132, 133, 164, 167, 168,
Saturation, 9, 136, 150, 152, 154–156, 161, 183–185, 187–189, 196–202, 219, 220,
268 256, 293, 294, 297, 341, 378–380
Scuba diving, 90, 91, 245 Spring bloom (phytoplankton), 119, 167, 199
Seasonal cooling, 196 Spring peak, 187, 199, 201, 202
Seasonal development, 108, 116, 117, 185, Standing crop, 75, 128, 135, 150, 191,
197 192, 232
See also Seasonality Standing stock (of bacteria), 233, 234, 238
Seasonal hysteresis, 196, 198 Stream ecology, 270
Seasonality, 58, 108, 117, 153, 181, 185, 187 Stomach and intestinal content, 279
Seasonal variation, 235, 236 Stomach content, 279
Seasonal warming, 196, 201 Strategy, 4, 96, 131, 288, 296, 361, 369, 379
Secchi depth, 46, 51, 52, 60, 61, 66, 93, 129, Stream ecology, 270
138, 165, 167, 178–180, 192, 195, Submerged plant biomass, 90
260–263, 288, 289, 295, 309 Submerged plants, 7, 90, 133, 258, 298, 300,
Sediment, 11, 17, 48–50, 52, 57, 58, 62, 308, 374
69–83, 134, 137, 166, 190, 232, 244, See also Underwater vegetation;
256, 258–264, 266–270, 279, 282, 283, Macrophytes; Underwater plants
286, 287, 299, 300, 356, 374–377 Submerged vegetation, 90, 93, 103, 109, 120,
Seepage, 3, 4, 20, 28, 33, 36, 40, 43, 130, 133, 134, 165, 181, 192, 202, 260, 263, 270,
150, 245, 374 298, 299, 308, 378
Selective-feeding, 138, 189, 202, 377 Sulfate, 58, 65, 78
Selective fish stocking, 46 Summer, 8, 23–25, 30–33, 38, 42, 46, 49, 52,
Sewage, 46, 133, 332, 338, 374 56–58, 63, 65, 90, 93, 108, 111, 113,
Shallow lake, 5, 7, 9, 10, 56, 60, 65, 129, 130, 116–119, 122, 125, 126, 129–133, 138,
150, 188, 193, 194, 197, 199, 360, 151, 164, 167–169, 183–185, 187, 189,
376, 383 196–198, 201, 202, 214, 219, 220, 245,
Shift 256, 268, 286, 289, 295, 335, 357, 360,
backward, 11 379, 380
forward, 378 depression, 185, 187, 201
Shore area, 337, 341, 350 warming, 197
396 Subject Index
Surface water temperature, 27, 29, 30, 33, 156, U

161, 164, 169, 183, 202, 379 Underwater-floodlight, 90, 91, 93
Sustainable, 82, 137, 164, 297, 309, 338, Underwater light, 47, 81, 131, 135, 152, 260
373–383 Underwater mowing, 193
Synechococcales, 112, 122, 131 See also Weed cutting
Underwater plants, 4, 90, 376, 383
Underwater vegetation, 128, 138, 164, 165,
T 181, 192, 193, 244, 260, 263
Taxa of macrozoobenthos/species inventory, Underwater weed cutting, 96, 103, 383
264 Unicellular centric diatoms, 112, 116, 117
See also Species composition; Species Untere Alte Donau (UAD), 62–66, 70, 72–74,
spectrum 78–80, 82, 110, 229–231, 314, 318
Technique Upgrading urban waterfronts, 359
carbon, 150 Urban development, 19–21, 23, 55, 357, 359,
oxygen, 150 360, 362, 365
Temperature, 9, 27–33, 37, 56, 57, 65, 130, Urban groundwater-seepage lake, 3, 4, 33,
131, 154–156, 161, 164, 165, 168, 169, 130, 150, 245
181–187, 195–198, 200–202, 229, 236, Urban lake, 3, 8, 11, 17–26, 33, 35, 36, 89,
270, 284, 286, 289, 324, 335, 379 129, 134, 138, 150, 160, 161, 164, 214,
air, 29–33, 56, 65, 195 351, 356, 360, 370, 373–383
increase, 164, 165, 197, 200–202
thermophilic species, 201
water, 27–33, 56, 57, 65, 155, 156, 161, V
164, 168, 169, 181–186, 200–202, 236, Vascular plants, 11, 150
284, 286, 289, 335, 379 Vertical profiles, 77, 78, 150, 151
Three-step investigation, 360, 368 Viral abundance, 236, 238
Top-down control, 8, 109, 190, 191, 194, 199, Viral burst size, 235–238
234, 235, 288 Viral counts, 237
Top-down-hypothesis, 288, 289, 296 Viral decay rate, 237, 238
Total nitrogen (TN), 60, 66, 133 Viral loop, 227–238
Total phosphorus (TP), 5, 6, 9–11, 50, 52, 60,
61, 66, 74–77, 108, 111, 113, 114, 122,
123, 125–127, 131, 133–135, 137, 138, W
164, 178, 181, 188–192, 194, 199, 230, Warming, 27, 129, 163–203, 375, 379
267, 378, 379 Wasserpark, 21, 36–41, 62, 66, 314, 317,
Toxins (cyanobacterial toxins), 129 320–322, 324, 325, 333, 380
Transparency, 46, 48, 50, 60, 61, 66, 90, 92, Water
93, 103, 109, 133, 152, 165, 167, 179, balance, 5, 28, 36–39, 41, 43
188, 190, 195, 244, 260, 263, 267, 269, clear, 299
288, 309, 377 content, 70, 72, 81
Traunsee, 152, 220–222 draw-down, 188
Trawl net, 279, 297 exchange, 52, 62–65, 79, 155, 171, 188
Triangular graphs, 168 fowl, 47, 50, 276
Trichoptera, 251, 257, 258, 260, 262, 281, 282 hygiene, 335
Trophic change, 107–138, 189, 192, 199, 201 level, 4, 8, 20, 21, 26, 37–43, 47, 51, 52,
Trophic chironomid index, 248 63, 64, 96, 97, 103, 104, 110, 120, 121,
Trophic classification, 108, 109, 113, 133, 170, 298, 341, 377, 378, 380
125–127, 135, 255 management measures, 20, 43, 53
Trophic gradient, 109, 136, 189 pollution, 374
Trophic state, 108, 109, 113, 114, 125, 127, quality, 11, 26, 36, 37, 46, 52, 53, 70, 74,
135, 190, 199, 232, 269 89, 90, 103, 109, 135, 168, 188, 200,
Turbidity, 7, 9, 10, 129, 152, 276, 214, 216, 245, 266, 277, 288, 309, 322,
289, 383 326, 332–334, 336, 338, 360, 369, 374,
Turbid state, 46, 160, 189, 377 376, 378, 381, 383
Subject Index 397
supply, 22, 374 White Bream, 46, 278, 280–285, 298,

turbid, 130, 260, 289 303, 308
Waterbird censuses, 319 Willow seeds, 279
Waterbirds, 63, 65, 313–326 Wind, 24, 32, 74, 81
Water purification (clearing-up water, water Winter, 29, 30, 42, 52, 56, 65, 75, 94, 108,
clearance rate), 288, 296, 299 116–119, 132, 155, 164, 168, 181, 185,
Water quality, 11, 26, 36, 37, 46, 52, 53, 70, 187, 194–198, 200–202, 214, 232, 268,
74, 89, 90, 103, 109, 135, 168, 188, 314–318, 320–322, 324, 379
200, 214, 216, 245, 266, 277, 288, 309, Winter climate index, 181
322, 326, 332–334, 336, 338, 360, 369, Winter NAO, 196
374, 376, 378, 381, 383
Water-sediment contact zone, 74, 75, 77, 78
Water-shed, 4, 374, 375 Y
Water temperature, 27–33, 56, 57, 65, 155, Yield, 114, 117, 119, 124, 128–130, 133–138,
156, 161, 164, 168, 169, 181–186, 169, 185, 190, 192, 200–202
200–202, 236, 284, 286, 289, 335, 379
Water transparency, 90, 92, 93, 103, 109, 133,
165, 167, 179, 188, 190, 195, 244, 260, Z
263, 267, 269, 288, 309 Zooplankton, 46, 48, 50, 51, 92, 108–111,
Waxing rates, 96 128, 134, 163–197, 216, 219, 234, 235,
Weed cutting (=underwater vegetation 238, 278, 279, 281, 286, 288, 289, 295,
mowing), 96, 103, 104 297–299, 309, 377, 382
Western Tubenose Goby, 280, 303, 308 abundance, 168–172, 175, 178–181, 188,
Wet weight of phytoplankton (elsewhere as 200, 234
freshweight of other biota), 136 biomass, 52, 167, 168, 178–181, 189, 194
Taxonomic Index
B Fulica atra, 315, 317, 320, 323, 324

Bird species Gallinula chloropus, 317
Aix Gavia
A. galericulata, 316 G. arctica, 316
A. sponsa, 316 G. immer, 316
Anas G. stellata, 316
A. acuta, 316 Ixobrychus minutus, 356
A. clypeata, 317 Larus
A. crecca, 316 L. argentatus, 318
A. penelope, 316 L. cachinnans, 318
A. platyrhynchos, 315–317, 323 L. canus, 318
A. querquedula, 317 L. fuscus, 318
A. strepera, 316 L. hyperboreus x L. argentatus, 318
Anser L. marinus, 318
A. albifrons, 316 L. ridibundus, 315, 318
A. anser, 316 Melanitta fusca, 317
A. fabalis, 316 Mergellus
Anser sp. hybrid, 316 M. albellus, 317, 323
Ardea M. albellus x M. serrator, 317
A. alba, 316 Mergus
A. cinerea, 314, 316 M. merganser, 317, 323
Aythya M. serrator, 317
A. ferina, 317, 323 M. serrator x M. albellus, 317
A. fuligula, 317, 323 Netta
A. marila, 317 N. rufina, 317
A. marila x A. ferina, 317 N. rufina x Aythya fuligula, 317
Branta canadensis, 316, 318 Nettapus auritus hybrid, 316
Bucephala clangula, 317, 323 Phalacrocorax
Cairina moschata, 316 P. carbo, 316, 323
Callonetta leucophrys, 316 P. pygmeus, 316
Clangula hyemalis, 317 Podiceps
Cygnus P. auritus, 316
C. atratus, 316, 324 P. cristatus, 316
C. olor, 315, 316, 318, 320, P. grisegena, 316
322, 323 P. nigricollis, 316

https://doi.org/10.1007/978-3-319-93270-5
400 Taxonomic Index
Bird species (cont.) Acipenser sp. x sp, 280

Somateria mollissima, 317 Alburnus alburnus, 46, 50, 278, 280–282,
Sterna caspia, 317 284, 286, 288, 289, 296, 297, 303
Tachybaptus ruficollis, 316 Anguilla anguilla, 280
Tadorna ferruginea, 316 Blicca bjoerkna, 46, 278, 280–282, 285,
287, 298, 303
Carassius
C C. carassius, 280, 298
Ciliate species C. gibelio, 280
Askenasia, 218, 219 Ctenopharyngodon idella, 104, 280
A. chlorelligera, 217 Cyprinus carpio, 46, 104, 280–282, 298
A. volvox, 217 Esox lucius, 46, 50, 279–282, 289, 296,
Balanion planctonicum, 216–219 298
Cinetochilum margaritaceum, 217 Gymnocephalus cernua, 280, 297, 303
Codonella cratera, 217–219 Lepomis gibbosus, 280, 282, 303
Coleps Leuciscus aspius, 280, 289, 292, 296
C. hirtus, 217 Perca fluviatilis, 46, 280, 282, 289, 297,
C. spetai, 217–219 298, 303, 308
Cyclidium glaucoma, 217–219 Proterorhinus semilunaris, 280, 303
Cyrtolophosis mucicola, 217, 218 Rhodeus amarus, 280
Didinium nasutum, 217 Rutilus rutilus, 46, 278, 280–282, 285,
Halteria 287, 298, 303
H. chlorelligera, 216, 217 Sander lucioperca, 46, 50, 279–282, 289,
H. grandinella, 216–218 291, 296
Lagynophrya acuminata, 217 Scardinius erythrophthalmus, 280, 298,
Mesodinium 303, 308
M. pulex, 217, 218 Silurus glanis, 280, 289
Monodinium balbiani, 217, 218 Squalius cephalus, 280
Pelagostrombidium Tinca tinca, 46, 280, 298
P. fallax, 217
P. mirabile, 217
Phascolodon vorticella, 217 H
Rimostrombidium, 217–219 Heterotrophic bacteria
R. brachykinetum, 217 Escherichia coli, 333–335
R. humile, 217 Salmonella, 333
R. hyalinum, 217 S. typhimurium, 333
R. lacustris, 217
Stobilidium humile, 216
Strobilidium hyalinum, 216 M
Strombidium viride, 217 Macrophyte and other plant species
Tintinnidium fluviatile, 217 Acorus calamus, 345, 346, 349
Tintinnopsis, 216, 218, 219 Alisma, 259
Uronema nigricans, 217, 218 A. lanceolatum, 350, 351
Urotricha, 216, 218, 219 A. plantago-aquatica, 341, 345, 346,
U. agilis, 217 349–350
U. apsheronica, 217 Butomus umbellatus, 341, 345, 346, 350
U. farcta, 217, 218 Carex, 343, 349, 351
U. matthesi, 217 C. acuta, 345, 346
Vorticella, 217, 218 C. elata, 345, 346
V. natans, 217 C. riparia, 345, 346, 366
Ceratophyllum demersum, 98, 259
Chara
F C. aspera, 98, 99
Fish species C. contraria, 98
Abramis brama, 46, 104, 280, 298 C. delicatula, 98
Taxonomic Index 401
C. globularis, 98 Populus
C. hispida, 46, 98 P. alba, 367
C. intermedia, 98 P. nigra, 367
C. tomentosa, 46, 98, 99, 259 Potamogeton spp., 94, 381
C. vulgaris, 97, 98 P. crispus, 98
Cyperus fuscus, 252, 350, 351 P. friesii, 98
Eleocharis acicularis, 98 P. lucens, 98
Elodea P. nodosus, 98
E. canadensis, 97, 98 P. pectinatus, 46, 98, 99
E. nuttallii, 98 P. perfoliatus, 98
Equisetum palustre, 350, 351 P. pusillus, 98
Eupatorium cannabinum, 345, 346, 350 P. x cooperi, 98
Glyceria Ranunculus
G. fluitans, 345, 346, 350 R. circinatus, 98, 299
G. maxima, 345, 346, 350 R. repens, 350, 351
Hippuris vulgaris, 97, 98 Robinia pseudacacia, 367, 369
Iris pseudacorus, 345, 346, 348, 349 Rumex hydrolapathum, 350, 351
Juncus, 341, 343 Sagittaria, 259
J. articulatus, 345, 346, 349 S. sagittifolia, 345, 346, 350
J. compressus, 345, 346, 350 Salicion albae, 367
J. effusus, 345, 346, 350 Salix, 299
J. filiformis, 350, 351 S. alba, 367
J. inflexus, 345, 346, 349 S. incana, 367
Lemna minor, 98 S. purpurea, 367
Lycopus europaeus, 351 S. rubens, 367
Lysimachia S. traindra, 367
L. nummularia, 350, 351 S. viminalis, 367
L. thyrsiflora, 345, 346, 350 Schoenoplectus, 341
Lythrum salicaria, 345, 346, 349 S. lacustris, 345, 346, 348, 350
Mentha, 341 S. mucronatus, 350, 351
M. aquatica, 345, 346, 349 Scirpoides holoschoenus, 345, 346, 350
M. longifolia, 345, 346, 350 Scirpus sylvaticus, 350, 351
Myosotis scorpioides, 350, 351 Solanum dulcamara, 350, 351
Myriophyllum spp., 94, 135, 258, 302, 304, Sparganium
381 S. emersum, 350, 351
M. spicatum, 46, 51, 91, 93–96, 98, 99, S. erectum, 346, 350
101, 103–105, 128, 135, 183, 192, Stratiotes aloides, 98
193, 259, 299, 300, 378–380 Tolypella intricata, 98
M. verticillatum, 97–99 Trapa natans, 99
Najas Typha sp., 341, 348
N. marina, 98, 99, 101, 192 T. angustifolia, 345, 346, 349, 351
N. minor, 98 T. latifolia, 345, 346, 350
Nitella mucronata, 98 T. laxmannii, 350, 351
Nitellopsis obtusa, 46, 94, 98, 101, 192, T. minima, 351
258 Utricularia vulgaris, 98
Nuphar lutea, 98 Veronica
Nymphaea alba, 98, 99, 101 V. anagallis-aquatica, 98
Nymphoides peltata, 98, 99 V. catenata, 98
Persicaria Zannichellia palustris, 98
P. dubia, 350, 351 Macrozoobenthic species
P. hydropiper, 351 Ablabesmyia
Phalaris arundinacea, 346, 349 A. longistyla, 252, 262
Phragmites australis, 338, 341, 346, A. monilis, 259
348–350, 366 Acricotopus lucens, 252
Polygonum amphibium, 98 Acroloxus lacustris, 250
402 Taxonomic Index
Macrozoobenthic species (cont.) C. carriana, 252

Aeolosoma, 250 C. coronata, 252
Alboglossiphonia hyalina, 250 C. gratias, 252
Amphichaeta leydigii, 250 C. scutellata, 252
Anodonta sp., 269 Craspedacusta sowerbii, 164, 169, 183,
A. anatina, 250 184, 197, 250
A. cygnea, 262 Cricotopus, 259
Arrenurus, 251 C. albiforceps, 252
Asellus aquaticus, 251 C. bicinctus, 252
Athripsodes cinereus, 251 C. cylindraceus, 252
Aulodrilus japonicus, 250 C. festivellus, 252
Axonopsis complanata, 251 C. flavocinctus, 252
Bezzia, 252 C. fuscus, 252, 350
Bithynia tentaculata, 250 C. intersectus, 252
Brachypoda versicolor, 251 C. pilitarsis, 252
Branchiura sowerbyi, 250 C. reversus, 252
Brillia bifida, 252 C. ricotopus, 252
Caenis C. sylvestris, 247
C. horaria, 251 C. trifasciatus, 252
C. lactea, 251 Criodrilus lacuum, 250
C. luctuosa, 251 Cryptochironomus, 252, 259, 261, 262
C. luctuosa/macrura, 251 C. albofasciatus, 252
Caenis rivulorum, 251 C. obreptans, 252
Casertiana nitida, 250 C. psittacinus, 252
Ceraclea dissimilis, 251 C. supplicans, 252
Chaetogaster diaphanus, 250 Cryptotendipes, 260
Chaoborus flavicans, 252 C. holsatus, 252
Chelicorophium curvispinum, 252 C. usmaensis, 252
Chironomus Demicryptochironomus vulneratus, 252
C. acutiventris/obtusidens, 252 Dendrocoelum lacteum, 250
C. balatonicus, 252, 259, 260, 263 Dero
C. longipes, 252 D. digitata, 250
C. longistylus, 252 D. obtusa, 250
C. luridus, 252 Dicrotendipes
C. nuditarsis, 252 D. lobiger, 252, 259
C. nudiventris, 252 D. nervosus, 252
C. plumosus, 259, 260, 263 D. notatus, 252
C. pseudothummi, 252 D. pulsus, 252
Cladopelma D. tritomus, 252, 259
C. bicarinata, 252 Dikerogammarus haemobaphes, 251
C. goetghebueri, 252 Dina punctata, 250
C. virescens, 252 Donacia sp., 251
C. viridula, 251 Dreissena polymorpha, 250, 264, 279
Cladotanytarsus Dugesia tigrina, 250, 262
C. atridorsum, 252 Echinogammarus ischnus, 251
C. lepidocalcar, 252 Ecnomus tenellus, 251
C. mancus, 259 Einfeldia pagana, 252
C. nigrovittatus, 263 Eiseniella tetraedra, 250
C. vanderwulpi, 252 Endochironomus albipennis, 252, 259
C. wexionensis, 251 Ephemera
Cloeon dipterum, 251 E. danica, 251
Coenagrion puella, 251 E. vulgata, 251
Corynoneura Ephemerella sp., 251
C. arctica, 251 Erpobdella octoculata, 250
Taxonomic Index 403
Erythromma najas, 251 Nais

Forelia curvipalpis, 251 N. christinae, 250
Frontipoda musculus, 251 N. communis, 250, 258
Galba truncatula, 250 N. elinguis, 250, 258
Gammarus roeselii, 251 N. pseudobtusa, 250, 258
Gerris sp., 251 N. simplex, 250
Glyptotendipes Nanocladius
G. mancunianus, 252 N. balticus, 253
G. pallens, 252, 260 N. bicolor, 253, 259
G. paripes, 252, 260 N. rectinervis, 247, 253
G. signatus, 252, 259 Nemoura sp., 251
G. viridis, 252, 259, 260 Neumania deltoides, 251
Gyraulus Nilotanypus dubius, 253
G. albus, 250 Oecetis
G. crista, 250 O. lacustris, 251
Haliplus sp., 251 O. ochracea, 251
Harnischia Ophidonais serpentina, 250, 262
H. curtilamellata, 252 Orthetrum sp., 251
H. fuscimana, 252 Orthocladius
Helobdella stagnalis, 250, 262 O. consobrinus, 253
Heterotrissocladius marcidus, 253 O. fuscimanus, 253
Hippeutis complanatus, 250 Orthotrichia costalis, 251
Hydra sp., 250 Oulimnius tuberculatus, 251
Hydrobaenus lugubris, 253 Oxyethira flavicornis, 251
Hydrochoreutes krameri, 251 Parachironomus
Hydrodroma despiciens, 251 P. arcuatus, 253, 259
Hydropsyche contubernalis, 251, 262 P. biannulatus, 253
Hydrozetes lacustris, 251 P. conversus, 253
Hygrobates longipalpis, 251 P. mauricii, 253
Ilyocoris cimicoides, 251 P. tenuicaudatus, 253, 258–259
Ilyodrilus templetoni, 250 P. varus, 253
Ischnura elegans, 251 P. vitiosus, 253
Kiefferulus tendipediformis, 253 Parakiefferiella
Labrundinia longipalpis, 253 P. bathophila, 253, 259, 262
Lebertia sp., 251 P. coronata, 253
Leptocerus tineiformis, 251 Paralauterborniella nigrohalteralis, 253
Limnesia undulata, 251 Paramerina cingulata, 253
Limnodrilus Paratanytarsus, 259, 260, 262
L. claparedeianus, 250 P. bituberculatus, 253
L. profundicola, 250 P. dimorphis, 253
Limnomysis benedeni, 251 P. dissimilis, 253
Limnophyes minimus, 253 P. inopertus, 253
Macrozoobenthic species P. laetipes, 253, 260
Limnodrilus P. tenellulus, 247
L. hoffmeisteri, 250 P. tenuis, 253, 259, 260
Microchironomus tener, 253 Paratendipes albimanus, 247, 253
Microtendipes Paratrichocladius rufiventris, 253
M. britteni, 253, 260 Pelmatohydra oligactis, 250
M. chloris, 253 Physella
Mideopsis orbicularis, 251 P. acuta, 250
Monohelea sp., 252 P. heterostropha, 250
Mystacides Piona
M. azurea, 251 P. discrepans, 251
M. longicornis, 251 P. pusilla, 251
404 Taxonomic Index
Macrozoobenthic species (cont.) Stenochironomus

Pisidium nitidum, 264 S. gibbus, 253
Planorbis planorbis, 250 Stictochironomus spp., 253, 259
Platycnemis pennipes, 251 Stylaria lacustris, 250, 258
Plumatella fungosa, 259 Stylodrilus heringianus, 250
Polypedilum Synorthocladius semivirens, 253
P. bicrenatum, 253 Tanypus
P. convictum, 253, 262 T. kraatzi, 253, 260, 262
P. cultellatum, 253 T. punctipennis, 253, 259, 260, 262
P. nubeculosum, 253, 259 Tanytarsus
P. sordens, 253, 260 T. bathophilus, 253, 263
Polypedilum sp., 247 T. chinyensis, 253
Porolohmanella violacea, 251 T. cretensis, 253
Potamopyrgus antipodarum, 250 T. ejuncidus, 253
Potamothrix T. eminulus, 253
P. bavaricus, 250 T. excavatus, 253
P. hammoniensis, 250 T. glabrescens, 253, 259, 260
P. moldaviensis, 250 T. gregarius, 253, 263
Pristina T. heusdensis, 253
P. aequiseta, 258 T. inaequalis, 254, 263
P. longiseta, 250 T. lactescens, 254
Procladius T. lestagei, 254
P. choreus, 253 T. mancospinosus, 254
P. lugens, 253 T. medius, 254, 260
P. rufovittatus, 253, 260 T. mendax, 254
P. sagittalis, 253 T. nigricollis, 254
P. signatus, 253 T. occultus, 254
Procladius spp., 259, 260, 262 T. pallidicornis, 254
Prodiamesa olivacea, 247, 253 T. signatus, 254
Propappus volki, 250, 262 T. smolandicus, 254
Psammoryctides T. sylvaticus, 254
P. barbatus, 250 T. verralli, 254
P. moravicus, 250 Thienemanniella sp., 254
Psectrocladius Tinodes waeneri, 251
P. limbatellus, 253 Trichobilharzia szidatii, 335
P. oxyura, 253 Trimalaconothrus novus, 251
P. psilopterus, 253 Tubifex
P. schlienzi, 253 T. ignotus, 250
P. sordidellus, 253, 259 T. tubifex, 250, 258
Pseudochironomus prasinatus, 253 Unio
Psilotanypus rufovittatus, 253, 260 U. pictorum, 250, 262, 269
Psychomyia pusilla, 251 U. pictorum latirostris, 250
Radix Unionicola
R. auricularia, 250 U. aculeata, 251
R. ovata, 250 U. crassipes, 251
Schineriella schineri, 253 Valvata
Sisyra sp., 251 V. cristata, 250
Slavina appendiculata, 250 V. piscinalis, 250
Specaria josinae, 250 V. studeri, 250
Stempellina Vejdovskyella
S. almi, 253 V. comata, 250
S. subglabripennis, 253, 260 V. intermedia, 250
Stempellinella minor, 253, 260, 263 Virgatanytarsus cf. arduennensis, 247
Taxonomic Index 405
Virgatanytarsus sp., 254 Diatoma, 112

Viviparus Dictyosphaerium, 112
V. acerosus, 250 Dinobryon, 112, 114–117
V. contectus, 250 D. divergens, 114–117
Xenochironomus xenolabis, 254 Dolichospermum, 112
Zavreliella marmorata, 254 Elakatothrix, 112
Encyonema triangulum, 115
Euglena acus, 115
O Fragilaria, 112, 115–117
Other animals (Vertebrata) F. construens, 116, 117
Bufo F. crotonensis, 116
B. bufo, 356 F. ulna, 116
B. viridis, 356 Gomphosphaeria, 112, 117, 122
Castor fiber, 356 Gymnodinium, 112
Pelophylax esculentus, 356 Hyalotheca dissiliens, 115
Pipistrellus pipistrellus, 356 Koliella, 112, 116
K. longiseta, 116
Lagerheimia, 112
P Limnothrix, 46, 112, 122
Phytoplankton species (cyanobacteria L. redekei, 46, 122, 130–132,
and algae) 189, 260
Ankyra, 112 Mallomonas, 112
Aphanizomenon, 112, 130 Melosira varians, 112
A. flos-aquae, 130 Merismopedia, 112, 117
Aphanocapsa, 112, 114, 131 Microcystis, 117, 122, 130, 131
Asterionella formosa, 116, 117 M. aeruginosa, 112, 114, 131
Aulacoseira, 116 M. aeruginosa f. aeruginosa, 112
A. granulata, 112 M. flos-aquae, 114, 131
Botryococcus, 112, 114, 115 M. novacekii, 114
B.braunii, 114, 115 M. viridis, 112, 114
Carteria, 112, 116, 117 Monoraphidium, 112
C. pseudomultifilis, 116, 117 Mougeotia, 112
Ceratium, 112, 116, 117 Navicula, 112, 116
C. hirundinella, 116, 117 Nitzschia, 112, 116
Chroococcus, 122 N.acicularis, 116, 117
Chrysomonas, 112 Ochromonas, 112
Closterium, 112, 116, 131 Oocystis, 112
C. aciculare, 131 Oscillatoria, 112, 130, 131
C. acutum, 116 Pediastrum, 112
Coelastrum, 112 Pennate diatoms, 112, 128, 131
C. astroideum, 116, 117 Peridinium, 112
C. reticulatum, 116 Planktolyngbya, 112
Coenochloris, 114, 115 P. circumcreta, 116, 117
Cosmarium, 112, 114, 115 P. limnetica, 131
C. botrytis, 114, 115 P. subtilis, 116, 117
Crucigenia, 112 Planktonema, 112
Cryptomonas, 117 Planktothrix, 112, 130
C. curvata, 112 P. agardhii, 130
C. erosa, 116, 117 Pseudanabaena, 112, 122
C. marssonii, 112 P. catenata, 116
Cylindrospermopsis raciborskii, 46, 116, Quadricoccus, 112
117, 122, 127, 129–132, 152, 165, Radiococcus, 112
189, 260 Radiocystis, 112, 122
406 Taxonomic Index
Phytoplankton species (cyanobacteria Cyclops vicinus, 172–174, 192, 201

and algae) (cont.) Daphnia
Rhodomonas, 112, 116, 117 D. cucullata, 172–174, 189–192, 194,
R. lacustris, 116, 117 281, 294
Scenedesmus, 112 D. hyalina, 172
Schroederia robusta, 116 D. pulex, 194
Snowella, 112, 116, 117, 122 Diaphanosoma
S. lacustris, 116 D. brachyurum, 172–174, 192
Staurastrum, 112, 116, 117 D. orghidani, 172
S. gracile, 116, 117 Disparalona rostrata, 172
Synura, 112 Encentrum, 172
Tabellaria, 112, 116, 117 Eubosmina coregoni, 46, 172–174, 281
T. flocculosa, 116 Euchlanis, 172
Testraselmis, 116 Eucyclops serrulatus, 172, 201
Tetraedron, 112 Eudiaptomus gracilis, 172–174, 189–192,
Tetraselmis, 112, 117 194, 201, 294
Tetrastrum, 112 Eurycercus lamellatus, 172
Trachelomonas, 112 Filinia
Unicellular centric diatoms, 112, 116 F. longiseta, 172
Uroglena, 112, 131 F. terminalis, 172
U. volvox, 114–116 Gastropus stylifer, 172, 173, 177
Woronichinia naegeliana, 112 Graptoleberis testudinaria, 172
Zygnema, 112 Hexarthra mira, 172
Kellicottia, 172, 173, 177, 192, 281
Kellicottia longispina, 172, 173,
Z 177, 192
Zooplankton species Keratella, 281
Acroperus harpae, 172, 174, 177 K.cochlearis, 172–174
Alona, 172, 181 K. quadrata, 172
A. affinis, 172 Lecane
Alonella nana, 172 L. closterocerca, 172
Anuraeopsis fissa, 172 L. cornuta, 172
Ascomorpha L. elongata, 172
A. ecaudis, 172 L. luna, 173, 177
A. ovalis, 172 L. lunaris, 172
A. saltans, 172 Lepadella, 172
Asplanchna priodonta, 172, 173 Leptodora kindtii, 172, 173, 281
Bosmina longirostris, 172–174, 198 Lophocharis, 172
Brachionus Macrochaetus subquadratus, 172
B. angularis, 172–174 Megacyclops
B. calyciflorus, 172 M. viridis, 172, 201
B. quadricornis, 172 Mesocyclops leuckarti, 172–174,
Camptocercus rectirostris, 172 192, 201
Ceriodaphnia quadrangular, 172–174 Monommata, 172
Chaoborus, 252, 281 Mytilina, 172
Chydorus sphaericus, 172–174, 189, Notholca
192, 281 N. acuminata, 172
Collotheca mutabilis, 172, 173, 177 N. jugosa, 172
Colurella uncinata, 172 N. squamala, 172
Conochilus unicornis, 172, 173, 177 Pleosoma hudsoni, 172
Craspedacusta sowerbii, 164, 169, 183, Pleuroxus truncatus, 172
184, 197, 250 Polyarthra dolichoptera-vulgaris, 172
Taxonomic Index 407
Polyphemus pediculus, 172 Thermocyclops

Pompholyx solcata, 172 T. crassus, 173, 174, 192, 201
Pseudochydorus globosus, 172 T. oithonoides, 172–174, 201
Scapholeberis mucronata, 172 Trichocerca
Sida crystallina, 164, 172, 174, 175, 193 T. birostris, 172
Simocephalus vetulus, 164, 172, 174, T. capucina, 173, 177
175, 193 T. insignis, 173, 177
Squatinella, 172 T. pusilla, 173, 177
Synchaeta T. similis, 172
S. pectinata, 172 Trichotria
S. tremula-oblonga, 172 T. pocillum, 172
Testudinella patina, 172 T. tetractis, 172

The Alte Donau: Successful Restoration and Sustainable Management

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Aquatic Ecology Series

Martin T. Dokulil · Karl Donabaum

The Alte Donau:

The Alte Donau: Successful

Aquatic Ecology Series

Library of Congress Control Number: 2018954812

© Springer International Publishing AG, part of Springer Nature 2018

Wien, Austria The Editors

Part II History, Physiography, Chemistry and Climate

Part III Biology, Ecology and Production

10 Phytoplankton Photosynthesis and Production�������������������������������������� 149

Part IV Recreation, Landscape Planning and Synthesis

Martin T. Dokulil was born 1943 in Vienna, Austria.

Karl Donabaum was born in Oberpullendorf, Austria,

Katrin Teubner born in Leipzig, East Germany, in

Marie-Curie Intra-European research grant at Biological

Keywords Eutrophication · Cyanobacteria · Blooms · Rehabilitation · Bistable

M. T. Dokulil (*) · K. Donabaum

© Springer International Publishing AG, part of Springer Nature 2018 3

acceleration of nutrient input, commonly termed cultural or anthropogenic eutro-

1.2 Nutrient Loading Reconsidered

Eutrophication modelling approaches are applicable only to waterbodies in which

Since no direct TP measurements are available for 1971–1973, approximate TP

1.3 Stable Equilibria Revisited

The existence of alternative stable states in natural ecosystems was hypothesized

Fig. 1.4 Total phosphorus 100

Chlorophyll-a [Chl-a, mg m-3]

and separates lakes Oligo Meso Eu Hypertrophic

considered as an alternative in situations where natural internal switching is unlikely.

function of freshwater systems. The theory of alternative stable equilibria however

Regime shifts in shallow lakes as a result of ecohydrological changes can be suc-

• Reduction of external nutrient loading (catchment restoration)

Keywords Urban lake · Historic development · Geology · Flood protection

© Springer International Publishing AG, part of Springer Nature 2018 17

2.2 The Major Regulation of the Donau

2.3 The Neue Donau (‘New Danube’)

2.4 The Hydroelectric Power Plant Freudenau

2.5 The Contaminated Site ‘Donaupark-Bruckhaufen’

2.6.2 First Leisure Time Activities

2.6.3 First and Second World War

2.6.4 Development Since the Second World War

2.6.5 Leisure Time Activities Today

Fig. 2.3 The Alte Donau today

Keywords Morphometry · Hydrology · Temperature · Warming · Climate ·

G. Kum (*) · M. T. Dokulil

© Springer International Publishing AG, part of Springer Nature 2018 27

Fig. 3.1 Depth contours for Alte Donau

Monthly mean surface water temperature (SWT) peaks in August at 25.3 °C. Average

Air temperature [°C]

Fig. 3.3 Climate diagram 30

Average air temperature [°C]

same period as in (a)

characteristics originating from the landscape, particularly from the location

Fig. 3.4 Diagram combining annual wind direction and speed

climate can be characterised by Cfb (Fig. 3.3a), which means moderately warm, no

Abstract The hydrological conditions in the Alte Donau and the appendix

Keywords Hydrology · Long-term trend · Numeric model · Nutrient balance ·

© Springer International Publishing AG, part of Springer Nature 2018 35

A network of groundwater probes, gauging posts and climate monitoring stations

annual variation is negligible. Groundwater levels in Wasserpark and Alte Donau

Such a relationship is exemplified in Fig. 4.6 for the inflow to the Wasserpark.

Keywords Restoration · Lake management · Alternative stable states · Water

K. Donabaum (*) · M. T. Dokulil

Part II History, Physiography, Chemistry and Climate

Part III Biology, Ecology and Production

10 Phytoplankton Photosynthesis and Production�� 149

Part IV Recreation, Landscape Planning and Synthesis

1.2 Nutrient Loading Reconsidered

1.3 Stable Equilibria Revisited

2.2 The Major Regulation of the Donau

2.3 The Neue Donau (‘New Danube’)

2.4 The Hydroelectric Power Plant Freudenau

2.5 The Contaminated Site ‘Donaupark-Bruckhaufen’

2.6.2 First Leisure Time Activities

2.6.3 First and Second World War

2.6.4 Development Since the Second World War

2.6.5 Leisure Time Activities Today

5.1 Eutrophication History

5.2 Restoration Measures

6.2 The History of Investigations in Alte Donau

6.6 Nutrient Balances

7.3 Main Constitutes and Iron-Content

7.4 Phosphor Components in the Sediment

7.5 Interstitial Water

7.6 Phytobenthic Biomass and Algal Classes

8.2.1 Determination of Plant Biomass

8.2.2 Mapping Procedure

8.2.4 Further Evaluations

8.3 Results and Discussion

8.3.1 Macrophyte Biomass

8.3.2 Species Spectrum

8.3.3 Dominance-Structure of the Vegetation