Acta Bot. Croat. 79 (1), 35–42, 2020
DOI: 10.37427/botcro-2020-009
CODEN: ABCRA 25
ISSN 0365-0588
eISSN 1847-8476
Alien water lettuce (Pistia stratiotes L.) outcompeted
native macrophytes and altered the ecological
conditions of a Sava oxbow lake (SE Slovenia)
Martina Jaklič1, Špela Koren2, Nejc Jogan3*
University Medical Centre, Zaloska 2, 1000 Ljubljana, Slovenia
1
2
Institute for Water of the Republic of Slovenia, Einspielerjeva ulica 6, 1000 Ljubljana, Slovenia
University of Ljubljana, Biotechnical Faculty, Department of Biology, Večna pot 111, 1000 Ljubljana, Slovenia
3
Abstract – Introduction of an invasive alien macrophyte water lettuce (Pistia stratiotes L.) radically changed the
oxbow lake in Prilipe (SE Slovenia) which has thermal springs that enables the winter survival of this tropical invader. About 10 years after the first record of P. stratiotes, the number, abundance and biomass of indigenous and
non-indigenous macrophytes as well as different abiotic parameters were measured. In that period, colonized
sections (~94% of the oxbow lake) were completely covered with water lettuce, and the only reservoirs of indigenous macrophyte species were the non-colonized areas (6%). Research in 2011 found only a third of the previously recorded indigenous macrophytes, but then only in small section without P. stratiotes. Three of the species
that disappeared were on the Red data list. In the colonized section a higher biomass was observed than in the
non-colonized section because of high abundance of water lettuce which remained the only macrophyte. Due to
the presence of P. stratiotes, the intensity of light penetrating into the depth and water circulation were reduced, as
was the oxygen saturation of the water. In addition to the well documented vegetative propagation of P. stratiotes,
a well-established and viable seed bank has been detected in the lake sediment and after winter floods also on lake
banks. In the future, special attention should be given to the thermal water ecosystems in temperate climates
since they can serve as stepping stones and recruitment centres for the establishment and spread of (sub-)tropical
invasive species. Facing predicted climate change such local populations of invasive species can act as stepping
stones for further dispersal.
Keywords: altering ecological conditions, non-indigenous macrophyte, outcompeting, thermal oxbow lake, transformer, tropical species, water lettuce
Introduction
Macrophytes have a great effect on nutrient cycling, the
concentration of dissolved oxygen, on the light conditions
in the water and on the biochemical reactions (Caraco and
Cole 2002, Chamier et al. 2012). Diversity of macrophytes
has been particularly affected by human activities and management in the last few decades. Introduction of non-indigenous macrophytes (i.e. alien plants) usually has consequences for ecosystem processes and results in changes of
biodiversity, reduction in the value of water for human activities and has a great influence on the water quality (Caraco and Cole 2002, DAISIE 2009). In addition, alien plants
can decrease or completely change indigenous macrophyte
biodiversity (DAISIE 2009, Ramey 2009). A good exam-
ple is the Canadian waterweed Elodea canadensis Michx.,
which outcompeted native macrophytes in Slovenian rivers (Jogan 2005). Tropical non-indigenous species can establish self-sustaining populations in temperate waters only
if water temperature and regime are appropriate. Thermal
water bodies have warmer water than non-thermal inland
waters and consequently contain unique native fauna and
flora (Castenholz 1973, Raab 1993, Kralj 2000, Zuellig et al.
2002, Laprida et al. 2006, Petutsching et al. 2008, Piazzini
et al. 2010).
Water lettuce (Pistia stratiotes L.) is a floating macrophyte originating in South America (Chamier et al. 2012).
It became a popular ornamental plant in aquaria and gar-
*Corresponding author e-mail: nejc.jogan@bf.uni-lj.si
ACTA BOT. CROAT. 79 (1), 2020
35
JAKLIČ M, KOREN Š, JOGAN N
den ponds and as a result of naturalization is one of the most
widespread aquatic weeds in almost all tropical and subtropical regions with extreme biomass production (Labrada and
Fornasari 2002, DAISIE 2009). In Europe, accidental release
of water lettuce is often noticed, but a low number of localities have stable populations (Šajna et al. 2007, Brundu et
al. 2012). The first record of water lettuce escape in Europe
was in 1973 in the Netherlands (Mennema 1977), while later it occasionally also occurred in other European countries
– France, south-west Spain and in Central Europe (Pyšek et
al. 2002, Šajna et al. 2007). In Eastern Europe (Moscow and
Astrakhan Oblast) it has been reported as an ephemerophyte
(Lisicyna et al. 2009). Recently it became established in water channels in northern Italy (Brundu et al. 2012), while it
has also been noticed in Serbia with casual occurrences in
some thermal waters and one record in non-thermal waters
in eastern Vojvodina but with very low frequency (Živković
et al. 2019). Despite many sporadic records in the temperate
region of Europe, winter survival has only been reported in
some localities in Austria (Hartl et al. 1992), Hungary (Simon
2000), Slovenia (Šajna et al. 2007) and Italy (Ercolini 2008).
In warmer regions worldwide different ways of controlling the populations of water lettuce are used including
chemical and physical methods, but no reports about total
local eradication could be found (Cilliers et al. 1996, Diop
and Hill 2009, Gherardi et al. 2011). Mitigation by biological control agents could be more successful (Cilliers et al.
1996, Ajuonu et al. 2009, Diop and Hill 2009, Baker et al.
2010), with the introduction of new competitive alien species, which might on the other hand, have additional impacts (usually negative) on native biodiversity (Lockwood
et al. 2007).
The oxbow lake in Prilipe (SE Slovenia) used to be an
ecosystem with high biodiversity and an important reservoir for freshwater fish. As such it has been recognized as a
natural value of state importance and as an area of ecological importance. During winter, numerous bird species stop
there, due to the lack of any ice covering in spite of the low
winter temperatures (Mirt 2009). Floristic inventories before
2001 were made on the whole area of the oxbow lake and the
first data of flora in the oxbow lake go back more than three
decades (Seliškar 1984) and later on some field records are
available from 1992 and 1997 when no alien macrophytes
were found (Jogan, unpbl.). Water lettuce was first observed
in this oxbow lake in 2001 (Hudoklin 2002, Haler 2005).
Since then water lettuce regenerates a large population every spring and rapidly grows luxuriantly to cover almost the
whole surface of the oxbow lake. In summer months only
about 4% of the water surface in the eastern part of the oxbow lake is devoid of lettuce (Šajna et al. 2007). Water lettuce reproduces by vegetative propagation and also by producing viable seeds (Haler 2005; Šajna et al. 2007). It is able
to survive winter conditions because of thermal springs (average temperature 15 °C) present in the oxbow lake (Haler
2005, Šajna et al. 2007). Only a small marginal part of the
oxbow lake was not yet colonized with water lettuce due to
low winter temperature (in average 8 °C) (Šajna et al. 2007).
Local communities and fishermen tried to reduce growth of
water lettuce using herbicide and physical removal. From
2003, there were several attempts to remove floating lettuce
biomass mechanically but without great success (Haler 2005,
Šajna et al. 2007). The fishermen used also floating wooden
barrier to fence water lettuce spread locally, but it turned out
to be only a short term solution until the first heavy rain.
The purpose of this study was to compare the Sava oxbow lake flora in Prilipe before and after colonization with
water lettuce. Moreover, objectives were also to examine
differences within colonized sections of the oxbow lake regarding water lettuce biomass, and physical and chemical
parameters, as well as to compare colonized areas covered
with water lettuce with non-colonized area with indigenous
macrophytes. Further on we checked viability of seeds to assume their potential role in lettuce population persistence
and spread.
Materials and methods
Study area
Slovenia is a country in the central part of Europe with
a mostly temperate climate. The study site is located in the
south-eastern part of Slovenia, near the village of Prilipe (vicinity of Brežice, 140 m a.s.l.); the oxbow lake is 4 km long
and the average water depth is approximately 1.5 m. It is
one of the very few remaining oxbow lakes along the Sava
River. In 1855, Čatež Spa has been established near the oxbow lake and today it is the largest natural spa in Slovenia
(Ivankovič and Nosan 1973, Shawish 2004). This oxbow lake
has a unique water temperature regime due to inflow from
underground thermal springs and the outflow of warm water from the pools inside the Čatež Spa resort. The study
area was divided into five sections A, B, C, D and E corresponding to water temperature gradient as already defined
by Jaklič and Vrezec (2011) (Fig. 1), sections A to D are referred as “colonized sections” in further text. Water temperature decreases downstream from section A (average annual
Tw = 35 °C) to section E (average annual Tw = 15 °C) (Tab. 1).
Tab. 1. Description of five sections of the Sava oxbow lake (data source: Jaklič and Vrezec 2011).
Section
Length (m)
Average width (m)
Average depth (m)
Area (m2)
Mean annual water temperature (°C)
A
247
30
1.2
4417
35
B
543
35
1.5
11285
30
C
252
7
0.4
6670
25
D
252
35
1.7
19344
20
E
142
16
1.3
2795
15
TOTAL
1436
44511
36
ACTA BOT. CROAT. 79 (1), 2020
PISTIA STRATIOTES ALTERING CONDITIONS IN SAVA OXBOW LAKE
Fig. 1. Study area placed in European and Slovenian perspective (upper sections), the oxbow lake is marked black (lower right section).
The Sava River is colored in grey. The Sava oxbow lake is divided into five sections A to E corresponding to a water temperature gradient
as defined by Jaklič and Vrezec 2011.
Sampling and analyses
In 2009 and 2010 a detailed study of macrophytes has
started. Sampling was performed seasonally in July and October 2009, and in January and May 2010. In each section
a census of macrophyte flora (emergent, floating and submerged) was done. Five biomass sub-samples were taken using a quadrate (1 m2). All macrophytes were identified according to Martinčič et al. (2007), counted and weighted.
Each subsample of each macrophyte species was separately
washed with distilled water and dried at 60 °С up to the constant weight. The density of macrophytes was expressed as
number of plants per m2, and fresh and dry biomass were
expressed in kg per m2.
As during the summer the water surface of the colonized
sections is completely covered with water lettuce, the available aerial photos from the past were used to check when the
lettuce population exploded.
Sub-samples for water quality were collected in four periods. In the middle of each section 1 L of water was sampled
along the depth gradient. Anions (Cl–, NO2–, NO3–, SO42–,
PO43–) and cations (Na+, K+, Ca+, Mg+, NH4+) were measured
by ion chromatography (Metrohm, 761 Compact IC). Abiotic parameters water temperature (°C), oxygen saturation
(%), concentration of dissolved oxygen (mg L–1) and pH were
measured in the middle of each section with a Multi 340i
measuring instrument (WTW GmbH, Weilheim). Transmitted daylight in the depth between surface and 1 m depth
was measured only once in July in section E and B with a
digital light meter (LI 1000; LI-COR; USA; range 0 – 1000
lux) and expressed as percentage of the daylight (%) measured just above the water.
ACTA BOT. CROAT. 79 (1), 2020
In March 2010 dried plants of water lettuce on the banks
of the oxbow lake in section D were collected (Fig. 2a, b) and
from them 25 seeds were taken and set for germination in
regular tap water in growth chambers with 20 °C and 16/8
h day/night photoperiod and 60% of air humidity (Fig. 2c).
Fig. 2. The dried plant of water lettuce collected from the bank
of the Sava oxbow lake in March 2010 (a), the seeds attached in
groups on the bottom side of the oldest leaves (b), typical barrel shaped seed (c), germinating seed after one month in growth
chambers (d).
37
JAKLIČ M, KOREN Š, JOGAN N
Statistical analysis
Parametric tests (analysis of variance, ANOVA with
post-hoc test) were used when the data met assumptions for
such tests (i.e. normality/ homoscedasticity). In cases when
both raw and transformed data violated these assumptions,
nonparametric tests were used instead.
Post-hoc tests, conducted after one-way ANOVAs,
showed significant effects of experimental temperature within each group (Species or Sections). In the case of equal variances, the Tukey HSD test was used and if Levene's test of
equality of variances indicated the variances were not equal,
the Dunnett T3 test was used. Tukey’s post-hoc test was used
to determine if significant differences occurred among sections from the same sampling sites at each sampling month
(season), which groups differed significantly (P < 0.05) in
their numbers and biomass.
After comparison of all biometrical measurements, we
tested differences within sections A to D (colonized with water lettuce) and a non-colonized section (section E) in biometrical measurements. Student t-test was used to test the
differences between colonized and non-colonized sections
of the oxbow lake.
All measured variables were divided into two groups:
biometrical variables (number of species, freshwater biomass, dry biomass and number of plants) and water quality variables (physical and chemical). The variations among
sampling sections was explored with forward stepwise discriminant analysis using the module in STATISTICA v. 12.0
software (STATISTICA 1995) with significance level for the
analyses set at P = 0.05.
Correlations between biometrical measurements of water lettuce and abiotic parameters were given, calculated
with Spearman correlations. Statistical analyses were performed using the statistical program SPSS 20.0 (SPSS Inc.
Chicago, IL, USA) with a level of significance of P < 0.05.
Results
Tab. 2. List of macrophytes in the Sava oxbow lake before (1984,
1992, 1997) and after (2009, 2010) the expansion of water lettuce
(+ = present; – = not present). Red data list (Anonymous, 2002)
status: V – vulnerable taxon, E – endangered taxon.
Macrophytes
Berula erecta (Huds.) Coville
Ceratophyllum demersum L.
Lemna gibba L.
Lemna minor L.
Myriophyllum spicatum L.
Najas marina L.
Pistia stratiotes L.
Potamogeton crispus L.
Potamogeton natans L.
Potamogeton pectinatus L.
Potamogeton trichoides
Cham. et Schltdl.
Trapa natans L.
Red data
Native Before After
list
.
Yes
+
V
Yes
+
+
V
Yes
+
.
Yes
+
+
V
Yes
+
V
Yes
+
.
No
+
.
Yes
+
.
Yes
+
.
Yes
+
E
Yes
+
-
V
Yes
+
+
the years before and after the introduction of P. stratiotes is
presented in Fig. 3, confirming that the lettuce population
exploded between 1995 and 2003, which corresponds to the
field records.
In this study, almost no indigenous macrophytes were
found in colonized sections. Indigenous macrophytes were
observed only in section E, the coldest part of the oxbow lake
(only a small population with negligible biomass of Lemna
gibba in section D (Tab. 2). Lemna gibba was discovered in
the oxbow lake for the first time in 2010 but there is a strong
possibility that it could have been overlooked previously due
to its similarity with L. minor, which had already been reported (Šajna et al. 2007). So today instead of 10 macrophyte
species (five of them at the Red data list) there are only five
species (three on the Red data list), the most abundant of
them water lettuce, which has obviously outcompeted several native taxa and caused local extinction.
Changes in floristic structure
Differences in quantitative biotic parameters among
sections
In the oldest floristic report of the discussed area (Seliškar
1984) four floating or submerged macrophytes were recorded: Ceratophyllum demersum, Potamogeton natans, P. pectinatus and Lemna minor but the results were incomplete
because the oxbow lake was not the main focus of the floristic mapping of the area. In 1984 (Seliškar 1984), 1992 and
1997 (unpbl. Jogan), no alien macrophytes were found, only
indigenous freshwater plants were widespread throughout
the whole oxbow lake (see Tab. 2). Out of 10 previously reported macrophytes, four were listed in the Red data list as
vulnerable and one as endangered (Anonymous 2002). The
first non-indigenous macrophyte, P. stratiotes was recorded
in the oxbow lake in 2001 (Hudoklin 2002, Haler 2005). In
2005, four native macrophytes (C. demersum, Myriophyllum
spicatum, Najas marina and Trapa natans) were still present in the oxbow lake (Šajna et al. 2007) (Tab. 2). A comparison of available aerial photos of colonized sections in
Number of plants and their biomass differed statistically among sections (F4.42 = 4.1, P = 0.006) and species (F4.42
= 93.8, P < 0.001). Post-hoc tests conducted after one-way
ANOVAs showed that plants were the most abundant in the
section E (Dunnett T3 post-hoc test, P = 0.003). The highest number of individual plants was counted for species L.
gibba and the lowest for T. natans, both in section E (Dunnett T3 post-hoc test, P < 0.003). Freshwater biomass differed among sections (F4.42 = 8.3, P < 0.001) and species (F4.42
= 7.9, P < 0.001). Statistically the highest freshwater and dry
biomass were measured in the sections C and D (Dunnett
T3 post-hoc test, P = 0.003, P = 0.04) as a result of only one
species, P. stratiotes (Dunnett T3 post-hoc test, P = 0.008).
Dry biomass differed significantly among sections (F4.42 =
5.1, P = 0.001), with the highest mass in section D (Dunnett
T3 post-hoc test, P = 0.001), as a consequence of the P. stratiotes abundance.
38
ACTA BOT. CROAT. 79 (1), 2020
PISTIA STRATIOTES ALTERING CONDITIONS IN SAVA OXBOW LAKE
Fig. 3. Section D of the Sava oxbow lake before water lettuce was introduced (A, year 1994) and complete coverage after colonization
(B, year 2003 and C, year 2013).
The number of individual plants did not differ among seasons (F3.43 = 1.2, P > 0.05), while differences were observed in
biomass (fresh biomass: F3.43 = 7.4, P < 0.001, dry biomass:
F3.43 = 4.9, P = 0.005). Biomass was higher in summer due to
the abundance of water lettuce and the lowest in the winter
months (Tukey HSD post-hoc test, F3.43 = 9.2, P < 0.001) (Fig. 4.
Mean freshwater biomass of all sampled macrophytes among
seasons and sections. Sections A to E are shown in Fig. 1.).
Within colonized sections, because the number of plant
individuals (Kruskal-Wallis tests, P > 0.05), and the biomass
(fresh and dry: Kruskal-Wallis test, P > 0.05) did not statistically differ, the data were pooled in further analyses. We
did not confirm differences in number of plant individuals
comparing the colonized and non-colonized sections (F1.40
= 0.02, P > 0.05), while the sections differed in biomasses
(F1.40 = 86.5, P < 0.001 for fresh biomass and F1.40 = 8.3, P =
0.003 for dry biomass). The average fresh biomass of water
lettuce in colonized sections was four times higher than the
average fresh biomass of native macrophytes in section E
(8.6 kg·m−2 and 0.63 kg·m−2). In section E, of the native macrophytes, C. demersum had the highest fresh biomass (1.5
kg·m−2) and L. gibba the lowest (0.01 kg·m−2 in the peak of
high growth season).
Differences in abiotic parameters among sections
According to discriminant function analysis the Mahalanobis distances between group centroids differed significantly between sections (Wilks' lambda = 0.019, F4.19 = 2.51,
P < 0.05). Only the first two functions were statistically sigACTA BOT. CROAT. 79 (1), 2020
Fig. 4. Mean freshwater biomass of all sampled macrophytes
among seasons and sections. Sections A to E are shown in Fig. 1.
nificant (P < 0.001), and only they were included in further
analysis. The first discriminant function explained 65% of
variation, showing the separation as a result of water temperature and the second discriminant function explained additional 29% of the variation with NO3– (Fig. 5). Both variables having significant discriminating effects (Partial Wilks’
lambda = 0.343, F4.19 = 4.8, P < 0.05 and Partial Wilks’ lambda
= 0.41, F4.10 = 3.5, P < 0.05).
Water temperature was significantly higher in the colonized section (annual average T = 27.6 ± 6.0 °C) than in the
39
JAKLIČ M, KOREN Š, JOGAN N
Seeds
We observed that larger plants of water lettuce died in
winter, when water and air temperature fell. With gradual
decay of dead plants, seeds were released and they accumulated in the sludge. In axils of oldest leaves where inflorescences developed during summer seeds were concentrated
almost without remnants of other inflorescence parts that
have already been completely decomposed (Fig. 2b). We
found that approximately 40% of seeds germinated in regular tap water in growth chambers (Fig. 2d). In the spring in
the oxbow lake we also noticed young floating seedlings derived from germinated seeds.
Fig. 5. Discrimination of the colonized (A-D) and non-colonized
section (E) in the oxbow lake by the first two discriminant functions. Input data matrix consisted of all measured abiotic parameters, projections of primary variables shown as bi-plot. Sections
A to E are shown in Fig. 1.
non-colonized section (annual average T = 15.8 ± 5.3 °C)
(F1.28 = 19.2, P < 0.001), with the highest water temperature
in section A (36.1 ± 2.5 °C). The colonized section had significantly lower oxygen saturation (average annual 41.3 ±
22.4% vs. 67.5 ± 14.5%) and concentration of dissolved oxygen (average annual 4.1 ± 1.2 mg L–1 vs. 6.0 ± 2.7 mg L–1)
than the non-colonized section (F1.28 = 43.1, P < 0.001 and
F1.28 = 11.9, P = 0.002). Concentration of dissolved oxygen
was on average below 5 mg L–1 in sections A-D and 7.8 ± 4.2
mg L–1 in section E. Variables pH, concentration of cations,
and concentration of two anions (Cl– and PO43–) were not
significantly different in the colonized and the non-colonized section. Higher concentration of NO2– and NO3– were
measured in the colonized section (especially in section C
and D), than in the non-colonized section (F1.28 = 2.9, P =
0.040 for NO2–, F1.28 = 6.3, P = 0.011 for NO3–). Concentration of SO42– was on average significantly higher in the colonized section (with the maximum in section A-C) than in
the non-colonized section (F1.28 = 10.6, P = 0.003).
Water temperature was correlated significantly with
number of water lettuce plants (rs = 0.583, P < 0.05), fresh
(rs = 0.550, P < 0.05) and dry biomass (rs = 0.532, P < 0.05),
while no correlation was found between ion concentrations
and number of plants, fresh or dry biomass of water lettuce (P > 0.05).
The non-colonized section statistically differed from
the colonized sections (Mann-Whitney U test, P < 0.001)
in water column light conditions (percentage of daylight
at a certain depth; Fig. 6a). The non-colonized section had
more than 50% higher percentage of light than the colonised (Fig. 2). In colonized sections the percentage of daylight was nearly 0% at 10 cm depth and deeper. Oxygen
saturation was not different down to the 50 cm water depth
(Fig. 6b), while a clearly lower saturation was measured in
the colonized section deeper than 50 cm (Fig. 2) obviously
due to the decay of biomass.
40
Fig. 6. Percentage of daylight (a) and oxygen saturation (b) in the
water column in a colonized (section D) and the non-colonized
section (section E) of the Sava oxbow lake in 2010.
Discussion
Prilipe oxbow lake in Slovenia is one of the few places in
continental Europe where overwintering of water lettuce has
been reported for almost two decades. Out of 10 previously
recorded macrophytes only a third have survived colonization by water lettuce but at the same time their populations
are reduced and limited to the 6% of the oxbow lake that has
not been colonized (section E, area of 0.28 ha). Haler (2005)
has already concluded that water lettuce outcompeted native
macrophytes in the lake and we can only reconfirm that in
2010. The presence of four previously recorded native macrophytes (Seliškar 1984, Šajna et. al. 2007), were confirmed
in our study. We also observed Lemna gibba, for the first
time, but populations of Berula erecta, Myriophyllum spicatum, Najas marina, Potamogeton crispus, P. natans, P. pectinatus and P. trichoides had became locally extinct. So obviously invasion by water lettuce has caused a degradation of
the local macrophyte community and damaged biodiversity,
particularly so because three of the mentioned native macrophytes are listed in the Red data list (Anonymous 2002).
Macrophyte beds have generally high primary production
and can thus be an important source of organic matter, supporting higher trophic levels and influencing the net metabolism of aquatic systems. As has been stated by Caraco and Cole
(2002), dense canopies of floating macrophytes can prohibit
ACTA BOT. CROAT. 79 (1), 2020
PISTIA STRATIOTES ALTERING CONDITIONS IN SAVA OXBOW LAKE
gas exchange and make oxygen depletion more severe, some
floating macrophytes can release much of their photosynthetically produced oxygen directly into the atmosphere rather
than into the water column. Exactly this can explain the situation with water lettuce in the oxbow lake, where after colonization the ecological conditions were radically changed, which
puts this species among invasive alien transformers (Pyšek
et al. 2002). During the vegetation period, when the studied
water lettuce population expanded exponentially, concentration of dissolved oxygen (DO) in the water decreased to less
than 5 mg L–1. That can have a negative impact upon sensitive
species of fish and invertebrates and can also impact nitrification/denitrification cycles (Urbanič and Toman 2003). Average concentration of DO was lowest in section C, where the
oxbow lake is the most shallow.
Comparison of oxygen saturation in the covered part and
the uncovered part of the oxbow lake showed that below 0.5
m depth low oxygen quantity was measured due to organic matter decay on the bottom and scarce light conditions.
Since the light conditions were completely changed due to
water lettuce, no other macrophytes were able to thrive below 10 cm of water depth.
In Europe, sites such as thermal water bodies and reservoirs for hydroelectric power stations have a high potential
for water lettuce naturalization. Nevertheless, in the last years
a few ephemeral occurrences of water lettuce were observed
about 20 km down the Sava River, in the Sava-Strmec Special
Reserve (Boršić and Rubinić 2018). But in the direct vicinity
of the oxbow lake (about 11 km towards NW), there is a warm
section of Sava River due to outflow from a nuclear power station. Further on, in the near future, close to the oxbow lake a
hydropower plant is planned on the Sava River, which might
provide a new potential habitat for water lettuce. In the warmer conditions, at least during the summer months, water lettuce can become a real pest of accumulation reservoir.
We may also assume that predicted climate change causing general warmer conditions might create new problems:
the species may extend its habitat range and in such case the
oxbow lake can serve not only as a stepping-stone to rivers
and other oxbow lakes of Central Europe, but also the source
of a water lettuce population partly adapted to the European
climatic conditions.
From our results we can conclude, that even in temperate
regions, in thermal water bodies, water lettuce establishment
is possible. It can outcompete all the native macrophytes,
radically change light penetrating conditions in water, its cyclic production of biomass is several fold bigger than of native macrophytes and in the following seasons decay of dead
biomass depletes oxygen and raises NH4+ concentration in
the deeper parts of the water, which can further influence
the living conditions of fauna. The oxbow lake, one of the
few remaining along the Sava River, which was once an important reservoir of local biodiversity of wetland organisms,
is today only a good example of the complete local destruction of biodiversity as a result of the irresponsible release of
a tropical aquarium plant into the nature.
Acknowledgements
We wish to thank those colleagues who have helped us
to collect plants. Many thanks also to the anonymous reviewers whose suggestions and corrections substantially improved the text.
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