Fundam. Appl. Limnol. Vol.

181/1, 59–72Article
Stuttgart, June 2012

The influence of different hydrological conditions on

periphytic invertebrate communities in a Danubian
floodplain

Jasna Vidaković 1, *, Ivana Turković Čakalić 1, Filip Stević 1 and Dubravka Čerba 1, 2

With 6 figures and 2 tables

Abstract: Changes in the hydrologic regime have been a driving force for maintaining biodiversity of floodplain
communities. Recent studies have shown that periphytic biomass is closely related to the hydrologic cycle and
disturbances caused by flood pulses. This study was conducted in Lake Sakadaš which is a part of one of the largest
conserved Danube floodplains, Kopački Rit in Croatia. The aim of our study was to determine colonization pat-
terns, invertebrate abundance and community structure within the periphyton community that developed on artifi-
cial substrates during two hydrologically different years. Low connectivity of the floodplain was present in 2008
while high connectivity of the floodplain and extreme flooding characterized 2009. There were two distinct phases
in the invertebrate colonization process. The early phase of colonization, which included the first four sampling
dates of both years, was characterized by Crustacea. The late phase of colonization differed between the research
years. Bivalvia (Dreissena polymorpha) was the most dominant taxonomic group in the late phase of colonization
in 2008, while Oligochaeta was the representative taxa in late phase of 2009. In 2008, food source and water tem-
perature were the most important factors influencing the late periphytic invertebrate community, while the most
important parameters influencing fauna in 2009 was the Danube water level, indicating the influence of the flood
pulse on the invertebrate community. Generally, the invertebrate community differed between the two years in both
composition and abundance. One of the most important differences was a substantial development and the domi-
nance of an invasive bivalve species, Dreissena polymorpha indicating a possible threat to the lake’s ecosystem
suggesting the need for further monitoring.

Key words: Flood pulse, floodplain connectivity, periphyton, invertebrate community, Dreissena polymorpha.

Introduction velopment and function in many lentic and lotic eco-

Periphyton is a complex community of bacteria, fungi,
algae, invertebrates, and associated organic and inor-
ganic detritus, developing on both natural and artificial
substrates immersed in the water (Wetzel 2001, Azim
et al. 2005). The importance (primary production, food
source, habitat, energy and nutrient transport) and
complexity of this type of community has driven many
scientists to study different aspects of periphyton de-
systems (Vadeboncoeur & Steinman 2002, Azim et al.
2005). Different materials can be used as an artificial
substrate such as glass and ceramic plates, artificial
plants and polyurethane foam. There are drawbacks
to using artificial substrates due to size restrictions,
avoidance of artificial substrates by some species and
lack of direct interactions between organisms and the
surface. Despite this, artificial substrates are often
used because we can determine the size and type of

Authors’ address:
  University of J. J. Strossmayer in Osijek, Department of Biology, Trg Ljudevita Gaja 6, HR-31000 Osijek, Croatia.
1 

  Corresponding author; dcerba@biologija.unios.hr

2 

* All authors have equally contributed to the realization of the manuscript.

© 2012 E. Schweizerbart’sche Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de

DOI: 10.1127/1863-9135/2012/0290 eschweizerbart_XXX

1863-9135/12/0290 $ 3.50
60 Jasna Vidaković, Ivana Turković Čakalić, Filip Stević and Dubravka Čerba
the substrate, time and depth of immersion, and can
avoid negative influence of the substrate e.g. where
some macrophytes produce allelopathic substances
(Moschini-Carlos et al. 2000, Czerniawska-Kusza
2004). Also, Pieczyńska & Spodniewska (1963) found
no significant differences in the dominance and com-
position structure of periphytic fauna on natural and
artificial substrate, with Nematoda, Oligochaeta and
Chironomidae the main representatives in both cases.
Artificial substrates allow algae and fauna to be sam-
pled where it is difficult to access natural substrata and
cause less disturbance to benthic and periphytic com-
munities (Borcherding & Strum 2002). Both abiotic
and biotic parameters influence the appearance and
development of periphyton, but in floodplain areas one
of the most important influences is the change between
dry and flood periods. Some recent studies (Leandrini
et al. 2008) showed that the periphytic biomass of dif-
ferent types of aquatic habitats along the floodplain is
closely related to the hydrologic cycle and disturbanc-
es caused by flood pulses (Rodrigues & Bicudo 2001).
The direct and indirect influence of the limno- and
potamophase on the floodplain biocenosis has been re-
corded at different trophic levels in Kopački Rit (Pali-
jan & Fuks 2006, Mihaljević et al. 2009, 2010).
Studying the colonization process of periphyton is
an important first step towards our understanding of
its functional role in aquatic ecosystems. Generally,
colonization of substrates starts with the deposition of
organic substances and attraction of bacteria followed
by colonization of algae and invertebrates (Azim et al.
2005). Representatives of invertebrate fauna are very
diverse and can differ depending on the water quality,
season or the substrate on which the periphytic com-
munity is developed. In eutrophic lakes, Rotatoria,
Nematoda, Oligochaeta, Gastropoda, Bivalvia, Bryo-
zoa, Crustacea and Insecta larvae, especially Chirono-
midae, can be very abundant in different periphytic
communities (Dvořák & Imhof 1998, Čerba et al.
2009, Bogut et al. 2010). In every ecosystem, invasive
species can play an important part in all communities
influencing autochthonous flora and their fauna. The
zebra mussel (Dreissena polymorpha Pallas, 1771) is
a Ponto-Caspian invasive species of temperate fresh-
water ecosystems (Qualls et al. 2007). Since this spe-
cies originated in the Balkan Peninsula, Poland and the
former Soviet Union (Çağlar 1952), but was stranded
in the Ponto-Caspian area after the last glaciation, we
can ask ourselves if we should call its dispersion to
East and Middle Europe invasion or re-colonization of
old habitats. According to several authors (e.g. Walz
1978, Garton & Johanson 2000) maximum growth of
eschweizerbart_XXX
D. polymorpha is usually early in the growing season
and corresponds with a peak in phytoplankton abun-
dance. The growth rates of D. polymorpha in the same
water-body vary significantly across years. Zebra
mussels are prolific filter-feeders and can effectively
filter a wide size range of particles (Ten Winkel &
Davids 1982). Due to their high filtration rate, high
densities of zebra mussels can significantly decrease
plankton densities and increase water clarity (Strayer
et al. 1999, Pillsbury et al. 2002) which can influence
invertebrate composition and abundance in the peri-
phyton (Ricciardi et al. 1997).
In this paper we present the data on invertebrate
communities in periphyton, developed on artifi-
cial substrate during two hydrologically contrasting
years, in the river-floodplain ecosystem (Nature Park
Kopački Rit). The aim of our study was to determine
the influence of lake’s connectivity on periphytic in-
vertebrate community. The specific issues addressed
in this paper are: (1) can the differences in coloniza-
tion patterns during two years be related to different
water-levels; (2) will the differences in invertebrate
abundance and community structure be visible and as-
sociated with changes of the water-level; (3) what is
the relationship between biotic and abiotic factors and
periphytic invertebrates; and (4) were there temporal
variations within invertebrate communities and meas-
ured biotic and abiotic parameters.
Material and methods
Study area
This study was carried out in a floodplain lake, Lake Sakadaš
within Kopački Rit Nature Park, one of the Danube’s largest
conserved floodplains (Fig. 1). Kopački Rit is situated in the
north-east of Croatia between the River Drava and the Danube.
It is included on the list of Important Bird Areas and on the
RAMSAR List of Wetlands of International Importance. The
protected floodplain area is located between 1383 and 1410
river km of the Danube. As the parent river, the Danube hy-
drologic regime has the prime influence on the floodplain eco-
system. Lake Sakadaš is an open water system and exchanges
water with the Csonakut Channel in the east, the Novi Channel
in the south and through the dam in the west. The lake is in di-
rect hydrologic connection with the main river channel through
the Hulovo and Csonakut Channels (total length ca. 10 km)
(Fig. 1). When the Danube water-level is 1.67 m (measured
at Apatin station at 1401.4 river km) its waters start to enter
the floodplain thus creating the flow pulse, but water-levels of
3.0 – 3.5 m mark the beginning of the flood pulse (Bognar 1990,
Palijan 2010). The lake has an average depth of about 5 m and
maximum depth of ca. 12 m, with a surface water area of about
0.15 km2. During the past few decades, the lake has been in
a eutrophic/hypertrophic state (Vidaković et al. 2002, Stević
et al. 2005, Mihaljević & Stević 2011). The first research of
 Influence of hydrological conditions on periphytic invertebrate communities
Fig. 1. Study area – Lake Sakadaš, a part of the Danube floodplain area of the Kopački Rit Nature Park (Croatia). Dashed line
represents the border of Nature Park Kopački Rit.
the periphyton communities within the Kopački Rit floodplain
was conducted on natural substrata (submerged macrophytes)
(Vidaković & Bogut 2007, Bogut et al. 2010, Čerba et al. 2009,
2011). Preliminary in situ experiments with glass plates as a
substrate were carried out in 2007 in the littoral area of Lake
Sakadaš, when nematodes dominated the periphytic inverte-
brate community (Vidaković et al. 2011).
Sampling
Colonization of periphyton developed on microscopic glass
plates was investigated during two hydrologically contrasting
years (from April till August 2008 and 2009). Sampling began
on April 22nd in 2008 and April 28th in 2009. Samples were taken
after 7 days of exposure and then every two weeks till August
19th 2008 and August 25th 2009. Prior to submergence, slides
were washed with detergent and soaked in 1 M HCl overnight.
They were positioned vertically within the water column at the
fixed depth of 25 cm in modified plastic slide-boxes. Sampling
was performed by taking nine slides on each sampling occasion
at the research site. Three slides were for fauna analyses fixed
eschweizerbart_XXX
61
in situ with the 4 % formaldehyde. Three slides were taken for
chlorophyll analysis and the next three slides were used to iden-
tify periphyton biomass. The data gathered from the three slides
were pooled and average values have been used for statistical
analysis of each parameter. Slides were transported to the labo-
ratory for further analysis. Samples of water were collected to
determine ammonia (NH4), nitrates (NO3), nitrites (NO2), total
nitrogen (TN) and total phosphorus (TP) (APHA 1985). Con-
centration of dissolved oxygen (DO), pH, electrical conductiv-
ity (COND) and water temperature (WTEMP) were determined
in situ with a portable multimeter (Multi 340 i/SET, WTW,
Weilheim, Germany). Transparency (SD) was measured with a
Secchi disc. The Danube water-level (DWL) was measured at
the Apatin gauge measuring station (at 1401.4 river km). Con-
centrations of chlorophyll-a in water (WCHLa) and periphyton
chlorophyll-a (PCHLa), -b (PCHLb) and -c (PCHLc) were de-
termined according to Komárkova (1989) and expressed in μg
per cm2. Periphyton biomass (mg per cm2) was determined as
follows; periphyton was scraped off the slide with a razor blade,
dried at 105 °C for 24 h, weighed, placed in an oven at 500 °C
for an hour and then weighed again.
62 Jasna Vidaković, Ivana Turković Čakalić, Filip Stević and Dubravka Čerba
Fauna samples
In the laboratory, periphyton for faunal analysis was scraped
off from both sides of the glass slide using a razor, stained with
Rose Bengal and washed through a 24 µm aperture sieve, prior
to inspection under stereoscopic microscopes (Carl Zeis Jena
and Olympus SZX9), under different magnifications (1.5 – 20×).
Fauna were identified using several standard taxonomic keys
for invertebrates such as Magda (1979), Nilsson (1996, 1997),
Pfleger (1999), Engelhardt (2003) and Wood & Okamura
(2005). Abundance was expressed as the number of individuals
per cm2 of glass slide except for Spongia and Bryozoa where
the numbers represent percentage per cm2 of the glass covered.
Statistical analysis
Several multivariate methods were used to explain the varia-
tions in the periphytic invertebrate communities and to assess
the relationship between invertebrate abundance and the meas-
ured environmental variables.
A clustering technique was used to join the samples in
hierarchical groups based on the similarity in invertebrate
abundance and is presented using a dendrogram. Non-metric
multidimensional scaling (nMDS) analysis was used to dis-
play invertebrate community similarity and temporal dynam-
ics in both investigated years in multivariate ordination space.
NMDS is an iterative ordination method which uses the ranks
of similarities between samples for a low-dimensional ordina-
tion. Clustering and nMDS were applied on the Bray-Curtis
similarity matrix based on the invertebrate abundance. To bal-
ance the contribution of common and rare taxa, invertebrate
abundance was square-root transformed prior to calculating
sample similarities. The contribution of different invertebrate
taxa to the average dissimilarity between the investigated years
was elucidated using the SIMPER routine. Hierarchical cluster-
ing, nMDS analyses and the SIMPER routine were performed
using the statistical program PRIMER version 5.0 (Clarke &
Warwick 2001).
Detrended Correspondence Analysis (DCA) was used to
determine whether the linear or unimodal statistical technique
would be most appropriate for the modelling of responses. DCA
suggested the use of the linear method (Redundancy analysis,
RDA) since the value of the longest gradient did not exceed 3.0
(Lepš & Šmilauer 2003). This method was used to assess the
relationship between invertebrate abundance and the measured
environmental variables (Ter Braak & Šmilauer 2002). The
invertebrate abundance data was log-transformed to meet the
normality requirements. The forward selection option using the
Monte Carlo unrestricted test involving 499 permutations iden-
tified the significant subset of environmental variables, retain-
ing only variables significantly (p < 0.05) contributing to the
total variance of each taxa. Environmental variables are repre-
sented by arrows with the maximum value located at the arrow
head. DCA and RDA were performed with the CANOCO for
Windows version 4.5 (Biometrics-Plant Research International,
the Netherlands).
The t-test was used to test for significant differences be-
tween two sampling seasons in environmental variables (Zar
1999). Correlation analysis was used to compare most frequent
and most abundant invertebrate taxa in both years. For the
analysis of fauna in 2008, Dreissena polymorpha, Nematoda,
Oligochaeta, Crustacea, Insecta larvae and Insecta pupae were
correlated, and in 2009, Hydra sp., D. polymorpha, Turbellaria,
eschweizerbart_XXX
Nematoda, Gastropoda, Oligochaeta, Crustacea, Insecta larvae
and Paludicella articulata. Correlations were considered to be
significant when p < 0.05. Calculations were performed using
Statistica v. 7.1 (StatSoft, Tulsa).
Results
Environmental parameters
During the investigation period (April to August) in
2008, the Danube water level was higher than 3.0 m
for 73 days but never reached 5.0 m; in 2009 during
the same period, Kopački Rit was flooded for 122
days with the highest recorded value of 6.25 m on the
84th day of periphyton exposure. In general, 2009 was
a year with long periods of flooding in Kopački Rit
(Fig. 2). Apart from the differences in the Danube and
Sakadaš water-level, the t-test indicated statistically
higher oxygen concentration (t = 3.065) and conduc-
tivity (t = 2.04) at p < 0.05 in 2008. Due to the higher
water level of the Danube in 2009, the mean value of
the Sakadaš water level, as well as transparency, were
higher than in 2008. The surface lake water was well
oxygenated in both years reaching the lowest value of
5.26 mg l–1 on the 14th day of periphyton exposure in
2009. The highest value of conductivity in 2008 was
803 µS cm–1, while in 2009 values above 475 µS cm–1
were not recorded. The water temperature was similar
during both years with maximal values being recorded
in June, 28.4 °C and 26.6 °C (on the 70th day in 2008
and 56th day in 2009). Chlorophyll-a concentration
in water varied between 7.2 – 42.4 µg l–1 in 2008 and
3.1– 38.4 µg l–1 in 2009. High concentrations of TP
(mean value 203.7 µg l–1 in 2008 and 176.1 µg l–1 in
2009) and TN (mean value 723.7 µg l–1 in 2008 and
691.9 µg l–1 in 2009) were found. pH values (7.4 – 8.7)
indicated slightly alkaline conditions (Fig. 3).
Periphyton biomass and chlorophyll
concentrations
There was a difference in both the periphyton bio-
mass and chlorophyll concentrations between research
years. The highest values of dry weight recorded in
2008 were 138.9 mg cm– 2, while in 2009 the peri-
phyton biomass was lower and maximal values for
dry weight were 46.6 mg cm– 2. During both years,
the minimum dry weight, found on the first sampling
date, did not exceed 0.04 mg cm– 2. The ash weight
and ash free weight followed the same trend during
the research periods (Fig. 3). Periphyton chlorophyll-a
concentrations in 2008 ranged from 0.051 µg cm– 2 to
4.38 µg cm– 2 and in 2009 from 0.59 µg cm– 2 to 6.19 µg
 Influence of hydrological conditions on periphytic invertebrate communities 63

Fig. 2. Water level of the Danube in 2008 and 2009, with extracted data for the research periods.

Table 1. Results of the SIMPER routine: contribution of invertebrates to average dissimilarities between investigated years based
on invertebrate abundance.
Species Average abundance Average Dissimilarity/ Species Cumulative
(ind. cm– 2) dissimilarity SD contribution (%)
(%)
Group 1 Group 2
Years 2008 and 2009
Average dissimilarity = 59.71
Bivalvia 4.93 0.31 16.23 1.54 27.18 27.18
Crustacea 1.39 1.89 11.94 0.98 20.00 47.18
Oligochaeta 1.69 2.73 11.74 1.28 19.67 66.85
Nematoda 0.82 0.08 5.98 1.23 10.01 76.86
Insecta larvae 0.58 0.34 5.57 1.34 9.33 86.18
Turbellaria 0.03 0.06 2.33 1.09 3.90 90.08

Note: Cut-off at 90 %; SD – standard deviation.

cm– 2. Higher values in 2009 were also recorded for
chlorophyll-c, 4.55 µg cm– 2 in relation to 4.80 µg cm– 2
(Fig. 3).
Fauna
In total, 14 invertebrate taxonomic groups were re-
corded during this study: Spongia, Cnidaria, Turbel-
laria, Nematoda, Gastropoda, Bivalvia, Oligochaeta,
Hirudinea, Hydracarina, Crustacea, Tardigrada, Insec-
ta larvae and pupae, and Bryozoa. Six of these groups
contributed most to the differences between the inves-
tigated years (Table 1, Fig. 4).
Macrostomum tuba (Leidy 1851) was recorded in
both years, while Microstomum caudatum (von Graff
1882) and Planaria torva (Müller 1774) were Turbel-
laria representatives in 2009. We found no Gastro-
poda in 2008, but in 2009 they were represented with
four species (Bithynia tentaculata (Linnaeus 1758),
Acroloxus lacustris (Linnaeus 1758), Planorbarius
corneus (Linnaeus 1758) and Physa sp.) and juvenile

eschweizerbart_XXX
64 Jasna Vidaković, Ivana Turković Čakalić, Filip Stević and Dubravka Čerba

Fig. 3. Variations in a) Danube water level (data source: daily recordings during the experiment period at river 1,401.4 km); b)
Lake Sakadaš water level; c) transparency; d) water temperature; e) dissolved oxygen; f) concentration of chlorophyll-a in the
water; g) pH; h) conductivity; i) total phosphorus; j) total nitrogen; periphyton biomass – k) dry weight, l) ash weight, m) ash free
weight; periphyton chlorophyll concentration – n) chlorophyll-a, o) chlorophyll-b, p) chlorophyll-c in the floodplain lake during
the experiment. Box-plots represent the median, the 25 and 75 % quartile and the range of the values; open circle outliers; open
square extremes.

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 Influence of hydrological conditions on periphytic invertebrate communities 65

Fig. 4. Changes in the invertebrate abundance of the most abundant and frequent groups, and pointed out by SIMPER analysis,
measured as number of individuals per cm2 during the research years.
Note: Data represent the mean value of three glass slides ± standard deviation. The abundance of given invertebrate taxonomic
groups is a sum of abundances of all its representatives (in brackets): Crustacea (Cladocera, Copepoda, Ostracoda), Bivalvia
(Dreissena polymorpha), Oligochaeta, Insecta larvae (Chironomidae, Ephemeroptera, Odonata, Ceratopogonidae, Polycentropodi-
dae), Nematoda and Turbellaria (Microstomum caudatum, Macrostomum tuba, Planaria torva).

stages. Bivalvia were represented with only one spe-
cies – Dreissena polymorpha (Pallas 1771). Two dif-
ferent species of Hirudinea were recorded, Helobdella
stagnalis (Linnaeus 1758) in 2008 and Erpobdella oc-
toculata (Linnaeus 1758) in 2009. Beside Cladocera
and Copepoda, Ostracoda were also recorded. Insecta
larvae were mostly represented by Chironomidae, al-
though we also found larvae of Ephemeroptera and
Ceratopogonidae in 2008. In 2009, in addition to these
taxa groups, Odonata and Polycentropodidae were
also recorded.
The first samples were taken a week after immer-
sion of the glass slides when representatives of Cla-
docera, Chironomidae larvae and Oligochaeta were
found as the first colonizers. Only chironomid larvae
displayed almost 100 % constancy in both sampling

eschweizerbart_XXX
66 Jasna Vidaković, Ivana Turković Čakalić, Filip Stević and Dubravka Čerba

Fig. 5. Dendrogram (a) and nMDS plot (b) based on invertebrate abundance. Numbers 7–126 represent number of days, 08 and 09
years, when slides were submerged in the water. Open circles (○) indicate 2008 samples and solid circles (●) indicate 2009 samples.

years. Taxa that were present on 70 % or more of the
sampling dates in both years were: Oligochaeta, Cla-
docera, Nematoda, Dreissena polymorpha and Co-
pepoda (Fig. 4). During both years, we recorded the
presence of Spongia and Bryozoa but with different
percent cover on the slides. In 2008, the sponge Ephy-
datia fluviatilis (Linnaeus 1759) was found on the
112th and 126th day while in 2009 Eunapius fragilis
(Leidy 1851) was present on the 84th and 98th day. We
also found traces of Spongia on the 56th day in 2009,
but without gemmules we could not determine the spe-
cies. In 2008, only one species of Bryozoa was found
– Plumatella emarginata (Allman 1844), on the 56th,
70th and 112th day, whereas in 2009 we found three
species: Paludicella articulata (Ehrenberg 1831),
Plumatella fungosa (Pallas 1768) and P. emarginata.
P. articulata appeared on the 56th day and was present
until the end of the trials.
In 2008, 98.5 % of the total fauna (excluding
Spongia and Bryozoa) was represented by five taxa:

eschweizerbart_XXX
 Influence of hydrological conditions on periphytic invertebrate communities 67

Fig. 6. Fauna ordination results from redundancy analysis (RDA) of invertebrate abundance data of the both years (2008 and 2009).
The plot shows the taxa scores along the first and second axes in relation to the statistically significant environmental variables:
DWL (Danube water level), WTEMP (water temperature), WCHLa (chlorophyll-a in water), COND (conductivity), DW (dry
weight) and PCHLa (periphyton chlorophyll-a). Environmental variables are represented with solid arrows and invertebrate taxa
vectors are represented with thick dashed arrows. The directions and lengths of the arrows indicate importance and correlation to
the respective axes. Arabic numbers indicate sampling day. Open circles (○) indicate 2008 samples and solid circles (●) indicate
2009 samples.

D. polymorpha, Oligochaeta, Cladocera, Nematoda
and Chironomidae larvae with the dominance of
D. polymorpha (52 %). In 2009, these taxa composed
88.2 % of the total fauna with a change in dominance
– Oligochaeta were the most abundant (49.4 %) while
D. polymorpha was not well-represented (5.6 %); sec-
ond in dominance in 2009 were Cladocera (27.5 %).
The highest diversity was found on the 70th day in
2008 and the 84th day in 2009 and although diversity
was greater in 2009 (28 vs. 14 taxa), the difference
was mostly due to representatives of Gastropoda and
Insecta larvae, despite invertebrate abundance being
1.7 times lower in 2009 than in 2008. Abundance
gradually increased in both years with the maximum
on the 70th day, although on day 14 in both years there
were high densities of cladocerans, and on the 112th
day in 2008 the abundance of D. polymorpha and Oli-
gochaeta was rather high.
In 2008 there was a positive correlation between
D. polymorpha and Insecta larvae (r = 0.80, p = 0.006),
Insecta larvae and pupae (r = 0.82, p = 0.014), and
Nematoda and Oligochaeta (r = 0.81, p = 0.016). In
2009, D. polymorpha and Insecta larvae were corre-
lated again (r = 0.80, p = 0.005). Turbellaria and Crus-
tacea (r = 0.84, p = 0.003), Nematoda with Gastro-
poda (r = 0.63, p = 0.049) and Palludicella articulata
(r = 0.95, p = 0.0001), Gastropoda and P. articulata
(r = 0.70, p = 0.024) were correlated as well.

eschweizerbart_XXX
68 Jasna Vidaković, Ivana Turković Čakalić, Filip Stević and Dubravka Čerba

Statistical analysis of the late colonization phase in two different phases

for each year. Thus, in 2008 (day 56 –126), the inver-
The results of hierarchical clustering and nMDS anal-
tebrate community was influenced by a high periphy-
yses based on invertebrate abundance indicate differ-
ton biomass, a high concentration of chlorophyll-a in
ences between the two sampling years (Fig. 5). The
water, a higher water temperature and a lower water
degree of similarity between invertebrate abundance,
level of the Danube. Bivalvia (D. polymorpha) and
at Bray-Curtis similarity level of approximately 50 %,
Nematoda were positively correlated with the first
revealed two distinctive groups. Group 1 represents
three and negatively with Danube water level (Fig. 6).
the early phase of colonization (April and May) of
On the other hand, in 2009 (day 70 –112), the inver-
both years (the first four sampling dates in 2008 and
tebrate community was influenced by a higher water
2009, with the exception of the fourth sampling date in
level of the Danube as well as higher concentrations of
2009). This period was mostly characterized by Crus-
periphyton chlorophyll-a. The most abundant inverte-
tacea. The other group (Group 2) was divided into two
brate taxa, Oligochaeta, was positively correlated with
subgroups, 2 a – late phase of colonization of 2008 and
water-level of the Danube (Fig. 6).
2 b – late phase of colonization of 2009, representing
the second part of the season (June, July and August).
Bivalvia (Dreissena polymorpha) were the most dom-
Discussion
inant in the late phase of colonization in 2008, while
Oligochaeta characterized the invertebrate community
Our study suggests that water temperature, conductiv-
in the late colonization phase in 2009. Dreissena poly-
ity, available food (periphyton chlorophyll and bio-
morpha, Crustacea and Oligochaeta contributed most
mass) and different hydrological conditions in this
to the differences between the investigated years (2008
floodplain lake were the most important factors in-
and 2009) with regard to invertebrate abundance (Ta-
fluencing and structuring the invertebrate community
ble 1). These three taxa accounted for a 66.85 % of
and this finding is in agreement with previous studies
cumulative species contribution, while the next three,
showing that the hydrologic regime of the Danube has
Nematoda, Insecta larvae and Turbellaria, contributed
a great influence on the communities within the flood-
23.23 % (Table 1).
plain area of Kopački Rit (Mihaljević & Stević 2011,
The species-environment correlations for RDA
Palijan 2010, Vidaković et al. 2011). Generally, low
axes 1 and 2 indicate a significant relationship between
connectivity of the floodplain characterized the year
environment variables and invertebrate abundance
2008, while high (almost permanent) connectivity
(Table 2). In the analysis based on both years’ data,
of the floodplain and extreme flooding characterized
the forward selection process identified 6 significant
2009. According to Ward et al. (1999), one of the most
environmental variables (Fig. 6). Dates representing
important factors structuring aquatic communities in
the early colonization phase of both years (day 7– 42
floodplains is connectivity since it is essential for the
of 2008 and 7– 56 of 2009) were grouped on the left
biodiversity of the floodplains.
side of the triplot. This period was characterized by in-
Periphyton growth and development can be influ-
creasing the conductivity and low periphyton chloro-
enced by different parameters such as temperature,
phyll-a concentrations. There was also a positive cor-
light, nutrients or a combination of these (Kiffney &
relation between Crustacea and conductivity (Fig. 6).
Bull 2000, Moschini-Carlos et al. 2000). The distinc-
The ordination of invertebrate taxa showed separation
tiveness of the two research years can be seen in the
importance of the environmental parameters influenc-
Table 2. Summary statistics (eigenvalues, species-environment ing invertebrate communities. The most important pa-
correlations, cumulative percentage variance of species data rameters were periphyton biomass, chlorophyll-a con-
and cumulative percentage variance of species-environment centration in the water and water temperature in 2008
relation) of redundancy analysis (RDA). (Fig. 6), whereas in 2009, the Danube water-level and
RDA axes 1 2 3 4 concentration of periphyton chlorophyll-a were the
Eigenvalues 0.509 0.138 0.051 0.029 most important (Fig. 6). Algal and invertebrate com-
position and richness in the periphytic communities
Species-environment correlations 0.937 0.795 0.625 0.860
are affected by floods due to physical disturbance, the
Cumulative percentage variance presence/absence of allochthonous particulate organic
– of species data 50.9 64.7 69.9 72.8 matter, the presence of predators or the appearance
– of species-environment relation 69.5 88.4 95.4 99.4
of new species (Algarte et al. 2009, Vidaković et al.

eschweizerbart_XXX
 Influence of hydrological conditions on periphytic invertebrate communities
2011). The impact depends on the frequency, magni-
tude and duration of the flood (Thoms 2003, Whited
et al. 2007).
Spongia, Nematoda, Rotatoria, Oligochaeta, Crus-
tacea, Bryozoa and Insecta larvae (mainly Chirono-
midae), are the main representatives of periphytic in-
vertebrate fauna of temperate freshwaters (Dvořák &
Imhof 1998). Dendrogram and nMDS plots of inverte-
brate abundance illustrate two separate periods of the
colonization and there is a difference in invertebrate
composition and abundance between the early (Group
1) and the later phase of the periphytic community
development (Groups 2 a and 2 b, Fig. 5). The repre-
sentative taxa for these phases were Crustacea, D. pol-
ymorpha and Oligochaeta (Fig. 5). SIMPER analysis
indicates the mentioned taxa as most important in con-
tributing to the dissimilarity between the years inves-
tigated. Cladocera, were often found in the periphyton
and successfully used periphytic algae as a primary or
alternative food source (e.g. Siehoff et al. 2009). The
initial phase of colonization is characterised by higher
densities of crustaceans and rotifers due to their swim-
ming ability compared to other invertebrates (Peters
et al. 2007).
Oligochaetes are often very abundant in periphytic
communities developed on natural (e.g. macrophytes,
sediment) or artificial substrates (e.g. glass slides,
plexiglass plates, plastic rubbish) (Mihaljević et al.
1998, Dumnicka 2007, Bogut et al. 2009). During this
study, Oligochaeta were mostly influenced by peri-
phytic chlorophyll-a concentrations, indicating food
as an important factor, and by changes in the Danube
water-level (Fig. 6). In periphytic communities, oli-
gochaetes can be influenced by the developed algae,
the amount of periphyton, or macrophyte architecture
(Botts & Cowell 1992, Alves & Gorni 2007). Eutroph-
ic lakes or organically enriched water-bodies can sup-
port high numbers of Oligochaeta (Mihaljević et al.
1998, Hirabayashi et al. 2011).
The change in the abundance of D. polymorpha was
most likely because of hydrologic differences in 2008
and 2009. According to RDA, in 2008D. polymorpha
abundance was directly proportional to water tempera-
ture, chlorophyll-a concentrations in water and peri-
phyton biomass (Fig. 6). Mussels play a special role in
the cycling of elements in aquatic environments and as
filter feeders they remove suspended matter and plank-
tonic algae from the water column (Stańczykowska
1977, Lewandowski & Stańczykowska 2000). One of
the ecological impacts of D. polymorpha is the for-
mation of complex 3 D-structures composed of their
shells, which create a new type of habitat. Dreissena
eschweizerbart_XXX
69
and other mussels can influence invertebrate commu-
nities through the increase of organic content by depo-
sition of its faeces and pseudofaeces (Griffiths 1993),
but this depends on the mussel size-frequency (Young
et al. 1996). Ricciardi et al. (1997) noted the enhanced
macroinvertebrate abundance in the presence of ze-
bra mussels, but also changes in composition of the
invertebrate community depending on the density of
colonized mussels. According to Wisenden & Bailey
(1995) and Ricciardi et al. (1997), dense communi-
ties of D. polymorpha on hard surfaces can have direct
or indirect negative effects on Gastropoda by fouling
them or competing for space or food, or by increasing
the number of smaller gastropods. When studying pe-
riphytic communities on natural substrates, we found
D. polymorpha on different macrophytes, but its abun-
dance was rather low in comparison with glass slides.
According to Lewandowski (1982) and Van Diepen
& Davids (1986), immersion and later coverage with
bacteria and algae of artificial substrates may be a
stimulating factor for the attachment of the postveliger
stage. We suspect that the larvae were brought from
the Danube and Hulovo channel and suitable condi-
tions allowed the abundant development of D. poly-
morpha. The hydrodynamic and environmental con-
ditions in 2009 could have been less favourable for
Dreissena larvae and adults. The cover by D. polymor-
pha could have negatively influenced the development
of Bryozoa even though it can utilize zebra mussels
as a colonizing substrate (Ricciardi et al. 1997). Small
spaces between the mussels and competition for food
could have prevented considerable development of
Bryozoa in 2008. Bryozoa are a very important part
of periphytic invertebrate communities since they in-
crease substrate complexity, represent an alternative
source of food and refugia for nematodes, Chironomi-
dae and other invertebrates (Wood & Okamura 2005,
Vidaković et al. 2011). The positive correlation be-
tween periphyton biomass and nematodes (Fig. 6) was
consistent with a previous study, conducted in 2007
at the same site (Vidaković et al. 2011). Also, in 2009
an increase in cover by bryozoans was followed by an
increase in Nematoda abundance, possibly as a result
of higher habitat complexity created by Paludicella
articulata, Plumatella fungosa and P. emarginata. Re-
corded, but not often found in Kopački Rit floodplain,
were representatives of Spongillidae. Eunapius fragi-
lis and Ephydatia fluviatilis have also been recorded in
the Danube floodplain in Austria, together with several
other species: Spongilla lacustris, Ephydatia mulleri
and Trochospongilla horrida (Dröscher & Waringer
2007). Sponges are active filterers and can filter large
70 Jasna Vidaković, Ivana Turković Čakalić, Filip Stević and Dubravka Čerba
amounts of water while feeding on plankton, small al-
gae, bacteria or organic matter, thus influencing water
quality (Frost 1991, Vohmann 2008).
Conclusions
In conclusion, our research indicated influence of the
flood pulse on the periphytic invertebrate community
in Kopački Rit. Its influence on the development and
structure of the invertebrates can be seen in the domi-
nance of D. polymorpha in 2008 vs. Oligochaeta in
2009 as well as lower diversity and higher abundance
of invertebrate fauna recorded in 2008. Aside from the
water-level, other important factors influencing peri-
phytic invertebrates were food source and the com-
plexity of habitat, which are often the limiting factors
in benthic communities. D. polymorpha, Bryozoa and
Spongia provided a suitable habitat for the develop-
ment of a more complex invertebrate community.
Large numbers of D. polymorpha on the glass slides
indicate the possible threat of it spreading further in
appropriate conditions which could lead to the distur-
bance of this ecosystem.
Acknowledgements
The authors would like to thank reviewers for their comments
and improvements. Many thanks to Tanja Žuna Pfeiffer, Bar-
bara Vlaičević, Dubravka Špoljarić, Anita Galir Balkić, Tonka
Kovačević and Dr. Goran Palijan for their time and efforts in
the field and laboratory work. Also, we are very grateful to Dr.
Ljiljana Čačić and the employees of Eco-laboratory in Vodo-
vod-Osijek Ltd. This study was financed by the Ministry of Sci-
ence, Education and Sport of the Republic of Croatia, as project
No. 285-0000000-2674.
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Submitted: 19 December 2011; accepted: 29 June 2012.

eschweizerbart_XXX