Cent. Eur. J. Biol. • 3(2) • 2008 • 177–187
DOI: 10.2478/s11535-007-0048-4
Central European Journal of Biology
The distribution and the abundance of hydrophytes
along the Danube River in Serbia
Research Article
Dragana Vukov*, Pal Boža, Ružica Igić, Goran Anačkov
Department of Biology and Ecology,
Faculty of Sciences and Mathematics,
University of Novi Sad,
21000 Novi Sad, Serbia
Received 16 May 2007; Accepted 19 November 2007
Abstract: T he Danube River runs through Serbia with flow 588 km long. Different hydrological, geological and ecological conditions occuring
along the Danube in Serbia enables its division into four sections: The upper region, Đerdap I accumulation, Đerdap II accumulation, and
the lower region. Each section is characterized by unique plant species diversity, frequency of occurrence and distribution pattern. The
species diversity ranged from low in the upper reach, to high or very high in the impoundments and decreased in the lower reach. This
paper provides a basis for a detailed study on the influence of large power plant impoundments over the aquatic vegetation of rivers.
Keywords: Aquatic vegetation • Macrophytes • Impoundments
© Versita Warsaw and Springer-Verlag Berlin Heidelberg.
1. Introduction
The Danube River is the Europe’s second longest
river. It originates in the Black Forest in Germany as
two smaller rivers, the Brigach and the Breg, which join
at Donaueschingen. The river flows eastwards for a
distance of some 2850 km, either passing through, or
forming parts of the borders of ten countries: Germany,
Austria, Slovakia, Hungary, Croatia, Serbia, Bulgaria,
Romania, Moldova, and Ukraine, before emptying
into the Black Sea via the Danube Delta in Romania
and Ukraine. In addition, the drainage basin includes
parts of ten more countries: Italy, Poland, Switzerland,
Czech Republic, Slovenia, Bosnia and Herzegovina,
Montenegro, Republic of Macedonia, Moldova, and
Albania. The flow of the Danube could be divided in to
three main sections. The upper section, from origin to
Devín Gate (at the border of Slovakia and Austria), where
it remains a characteristic mountain river until Passau
(Germany), with an average bottom gradient of 0.0012%.
From Passau to Devín Gate the gradient lessens to
0.0006%. The middle section, from Devín Gate to Iron
Gate (at the border of Serbia and Romania), where
the riverbed widens and the average bottom gradient
becomes only 0.00006%. The lower section, from Iron
Gate to Sulina (Romania), with an average gradient as
little as 0.00003% [1]. The Danube River enters Serbia
in its middle section, 1433 kilometers upstream of its
mouth, and it leaves Serbia at the mouth of the river
Timok to the Danube (river km 845.5; lower section). Its
flow through Serbia is 588 km long. Left riverside mainly
belong to the ecoregion of Hungarian lowlands, while
the right riverside of the river in Serbia belong to the
ecoregions of Dinaric western Balkan (river km 1296
and 1104.5), and the Eastern Balkan (river km 1104.5
and 845.5). During the year of 1972, the construction of
the Power Plant and the dam Đerdap I, on the river km
943, was completed. Under conditions of mean water
level, the reservoir Đerdap I reaches upstream to the
river km 1155 (which is the mouth of river Tamiš). During
the 1980s a second dam and Power Plant Đerdap II
was constructed. It is situated on the river km 863.4.
Its influence on water hydrology can be detected as
far as 80 river km upstream. Due to the undertaken
hydro technical measures and their influence on the
river, Danube in Serbia could be divided in to four
subsections: the upper region (river km 1433 to river
km 1155), Đerdap I run of the river reservoir (river km
1154 to river km 943), Đerdap II run of the river reservoir
* E-mail: [email protected]
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The distribution and the abundance of hydrophytes
along the Danube River in Serbia
(river km 942 to river km 863) and the lower region
(river km 862 to river km 845.5). The construction of
the Danube dams in Serbia caused severe changes in
the river hydrology. The flow velocity was reduced, the
water level rose, and the sedimentation was increased.
Those ecological changes have been observed in both
of the Danube reservoirs and are manifested in the
occurrence and the abundance of aquatic plant species
[2–7]. Hydrophytes are a specific group of plants
that are adapted to grow in water. Since the water is
their habitat, their presence, species composition,
abundance, and distribution can be considered as
important indicators of ecological conditions occuring
in water bodies. In this case, hydrophytes can be
considered as important indicators of the Danube
subsections in Serbia. Different studies of aquatic
vegetation have been conducted in various water
bodies in the Danube flood plane [8–10]. The results
gained in this study give insight in to the hydrophytes
composition, abundance and the distribution pattern
in the main channel of this large river, as well as the
influence that the newly created impoundments have
on them.
2. Experimental Procedures
The survey of aquatic plants was done during the
2002-2005 period. Investigations included 358 river
km along the left riverside of the main channel and 451
river km along the right riverside. The occurrence and
the abundance (estimated according to a descriptive
five-level scale: 1-rare, 2-occasional, 3-frequent,
4-abundant, 5-very abundant to massive occurrence
of plant species) of species were assessed in survey
units of one river km length. The methodology applied
has a long tradition in the Danube catchment [11–30]
and includes calculation of Relative Plant Mass (RPM
%; Eq. 1), Mean Mass Indices (MMT; Eq. 2, MMO; Eq.
3) and Distribution Ratio (d; Eq. 4 )
Regarding the Mean Mass Indices (MMT, MMO)
n
RPM [% ] =
∑ (PM
i =1
k
i
⋅ Li ) ⋅ 100
 n

 ∑ PM ji ⋅ Li 
∑
j =1  i =1

(1)
RPM – Relative Plant Mass
PM – estimated abundance of the species i rasied to the third power
L – length of the survey unit where the species i occurs, in this case one
river kilometer
and the Distribution Ratio (d) of the aquatic plants,
in principle, there are three possible situations. The
first one is that the plant species has high and equal
n
MMT =
3
∑ PM
i =1
i
⋅ Li
i
⋅ Li
L
n
MMO =
∑ PM
3
i=x
(3)
n
∑L
i=x
d=
(2)
MMT 3
MMO 3
i
(4)
MMT – Mean Mass Index calculated for the whole length of the surveyed
area
MMO – Mean Mass Index calculated for the length of the survey units
where the species i occurs
L – total length of the surveyed area
Li – length of the survey unit where the species i occurs
or approximately equal values for MMT and MMO,
and that will result in high, or maximum value of the
distribution ratio. In the second case, the MMT and
MMO values are equal, or almost equal, but low,
which will again result in high or the maximum value of
distribution ratio. In the third case, values of MMT and
MMO are different, and the more different they are, the
lower is the distribution ratio value. Accordingly, it is
possible to describe three different distribution types of
the aquatic plants. In the first case plant occurrence is
frequent or very abundant regarding the whole length of
the surveyed area (MMT≥3), and regarding the length
of the survey units where it occurs (MMO≥3). So, its
distribution ratio will gain the maximum value (d=1),
in other words, this plant is abundant along whole
water body and it is homogeneously distributed. In the
second case plant is rare or occasional along the whole
surveyed area (MMT<3) and it is rare or occasional in
the survey units where it grows (MMO<3). Its distribution
ratio value will be high or maximal, so the distribution of
the plant will be described as homogeneous. The third
option is that, regarding the whole length of surveyed
area, plant occurs rarely or occasionally (MMT<3),
but in the survey units where it grows it appears to
be ranging from frequent to very abundant (MMO≥3).
Since the distribution ratio is the ratio of MMT and
MMO, it will gain small values. The distribution of plant
will be described as heterogeneous as it is localized
only on certain parts (survey units) of surveyed area,
and does not grow along whole studied water body.
Field data were collected and processed according
to the guidelines established during the “Multifunctional
Integrated Study Danube: Corridor and Catchment”
project (www.midcc.at, and according to EU Water
Framework Directive [31]. Plant species were
determined according to the Flora Europaea [32,33].
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Species
left riverside
1
right riverside
2
1
2
Abbrev.
3
4
1.
Acorus calamus L.
Aco cal
2.
Alisma gramineum Lej.
Ali gra
3.
Alisma plantago-aquatica L.
Ali pla
4.
Azolla filiculoides Lam.
Azo fil
5.
Butomus umbellatus L.
But umb
6.
Ceratophyllum demersum L.
Cer dem
7.
Elodea canadensis Michx
Elo can
8.
Elodea nuttallii (Planchon) St John
Elo nut
9.
Hydrocharis morsus-ranae L.
Hyd mor
10.
Iris pseudacorus L.
Iri pse
11.
Lemna gibba L.
Lem gib
12.
Lemna minor L.
Lem min
13.
Mentha aquatica L.
Men aqu
14.
Myriophyllum spicatum L.
Myr spi
15.
Myriophyllum verticillatum L.
Myr ver
16.
Najas marina L.
Naj mar
17.
Najas minor All.
Naj min
18.
Nuphar lutea (L.) Sibth. & Sm.
Nup lut
19.
Nymphaea alba L.
Nym alb
20.
Nymphoides peltata (Gmelin) Kuntze
Nym pel
21.
Oenanthe aquatica (L.) Poiret.
Oen aqu
22.
Plantago lanceolata L.
Pla lan
23.
Polygonum amphibium L.
Pol amp
24.
Potamogeton crispus L.
Pot cri
25.
Potamogeton gramineus L.
Pot gra
26.
Potamogeton lucens L.
Pot luc
27.
Potamogeton natans L.
Pot nat
28.
Potamogeton pectinatus L.
Pot pec
29.
Potamogeton perfoliatus L.
Pot per
30.
Potamogeton pusillus L.
Pot pus
31.
Potamogeton x fluitans Roth.
Pot flu
32.
Potamogeton zizii Koch ex Roth
Pot ziz
33.
Rorippa amphibia (L.) Besser
Ror amp
34.
Sagittaria sagittifolia L.
Sag sag
35.
Salvinia natans (L.) All.
Sal nat
36.
Sparganium erectum L.
Spa ere
37.
Spirodela polyrhiza (L.) Schleiden
Spi pol
38.
Trapa natans L.
Tra nat
39.
Vallisneria spiralis L.
Val spi
40.
Zannichellia palustris L.
Zan pal
Table 1.
List of aquatic plant species recorded and their absence/presence in the investigated river sections: 1-upper region, 2- Đerdap I, 3Đerdap II, 4-lower region, Abbrev.-abbreviations of the species names.
3. Results
In total 40 plant species were recorded and many
species showed a specific pattern of occurrence: 24 in
the upper region, 37 in the impoundment of Đerdap I,
17 in Đerdap II, and 11 in the lower region of the river.
Mentha aquatica was found only in the upper part, Alisma
gramineum, Azolla filiculoides, Lemna gibba, Najas
minor, Nuphar lutea, Nymphaea alba, Nymphoides
peltata, Oenanthe aquatica and Sparganium erectum
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The distribution and the abundance of hydrophytes
along the Danube River in Serbia
14331430
Aquatic Plants
Aco cal
Ali pla
Azo fil
But umb
Cer dem
Elo can
Elo nut
Hyd mor
Iri pse
Lem gib
Lem min
Men aqu
Myr spi
Naj mar
Naj min
Nym alb
Nup lut
Oen aqu
Pol amp
Pot cri
Pot flu
Pot gra
Pot luc
Pot nat
Pot pec
Pot per
Pot pus
Pot ziz
Ror amp
Sag sag
Sal nat
Spa ere
Spi pol
Tra nat
Val spi
1420
1410
1400
1390
1380
1370
1360
1350
1340
1330
1320
1310
1300
1290
1280
1270
1260
1250
1240
1230
1220
1210
1200
1190
1180
1170
1160
1150
1140
1130
1120
1110
1100
1090
10801076
Figure 1.
Distribution of aquatic plant species, left riverside. The vertical lines indicate the investigated river km. The abundance of each species
is shown semi quantitatively by the black columns differing in height (low height, rare and occasional occurrence; medium height,
frequent occurrence; high columns, abundant and very abundant).
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were recorded in Đerdap I reservoir only. Zannichellia
palustris occurred only in Đerdap II, and the amphibian
form of Plantago lanceolata was only found in the
lower region of the river. Aquatic species recorded
exclusively in the impoundment areas were Acorus
calamus, Elodea nuttallii, Potamogeton natans and
Potamogeton zizii. Although the number of the species
was comparable between the left (35) and the right (36)
riversides, both sides differed significantly in the species
composition and abundance (Table 1, Figures 1, 2).
Lemna gibba, Mentha aquatica, and Nymphaea alba
were recorded on the left riverside only, while Alisma
gramineum, Myriophyllum verticilatum, Nymphoides
peltata, Plantago lanceolata, and Zannichellia palustris
were found exclusively on the right riverside.
With regard to the species diversity, the left riverside
showed two distinct regions. The upper region was
characterised by general low species diversity. Except
of Rorippa amphibia which is almost continually
distributed, all other species occurred in low abundance
and only at a rather few sampling sites. In contrast, the
region downstream of river km 1155 showed a much
higher species diversity with many species occurring in
high abundance and continuously throughout the whole
region. This threshold coincides with the beginning
of the Đerdap I impoundment area. Consequently,
the high species diversity and high abundance of the
aquatic plants in the downstream region of river km
1155 was caused by the hydrological changes induced
by the impoundment, i.e. a decreased flow velocity,
larger littoral areas and higher sedimentation rates. In
contrast to the left riverside, the species diversity and the
abundance of aquatic plants on the right riverside was
high throughout the whole study area (Figure 2). Some
regions showed an extremely high species diversity,
such as the impoundment areas of Đerdap I and II.
Along the left riverside 35 aquatic plant species
were recorded. All of them had very low values
of MMT. Only for Potamogeton pectinatus and
Ceratophyllum demersum it was around level 2
(Figure 3). Potamogeton pectinatus occurs frequently
along the river (MMO>3), while Ceratophyllum
demersum, Elodea canadensis, Mentha aquatica,
Najas marina, Potamogeton gramineus, Potamogeton
lucens, Potamogeton perfoliatus and Trapa natans
occur occasionally to frequently where they are growing
along their habits. Since there are big differences
between MMT and MMO values of recorded species,
their values of distribution ratio are much smaller than
0.5, so their distribution is very heterogeneous. Only
Rorippa amphibia has low, but equivalent values of MMT
and MMO. Its distribution ratio is near 0.5, so it appears
to be rare but homogeneously distributed in almost half
of the surveyed river kilometres along the left riverside.
Along the right riverside, 36 aquatic plant species
were found. Potamogeton pectinatus, Potamogeton
perfoliatus, Potamogeton lucens have MMT value
over 2, so along the whole 451 river km of the right
riverside they occur occasionally or frequently. MMT
values for Ceratophyllum demersum, Potamogeton x
fluitans, Potamogeton gramineus, and Potamogeton
natans are near the level 2, while for the other
recorded species they are well below 2 (Figure 4). The
highest MMO values, near the level 3, are recorded
for Plantago lanceolata, Potamogeton pectinatus
and Potamogeton perfoliatus. They are followed
by Ceratophyllum demersum, Elodea canadensis,
Elodea nuttallii, Myriophyllum spicatum, Potamogeton
x fluitans, Potamogeton gramineus, Potamogeton
lucens and Potamogeton natans. Their MMO values
are over the level 2. The other recorded species occur
rarely in the survey units were they are growing. The
highest distribution ratio was recorded for Potamogeton
pectinatus, Ceratophyllum demersum,Potamogeton
lucens and Potamogeton perfoliatus which means that
they are almost homogenously distributed over the
half of surveyed river km (d>0.5). They are followed
by Butomus umbellatus, Potamogeton x fluitans,
Potamogeton gramineus and Spirodela polyrhiza with
the distribution ratio value just below 0.5. Other recorded
species have distribution ratio value well below 0.5,
and they are distributed heterogeneously along the
right riverside of the Danube River in Serbia.
Potamogeton pectinatus is the dominant plant
species in aquatic vegetation along both surveyed
riversides. Along the left and right riversides of the
Danube River in Serbia, it is present with 24.96%
(Figure 5) and 20.6% RPM, (Figure 6) respectively.
Along the left riverside it is followed by Ceratophyllum
demersum (15.5%), Potamogeton gramineus (12.3%)
and Potamogeton lucens (11.1%) and on the right
riverside by Potamogeton perfoliatus (16.1%),
Potamogeton lucens (11.7%). Other recorded species
are present in the aquatic vegetation of the Danube
River with less than 10% of Relative Plant Mass.
4. Discussion
The 40 recorded species showed specific occurrence
and the distribution patterns depending on the
subsections of the Danube in Serbia sampled. In the
surveyed river kilometres of the upper region, where
there is no influence of the impoundment, 24 recorded
species occur rarely and in quite few survey units,
except for Rorippa amphibia, which is continually
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The distribution and the abundance of hydrophytes
along the Danube River in Serbia
1296
Aquatic Plants
Aco cal
Ali gra
Ali pla
Azo fil
But umb
Cer dem
Elo can
Elo nut
Hyd mor
Iri pse
Lem min
Myr spi
Myr ver
Naj mar
Naj min
Nym pel
Nup lut
Oen aqu
Pla lan
Pol amp
Pot cri
Pot flu
Pot gra
Pot luc
Pot nat
Pot pec
Pot per
Pot pus
Pot ziz
Ror amp
Sag sag
Sal nat
Spa ere
Spi pol
Tra nat
Val spi
Zan pal
1290
1280
1270
1260
1250
1240
1230
1220
1210
1200
1190
1180
1170
1160
1150
1140
1130
1120
1110
1100
1090
1080
1070
1060
1050
1040
1030
1020
1010
1000
990
980
970
960
950
Dam I
930
920
910
900
890
880
870
Dam II
850 846
Figure 2.
Distribution of aquatic plant species, right riverside. The vertical lines indicate the investigated river km. The abundance of each
species is shown semi quantitatively by the black columns differing in height (low height, rare and occasional occurrence; medium
height, frequent occurrence; high columns, abundant and very abundant).
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Aco cal
Ali pla
Azo fil
But umb
Cer dem
Elo can
Elo nut
Hyd mor
Iri pse
Lem gib
Lem min
Men aqu
Myr spi
Naj mar
Naj min
Nym alb
Nup lut
Oen aqu
Pol amp
Pot cri
Pot flu
Pot gra
Pot luc
Pot nat
Pot pec
Pot per
Pot pus
Pot ziz
Ror amp
Sag sag
Sal nat
Spa ere
Spi pol
Tra nat
Val spi
1
Figure 3.
2
3
4
5
0
0.5
1
Mean Mass Indices and Distribution Ratio, left riverside. The left diagram represents the values of Mean Mass Indices: MMT is shown
using black bars, and MMO with white bars. The right diagram shows the values of the Distribution Ratio.
distributed. Mentha aquatica is specific for the upper
region. In the following subsection of Đerdap I, 37 plant
species were recorded. This reservoir is older than
Đerdap II, and characterized by the high diversity of
aquatic plants, their high abundance and homogenous
distribution. Đerdap II accumulation is similar to the
Đerdap I but with lower number of recorded species.
Hydrophytes characteristic for the impoundments
are: Alisma gramineum, Azolla filiculoides, Lemna
gibba, Najas minor, Nuphar lutea, Nymphaea alba,
Nymphoides peltata, Oenanthe aquatica, Sparganium
erectum Acorus calamus, Elodea nuttallii, Potamogeton
natans, Potamogeton zizii and Zannichellia palustris.
High diversity and the abundance, as well as the more
or less continual distribution of the hydrophytes in the
surveyed impoundments indicate more lacustric type
of habitat. Similar influence of the impoundment was
recorded in different water bodies influenced by the
Gabčíkovo hydropower station in Slovakia [34], although
the higher diversity of hydrophytes was recorded in
the backwaters and not in the main channel of the
river. In Gabčíkovo, as well as in Đerdap I and II, the
participation of the Potamogeton species in the aquatic
vegetation of the sections influenced by the dam, is
quite significant and worth further studying. In the lower
region, eleven plant species occur heterogeneously
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The distribution and the abundance of hydrophytes
along the Danube River in Serbia
Aco cal
Ali gra
Ali pla
Azo fil
But umb
Cer dem
Elo can
Elo nut
Hyd mor
Iri pse
Lem min
Myr spi
Myr ver
Naj mar
Naj min
Nym pel
Nup lut
Oen aqu
Pla lan
Pol amp
Pot cri
Pot flu
Pot gra
Pot luc
Pot nat
Pot pec
Pot per
Pot pus
Pot ziz
Ror amp
Sag sag
Sal nat
Spa ere
Spi pol
Tra nat
Val spi
Zan pal
1
Figure 4.
2
3
4
5
0
0.5
1
Mean Mass Indices and Distribution Ratio, right riverside. The left diagram represents the values of Mean Mass Indices: MMT is
shown using black bars, and MMO with white bars. The right diagram shows the values of the Distribution Ratio.
and with low abundance. This part of the river is quite
similar to the upper region, and could be considered
by its characteristics as typical for a free flowing river.
Although the study deals only with the Hydrophytes and
Plantago lanceolata does not belong to that group of
plants, it was included because of its specific occurrence
in the lower part of the Danube in Serbia. It grows on
the flat slopes of the fine material on the bank and in
the water submerged areas, even during the low water
level. Its morphological characteristics put this plant
between Plantago lanceolata and P. altissima. Since its
taxonomical status is yet to be revealed, for the purpose
of this study it is named as Plantago lanceolata, and its
taxonomical status is under investigation. Although the
list of recorded hydrophytes is more or less similar to
the lists gained in other countries along the Danube, it
is important to notice the absence of aquatic mosses in
Serbian reach, while in the upper parts of the Danube
they are quite common [11-25,34].
In the River Danube in Serbia, as well as in other
parts of the Danube [11-25,34], in the whole length
of surveyed area plants occur rarely or occasionally
(MMT<3). In the survey units where they actully
were recorded, they are usually frequent to abundant
(MMO>3).
Since the distribution ratio is the ratio of MMT to
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Pot pec
Cer dem
Pot gra
Pot luc
Pot per
Spi pol
Ror amp
Naj mar
Elo can
Pot cri
But umb
Tra nat
Pot nat
residual (22)
0
Figure 5.
10
20
%
Relative Plant Mass, Danube-main channel, left riverside.
Pot pec
Pot per
Pot luc
Cer dem
Pot gra
Pot flu
Pot nat
Myr spi
But umb
Spi pol
Tra nat
Pot cri
Elo can
residual (24)
0
Figure 6.
10
%
Relative Plant Mass, Danube-main channel, right riverside.
MMO, its value decreases, so the distribution of plants
in the Danube can be described as heterogeneous
because recorded species are localized only on certain
parts (survey units) of the surveyed area.
Comparing the Relative Plant Mass values gained
for the aquatic plants that are growing on the left and
the right riversides, some differences can be seen.
First of all, the number of recorded aquatic plant
species are different. On the left riverside 35 plant
species were found, while on the right riverside this
number is 36. Potamogeton pectinatus dominates the
aquatic vegetation along the both riversides. Until 2002
Ceratophyllum demersum was the dominant species
not only in the Serbian reach of Danube but also in
the whole river [12]. The order of dominance according
to their RPM values, among other recorded species
also differs among left and right riverside. Other than
the differences between the surveyed subsections of
the Danube in Serbia, there are differences between
the left and right riversides in general. The data is
not completely comparable because of the lack of
information about the aquatic vegetation in sections of
the main channel where it is a natural border between
Serbia and Croatia (river km 1433 to 1296) and Serbia
and Romania (river km 1075 and 845.5). Although
the number of recorded species is comparable, their
composition, abundance and distribution is different.
These differences can be explained by the differences
in habitat conditions and ecological characteristics
between the left and right bank.
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The distribution and the abundance of hydrophytes
along the Danube River in Serbia
This study provides a basis for studying the
influence of large power plant impoundments on the
aquatic vegetation of rivers in more detail. Also it will
be important to survey other large rivers in Serbia and
in surrounding region using the same methodological
approach so that the obtained results could be
comparable.
done in frame of the project No. 143037, financed by
the Ministry of Science and Environmental Protection,
Republic of Serbia. Authors would like to thank Prof.
Dr. Georg A. Janauer and Mr. Norbert Exler, from the
Department of Limnology and Hydrobotany, University
of Vienna for the help in processing and discussing the
data.
Acknowledgements
Basic data for this study was collected during the
realization of the MIDCC project funded by the Austrian
Federal Ministry of Education, Science and Culture
(bm:bwk, www.midcc.at), while further analyses was
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Moog O., Stubauer I, et al., Developing the tipology
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