Accepted Manuscript

Title: Long-term changes in copepods assemblages in the area

of the Danube floodplain (Slovak–Hungarian stretch)

Authors: Marta Illyová, Pavel Beracko, Marián Vranovský,

Igor Matečný

PII: S0075-9511(16)30186-4
DOI: http://dx.doi.org/doi:10.1016/j.limno.2017.05.001
Reference: LIMNO 25584

To appear in:

Received date: 9-11-2016

Revised date: 6-3-2017
Accepted date: 3-5-2017

Please cite this article as: Illyová, Marta, Beracko, Pavel, Vranovský,
Marián, Matečný, Igor, Long-term changes in copepods assemblages in
the area of the Danube floodplain (Slovak–Hungarian stretch).Limnologica
http://dx.doi.org/10.1016/j.limno.2017.05.001

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Long-term Changes in Copepods Assemblages in the Area of the Danube Floodplain
(Slovak-Hungarian Stretch)

Marta Illyováa, Pavel Berackob, Marián Vranovskýc, Igor Matečnýd

a
Plant Science and Biodiversity Centre, Institute of Botany, Slovak Academy of Sciences,
Dúbravská cesta 9, 845 23 Bratislava, Slovakia. email: marta.illyova@savba.sk

b
Department of Ecology, Faculty of Natural Sciences, Comenius University in Bratislava.
Slovakia. email: beracko@fns.uniba.sk

c
Račianska 95, 831 02 Bratislava, Slovakia. email: marian.vranovsky@gmail.com

d
Department of Physical Geography and Geoecology, Faculty of Natural Sciences, Comenius
University in Bratislava. Slovakia.

email: matecny@fns.uniba.sk

Abstract
The present study investigates the relationship between hydrological connectivity and species
diversity in a by-passed channel section, and in the adjacent water bodies of the Slovak-
Hungarian Danube section (1,840.5-1,804.4 rkm). The study was designed to assess long-term
temporal trends (from 1991 to 2013) in freshwater copepods assemblages and their ecological
indices in different habitats of the Danube floodplain area. One of the purpose of this study
was to monitor the species composition of copepods communities and ascertain their shift in
various biotopes of the Danube floodplain system. Based on 23 years data, in the first step,
the copepods habitat preferences using habitat values (HV) and indicator weights (IW),
calculated from data collected over all the years of monitoring of planktonic communities of
the Danube floodplain, were quantified. Subsequently, the floodplain index (FI) from a
summary of the habitat values and indication weights of the current species was calculated, to
evaluate changes in the connectivity of the anabranched section of the Slovak-Hungarian
Danube. This confirmed the loss of active hydrological connectivity within the main river
channel, ranging from the eupotamal to more or less isolated floodplain water bodies. Out of
50 recorded copepods species, 11 species manifest a preference for eupotamal habitats, 18
taxa preferred eupotamal B/parapotamal habitats and 21 species were found to prefer the
plesiopotamal habitat type. The statistical analyses demonstrated that the structure of
planktonic copepod communities in this area has changed since the Gabčíkovo hydropower
plant was placed into operations. The NMDS analysis revealed shifts in the proportion of
euplanktonic and tychoplanktonic species. Reversible community changes were found in the
old river bed and in the eupotamal-B side arms.

Keywords: Biological assessment, Floodplains, Copepoda, Connectivity, Index

1. Introduction

In recent years, a great deal of theoretical and applied limnological research has focused
on studies of floodplain rivers (e.g. Amoros and Roux, 1988; Junk et al., 1989; Ward and
Stanford, 1995; Ward et al., 2002; Fantin-Cruz et al., 2016) and in particular the importance
of lateral connectivity and exchanges between rivers and floodplains (Chovanec et al., 2005).
Damming for hydroelectric power can have a significant negative impact on the hydrological
relationship between a river and adjacent water bodies (Ward et al., 1999; Pringle et al.,
2000), with the consequences including fragmentation of lateral connectivity and exchange
pathways, reduction of habitat heterogeneity and declining biodiversity (Tockner et al., 1998).
The current study investigates changes in the hydrological regimes of water bodies in the
bypassed river-floodplain system. The floodplain index approach, developed by (Waringer et
al., 2005), assesses the ecological status on the basis of the habitat preferences of species
present. Previous schemes for assessing floodplain areas have focused on representatives of
various aquatic fauna, e.g. dragonflies (Chovanec and Waringer, 2001; Chovanec et al.,
2004), caddisflies (Waringer and Graf, 2002) and other indicator groups (Chovanec et al.,
2005; Waringer et al., 2005). We present herein a new scheme developed to assess the
hydrological status of a bypassed section of the Danube using long-term monitoring data
pertaining to communities of planktonic copepods. Although not listed as an indicator group
by the EU Water Framework Directive, Copepoda are characterized by a strong affinity to a
habitat type based on their feeding preferences (Gaviria et al., 2002a, 2002b, 2002c), play a
pivotal role in the trophic chain (Dussart and Defaye, 1995) and are already used as indicators
of the saprobic index (Sládeček, 1964). All this led us to expect that monitoring of copepods
assemblages would prove useful in assessing the ecological integrity of the river-floodplain
system. The floodplain index, as an excellent tool for indicating landscape level hydrological
changes, was applied to other representatives of water fauna in the Danube river inundation
area (Illyová and Matečný, 2014; Beracko et al., 2016; Šporka et al., 2016). Investigation of
Copepoda fauna has a long tradition for the middle section of the Danube River and adjacent
water bodies, as part of the study of zooplankton or planktonic crustaceans, on the left site in
Slovakia (e.g. Vranovský, 1974, 1985, 1991, 1995a, 1995b; Illyová and Némethová, 2002),
and on the right site, in Hungary (Bothár, 1973, 1979; Bothár and Ráth, 1994; Gulyás, 1995;
Kiss et al., 2015). Intensive hydrological investigation in the region began in 1990 and has
continued up to the present in order to monitor the environmental impact of the river
regulation. As a result of a barrage system implementation at Gabčíkovo, a number of
ecological changes occurring in aquatic communities have been predicted (Holčík et al.,
1981). Longtime extensive environmental monitoring of the water biota has been
implemented to optimize the water regime in the affected area (Mucha, 1995, 1999; Lisický
and Mucha, 2003). Over the years, changes in the species composition and the dominance
hierarchy of planktonic crustacean assemblages have been observed after damming, both at
the Slovakian left bank (e.g. Illyová, 1996; Vranovský, 1995b, 1997; Illyová and Némethová,
2005) and at the Hungarian right bank in the Szigetköz floodplain (Bothár, 1988, 1998;
Gulyás, 1994; Kiss, 2004; Kiss et al., 2014).

The purpose of this study was to monitor the species composition, diversity, and dominance
of copepods communities and ascertain their shift in various biotopes of the Danube
floodplain system over a 23 years period. A secondary aim was to use the calculated values to
assess the ecological integrity of the floodplain system and to evaluate changes in the
connectivity of the anabranched section of the Slovak-Hungarian Danube using floodplain
index (FI) calculation based on the Copepoda community.

2. Material and methods

2.1 Study area

The study section is situated in the upper part of the Slovak-Hungarian section of the
Danube River, on the left-bank floodplain (1,840.5 - 1,804 rkm). The study area is a once
pristine, anabranched section of the Danube characterized by hydrologically dynamic side
arms, now substantively bypassed by the head- and tail-races of the Gabčíkovo hydropower
plant. The area incorporates a gradient of laterally connected habitat types ranging from
highly dynamic waters in the main channel, through less dynamic waters of the side arms, to
stagnant zones. The main area of interest was a bypassed section between the old Danube,
with an anabranched system of side arms and the new Danube (Fig. 1, Table 1). The typology
of the habitat analysis was based on the ´functional sets´ concept (Roux et al., 1982; Amoros
et al., 1987). We used a modified system (eu/parapotamal A, B) which is system of
Hohensinner et al. (2005). They used this definition in the Austrian section of the middle
Danube, in the natural anabranched-type section, where followed with minor modifications
the system of Amoros et al. (1987).

The comprehensive characteristics of the monitoring sampling localities have already been
published by Vranovský (1997) and Illyová and Némethová (2005). The general channel
types present in the study area are as follows:

Eupotamal-A

The wide principal channel of the old Danube, from the reservoir to the confluence with the
tail-race channel, conforms to the eupotamal-A type. Since damming, the main channel
between rkm 1851 and 1811 has been referred to as the old river bed. The average annual
discharge in the bypassed section (measured at rkm 1,848.4) has decreased from 1,900 m3s-1
prior to the diversion to about 350 m3s-1 in 1993 and 220 m3s-1 in 1994. The flow rate in the
old river bed during normal operations at present is only 250 in winter and 600 m3s-1 in the
growing season (Mucha, 1995). Maximum current velocities decreased from 2.0-3.5 ms-1 to
around 1 ms-1. In the upper part of the old Danube (rkm 1,840.5), however, the decrease in
water level was even more significant than in the lower part (rkm 1,819.5). The bottom is
composed of gravel and littoral without macrophytes.

Eupotamal-B

The two main branches, originally parapotamal-type side arms situated in the upper part of the
bypassed region (rkm 1,830), i.e. upstream from the Gabčíkovo power plant, are classified as
eupotamal-B. Prior to damming, these branches were connected with the main channel at their
downstream ends. When the river was diverted, this direct connection with the former main
channel was abolished, and as of May 1993, the main branches of the side arm system
between rkm 1,840 and 1,820 have been fed from the bypass canal by means of an intake
structure built at Dobrohošť village. Currently, due to artificial feeding, the arms flow
permanently. The artificial water replenishment from the main arm of the Danube River has
been providing a permanent water supply to the branch-system with a capacity of 40-180 m3s-
1
, although the major connections between the main arm and the branches have been blocked
by weirs (Guti, 2002). The channels are subject to permanent aggradations of fine-grained and
clay sediments. Due to the sluggishness of the current, macrophytes biomass has increased
significantly in the littoral zone.

Parapotamal-A

Parapotamal-A side arms are relatively dynamic backwaters joined by the downstream outlets
of previous branching of the channel system. The partially abandoned side arm near Istragov
is situated 5 km upstream from the confluence of the tail-race canal and the old Danube (rkm
1,815.5). Prior to damming, its maximum depth varied from approximately 1.5 to 5.6 meters,
although it underwent significant swallowing after the main flow was diverted. It can still
flood, however, along with the surrounding floodplain, in periods of high discharge when
water backs up from the bypass-old Danube confluence at rkm 1,811. Its bottom is composed
of gravel, in which a limited macrophyte community develops during the stagnant period,
although the contribution of macrophytes is low.

Plesiopotamal

One plesiopotamal-type arm (KL) is relatively deep (up to 3 m), with isolated abandoned
channels or floodplain lakes, which in the context of the current study area are only flooded
when levels in the old Danube channel are high (about 650 cm at the Dobrohošť gauging
station). Connectivity with the old river channel is gradually weakening as flooding declines
in frequency. Two plesiopotamal-type side arms (SS and SLE) are located approximately 7
km downstream from the confluence of the power plant tail-race canal with the old river bed,
i.e. outside the section directly affected by the power plant. The localities are less than 1 m in
depth, have a muddy bottom and rich macrovegetation. They are flooded in periods of high
discharges as well as after the damming.

The geographical coordinates for each site are DK: 47.987139N, 17.343854E; GAB:
47.861314N, 17.533889E; BB: 47.921151N, 17.442008E; SUL: 47.9214889N, 17.4419389E;
IST: 47.8283961N, 17.559711E; KL: 47.903632N, 17.488698E; SS: 47.7863450N,
17.6789231E; SLE: 47.7650131N, 17.7269750E.

2.2 Sampling procedure and data analysis

Qualitative samples of copepods assemblages from eight sampling sites were collected
from 1991 to 2013 and analyzed three times a year: spring, summer and autumn. The
qualitative samples were collected from the medial, open-water zones, using a 70 µm mesh
plankton net, either from a boat by vertical tows or from the bank by tows from the bottom to
the surface. Samples of littoral plankton were also obtained. All the samples were preserved
in 4% formaldehyde. The adult and final copepod stage were taken into consideration in
dominance determination. All of the 787 samples collected and analyzed over the 23 years
period (15 years from SLE) of investigation, 496 were taken from the medial zone and 291
from the littoral zone.
Floodplain index (FI), which indicates the extent of disruption to lateral connectivity in
the floodplain area, was calculated from the ecological indices. Floodplain index values were
calculated for each study site and every year of monitoring according to Waringer et al.
(2005), using the equation: FI = Σ (HV*IW)/ Σ IW, where HV is the habitat value and IW is
the indication weight of all the species present.
The calculation of HV required a list of the Copepoda species recorded in the Danube
floodplain area and the frequency of their occurrence (%). The first step was the valency point
calculation. In order to describe the habitat preferences numerically, a 10-point system was
used, in which, for example, to a species exclusively inhabiting temporary pools would be
given 10 valency points in the relevant category. For eutrophic species occurring in a wide
range of waters of the floodplain system, the 10 points will be more or less evenly distributed
between different categories (Chovanec and Waringer, 2001). 10 valency points were
distributed among four habitat types (H1-H4) based on the records from the investigated area.
For the valency point calculation, averages from the three samplings in each year (283
records) were taken into account.
The habitat type assessment is based on the frequency of occurrence (%) of Copepoda
species in the river-floodplain system within specific habitat types. These habitat types (H1,
H2, H3 and H4) follow a gradient of lateral connectivity with the main river channel, ranging
from H1 (eupotamal A), H2 (eupotamal B), H3 (parapotamal), to more isolated floodplain
waters in H4 (plesiopotamal).
Individual species-specific habitat values (HV) were then calculated according to
Chovanec and Waringer (2001) and Chovanec et al. (2005), with the equation: HV =
(1*H1+2*H2+3*H3+4*H4)/10. Where H1-H4 are individual habitat types to which valency
points are assigned based on the occurrence (%) of river-floodplain Copepoda species within
each specific habitat (H1-H4; H1 eupotamic, that is high connectivity between floodplain
waterbody and main river, H4 – that is no connectivity and terrestrialisation process).
Indication weights (IW) range from 1 for eurytopic species to 5 for stenotopic species
have been allocated to each species in order to identify sensitive species. The gradient of the
indicator features ranging from excellent indicator (5), mean indicator (3), to weak indicator
(1). The allocation of indication weights was carried out according to the scheme of Sládeček
(1964) and the ratios were calculated using values of occurrence (%) for each species in the
habitat.

2.3 Statistical analysis

Correspondence analysis (CA) on log(x+1) transformed dominance data was used for
determination of community positions in ordination space over the study period. There were
the medial samples used for the statistical analyses. Significant changes in the copepod
community between the two following years were identified by a cluster analysis based on
chi-squared distance. A significant change meant that these years were clustered amongst the
significantly different clusters. The significance of clusters was tested with a permutation test
with 999 permutations and at a 0.05 significance level. Cluster analysis and correspondence
analysis were performed by “Clustsig” library (Whitaker and Christman, 2010) and “Vegan”
library (Oksanen et al., 2012) in the R 3.1.0 (R Core Team, 2014; Killick and Eckley, 2014)
software environment.

3. Results

3.1 Copepoda community composition and species-specific habitat preferences

50 taxa of Copepoda (33 Cyclopoida, 12 Harpacticoida and 5 Calanoida), were identified

in total over the study period from 1991 to 2013 in eight localities. The number of species in
the individual years and individual sampling stations is shown in Fig. 2. A list of species
including species-specific habitat valencies, habitat values, and indication weights is provided
in Table 2. The highest dominancy was achieved by the species Thermocyclops oithonoides
(23%), throughout the investigated area, followed by Eurytemora velox, Acanthocyclops cf.
robustus, Nitocra hibernica, Eucyclops serrulatus and Eudiaptomus gracilis.
11 of the recorded copepods species manifested overall preferences for habitat H1,
eupotamal-A, with HV value ranging from 1 to 1.6 (Table 2). Five species, two cyclopoids
Diacyclops languidoides, D. crassicaudis, and three harpacticoids Bryocamptus (B.) mrazeki,
Bryocamptus (Rh.) zschokkei and Echinocamptus pilosus were recorded exclusively in the
main channel. As many as all 11 species in this group belong to the littoral and benthic
species, with three quarters of them being harpacticoids (Table 2). Out of representatives of
cyclopoids, only two species of Diacyclops and two species of Paracyclops (P. fimbriatus and
P. poppei) revealed a higher affinity to the eupotamal-A habitat type.
There was a considerable overlap in the species composition of H2 and H3, between the
eupotamal-B and parapotamal habitat types. All in all, 18 recorded species of Copepoda
preferred these habitats, with HV values in a range from 2.0 to 3.2. Half of these species were
a pelagic and herbivorous one: Cyclops vicinus, Eudiaptomus gracilis, Eurytemora velox,
Diacyclops bicuspidatus, Acanthocyclops cf. robustus, Thermocyclops oithonoides,
Thermocyclops crassus and Mesocyclops leuckarti. With the possible exception of
Elaphoidella bidens, no species appeared to exhibit any exclusive affinity to the
eupotamal/parapotamal channel type. A higher affinity of the eupotamal habitat type was seen
with Nitocra hibernica. Genus Eucyclops, with five recorded taxa, was the most frequently
recorded in these side arms.
Overall, 21 copepods species were found to prefer habitat H4, the plesiopotamal type,
with HV values ranging from 3.4 to 4. As many as 12 species were recorded exclusively in
these habitat types, achieving valency point scores of 10 (Table 2). Almost all the species
associated with the plesiopotamal belonged to phytophilous or benthic taxa. Only one species,
Macrocyclops distinctus, was recorded in all three plesiopotamal side arms. Two species,
Eudiaptomus transylvanicus and Mixodiaptomus kupelwieseri, were only recorded in Sporná
sihoť (site 7) and Metacyclops minutus and Elaphoidella gracilis were recorded exclusively in
Kráľovská lúka (site 4).

3.2. Indication of habitat types and the differences between the sampling sites

Significant overlap in FI values between same habitat types causes some difficulty in
assigning them confidently to a particular type.

Changes in the FI values recorded over 23 years (or 15 years in case of SLE) at the
monitoring sites indicates Fig. 3.
A decrease in FI values (Fig. 3 DK; Fig. 3 GAB) was recorded in the H1 eupotamal
habitat type. This trend reflects a declining proportion of euplanktonic species in the copepods
community in the second decade of the reporting period. This reflects the fact that the
copepods community profile has shifted to one adapted to flowing water. Benthic littoral
harpacticoids appeared in the potamoplankton and their relative abundance has increased over
this time.
The FI values calculated for the H2 and H3 sites documented distinct differences in the
hydrological regimes of floodplain river arms situated upstream and downstream of the port
of Gabčíkovo. The FI value of the arms situated upstream indicate an unstable hydrological
regime because the arms are artificially supplied with water. Over the course of the first years
after damming, the FI values decreased and responded to the eupotamal-B habitat type. The
trend involving an increase in FI values is well illustrated for both eupotamal type side arms,
especially in the second half of the monitored period (Fig. 3 SUL; Fig. 3 BB). This means that
the species richness of phytophilous species increased. The parapotamal side arm situated
downstream (Fig. 3 IST) is not artificially supplied with water, but is sporadically (and
continually less) influenced by water backing up from the confluence of the old Danube and
the Gabčíkovo tail race. The increase in FI after 1997, the decrease in the connectivity of this
arm and the main river channel are illustrated.
The decrease in connectivity between the main river channel and the floodplain area is
also well documented by the trend of increasing FI values in the plesiopotamal habitat type
(Fig. 3 KL, Fig. 3 SS and Fig. 3 SLE). Dense macrophyte growth has contributed to an
increasing proportion of phytophilous copepods in the communities and species usually
associated with temporary shallow waters were recorded.
Based on changes in the copepods community and FI values calculated during the
monitoring period, long-term trends can be determined and a decrease in connectivity in the
hydrological regime for the entire area (Fig. 4) nested F (3,4) = 30.8, p< 0.01.

3.3 The temporal trajectories and species composition changes

The spatio-temporal analysis of copepods communities showed that the first axis of
correspondence analysis represent difference between eupotamal sites and para-plesiopotamal
sites (Fig. 5). The second axis of correspondence analysis reflects the heterogeneity in para-
plesiopotamal sites. During the study period, the highest changes of copepod communities
were observed at the eupotamal sites, while the lowest at the plesiopotamal sites.

3.3.1 Main channel/Old river bed of the Danube

The main community changes were led in two different ways in the upper part of the
main channel (Fig. 6, DK). The first (vertical) way represented reversible changes in the time
period between 1991 and 1999. The shift in proportion of euplanktonic in favour of the
tychoplanktonic species were recorded between 1992 and 1993. A reversible change took
place between 1997 and 1998. This occurred with the “euplanktonic period” species such as
A. cf. robustus, E. gracilis, E velox, C. vicinus and both Thermocyclops species, while in
“tychoplanktonic period” an increase in abundance in Eucyclops serrulatus, N. hibernica and
Paracyclops fimbriatus was observed. The second (horizontal) way represented the time
period between 1999 and 2006, when partial taxonomic change of the copepod community
(with some returns to the original community) took place. In this period Ectinosoma abrau
and Eurytemora velox began to appear regularly in the copepod community. Euplanktonic
species were gradually eliminated as of 2006 and the relative abundance of tychoplanktonic
species N. hibernica and E. abrau increased.
Two different periods can be determined, from 1991 to 2004 and from 2005 to 2013, in
the lower profile of the main channel (Fig. 6 GAB). During the first period, similar to the
upper profile, the euplanktonic species A. cf. robustus, E. velox, C. vicinus, Th. oithonoides
and T. crassus dominated. During the second period, the dominance of all euplanktonic
species decreased, apart from E. velox, and the dominance of both harpacticoids N. hibernica
and E. abrau increased. Certain phytophilous and benthic species (E. serrulatus, E. speratus,
E. macruroides, E. macrurus, P. affinis, P. fimbriatus and Macrocyclops albidus) were
identified in both profiles of the main channel (DK and GAB) over the years from 1996 to
2004. The highest number of species was recorded at that time (Fig. 2), but after 2004, a
number of these species disappeared, subsequently a decrease in the number of species was
recorded.

3.3.2 Eupotamal-B/Parapotamal (partially abandoned) type side arms

The major community shift from the period before damming (1991, 1992) up to 2013 is
well documented in the both eupotamal-B side arms situated in the upper part of the by-
passed region (Fig.6 BB; Fig. 6 SUL). Prior to damming, euplanktonic species T. oithonoides,
dominated, followed by C. vicinus and E. gracilis. Subsequently, four years after damming, a
decrease in pelagic species was recorded and the harpacticoid N. hibernica dominated. Over
the next nine years (from 1996 to 2005), euplanktonic species E. velox, less A. cf. robustus,
dominated. All these euplanktonic species disappeared from the medial zone after 2010 and
only tychoplanktonic species were present. Mainly harpacticoids N. hibernica and E. abrau,
cyclopoids Eucyclops serrulatus and E. macruroides dominated out of them.
In the parapotamal side arm (Fig.6 IST) no noticeable trend in a shift in the copepod
community was recorded. Prior to damming (1991, 1992), and two years after damming
euplanktonic species, E. gracilis, E. velox, C. vicinus and A. cf. robustus dominated. Over the
following years, the relative abundance of T. oithonoides increased. All these euplanktonic
species occurred in the medial zone of this arm about thirteen years, but after 2005 their
relative abundance gradually decreased, apart from E. velox and E. gracilis (in 2009) and the
dominance of tychoplanktonic species N. hibernica, Eucyclops speratus, E. serrulatus and M.
albidus slightly increased.

3.3.3 Plesiopotamal type side arms

A weak species copepods community shift was recorded in the deeper, more isolated
plesiopotamal type side arm (Fig. 6 KL), because euplanktonic species dominated over all the
monitored period. Prior to and shortly after damming, T. oithonoides, E. gracilis, E. velox, C.
vicinus and T. crassus reached a higher relative abundance. Followed a twenty-year long
period, T. oithonoides dominated and the species richness and higher relative abundance of
tychoplanktonic species, E. serrulatus, E. macruroides and E. macrurus, increased in the
medial zone.
In the shallow plesiopotamal side arm with a fluctuation in the water level (Fig. 6 SS), a
shift in copepods communities was recorded over two periods. During the first period (from
1991 to 1999), euplanktonic species E. gracilis, Cyclops vicinus, T. oithonoides and
Mesocyclops leuckarti, dominated. During the second period (from 2000 to 2013), a reach
community of phytophilous species was recorded at this site: five species of Eucyclops, three
species of Macrocyclops (M. albidus, M. fuscus and M. distinctus), and some rare species
Cyclops heberti, Diaptomus castor, Tropocyclops prasinus, Thermocyclops dybowskii,
Eudiaptomus transylvanicus and Mixodiaptomus kupelwieseri. A higher relative abundance
was also reached for E. velox, E. serrulatus, T. crassus and T. oithonoides.

4. Discussion

Once the hydropower plant was placed into operations, two major changes in the
hydrological regime of the Danube floodplain area were recorded: the discharge in the old
main channel declined and the water level in the adjacent side arms started fluctuating (Krno
et al., 1999). The direct impact of Danube damming includes morphological changes to the
river banks and oxbow lakes, changes in river sediment transport, discharge and water level
changes in the river and the side arms as well as the alterations in the lateral and longitudinal
hydrological connectivity of the floodplain water bodies (Mucha, 1995; Farkas-Iványi and
Guti, 2014). Based on the Hungarian Danube section, the active connection between the
abandoned Danube stretch and the side-arm system in the floodplain disappeared and the
water supply of the protected floodplain was materialized through an artificial water recharge
system (Kiss and Schöll, 2009). Damming of the river has had a negative impact on the
hydrological relationship between the river and adjacent water bodies (Pringle et al., 2000),
with consequences including the disruption of lateral connectivity. The importance of
connectivity between aquatic habitats in the river floodplain systems is well documented
(Tockner et al., 1998). The floodplain index is useful in characterizing the long-term
hydrological changes occurring in the floodplain area (Chovanec and Waringer, 2001;
Chovanec et al., 2004, 2005). This multi-species approach was used to evaluate the changes
occurring in the bypassed section of the Danube and adjacent water bodies since the
activation of the Gabčíkovo hydroelectric dam. Quality indicators of long term change in
hydrological connectivity are seen with cladocerans (Illyová and Matečný, 2014), molluscean
fauna (Beracko et al., 2016), or macroinvertebrates of the Danube floodplain area (Šporka et
al., 2016).
As a result of the damming of the Danube in October 1992, the main channel of the river
between km 1,851 and 1,811 has become an old river bed. Since the diversion, the average
annual discharge of the bypassed section has decreased significantly (CEC, 1992), the water
level has decreased and the flow velocity slowed from 2.0-3.5 to 1 m s-1 (Vranovský, 1997).
The average winter flow rate was reduced from about 2,025 m3 s-1 to 250 m3 s-1 and the
summer flow rate dropped to 600 m3 s-1. Averaged over a year, the new flow rate is around
400 m s-1. At the same time, there was a significant narrowing of the old river channel, and the
water level dropped significantly by several meters in the upper part of the study area.
Connectivity between the main channel and the arm system was cut and flooding ceased over
the entire former inundation area. These factors have led to long term changes in the
Copepoda community at the entire monitored area.
These changes consequently changed the trophic relations in the eupotamal habitat
type/old river bed of the Danube. Loss of the lateral connectivity and the missing lentic
habitats in the littoral zone of the main channel as consequences of a decrease in water level
and deceleration of the flow to the main riverbed were in all probability responsible for the
disappearance of the planktonic crustacean community (Illyová and Némethová, 2005).
A decline in dominance of E. gracilis, E. velox, A. cf. robustus was recorded in the first years
after damming (Gulyás, 1994; Vranovský, 1995b, 1997), with the littoral species N.
hibernica, E. abrau, E. serrulatus and others, achieving dominance in the potamoplankton
over the long term. Fine sediments accumulating in gravel beds created a suitable biotope for
species of harpacticoids (Nitocra, Ectinosoma) and Paracyclops which are most often grazers
and are predominantly detritivores (Dussart and Defaye, 1995). In contrast, the disappearance
of limno-rheophilic and limnophilic species from the old main channel was recorded in
communities of snails (Beracko et al., 2016), and similar changes were observed in the fish
community of the main channel of the Volga River after damming (Górski et al., 2012).
Significant changes in copepods community composition and in the ecological parameters
of communities were observed at the partially abandoned side arms between the river km
1,840 and 1,820. The greatest changes were observed in the upstream part of the Danube arm
system, which is fed artificially with water from the head-race channel, which provides a
continuous minimum flow of 20 to 30 m3s-1, and has been tested to 140 m3s-1 to simulate
flood conditions. Prior to damming, the local copepods assemblages of the medial zone
comprised almost all euplanktonic species (Vranovský 1974, 1985). Over the following years,
the number of euplanktonic Copepoda species dropped and the number of tychoplanktonic
ones increased as a consequence of the artificial system. This trend was observed for
cladocerans assemblages in the early years after damming (Illyová and Némethová, 2005) and
continues up until the present. Prior to damming, these arms were flowing at a discharge
higher than about 2,040 m3 s-1 (Vranovský, 1997) and were stagnant at a lower discharge.
These were good conditions for the development of euplanktonic fauna. Due to artificial
feeding they currently flow permanently. Additionally, a littoral zone was covered with
macrovegetation and the dominance of the macrophyte-associated species (Eucyclops,
Macrocyclops and Megacyclops) increased.
A very different situation was recorded in an arm not artificially supplied with water from
the bypass canal (Istragov, Site 5). This arm became shallow after the main flow was diverted,
and partially lost connectivity with the main channel. The conversion to a stagnant
hydrological regime is well documented by the copepods community, with calculated FI
values increasing over the monitoring period. This arm is steadily becoming shallower after
diverting the Danube. Euplanktonic species, primarily Thermocyclops oithonoides,
predominated there almost over the entire period of monitoring. Similarly, Kiss (2006) found,
that Thermocyclops oithonoides has the highest relative abundance in the parapotamal side
arms. This is related to the lentic character of the habitat and poor littoral macrovegetation.
Over recent years, we have noticed a slight growth in the relative number of tychoplanktonic
species, which is probably caused by shallowing in the arm in recent years, since the arm is
not well connected at the surface with the main channel, with the exception of the floods in
the years of 1997-2003.
Changes have been recorded in the plesiopotamal-type side arms, where the proportion of
phytophilous species in the copepods community has increased. The decrease of the
dominance of filter feeding copepods in plesiopotamal wetlands was caused by the decrease
of phytoplankton community. Considering the much increase of littoral macrophytes cover, it
is possible that food also may have played some role in our case. Herbivorous calanoid,
Eudiaptomus gracilis was dominant in the arms type plesiopotamal over the years 1991-1992.
At this time, there was in all probability a higher occurrence of phytoplankton (Štefková,
1998), due to more frequent connection of the arms to the side arms of the inundation, so
more remote arms also received nutrients (Thomaz et al., 2007). After damming, these arms
lost their periodical connections with the adjacent arm-system. It has been found, that the
average phytoplankton abundance in the isolated lake was over two times lower than in the
lake connected to the river (Dembowska and Napiórkowski, 2015). This situation has
developed due to a decrease in its depth and increase of macrophyte vegetation grow. Dense
macrophyte vegetation has supported the increase in the phytophilous copepods. There were
also rare species recorded which inhabit eutrophic shallow waters, e.g. Cyclops heberti,
Mixodiaptomus kupelwieseri, Diaptomus castor, or Tropocyclops. In the water-bodies densely
overgrown with stands of macrophytes, the number of euplanktonic species was low.
According to Gulyás (1994), mainly species living among the vegetation, or in the surface
layer of the bottom sediment were found in this type of water-body.
There has been a clear profiling of habitat valency of the species of certain types towards
a certain type of habitat in the copepods community. We have noticed exclusively three
species of harpacticoids and two species of genus Diacyclops in this type of flow. Truly
interesting in the occurrence, since the languidus-languidoides are stygobionts or
stygophilous species, is that they are adapted to an interstitial and mossy environment (Pesce,
1994). A higher affinity to the flowing water was revealed for two Paracyclops species. The
high affinity of Paracyclops fimbriatus to the main channel of the Danube River was
confirmed by additional authors (Gulyás, 1994; Kiss and Schöll, 2009; Kiss et al., 2014). The
occurrence of these species in potamoplankton of the Danube is related to the food demands,
since P. fimbriatus is a grazer and D. languidoides is detritivore (Gaviria et al., 2002a).
Harpacticoids are also most often grazers and predominantly detritivores. They eat what they
find on the bottom, mainly bacteria, Protozoa, decomposing plant and animal matter (Dussart
and Defaye, 1995; Gaviria, et al., 2002c). In the arms type eupotamal-B and parapotamal, the
largest occurrence of the species was of phytoplankton filtrators. Representatives of the
genera Acanthocyclops, Thermocyclops, Mesocyclops, Eudiaptomus, are active filter-feeding
copepods of phytoplankton. (Gaviria et al., 2002b). They have the highest food offering of
algae in this type of habitat. During the higher water levels and floods, under ideal conditions,
these biotopes are at the surface interconnected with the main channel. The arms are provided
with nutrition which promotes the development of phytoplankton (Thomaz et al., 2007). Most
of these species in case of an insufficiency of food become predators (Dussart and Defaye,
1995). Exclusively in the arm type plesiopotamal, rare species occurred which inhabit
eutrophic shallow waters, e.g. Tropocyclops prasinus, Diaptomus castor and Mixodiaptomus
kupelwieseri. In addition, there were species Cyclops heberti, typical for ephemeral ponds
(Einsle, 1996).
A total of 50 taxa of Copepoda were encountered over the 23 year period in the Danube
floodplain area (1,840.5 - 1,804 rkm). At the beginning of the monitoring period, Vranovský
(1997) recorded 30 copepods species. Over long term monitoring, the same rare species
(Cyclops heberti, Diaptomus castor, Mixodiaptomus kupelwieseri, Tropocyclops prasinus,
and Tisbe furcata) were found. Among the neozoans, harpacticoid Ectinosoma abrau
Kričagin and alien calanoid Eurytemora velox Lilljeborg enriched the potamoplankton of the
Danube. This may be related to the stoppage of the main channel, similar to that of fish (Liew
et al., 2016). In the various water-bodies of Szigetköz, Gulyás (1994) recorded 29 species (6
Calanoida and 23 Cyclopoida) and later Kiss (2004) recorded 26 species. The most frequent
species in this area were Acanthocyclops cf. robustus and Eucyclops serrulatus. Later, during
a long-term (1999-2008) zooplankton investigation focused on three parts of the Hungarian
Danube section (including Gemenc), 25 copepods species were identified (Kiss and Schöll,
2009), although the latest data concerning this area documented a slight decline in 18 species
(Kiss et al., 2014).

5. Conclusion

The aim of this study was an examination of the influence of changes to the hydrological
regime on the copepods community over an extended period. The results indicate that
Copepoda species are good indicators of long-term changes in hydrological connectivity.
Although not listed as an indicator group in the European Union Water Framework Directive,
copepods are revealed to be apposite in the ecological assessment of the river-floodplain
system. Floodplain index (FI), based on the combined habitat values and indication weight of
all the Copepoda community, confirmed the dramatic rates of habitat degradation resulting
from loss of an active connection between the abandoned main channel of the Danube and the
side-arm system since the construction of the Gabčíkovo dam.

Acknowledgements

The study was supported by the project VEGA 1/119/16, VEGA 2/0030/17 and ITMS:
26240220049. We are also grateful to Dr. David Livingston (UK) for correcting the English
and for valuable suggestions. We are also thankful to both anonymous reviewers for their
invaluable criticism, which markedly contributed to the improvement of this manuscript.
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Figure 1.
Map of the Gabčíkovo barrage system with studied sites indicated (1-DK, 2-BB, 3-SUL, 4-
KL, 5-GAB, 6-IST, 7-SS, 8-SLE).
Figure 2.
The number of species at each site over the years 1991 – 2013; DK-site 1, GAB-site 5, BB-
site 2, IST-site 6, KL-site 4, SS-site 7.
Figure 3. Changes in the floodplain index values recorded over 20 years at the monitor sites
(H1: DK, GAB; H2: BB, SUL; H3: IST; H4: KL, SS, SLE.
Figure 4. Comparison of values in Floodplain indices between the studied habitats. Nested F
(3,4) = 30.8, p< 0.01.
Figure 5. CA (correspondence analysis) ordination plot based on log-transformed abundance
data. The position of square represents polygon centroid for each site. The size of square
showed the scaled size of polygon area (within site heterogeneity in species composition).
The number in legend represents the real size of the polygon area.
Figure 6. Results of a correspondence analysis (CA) showing the trajectories of shift in
species composition of the copepod community at each sampling site. The log(x+1)
transformed abundance data. Dashed lines connect the years from a significantly different
cluster from a cluster analysis based on chi-squared distance. The significance of clusters was
tested in a permutation test with 999 permutations and at a 0.05 significance level.

DK TI= 3.94 DK Bpyg

2
2

12
Nhib
1

1
10
CA2 (14.39%)

11 94
07 96
03 AcraEpha

CA2
97 93
09 95
08 Mleu
Tcra
0

Pfim

0
Eser Arob
MvarDbic
02 Epro
01 92 Malb Emac
00 05 99 91 EgraToitCvic
04 98 Eabr Bmra Epil
Evel
-1

-1
06
13

-3 -2 -1 0 1 2 -3 -2 -1 0 1 2
CA1 (15.25%) CA1
2.0
2.0

GAB TI= 4.69 GAB

Eser
Espe
1.5

1.5

Dcra
94
1.0

1.0

99 Emac
CA2 (10.77%)

91 Arob
08 Cvic
0.5

Tfur Tcra
0.5

96 02 Dlan Malb
CA2

93 97 04 Pfim Dbic Mleu

03
0.0

0.0

98 95 Csta
01 Nhib
07 13 05 Paff Toit
BpygEvel
-0.5

-0.5

09 00 10 Mvar
Emaci Eabr
12
11
-1.0

-1.0

06 Epro
92
Egra
-1.5

-1.5

-3 -2 -1 0 1 2 3 -3 -2 -1 0 1 2 3
CA1 (11.94%) CA1
 SUL TI= 7.08 SUL Pfim
2

2
Epro

Cbic
DCA2 (10.00%)

Tcra
1

1
Cstr
07 Eser

DCA2
96 11 09 Csta
Egra Malb
93 94 04 12 13 Nhib Emaci
95 08 10 06 Mleu Espe
05
0

0
91 97 Emac
92 01
02 99 Cvic
03 Mrub
Acra Arob
00
Eabr
-1

-1
Toit
98
Ebid Evel
-2 -1 0 1 2 3 4 -2 -1 0 1 2 3 4
DCA1 (11.09%) DCA1
1.0

1.0
BB TI= 6.43 BB Acra
96 94 Mleu
09 07 93 Nhib

0.5
0.5

03
01 Ebid
02 97 Eabr
Csta Arob
08 00 95 98 Malb Cbic Pfim Cvic
0.0

0.0
12 05
92 Emaci Tcra Evel Mvar
CA2 (10.46)

06 Eser Espe
Toit
99 91 Epro Egra
CA2
-0.5

-0.5
13
11
-1.0

-1.0
Dbic Cstr

04
-1.5

-1.5

10
-2.0

-2.0

-2 -1 0 1 2 -2 -1 0 1 2
CA1 (11.93%) CA1
2.0
2.0

IST Egra
IST TI= 5.61 Epro
1.5
1.5

Esie
Cvic
Eser
1.0
1.0

93
DCA2 (10.94%)

Dbic
0.5

97
0.5

91 03 Emac
DCA2

02 94 Eabr Bpyg
10 98 95 Toit
0.0

09
0.0

06 00 05 99 Arob
12 96 92
07
-0.5

08 11
-0.5

04 01
Mleu Pfim Malb
13
-1.0

Tcra Nhib
-1.0

Dbis Csta
Evel Emaci Cbic
-1.5
-1.5

-1 0 1 2 3
-1 0 1 2 3
DCA1
DCA1 (11.29%)
 Arob

2.0
2.0

KL TI= 4.23 KL
Espe

1.5
1.5

Evel
93

1.0
1.0

Egra
DCA2 (14.11%)

Egrac
Mvir Mleu

0.5
DCA2
0.5

98 07 Eser Cfur
97 00 02 94 Nhib Cstr
03 09 Mfus Cvic

0.0
Malb Mgra
0.0

04 01
05 08 99 92 Emac Esie
12 06 95 91 Cbic

-0.5
96
-0.5

10 13 Tcra
Toit

-1.0
11
-1.0

Epro
Emaci

-1.5
Mrub
-1.5

-2 -1 0 1 2 3
-2 -1 0 1 2 3
DCA1
DCA1 (16.34%)
SS Atri Aver

1
05
1

SS TI= 4.43 Nhib

12 06 13 Epro Mfus Tdyb Evel Cfur
11 00 99 97 Mleu Espe Mdis MvirEmaci Cvic
10 03 Emac
92 Eser Cbic Malb Arob
09 04 Bmra Csta Cstr

0
94
0

Tcra
01 96 Mvar
08
CA2 (12.68%)

07 Tpra
Dcas

CA2
98 Dbic
02

-1
95
-1

93 Cheb

91 Toit
Egra
-2
-2

Dbis

-1 0 1 2 3 4 -1 0 1 2 3 4
CA1 (16.00%) CA1
Table 1 List of sampling stations and potamon types

Site Station name Mark Potamon type

1 Dunajské kriviny DK Eupotamal A, main river channel/old r. bed,

upper part

2 Bodícka brána BB Eupotamal B, side arm in the upper part

3 Šulianske rameno SUL Eupotamal B, side arm in the upper part

4 Kráľovská lúka KL Plesiopotamal-type water body

5 Gabčíkovo GAB Eupotamal A, main river channel/old r. bed,

lower part

6 Istragov IST Parapotamal, partially abandoned side arm

7 Sporná sihoť SS Plesiopotamal-type water body

8 Starý les SLE Plesiopotamal-type water body

Table 2: Copepoda species of the river/floodplain systems of the Hungarian-Slovak stretch of

the Danube considered in the Floodplain Index. H1-H4: habitat types, HV: habitat values, IW:
indication weights, short used in Figure 6.

Taxon short H1 H2 H3 H4 HV IW
Diacyclops languidoides (Lilljeborg) Dlan 10 1 5
Diacyclops crassicaudis (Sars) Dcra 10 1 5
Bryocamptus (B.)mrazeki (Minkiewicz) Bmra 10 1 5
Bryocamptus (Rh.) zschokkei (Schmeil) Bzsc 10 1 5
Echinocamptus pilosus (Van Douwe) Epil 10 1 5
Paracyclops fimbriatus (Fischer) Pfim 6 3 1 1,5 3
Attheyella crassa (Sars) Acra 5 5 1,5 3
Ectinosoma abrau (Kritschagin) Eabr 5 5 1,5 3
Tisbe furcata (Baird) Tfur 5 5 1,5 3
Bryocamptus (Rh.) pygmaeus (Sars) Bpyg 7 3 1,6 4
Paracyclops poppei (Rehberg) Ppop 3 7 1,6 4
Elaphoidella bidens (Schmeil) Ebid 10 2 5
Nitocra hibernica (Brady) Nhib 3 5 1 1 2 1
Cyclops vicinus Uljanin Cvic 3 2 4 1 2,3 1
Eudiaptomus gracilis (Sars) Egra 3 2 3 2 2,4 1
Eurytemora velox (Lilljeborg) Evel 2 3 3 2 2,5 1
Diacyclops bicuspidatus (Claus) Dbic 3 1 4 2 2,5 1
Eucyclops serrulatus (Fischer) Eser 2 4 1 3 2,5 1
Eucyclops s. var. proximus (Lilljeborg) Esvp 2 4 1 3 2,5 1
Eucyclops speratus (Lilljeborg) Espe 1 5 1 3 2,6 1
Eucyclops macruroides (Lilljeborg) Emaci 1 4 2 3 2,7 1
Acanthocyclops cf. robustus (Sars) Arob 2 2 3 3 2,7 1
Microcyclops rubellus (Lilljeborg) Mrub 6 4 2,8 3
Macrocyclops albidus (Jurine) Malb 1 3 2 4 2,9 1
Thermocyclops oithonoides (Sars) Toit 1 1 5 3 3 1
Eucyclops macrurus (Sars) Emac 4 2 4 3 2
Mesocyclops leuckarti (Claus) Mleu 1 1 3 5 3,2 1
Thermocyclops crassus (Fischer) Tcra 1 1 3 5 3,2 1
Canthocamptus staphylinus (Jurine) Csta 3 2 5 3,2 2
Microcyclops varicans (Sars) Mvar 3 7 3,4 4
Cryptocyclops bicolor (Sars) Cbic 2 1 7 3,5 3
Diacyclops bisetosus (Rehberg) Dbis 5 5 3,5 3
Ectocyclops phaleratus (Koch) Epha 2 1 7 3,5 3
Macrocyclops fuscus (Jurine) Mfus 2 1 7 3,5 3
Paracyclops affinis Sars Paff 5 5 3,5 3
Eucyclops denticulatus (Graeter) Eden 2 8 3,6 4
Megacyclops viridis (Jurine) Mvir 1 1 8 3,6 4
Cyclops strenuus Fischer Cstr 1 9 3,8 5
Eudiaptomus transylvanicus (Daday) Etra 10 4 5
Cyclops furcifer Claus Cfur 10 4 5
Cyclops heberti Einsle Cheb 10 4 5
Diaptomus castor (Jurine) Dcas 10 4 5
Macrocyclops distinctus (Richard) Mdis 10 4 5
Metacyclops gracilis (Lilljeborg) Mgra 10 4 5
Metacyclops minutus (Claus) Mmin 10 4 5
Mixodiaptomus kupelwieseri (Brehm) Mkup 10 4 5
Tropocyclops prasinus (Fischer) Tpra 10 4 5
Thermocyclops dybowskii (Lande) Tdyb 10 4 5
Attheyella (M.) trispinosa (Brady) Atri 10 4 5
Elaphoidella gracilis (Sars) Egrac 10 4 5

Table 3. Classification of the habitat types according to the Floodplain index (FI).
FI range Average Habitat type Potamal type
1.5-2.8 2.2 H1 Eupotamal-A
2.0-3.2 2.55 H2 Eupotamal-B
2.4-3.1 2.76 H3 Parapotamal
2.1-3.6 3.2 H4 Plesiopotamal