Cent. Eur. J. Biol. • 5(2) • 2010 • 240–255
DOI: 10.2478/s11535-009-0062-9
Central European Journal of Biology
Zooplankton communities of two lake outlets
in relation to abiotic factors
Research Article
Robert Czerniawski*, Józef Domagała
Department of General Zoology, University of Szczecin,
71-412 Szczecin, Poland
Received 9 June 2009; Accepted 1 October 2009
Abstract: W
e examined the quantitative and qualitative zooplankton community structure in two small rivers flowing out from lakes differing in
trophic conditions. Within each river, three sites were chosen for the collection of drifted zooplankton: one at the outflow, and two at
distances of 0.2 km and 1 km from the outflow. The most significant difference in zooplankton community between the outflow and the
lower course of the river occurred in the first section directly after the outflow. These differences in the zooplankton community were
driven largely by crustaceans, which declined faster in the river flowing out from the mesotrophic lake. Physical parameters mainly
impacted the zooplankton community found in the river flowing from the mesotrophic lake; however, chemical parameters also had an
impact in the river discharging from the strongly eutrophic lake.
Keywords:  Zooplankton drift • Small river • Outflow • Downstream • Trophic status
© Versita Sp. z o.o.
1. Introduction
Different types of water reservoirs have significant
effects on the quantitative and qualitative structure of
the zooplankton of the rivers flowing from them [1-3].
This phenomenon concerns both small and large
rivers. Little is known about zooplankton communities
found in flowing water bodies. The majority of authors
concerned with this subject studied large or relatively
large rivers [4-7] in which the density of zooplankton is
greater than in small rivers. In lake outlets, the number
of zooplankton species, their density and their body
length decreases with increasing distance downstream
along rivers [8,9]. This overall reduction in zooplankton
community parameters in rivers is believed to arise
from hydromorphology (depth, width, velocity) and the
activity of predatory fish and other invertebrates in the
rivers. Biological conditions such as predation by fish or
invertebrates [8,9] are mainly thought to be responsible
for this reduction and authors have found stronger
correlations between zooplankton features and biotic
rather than abiotic factors. Perhaps predatory activity
of fish should be greater in the rivers flowing out from
240
the lakes of lower eutrophication because of smaller
amounts of seston in water and better visibility. Therefore,
the reduction of zooplankton in the rivers of lower trophy
should be expected to be greater than in those with higher
amounts of seston. The relationship between the trophy
and predatory fish is characteristic of lakes, in which
the zooplankton is closely related to the abiotic factors
[10-12]. Although higher primary productivity in
eutrophic lakes may be the main reason for decreased
activity of predatory fish, lower visibility may also play
a significant role. In view of this, three questions arise:
(1) Are there significant differences of some properties
of the zooplankton community between the outflow and
the lower course of the river? (2) What are differences in
the zooplankton community between two rivers flowing
out of the lakes with different trophy? (3) What are the
relationships between zooplankton community and
physical and chemical conditions in small rivers?
Differences in the trophic state of lakes can also
affect characteristics of the river bottom and the degree
of its cover with macrophytes. This may indirectly affect
the flow rate of the river, leading to formation of small
floodplains and slack waters, and favouring zooplankton
* E-mail: [email protected]
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R. Czerniawski, J. Domagała
Figure 1.
Map of study sites.
reproduction yielding an increase its density, especially
those of Cladocera [13,14].
To answer to the question above, we analysed
differences in the qualitative and quantitative structure
of zooplankton communities in two small rivers flowing
out of lakes of different trophic status, relating to physical
and chemical conditions.
2. Experimental Procedures
The study was performed on two small polish rivers:
Słopica (GPS: N 53° 12’ 30”, E 15° 50’ 17”) and Korytnica
(GPS: N 53° 12’ 31”, E 15° 58’ 8”), flowing out from lakes
of significantly different trophic status. At each river, three
sampling sites were selected, the first at the outflow; the
second at a distance of 0.2 km below the outflow and
the third at a distance of 1 km from the outflow. The sites
on the River Słopica were marked as S1, S2, S3, while
those on the River Korytnica as K1, K2, K3 (Figure 1).
The Słopica is 14.0 km long. It flows through three
lakes with a low degree of eutrophication. The last of
the lakes is Dominikowo Wielkie, below which the
zooplankton samples were collected. The area of the
lake is 70.9 ha, its mean depth is 9.3 m and its maximum
depth is 16.5 m. Isoteids grow over the bottom of the
lake, even to a depth of 10 m. The lake is not susceptible
to phytoplankton blooming. The river selected for the
study has a mean depth of 0.3 m and a mean width of
2.5 m. The bed of the river is regularly shaped and the
bottom of this section of the river consists of sand and
gravel.
The River Korytnica is 35.2 km long. It flows through
two lakes; the first of them has an area of 12 ha and
mean depth of 0.5 m. The lake substantially modifies the
biological and physico-chemical conditions of the river;
its trophy increases and its water quality deteriorates.
The other lake is Nowa Korytnica. Below this lake
zooplankton samples were collected and analysed. The
area of the lake is 97.5 ha, its mean depth is 2.4 m with
a maximum depth of 4.7 m. Nowa Korytnica Lake is very
susceptible to long lasting blooms of phytoplankton from
April to October, and does not show the presence of
isoetids. The section of the river selected for study has
a mean depth of 0.6 m and a mean width of 5 m. The
bottom of the studied section of the river is covered with
over 90% mud and only sporadically covered with sand.
Between sites K1 and K2, c.a. 60% of the river bottom
is covered by macrophytes, mainly Typha angustifilia L.
Nyphaea alba L. and Myriophyllum verticillatum L. The
river makes a 20 m2 large flooded plain area of 0.2 m
depth fully covered with Glyceria maxima (Hartm.).
Between sites K2 and K3 the river has a regular course
in a narrow 4 m wide bed, with Phragmintes communis
Trin. growing on the banks.
Although the characteristics of the river catchment
area may be an important factor influencing the
zooplankton community [15], there are no differences in
the physical structure of the land, or in the agricultural
usage along these two rivers. At the outflows of both
rivers, large aggregations of fish fry (mostly cyprinids)
were observed from April to September.
The zooplankton samples were collected every
month from February 2007 to January 2008. At each
site 50 l of water were collected from the river drift, the
water was filtered through a 25 mesh – sized plankton
net and then fixed in a 4-5% solution of formalin. A
Glass Sedgewick Rafter Counting Chamber was used
for counting. Zooplankton specimens were identified
and counted in five subsamples. For identification a
Nikon Eclipse 50i microscope was used. These species
identifications were made using the keys of Wagler [16],
Kutikova [17], Harding and Smith [18] and Rybak [19,20].
In each sample, the body length of at least 30 individuals
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Zooplankton communities of two lake outlets
in relation to abiotic factors
9
8
Secchi depth (m)
7
6
5
4
3
2
1
0
F
M
Figure 2.
A
M
J
J
Dominikowo Wielkie
A
S
O
N
D
Nowa Korytnica
J
Seasonal variations of Secchi depth in the study flowthrough lakes.
from each species was measured by the Pixelink
Camera Kit 4.2 computer program. If the number of
individuals representing a given species was lower than
30, the body lengths of all individuals were measured.
The body length conversion to wet mass was made with
the use of the Ejsmont-Karabin [21], Ruttner-Kolisko
[22], McCauley [23] tables.
At sites S1, K1 and S3, K3 measurements of
temperature, pH, conductivity, dissolved oxygen, and
BOD5 were made by an oxygen content meter and pH
meter CX-401 made by Elmetron (Poland). The contents
of ammonium, nitrites, nitrates, and ortophosphates
were measured by a photometer DR-850 made by Hach
Lange (USA). At sites S2 and K2 no such measurements
were made because the differences in the parameters
measured over a distance of 0.2 km were expected to
be very small. At each site the width, depth and rate
of the river flow were measured (water flow sensor
OTT, Germany) to determine the discharge of water
and the mass of the zooplankton carried. Each month
water transparency was measured by Secchi-disk. In
the month when the Secchi depth was the lowest, the
trophic status of the lake was determined and expressed
in terms of the Carlson index [24]:
TSISD = 10 (6 – log2 SD),
where: SD – visibility of Secchi disc in metres.
In order to examine zooplankton diversity, we
calculated Shannon-Weaver index using the quantitative
zooplankton sample [25].
The statistical significance in the differences of some
properties of the zooplankton community between the
outflow and the lower course of the river between and
in the zooplankton community between the two rivers
with different trophy was tested using a pairwise t-test
(P<0.05). For the comparisons among different points
along the same river and between the same points
on different rivers, values of percentage decrease of
species number, abundance, biomass and body-size in
the 12 different months were used. Two sites form two
base sets with twelve data.
In order to determine the influence of the chemical
and physical parameters on the abundance of particular
zooplankton species, the Canonical Correspondence
Analysis (CCA) was applied with the software Vegan
1.15.1 [26].
3. Results
The value of TSISD calculated for Dominikowo Wielkie
Lake was 40, which corresponds to mesotrophic
water. In Nowa Korytnica Lake TSISD was 65.1, which
corresponds to extremely eutrophic or even polytrophic
water. In Dominikowo Wielkie Lake the mean value of
Secchi depth was 6.3 m and varied from 4 m (in June
and July) to 8 m (from November to February). In Nowa
2,6
1,3
2,4
Discharge (m s )
1,1
2,2
-1
1
3
3
-1
Discharge (m s )
1,2
0,9
0,8
0,7
2
1,8
1,6
1,4
0,6
1,2
0,5
0,4
1
F
M
A
M
J
S1
Figure 3.
J
A
S2
S
O
N
D
J
F
M
A
S3
M
J
K1
J
A
K2
S
O
N
D
J
K3
Seasonal variations of discharge in Słopica (S1. S2, S3) and Korytnica (K1, K2, K3).
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R. Czerniawski, J. Domagała
Figure 4.
Seasonal changes in zooplankton community in Słopica river.
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Zooplankton communities of two lake outlets
in relation to abiotic factors
Figure 5.
Seasonal changes in zooplankton community in Korytnica river.
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R. Czerniawski, J. Domagała
Korytnica the mean value of Secchi depth was 1.8 m
and varied from 0.7 m (in June and July) to 3 m (from
December to February) (Figure 2). Discharge values
were higher in Korytnica than in Słopica (Figure 3).
Słopica river was characterised by higher values
of the Shannon-Weaver index in all sites compared to
Korytnica river (Figure 4,5). These values generally
declined downstream, but differences between sites
and between rivers were insignificant.
Greater changes between sites in the decrease
of zooplankton community were observed in Słopica,
flowing out from mesotrophic lake (Figure 4). The
smallest differences were found in Rotifera, of which
the significant reduction of biomass was noted at S2
and S3 (P=0,0170; P=0,0455 respectively). The highest
decrease in rotifer biomass, always over 70%, at S2 was
observed in March, June, August and in January, however
at S3 in every month, except January. Other parameters
of rotifers in Słopica decreased insignificantly.
The greatest changes when considering all
parameters were found in the Cladocera. At Słopica, in
all months except March and November, cladocerans
were not noted at last site (Table 1). At S2 reduction
of the cladocerans species number was also high,
and in June, August and January they were not noted
at this site. In most other months the species number
of cladocerans decreased at least by 50% (Figure 4).
A significant decrease of cladocerans abundance
was noted at S2 (P=0,0455). The highest reduction of
cladocerans abundance occured in April (over 70%). A
significant reduction in cladoceran biomass in Słopica
was noted also at S2 (P=0,0421). The highest reduction
of this parameter was observed in four months, February,
April, July and December, and was always over 90%. The
body-length of cladocerans decreased significantly at S2,
with the highest reduction occurring in February (79%).
Copepods in Słopica in June were present only
at S1, in October and in May at S1 and S2, and in
September they were not noted at all sites (Table 1).
A significant reduction in the abundance, biomass
and body size of copepods was observed only at S2
(P=0,0068; P=0,0402; P=0,0127 respectively). The
greatest difference in copepods abundance was noted
between S1 and S2 in December (52%) and in January
(57%). The greatest reduction in copepod biomass was
noted also in January (75%), whereas the greatest
reduction in body size occurred in April 56% (Figure 4).
In Korytnica, which flows out from the strongly
eutrophic lake, differences found in some parameters of
the zooplankton community were much lower between
sites than found in Słopica. At the last site (K3) Cladocera
and Copepoda were not observed for six months and
five months respectively (Table 2). In Korytnica, a
significant reduction of abundance, biomass and body
size of zooplankton was noted only at S3 and concerned
only crustaceans (Table 3). In May at K2, cladoceran
biomass was 63% higher than at the outflow (K1)
(Figure 5). However the body-length of cladocerans
and rotifers was higher at K2 than at K1 for six months
yielding the higher annual average body size at K2
compared to K1. Reduction in mean cladoceran body
size did not occur at the second site (K2).
The comparisons between the two rivers in percent
reduction in mean species number, abundance,
biomass and body size for the main taxonomic groups
demonstrated that in the first section of the river the
decline was faster in Słopica than in Korytnica (Table 3).
Significant differences between both rivers in decrease
of some parameters of the zooplankton community were
only observed in these sections of rivers (S1-S2 vs. K1K2) and concerned biomass of rotifers (P=0,0198) and
cladocerans (P=0,0193), as well as body size of rotifers
(P=0,0109), cladocerans (P=0,0001) and copepods
(P=0,0462) (Table 3). Between river comparisons in
the second section (0.2-1 km) revealed no significant
differences in some parameters of zooplankton
community. Thus, in the first section, differences
between rivers were significant, however in the second
section differences between rivers were insignificant.
The number of species in both rivers declined faster
in the second section (0.2-1 km), except in rotifers,
although this difference was small. The extent of the
reduction in abundance, biomass and body size of
zooplankton differed between Słopica and Korytnica.
In Słopica abundance, biomass and body size of
cladocerans decreased faster in first section of the river
(0-0.2 km) than in second section, however in Korytnica
the pattern was inverted. Abundance and biomass of
rotifers was reduced faster in second section of both
rivers than in first section (Table 3).
To sum up, in the first section of river (0-0,2km),
higher and significant changes in the reduction of the
parameters of zooplankton community occurred in
Słopica river, which flows out from the mesotrophic
lake, than in Korytnica river, which flows out from the
eutrophic lake. However, in the second section of the
rivers (0.2-1 km) the decline in several zooplankton
community parameters were similar in both rivers and
there was a lack of significant differences between
them. Only species number of every taxonomic group
decreased faster in the second section of rivers than in
the first section. Quantitative reduction of crustaceans
was faster at the first section in Słopica, but faster in
the second section in Korytnica. Rotifer communities
declined faster at the second section in both rivers,
irrespective of the lake’s trophy.
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Zooplankton communities of two lake outlets
in relation to abiotic factors
Taxa
Anuraeopsis fissa (Gosse,1851)
S1
S2
S3
M;N
Ma;O
Ma
Ascomorpha ecaudis Perty, 1850
Ma;Jy
Ma;Jy
Ascomorpha ovalis (Bergendal, 1892)
Jy;A;S
Jy;Au;S
Au;S
Ascomorpha saltans Bartsch, 1870
A;D
Au
Au
Asplanchna brightwellii Gosse,1850
Ma
Asplanchna herricki De Guerne, 1888
A
Asplanchna priodonta Gosse,1850
F;M;A;Ma;Au;O;N;J
Asplanchna sieboldi (Leydig, 1854)
M;Au
Brachionus calyciflorus Pallas,1766
O
Cephalodella sterea (Gosse, 1887)
Conochilus unicornis Rousselet, 1892
F;M;A;N;D;J
Jy
A;Ma;O;N
A;Ma
S
S
Conochiloides natans (Seligo, 1900)
Au;D
A;O
Colurella adriatica Ehrenberg, 1831
Jy;S;J
Jy;Au;S;N;J
Conochiloides coenobasis (Skorikov, 1914)
Euchlanis lyra Hudson, 1886
Au
Euchlanis triquerta Ehrenberg, 1838
Ma
Filinia longiseta (Ehrenberg,1834)
Filinia terminalis (Plate, 1886)
Kellicotia longispina (Kellicott, 1879)
Keratella coclearis cochlearis (Gosse, 1851)
Keratella c. hispida (Lauterborn, 1898)
F;A;N;D;J
A;Ma
Jy;Au;S;N;J
A
A
A
A
M;A;Ma;Je;Au;J
A;Ma;Je;Jy
A;Ma;Je
100%
100%
100%
Ma
Ma
Kerataella c. tecta (Gosse, 1851)
Jy;Au;S;N;J
M;S;O
M
Keratella quadrata (Müller, 1786)
A;Ma;Jy;Au;S;O;N;D
A;Ma;S;N
A;Ma;N
Jy;S
S
Lecane aculeata (Jakubski, 1912)
Lecane closterocerca (Schmarda, 1859)
Lecane luna ( Müller ,1776)
Lepadella acuminata (Ehrenberg, 1834)
N
Jy;S;O;N
Au
Je;Jy;N
Lepadella astacicola Hauer, 1926
Lepadella costata Wulfert, 1940
O
Au
Au
Lepadella ovalis ( Müller ,1786)
N
F;N
Mytylina mucronata ( Müller ,1773)
Au
Notholca foliacea (Ehrenberg, 1838)
M
Notholca labis Gosse, 1887
A
Polyarthra dolichoptera Idelson, 1925
A;Ma;N;D;J
Polyarthra euryptera Wierzejski, 1891
J
M;A
M
M;A;Ma;N;D;J
M;A;Ma;N;D;J
Polyarthra longiremis Carlin, 1943
Ma;Je;Jy;Au;S
Ma;Au;S
Ma;Au;S
Polyarthra major Burckhardt, 1900
M;Ma;Je
M;A
M
Polyarthra remata Skorikov, 1896
Ma;Je
Polyarthra vulgaris Carlin, 1943
A;Je;Jy;Au;S
A;Ma;Je;Jy;Au;S;N
A;Je;S
Pompholyx complanata Gosse, 1851
Je;Jy;Au;N;D
Jy;Au;O;N
Au;N
M;Je;Jy;Au;O;N
M;Au;N;D
Au;N
Jy
O
Pompholyx sulcata Hudson,1885
Proales daphnicola Thomson, 1892
Squatinella rostrum (Schmarda, 1846)
Synchaeta kinina Rousselet, 1902
Au
Je;Jy;S;O
Jy
Synchaeta lakowitziana Lucks, 1930
Je
Je
Synchaeta oblonga Ehrenberg, 1831
M;A;N;J
M;A
Testudinella patina (Hermann, 1783)
Ma
Ma
Table 1.
Jy
M;A
Taxonomic composition and occurrence data of zooplankton in Słopica River. F-February, M-March, A-April, Ma-May, Je-June, Jy-July,
Au-August, S-September, O-October, N-November, D-December, J-January.
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Taxa
Testudinella truncata (Gosse, 1886)
Trichocerca capucina (Wierz. et Zach. 1893)
Trichocerca dixon-nuttalli (Jennings, 1903)
S1
S2
Au
Au
Jy;Au;S
Au;S
O
O
Trichocerca elongata (Gosse, 1886)
Au
O
Trichocerca pusilla (Lauterborn, 1898)
Au
Trichocerca similis (Wierzejski, 1893)
M;Je;Jy;Au;
M;Je;Jy;Au
M;Jy;Au
N
D
D
F;Ma;Jy;Au;S;O;N
F;A;Ma;Au;S;D;J
F;M;A;D
A;D
Trichortia pocillum ( Müller, 1776)
Rotatoria non det.
Alonella nana (Baird,1843)
Ma
Bosmina coregoni Baird, 1857
A;Je;J
F;A;Ma;D
Ceriodaphnia quadrangula (Müller, 1785 )
N
O
Chydorus gibbus (Lilljeborg, 1880)
A
A
Chydorus sphaericus ( Müller, 1785)
M;Jy;Au;S
A;S
A
Daphnia cucullata Sars, 1862
F,M;Je;Jy;Au;N;D
M;A;Jy;N
A
Leptodora kindtii (Focke,1844)
Au
except Je;S;O
Nauplii Cyclopoida
except S
except Je;S
Nauplii Calanoida
M;Ma;Je;Jy;Au
M;Jy
Au
Copepodites Cyclopoida
except S
F;M;A;Jy;O;N;D;J
F;M;A;N;D
Copepodites Calanoida
F;Ma;Au;N
F;Au;J
F;J
F;M;Je;N
F;J
F;J
F;M;Au;N;D;J
F;M;D
D
F;M;N;D;J
F;M;D;J
F;M;D;J
F;M;D;J
F;M;D;J
F;D;J
Eudiaptomus gracilis (G.O.Sars, 1863)
Eudiaptomus graciloides (Lilljeborg, 1888)
Cyclops kolensis Lilljeborg, 1901
Cyclops vicinus Uljanin, 1875
Diacyclops bicuspidatus (Claus, 1857)
Au;O
Eucyclops serrulatus (Fischer, 1851)
Au
Metacyclops gracilis (Lilljeborg, 1853)
O
Thermocyclops crassus (Fischer, 1853)
A
Thermocyclops emini (Mrazek, 1895)
O
A
Thermocyclops oblongatus (Sars, 1927)
M;O
M;N
Thermocyclops oithonoides (Sars, 1863)
Au;O;N
N
N
D
M
Harpacticoida
continued
S3
Table 1.
Taxonomic composition and occurrence data of zooplankton in Słopica River. F-February, M-March, A-April, Ma-May, Je-June,
Jy-July, Au-August, S-September, O-October, N-November, D-December, J-January.
Analysis of CCA revealed that conductivity, Secchi
depth and PO4 correlated best with the first axis. NO3
and temperature correlated little bit less with this axis.
However, PO4 and discharge correlated best with
the second axis (Figure 6). Thus, all physical factors
influenced the abundance of zooplankton. CCA showed
that abundance values of the two rivers were clearly
divided along the first and second axis. Points of months
in every river located close to each other, irrespective of
sampling site. Thus, in the two rivers the abundance of
zooplankton was correlated with different parameters.
Samples from Słopica showed a greater correlation
with the Secchi depth, particularly in the winter months.
In contrast, samples from Korytnica correlated more
closely to conductivity, discharge, temperature, PO4 and
NO3. The Secchi depth seemed to have impact on the
abundance of big copepods such as Eudiaptomus sp.
and Cyclops sp. in the winter months in Słopica. In
Korytnica, however, the influence of Secchi depth on
zooplankton abundance was observed in December
only, and was associated with Cyclops sp. and
Asplanchna priodonta. Additional parameters had strong
impact on zooplankton abundance only in Korytnica.
Orthophosphates, nitrates, temperature and conductivity
were all correlated with the abundance of rotifers (mainly
small species) in the summer months. Discharge was
correlated with the abundance of Chydorus sp. and
small rotifers in September. It is possible that additional
unmeasured abiotic parameters in the two different rivers
had significant impact on the zooplankton community.
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Zooplankton communities of two lake outlets
in relation to abiotic factors
Taxa
Anuraeopsis fissa
Ascomorpha ovalis
Ascomorpha saltans
K1
K2
K3
Ma;Je;Jy;Au;S;O
Ma;Je;Jy;Au;S;O
Ma;Je;Jy;Au;S;O
D
D
Ma;Jy;Au
D
Ascomorphella vovociola (Plate, 1886)
S
Asplanchna brightwellii
Asplanchna priodonta
Brachionus angularis Gosse, 1851
D
F;A;N;D
F;M;A;Ma;N;D
M;A;D
F;M;A;Ma;Au;N;J
F;M;Je
M;Je
Brachionus budapestinensis Daday, 1885
Ma;S
Brachionus calyciflorus
M;A
F;M;A;D
M;A
A;Je;Au
Jy;Au
Au
Brachionus diversicornis (Daday, 1883)
Brachionus quadridentatus Hermann, 1783
Brachionus urceus
Cephalodella apocolea Myers, 1924
Conochilus unicornis
Colurella adriatica
M;Je;Jy;Au
M
S
Je;Au
A;Ma;S;O;N
A;S;O;N
A;S;O;N
Jy;S;N
Je;S
Je;S
Au
Au
Colurella colurus (Ehrenberg, 1830)
S
Elosa worallii Lord, 1891
Au
Euchlanis deflexa Gosse, 1851
A
Euchlanis dilatata Ehrenberg, 1832
Ma
Je
Je
Filinia longiseta
M;A
A;Ma
A
Filinia terminalis
A
A
A
Gastropus stylifer Imhof, 1891
Je
Itura aurita (Ehrenberg, 1830)
S
M;S
S
Kellicotia longispina
except F;J
A;Ma;Je;Jy;S;O;N;D
Ma;Je;Jy;S;O;N
Keratella coch. cochlearis
except Au
except Jy;Au
except Jy;Au
Keratella coch. hispida
Kerataella coch. tecta
Keratella quadrata
Je
Je
except D;J
Ma;Je;Jy;Au;S;O
Ma;Je;Jy;Au;S;O
except Je;Jy
except M;J
except F;M;J
Keratella ticinensis (Callerio, 1920)
Ma
Lecane arcuata (Bryce, 1891)
Au
Lecane clara (Bryce, 1892)
Lecane closterocerca
Lecane lunaris (Ehrenberg, 1832)
Ma;Je;Au;O
Au
Au
O
Au
Lepadella ovalis
Mytilina crassipes (Lucks, 1912)
Jy
J
Lecane tenuisecta (Harring, 1914)
Ma
Au
Au
Au
Mytlina mucronata
Au;S
Au;S
Au;S
Notholca foliacea
M
Notholca squamula (Müller, 1786)
M
M
M
Polyarthra dolichoptera
F;M;A;Ma;O;N;J
A;Ma;N;J
A;Ma;N;J
Polyarthra major
M;A
M;A;Ma
M;A
Polyarthra remata
M
Polyarthra vulgaris
M;A;Ma;O
M;A;Ma;S
M;A;Ma;S
F;M;Ma;Je;Jy;Au;S;O
M;Je;Jy;Au;S;O
Ma;Je;Jy;Au;O
Je;Jy;Au;O
Pompholyx complanata
Pompholyx sulcata
Squatinella rostrum
Synchaeta kitina
Table 2.
S
Ma
F;M;Ma;Je;Jy;Au;S;O
Je
Ma;Je;Au;O
Taxonomic composition and occurrence data of zooplankton in Korytnica River. Month symbols see Figure 1.
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R. Czerniawski, J. Domagała
Taxa
K1
K2
F;M;A;Ma;O;N;D;J
F;M;A;Ma;Je;N;J
Synchaeta lakowitziana
Synchaeta oblonga
D
Synchaeta pectinata Ehrenberg, 1832
F;M;A;Ma;N;J
N
Testudinella mucronata (Gosse, 1886)
O
Testudinella patina
M
M
Jy;Au
Au
Au
Trichocerca brachyura (Gosse, 1851)
S
S
S
Trichocerca dixon-nuttalli
S
S
S
Je;Jy;Au
Je;Jy;Au
Je;Jy;Au
Testudinella truncata
Trichocerca pusilla
Trichocerca rattus (Müller, 1776)
Je
Trichocerca rousseleti (Voight, 1902)
O
O
O
Trichocerca similis
F;Je;Jy;Au
Je;Jy;Au
Je;Jy;Au
Rotatoria non det.
A;Ma;Je;Au;S;O;N
M;Je;Jy;S;O
Ma;S;O
Alona rectangula Sars, 1861
Bosmina longirostris
Bosmina coregoni
F;Je
Ma;Je;N
A;Ma
Ma
A;Ma;Je;Jy;O;N;D
M;A;Ma;Je;Jy;O;N;D
Ma;Je;Jy,N
Jy
Je
Ceriodaphnia laticaudata (P. E. Müller, 1867)
Ceriodaphnia quadrangula
Jy
Chydorus gibbus
A;S;O
Je;S
M;A;Jy;S;D
Jy;S
S
Ma;Je;Jy;S;O;N;D
Ma;Je;Au;S;O;N;D
N
D
D
Nauplii Cyclopoida
100%
100%
except F
Nauplii Calanoida
O
except J
except J
Ma;Je;Jy;Au;N;D
Je
Je
Cyclops kolensis
N;D
N;D
D
Cyclops vicinus
D
D
D
Eucyclops serrulatus
Ma
Ma
Metacyclops gracilis
Je
Chydorus sphaericus
Daphnia cucullata
Daphnia longispina (Müller, 1785)
Copepodites Cyclopoida
Acanthocyclops robustus (G. O. Sars, 1863)
Thermocyclops crassus
Thermocyclops oithonoides
Table 2.
Je;S
Au;S;O
S;O
Ma;Je;Au
Je;Au
Je
Je
Au
Ma
Harpacticoida
continued
K3
Taxonomic composition and occurrence data of zooplankton in Korytnica River. Month symbols see Figure 1.
4. Discussion
Differences in some properties of the zooplankton
community between the outflow and the lower course
of a river are typical for lake-river systems. A greater
reduction in the number of species, abundance,
biomass and body size of the zooplankton community
was observed in the river Słopica. In shallow rivers, like
in Słopica, a rapid decrease of zooplankton abundance
has been noted: rotifers are often reduced the least,
while cladocerans show the greatest decline [8,9,27].
In Słopica no significant reduction in the abundance,
biomass or size of rotifers was observed, in any section
of the river. In small rivers, especially in their lower
sections, the abundance of rotifers is usually large
[9,14,28]. Perhaps planktivorous fish, which mainly
reduce the abundance of the zooplankton [2,9,27,29],
selected for larger specimens and did not influence the
abundance of small rotifers. Chang et al. [9] found very
small quantities of small rotifers in stomachs of fish,
even on sites far from the outflow. However, big rotifers
(e.g. Asplanchna sp. or Brachionus calyciflorus), for
which abundance at S3 was very small, could possibly
be eaten by fish [9].
The cladoceran community was strongly and
significantly reduced abruptly in the first section of the
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Zooplankton communities of two lake outlets
in relation to abiotic factors
S1 - S2
K1 - K2
S2 - S3
K2 - K3
Species number
Rotifera
21
19
34
17
Cladocera
44
24
54
58
Copepoda
24
10
43
62
Abundance
Rotifera
39
32
57
48
Cladocera
59*
34
56
78*
Copepoda
66*
46
57
51*
Biomass
Rotifera
54*
17
67*
50
Cladocera
77*
4
65
80*
Copepoda
72*
57
65
86*
Body-length
Rotifera
21
-1
20
10
Cladocera
56*
-3
56*
61*
Copepoda
35*
13
33
43*
Table 3.
Percentage decrease of four parameters of the zooplankton
community between sites in the two rivers.
Significant differences in the decrease of the parameters between
sites are marked with an asterisk;
Significant differences in the decrease of the parameters between the
same sections of two different rivers are marked with bold.
Słopica river. Similar results were reported by Chang
et al. [9] who noted the greatest quantitative reduction
in cladoceran community at the river section between
the outflow and the first site below it. Additionally Chang
et al. [9] revealed a high amount of Cladocera in the
stomach of fish, suggesting that cladocerans may be
important prey for planktivorous fish. Shallow lake
outlets, such as outlets of Słopica river, have often
been observed as areas where fish fry accumulate to
fed on the zooplankton carried out from lakes [30]. At
these sites the reduction of zooplankton is the greatest
[2,9]. Copepods were found much more often at the
last site compared to cladocerans, because of the
presence of small copepods naupliis. Chang et al. [9]
also noted a relatively high amount of nauplii even at a
site situated 1.4 km away from outflow. The reduction
of copepods abundance and biomass observed at S2
was considerable and significant only in cold months,
when mainly large and adult specimens dominated
the community. To sum up, the higher decrease of
crustacean parameters was observed at our second
site, however parameters of rotifers community reduced
faster at third site.
Significant differences in some parameters of the
zooplankton community were observed between two
rivers flowing out of separate lakes differing in trophy.
Significant reductions in the biomass of rotifers and
cladocerans, and in the body size of every taxonomical
group were only noted between the first sections of
the rivers (S1-S2 vs. K1-K2). These differences could
be related to different conditions of the lakes from
which the rivers flow out, at least in their first sections.
In first section of Korytnica the decline in zooplankton
community parameters proceeded much more slowly
than in Słopica. This could be due to differences in
the transparency of water, hydrogeological conditions
of the rivers, or the degree of macrophytes coverage
along the bottom. Perhaps planktivorous fish attacked
plankters ineffectively, because of low transparency
and high number of zooplankton refugees. Although
planktivorous fish and macro-invertebrates control the
zooplankton community in the rivers, the most important
controlling factors of zooplankton production are
physical, particularly river hydrology [4,31].
In Korytnica river, particularly in zones with abundant
vegetation and in the small floodplain between the
outflow and site K2, the current speed of the river
was reduced such that, at a few sites near the banks
and in the floodplain, the water velocity was strongly
inhibited or the water current stopped. Such conditions
may have favoured the reproduction of zooplankton
between the two sites K1 and K2; in particular the
species of Cladocera, which were not noted at the
river outflow (K1). According to Estlander et al. [12],
in lakes the abundance of Cladocera increases with
increasing development of vegetation. Czerniawski [14]
also reported from floodplain outflow other species of
Cladocera and Rotifera than those found at the above
outflows of the rivers from the lakes. In Słopica, where
the current speed could not slow down, the species
observed at the outflow and the subsequent sites were
generally the same. Similar results were reported by
Chang et al. [9]. In Korytnica, the type of water vegetation
growing between the first two sites could be responsible
for the greater body length of Cladocera at K2 than
at K1. It is well known that submerged macrophytes
can be an effective hiding place for zooplankton from
predatory fish [32,33]. Thus, the dense vegetation over
this section of Korytnica could protect larger Cladocera
well. Accordingly, species of Cladocera reached high
abundance between K1 and K2, where macrophytes
covered the bottom of the river [34,35]. The smaller
reduction of the investigated parameters of Copepoda
community at K2 than at S2 could also be related to the
differences in the hydrological conditions of the river. At
the outflow of Korytnica, which was densely covered with
macrophytes, Copepoda could hide in macrophytes and
avoid being washed away by the river current. Gliwicz [36]
and O’Brien [37] report that Copepoda are able to swim
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Figure 6.
CCA constrained ordination of the samples and taxa from different rivers (filled symbols: Słopica, open symbols: Korytnica), sites
(triangles, circles and squares: sites 1, 2 and 3, respectively) and sampling months (numbers). Abiotic variables: temp –
temperature; SD – Secchi depth; cond – conuctivity; disch – discharge; NO2 – nitrites; NO3 – nitrates; NH3 – ammonia;
O2 – dissloved oxygen; PO4 – orthophosphates; BOD5 - Biochemical Oxygen Demand. Taxa: Anu. fis. – Anuraeopsis
fissa; Asc. ova. – Ascomorpha ovalis; Asc. sal. – A. saltans; Asp. pri. – Asplanchna priodonta; Bos. cor. – Bosmina
coregoni; Bos. lon. – B. longirostris; Bra. ang. – Brachionus angularis; Bra. bud. – B. budapestinensis; Bra. cal. –
B. calyciflorus; Bra. div. – B. diversicornis; Bra. qua. – B. quadridentatus; Cal. cop. – calanoids copepodids; Cal. nau. – calanoids
nauplii; Chyd. Gib. – Chydorus gibbus; Chyd. sph. – Ch. sphaericus; Col. adr. – Colurella adriatica; Con. nat. – Conochiloides natans;
Con. uni. – C. unicornis; Cyc. cop. – cyclopids copepodits; Cyc. kol. – Cyclops kolensis; Cyc. nau. – cyclopoids nauplii; Cyc. vic. –
C. vicinus; Dap. cuc. – Daphnia cucullata; Euc. dil. – Euchlanis dilatata; Eud. gra. – Eudiaptomus gracislis; Eud. grac. – E. graciloides;
Elo. wor. – Elosa worali; Euc. ser. – Eucyclops serrulatus; Fil.lon. – Filinia longiseta; Fil. ter. – F. terminalis; Harpac. – Harpacticoida;
Itu. aur. – Itura aurita; Kel. lon. – Kellicotia longispina; Ker. c. coc. – Keratella cochlearis cochlearis; Ker. c. his. – K. c. hispida;
Ker. c. tec. - Ker. c. tecta; Ker. qua. – K. quadrata; Lec. clo. – Lecane closterocerca; Lep. acu. – Lepadella acuminata; Myt. cra. – Mytilina
crassipes; Myt. muc. – M. mucronata; Not. squ. – Notholca squamula; Pol. dol. – Polyarthra dolichoptera; Pol. lon. – P. longiremis;
Pol. maj. – P. major; Pol. rem. – P. remata; Pol. vul. – P. vulgaris; Pom. com. – Pompholyx complanata; Pom. sul. – P. sulcata;
Rot. nd. – rotifers not identified; Syn. kit. – Synchaeta kitina; Syn. obl. – Synchaeta oblonga; Tes. tru. – Testudinella truncata; The. cra.
– Thermocyclops crassus; The. oit. – T. oithonoides; Tri. bra. – Trichocerca brachyura; Tri. cap. – T. capucina; Tri. dix. – T. dixon-nuttalli;
Tri. pus. – T. pusilla; Tri. rou. – T. rouseletti; Tri. sim.– T. similis; Tri. puc. – Trichotria pocillum.
faster than Cladocera and can more effectively avoid
the attack of predatory fish. Because of the probable
decrease in the current speed of the River Korytnica
favouring the escape of large Copepoda, the Copepoda
biomass was larger in Korytnica than in Słopica.
Between the second parts of the two rivers (S3-S3
vs. K3-K3) a significant difference was not observed in
the reduction of the investigated parameters of every
taxonomic group. An explanation for this phenomenon
may be that in the last section of the rivers, the reduction
of the investigated parameters of the zooplankton
community proceeded similarly and the effect of the
lake on the community decreased. In exception, the
parameters of the rotifer community were reduced at
their greatest in the second section. At the third sites,
fish could eat small rotifers and could not select for larger
crustaceans as they were practically absent. Chang
et al. [9] analysed the content of the alimentary track of
fish feeding below the lake outflow and reported finding
crustaceans much more often than small rotifers.
Primarily at site 3 in both rivers, rotifers dominated
crustaceans in S-W index, the number of species and
abundance. Similar observations were reported by
[2,9,14]. The highest relative abundance at all sites in
both rivers was found for the most commonly occurring
species, K. c. cochlearis and nauplii ciclopoida, both
of which are characteristic of a large number of rivers
flowing out from lowland lakes [9,14,28]. In Słopica,
large sized components of zooplankton were relatively
frequent, which is typical of waters with a low degree of
eutrophication. In contrast, in Korytnica K. c. tecta and
P. sulcata were relatively frequent, and characteristic for
waters with high trophic status [10,38,39].
In sum, the different trophic conditions found in lakes
influenced the changes of the zooplankton community
structure in the outflowing rivers, but only in close
proximity to the lakes. The flow through lakes can exert
considerable influence on the character of the river
below its outflow [40] and disturb the river continuum
shaped along its course. This may occur through the
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Zooplankton communities of two lake outlets
in relation to abiotic factors
elimination of typical river organisms, the appearance
of large amounts of plankton and predators, or rapid
decrease in the oxygen content in the river water [41,42].
Finally, we conclude that as distance from the lake
increases the qualitative and quantitative composition
of the zooplankton becomes more similar, irrespective
of the character of the lakes from which the rivers flow
out.
Moreover, it seems that the chemical factors of
rivers did not have a significant effect on differences
in zooplankton community between our chosen rivers.
In both rivers, the most probable factors limiting the
structure of zooplankton were physical factors, which
could be correlated with the efficiency of predatory fish.
According to Chang et al. [9] the zooplankton produced
at the lake outlets are the main component of the diet of
fish and fish fry found in the lower courses of the rivers.
Analysis of CCA revealed that all physical parameters
had an impact on zooplankton abundance in both rivers,
but particularly in Słopica. In these rivers, positive
correlations were found between the abundance of large
copepods and transparency only in cold season. This
fact can be explained by the presence of large Calanoida
and Cyclopoida in the seasons when the transparency is
greater and when small numbers of fish were observed.
The Secchi depth in lakes was negatively correlated with
zooplankton abundance in both rivers. An explanation for
this may be that greater water transparency facilitated
the zooplankton consumption by fish. According to
Wissel et al. [43] smaller abundance of zooplankton can
be a consequence of increased water transparency as
the latter facilitates predation by fish.
In lakes, stronger correlations between the
zooplankton community and abiotic factors are observed,
in particular those characterising the water chemistry
[11,44,45]. In contrast, in water courses, zooplankton
abundance is correlated more strongly with physical
parameters, mainly river regime [31,46]. According to
our results, these generalisations fit only on river Słopica,
which was characterised by good transparency, lack of
macrophytes, floodplains or slack waters. In Korytnica
the influence of chemical parameters, such as nutrients
and conductivity was visible, particularly on rotifers in
warm seasons when the trophic status was the highest.
The high trophic status of Nowa Korytnica lake had a
significant impact on the rotifer community in the outflow.
Additionally, rotifers play a significant role in phosphorus
recycling [47], so they were strongly correlated with
orthophosphates in the water of Korytnica.
In Korytnica we observed the positive influence of
discharge on the abundance of small cladocerans and
small rotifers, mainly in September when the discharge
rapidly increased. Campbell [48] claims that high flow
rate and high current speed favour increasing plankton
abundance. Perhaps, the pressure of planktivorous fish
on zooplankton exported from lakes is smaller, because
of high current speed that drifts zooplankton faster,
making it hard to reach. Although, according to Chang
et al. [9] and Nielsen et al. [49], the relationship between
flow rate and the quantitative structure of the zooplankton
downstream has yet to be clearly explained.
In conclusion, our results show that: (1) the most
significant differences in the zooplankton community
between the outflow and the lower course of the river
happen in the first section directly after the outflow, (2)
significant differences in the zooplankton community
between the two rivers flowing out of the lakes with
different trophy was only found only crustaceans, which
declined more quickly in the river flowing out from the
mesotrophic lake, (3) only physical parameters had an
impact on the zooplankton community in the river flowing
out from the mesotrophic lake, however in the river
discharging from the strongly eutrophic lake chemical
parameters also had an impact.
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254
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R. Czerniawski, J. Domagała
Appendix
The mean content ± SD, range of physical and chemical parameters in both rivers Słopica (S 1; S 3)
and Korytnica (K 1; K 3).
Station
T
(°C)
pH
conduct.
(µS)
O2
(mg l-1)
BOD5
(mg l-1)
N-NO3
(mg l-1)
N-NO2
(mg l-1)
N-NH3
(mg l-1)
P-PO4
(mg l-1)
S1
Range
12.3 ± 8.1
1.4 – 23.8
8.07 ± 0.46
7.01 – 8.66
244.3 ± 54.7
182.0 – 348.1
10.19 ± 1.72
8.57 – 12,86
3.99 ± 0.98
2.60 – 5.02
0.4 ± 0.2
0.2 – 0.7
0.010 ± 0.010
0.001 – 0.030
0.06 ± 0.06
0.01 – 0.18
0.31 ± 0.35
0.01 – 1.12
S2
Range
12.1 ± 7.7
1.9 – 23.2
8.06 ± 0.48
7.12 – 8.55
232.1 ± 49.8
164.3 – 324.2
9.00 ± 1.42
7.55 – 11.41
3.67 ± 0.81
2.41 – 4.64
0.2 ± 0.15
0.1 – 0.5
0.004 ± 0.003
0.001 – 0.008
0.03 ± 0.04
0.01 – 0.11
0.18 ± 0.25
0.01 – 0.82
K1
Range
12.8 ± 8.0
1.6 – 23.8
8.61 ± 0.32
8.01 – 9.05
296.7 ± 52.1
210.9 – 399.2
10.09 ± 2.54
5,71 – 14,51
4.29 ± 0.88
3.21 – 5.55
0.5 ± 0.3
0.1 – 0.9
0.011 ± 0.001
0.006 – 0.020
0.11 ± 0.07
0.02 – 0.21
0.36 ± 0.57
0.04 – 2.09
K2
Range
12.7 ± 7.7
1.8 – 23.0
8.33 ± 0.33
7.72 – 8.96
289.7 ± 45.8
206.3 – 392.5
9.84 ± 2.55
5.11 – 14.62
4.08 ± 0.82
3.11 – 5.09
0.4 ± 0.25
0.1 – 0.8
0.009 ± 0.005
0.004 – 0.020
0.07 ± 0.07
0.04 – 0.17
0.30 ± 0.48
0.02 – 1.77
255
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