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Key environmental variables affecting the distribution
of Elatine hungarica in the Pannonian Basin
Faktory prostředí ovlivňující rozšíření Elatine hungarica v Panonské pánvi
Attila T a k á c s1, András S ch m o t z e r2, Gusztáv J a k ab3, Tamás D e l i4,
Attila M e s t e r h á z y5, Gergely K i r á l y6, Balázs András L u k á c s7,
Boglárka B a l á z s8, Ranko P e r i ć9, Pavol E l i á š jun.10, Gábor S r a m k ó11,
Jácint T ö kö l y i12 & Attila M o l n á r V.1
1Department of Botany, Faculty of Sciences & Technology, University of Debrecen, H-4032
Debrecen, Egyetem tér 1., Hungary, email: mva@science.unideb.hu; 2Bükk National Park
Directorate, H-3304 Eger, Sánc u. 6., Hungary; 3Department of Environmental Sciences,
Faculty of Water and Environmental Management, Szent István University, Szabadság u.
1–3., H-5540 Szarvas, Hungary; 4Directorate of Békés County Museums, H-5600
Békéscsaba, Széchenyi u. 7., Hungary; 5Hunyadi u. 55., H-9500 Celldömölk, Hungary;
6West Hungarian University, Faculty of Forestry, Institute of Sylviculture and Forest
Protection H-9400 Sopron, Ady E. u. 5, Hungary; 7Department of Tisza Research, MTA
Centre for Ecological Research-BLI-HAS), Bem tér. 18/C, H-4026, Debrecen, Hungary;
8Physical Geography and Geoinformatics, University of Debrecen, H-4032 Debrecen,
Egyetem tér 1., Hungary; 9Institute for Nature Conservation of Vojvodina province, Radnička
20a, 21000 Novi Sad, Serbia; 10Department of Botany, Slovak University of Agriculture,
Tr. A. Hlinku 2, SK-949 76 Nitra, Slovak Republic; 11MTA-ELTE-MTM Ecology Research
Group, H-1117 Budapest, Pázmány Péter sétány 1/C, Hungary; 12MTA-DE “Lendület”
Behavioural Ecology Research Group, University of Debrecen, H-4032 Debrecen,
Egyetem tér 1, Hungary
Takács A., Schmotzer A., Jakab G., Deli T., Mesterházy A., Király G., Lukács B. A., Balázs B.,
Perić R., Eliáš P. jun., Sramkó G., Tökölyi J. & Molnár V. A. (2013): Key environmental variables
affecting the distribution of Elatine hungarica in the Pannonian Basin. – Preslia 85: 193–207.
Elatine hungarica Moesz is a small wetland ephemerophyte that occurs and is classifiedas extinct,
data deficient or a very rare and endangered taxon in most countries in eastern and central Europe.
Based on literature and herbarium data, supplemented by 160 field records collected between 1998
and 2011, we present the currently known distributionof this species in the Pannonian Basin, which
mostly but not exclusively includes Hungary. Within the Pannonian Basin this species is distributed
throughout Hungary, with sporadic occurrences in Romania, Serbia and Slovakia. The temporal dis-
tribution of floristic records is very uneven. This species was recorded only in 27 years during the
last 213 years (1798–2011). When examining presence/absence data for most of the 20th century,
we found a significant correlation between the number of records of this species in a given year and
two key, but not independent, environmentalvariables: rainfall and the extent of the area inundated
in the same year. In the more intensively documented period between 1998 and 2010, there is only
a significant correlation between the numbers of records of this species and the extent of flooding,
which is because there is a delay in the effect of an increase in rainfall. The peak occurrence of
records in the 1940s and 1950s is associated with extensiverice productionin Hungary. Today, most
records are for agricultural fields that are subject to flooding and becoming temporary wetlands.
The comparison of recent and past distributions of E. hungarica reveals a consistent and marked
regional difference; whereas this species is not rare along the Tisza river and its tributaries, it is
markedly scattered in similar habitats near the Danube.
K e y w o r d s: ephemerophyte, flooding, Hungary, inland water, Isoëto-Nanojuncetea,Elatina-
ceae, mapping, rainfall, rice weeds, Romania, Serbia, Slovakia, temporary ponds
Preslia 85: 193–207, 2013 193
Introduction
All members of the family Elatinaceae are poorly known, especially in terms of their tax-
onomy and phytogeography (Popiela & Łysko 2010, 2011, Popiela et al. 2011, 2012).
Elatine hungarica Moesz is probably one of the most data deficient vascular plant species
in Europe, which is indicated by the low number of localities documented by herbarium
specimens. A further telling fact is that, to our knowledge, no photographs of this species
were published before that in Molnár V. et al. (1998). This species was described by
Gusztáv Moesz (1908) and the lectotype for this name has recently been designated
(Molnár V. et al. 2013). The only illustration of E. hungarica by Moesz (1908) appeared
later as a redrawn figure in numerous other works (e.g. Hegi 1927, Jávorka & Csapody
1929–1934, Ţopa 1955, Felföldy 1990, Dihoru & Negrean 2009).
Elatine hungarica is an ephemerophyte growing on wet mud or on the bottom in shal-
low water. The individuals are minute, reaching only a few centimeters in diameter, but
under favourable conditions dense colonies may arise that are up to 60–70 cm in diameter.
Shoots are branched, rooting at nodes and creeping on the mud. The most remarkable dif-
ferentiating characters of this species are seed curvature and testa reticulation (Molnár V.
et al. 2013). The measurements reported indicate that the seeds of the lectotype of
E. hungarica are much more curved (n = 50, mean ± SD = 188 ± 22°, range: 144–236°)
than those of E. orthosperma (n = 30, mean ± SD = 40 ± 20°, range: 0–76°), but somewhat
less curved than those of E. hydropiper (n = 40, mean ± SD = 230 ± 20°, range: 171–276°).
The seed testa of E. hungarica is characterized by regular hexagons. In this respect
E. orthosperma and E. hydropiper are very different from E. hungarica because the reticu-
lation on their seed is characteristically covered by long-hexagons.
Elatine hungarica is reported occurring in central Europe in Hungary (Moesz 1908),
Romania (Moesz 1908, Oprea 2005), Serbia (Jávorka 1924–1925) and Slovakia
(Margittai 1927, 1939, Ťavoda & Goliašová 2008). Outside central Europe this species is
known from Moldova (Gejdeman 1986), Ukraine (Prokudin 1987, Kricsfalusy et al. 1999,
Mosyakin & Fedoronchuk 1999, Shapoval 2006) and Russia, where it grows from the
coastal area of the Black Sea to western Siberia (Gorshkova 1949). In Portugal, it is
recorded as an introduced (naturalized) species (Walters 1968, Uotila 2009), although data
in Flora Europaea are highly unreliable for evaluating the invasive status of species at the
country level (Pyšek 2003). In most of the countries where E. hungarica occurs it is listed
in national Red Lists or in Red Data Books. The threat status varies between the countries
and includes e.g. “critically endangered” (Romania; Dihoru & Negrean 2009) or “vulnera-
ble” (in Transcarpathian Ukraine; Kricsfalusy et al. 1999), though in some cases Red List
data do not reflect the real status of this species (e.g. in Slovakia, see Holub et al. 1999). In
the IUCN Red List E. hungarica is categorized as a Data Deficient (DD) taxon (Bilz et al.
2011). The threat status and exact distribution of E. hungarica are insufficiently known for
the eastern part of its distribution; although the species is listed in the Red Data Book of
Ukraine (Mosyakin 2009) the published photograph of the species shows E. triandra
Schkuhr.
Elatine hungarica grows only in periodically flooded habitats (i.e. temporarily inun-
dated arable fields, rice paddy fields and natural marshes). In Hungary, this species,
together with Alisma lanceolatum,Alopecurus aequalis,Echinochloa crus-galli,Elatine
alsinastrum,Lindernia procumbens,Peplis portula,Ranunculus sardous,Schoenoplectus
194 Preslia 85: 193–207, 2013
supinus and Typha latifolia are classified as diagnostic species of waterlogged arable
fields with characteristic Isoëto-Nanojuncetea vegetation (Lukács et al. 2013). The spe-
cific habitat of E. hungarica (3130: Oligotrophic to mesotrophic standing waters with veg-
etation of Littorelletea uniflorae and/or of Isoëto-Nanojuncetea) is considered to be of
community interest in the European Union (Annex I of Habitats Directive), which implies
the designation of special areas for the conservation of this species (Council of the Euro-
pean Communities 1992). Hence, investigations on the distribution and ecology of
E. hungarica, identification of threatening factors and revision of the IUCN threat status
are desirable.
This paper has four main aims: (i) to summarize the data on its distribution scattered in
the literature and herbaria in the Pannonian Biogeographic Region; (ii) to present data col-
lected in the field between 1998–2011, which is a significant contribution to the present
distribution of this species; (iii) to examine environmental factors influencing its erratic
appearance; and (iv) to evaluate threats to this species.
Materials and methods
Study area
The study area is located in the Pannonian Basin, which corresponds to the Pannonian
Biogeographic Region of the EU (EEA 2008) (Fig. 1). Basically the region extends over
eight countries (Austria, Czech Republic, Croatia, Hungary, Slovakia, Serbia, Ukraine and
Romania), with the overwhelming majority of the region in Hungary. This country is in the
centre of the basin and has the largest area of lowland in central Europe. Located in the
temperate zone it is at the border of three climate zones and experiences a very variable cli-
mate (Mezősi 2011). The annual average temperature is ~10°C, but partly because it is in
a basin the amplitude of the temperature fluctuation is high (24.5°C in the central part).
Annual precipitation decreases from 800 mm in the west to 500 mm in the east, depending
on oceanic, orographical and Mediterranean influences (Péczely 1979). The area is rich in
surface and ground water and the main rivers are the Danube and Tisza. Before the rivers
were regulated in the 19th century the area was mostly flooded and swampy (Ihrig 1973).
Nowadays, river regulations prevent large-scale flooding but a significant part of Hungary
is still subject to inland water inundation caused mainly by a significant increase in ground
water level. Two-thirds of the region is lowland (< 200 m), with most of the remaining area
at mid-altitudes (200–500 m) and only a small proportion in the submontane region (> 500
m). This lowland area functions as a real basin where the incoming large rivers (Danube,
Tisza and its tributaries) create large, alluvial river valleys with restricted active flood-
plains due to extensive river regulations.
Distribution of the species
The distribution of Elatine hungarica presented here is based on recent (1998–2011) field
observations (n = 160), critically evaluated herbarium data (n = 30, from herbaria BP, DE,
CL – herbarium acronyms according to Holmgren & Holmgren 1998) and published
floristic records (n = 79) for 21 of which there were voucher specimens.
Takács et al.: Elatine hungarica in the Pannonian Basin 195
The taxonomic identity of herbarium specimens and data collected by the authors were
verified based on seed morphology. Only records that could be located unequivocally
were used to prepare the distribution map. The complete list of localities is given in Elec-
tronic Appendix 1.
Distribution data are displayed on a map using the central-European floristic mapping
system (Niklfeld 1971). The distribution map summarizing all records for this species in
the region is presented in combination with the map (Hydrographic Institute 1938) of
waters and wetlands in the Pannonian Basin before the large-scale regulation of the rivers
(i.e. before 1860s) to visualize the possible coincidence between the extent of historical
wetlands and the species’ distribution. The nomenclature of vascular plants follows Király
(2009).
Environmental data
To evaluate the possible effects of environmental factors on the occurrence of E. hungarica
we used rainfall data (including snowfall) measured between 1951 and 2010 by the Lower
Tisza Regional Water and Environmental Directorate, Hungary (Pálfai 2011). The maxi-
mum extension of the area inundated each year (between 1936 and 2010) was obtained
from hydrological sources (Szlávik 2003, Hungarian Hydrological and Environmental
Central Directorate 2011). As the increase in the area used for producing rice in Hungary
might have influenced the distribution of this species, data on rice production obtained
from Hajdú (2006) was used to indicate possible effects.
196 Preslia 85: 193–207, 2013
Fig. 1. – The location of the study area.
Statistical analysis
Long-term inundation and rainfall data available for Hungary, which are representative for
the entire region, were used to statistically evaluate the role of these factors in influencing
the occurrence of E. hungarica. Since there were many years with no records (i.e. this
variable was strongly zero-inflated), we coded each year based on whether E. hungarica
was recorded or not. This variable was used to test the null hypothesis (by means of
ANOVA) that rainfall and inundation do not differ between years with or without records
of E. hungarica. The variable “inundation” was log-transformed to improve normality.
Since there were fewer years with zero records after 1997 (5 out of 13), we also tested
whether there is a correlation between the number of records, and inundation and rainfall,
in the period 1998–2010. This analysis allowed us to evaluate the effect of these two envi-
ronmental variables on the population size of E. hungarica (not only its presence/absence,
as in the first analysis). Since the number of records per year was not normally distributed
Spearman rank correlation was employed. Analyses were done in the R Statistical Envi-
ronment (R Development Core Team 2010).
Results
Altogether, 269 floristic records for 105 mapping grid units from Hungary, Romania, Ser-
bia and Slovakia were collected. The earliest record was for 1798, the latest 2011. The
recent and historical distribution of E. hungarica is shown in Figs 2 and 3 (see Electronic
Appendix 1 for source data). The majority of all the records (86%) of E. hungarica are for
Hungary, but within the Pannonian Basin the species is more widespread occurring in
Romania, Slovakia and Serbia, at least in the areas bordering on Hungary (Fig. 3).
The distribution of E. hungarica records over time in the region is very uneven (Figs 4,
5). Over a period of 213 years between 1798 and 2011 E. hungarica was recorded only in
27 years. Between the 1790s and 1930s the species was recorded only in 10 years in 13
grid units. Between 1940 and 1960 it was collected in 13 years, mostly in rice paddy fields.
In Hungary, disregarding the artificially flooded rice paddy fields, the plant was found
only in 18 years. Between 1951 and 2010 E. hungarica was recorded only in 9 years,
mostly after 1998 (Fig. 3).
Annual rainfall and the maximum extent of the area inundated were significantly
higher in years when E. hungarica was recorded (Table 1). The annual number of records
between 1998 and 2010 correlated with the maximum extent of yearly inundated area
(Spearman rank correlation, n = 13, ρ= 0.82; P < 0.001), but not with annual rainfall (ρ=
0.41; P = 0.17).
The distribution of this species shows a notable geographic disparity in the Pannonian
Basin (Fig. 3). Although most of the basin is a large floodplain formed by the rivers Dan-
ube and Tisza, the majority of the records of E. hungarica are for the eastern part of the
area, in the region of Tisza and its (sub)tributaries (Bodrog, Berettyó, Zagyva, Körös; Figs
2, 3). Although suitable habitats and typical accompanying species, such as Schoeno-
plectus supinus, Lindernia procumbes, Elatine alsinastrum and Eleocharis acicularis, are
relatively common along the Danube river E. hungarica is a rather rare species there.
Takács et al.: Elatine hungarica in the Pannonian Basin 197
198 Preslia 85: 193–207, 2013
Fig. 2. –Records of Elatine hungarica in the Pannonian Basin before 1989. In the case of multiple records in the
same grid the symbol displayed is for the latest record. Note that between 1900 and 1959 in Hungary, the species
was recorded mostly in rice paddy fields.
Fig. 3. –Complete distribution of Elatine hungarica based on records from several periods, overlaid with the pat-
tern of inundation. Areas covered permanently or temporarily with water in the Pannonian Basin before the rivers
were regulated are distinguished (according to the Hungarian Hydrographic Institute 1938). In the case of multi-
ple records in the same grid the symbol displayed is for the latest record.
Takács et al.: Elatine hungarica in the Pannonian Basin 199
Fig. 4. – The number of records of Elatine hungarica in the Pannonian Basin recorded in each decade.
Fig. 5. – The annual number of records of Elatine hungarica recorded in the Pannonian Basin between 1998 and
2010, related to annual rainfall and maximum area inundated in Hungary. Note that records from rice fields are
Table 1. – Comparisonof the annual rainfall (1951–2010) and the maximumarea inundated yearly (1936–2010)
in Hungary between years with and without records of Elatine hungarica in the Pannonian Basin.
Records of
Elatine hungarica
n Mean Median SE ANOVA
Annual rainfall (mm/year) Yes 9 638 665 46 F1,58=8.0;
P = 0.006No 51 536 522 13
Maximum area inundated each year
(1000 hectares)
Yes 13 231 186 50 F1,74 =14.611;
P < 0.001No 63 78 42 12
The historical longevity of this geographic disparity can be proven by the historical
data on this species’ occurrence. Elatine hungarica was recorded long before the regula-
tion of the Tisza river in the1860s at four and three localities by two botanists, Pál Kitaibel
and Anton Kerner, respectively. All these localities are on the Tisza floodplain. Nonethe-
less, other Isoëto-Nanojuncetea-species (e.g. Cyperus fuscus,Dichostylis micheliana,
E. alsinastrum,Eleocharis acicularis,Gnaphalium uliginosum,Juncus bufonius,Limosella
aquatica,Lythrum hyssopifolia,Marsilea quadrifolia,Peplis portula) were also reported
by these authors from 12 and six localities, respectively, including the Danube floodplain
(Fig. 6).
Discussion
Our paper describes the geographic distribution of the poorly known Red Listed species
Elatine hungarica in the Pannonian Biogeographic region. We present its current and his-
torical distribution based on the literature, herbarium data and 160 field records reported
here, which significantly enhance our knowledge of this species distribution. The erratic
distribution of E. hungarica may be influenced by several factors. Danihelka et al. (2009)
suggest that the number of botanists specialized on a certain group of plants has an in-
disputable role in determining the extent of the knowledge of a species distribution and
200 Preslia 85: 193–207, 2013
Fig. 6. – Records of Elatine hungarica (full symbols) and other Isoëto-Nanojuncetea species (empty symbols)
recorded by Pál Kitaibel (circles) and Anton Kerner (squares) before the beginning of the regulation of the Tisza
river in the 1860s.
ecology. This might be especially pronounced in the case of aquatic plants, for which the
number of experts is relatively low (Kaplan 2010). Nevertheless, we think that the differ-
ences in the intensity of floristic research cannot be the only reason for the conspicuous
temporal unevenness we have documented in this study (Figs 4, 5) as climatic conditions
in the region, such as highly fluctuating annual rainfall and annual extent of the area inun-
dated, also play an important role in determining the occurrence of E. hungarica.
Our results indicate that the occurrence of E. hungarica significantly increased with
increase in annual rainfall and extent of the area inundated. These two key environmental
variables are not independent as the former factor evidently influences the latter and due to
the basin’s geographic characteristics large-scale inundations by direct flooding and lift in
the level of ground water are usually associated with increased rainfall. Therefore, the indi-
rect effect of annual rainfall may be crucial. However, in order to understand why we failed
to find a correlation between annual rainfall and records of this species in 1998–2010 we
need to understand the inundation dynamics of the region. Within this period the rainfall in
the wettest year in Hungary (2010: 876 mm) was more than twice that in the driest year
(2002: 388 mm). E. hungarica was recorded in relatively wet years in which the average
rainfall was 141 mm greater than in years when it was not recorded. But, rainfall also has
a delayed effect on the extent of the area inundated that significantly contributes to the
appearance of E. hungarica. This can be exemplified by the period 1998–2002 (Fig. 5), in
which the annual rainfall in the firsttwo years, was relatively high (~700 mm) and caused an
expansion in the area temporarily inundated to about350,000–450,000 hectares, which pro-
vided suitable habitats for mudflat species, hence the number of records of E. hungarica
suddenly increased in 1999 and 2000. In the third year (2000) rainfall decreased to only 411
mm, which implied there should have been a decrease in the occurrence of E. hungarica in
2001. In 2000, annual rainfall was far below the average, but the number of records was still
high because conditions for this plant continued to remain favourable. In contrast, despite
the significant rainfall (623 mm) in 2001 there was no extensive flooding causing an absence
of records for this plant. In summary, we conclude that the lack of a significant correlation
between annual rainfall and the occurrence of this species is caused by the delayed effect of
rainfall on the extent of inundation.
An additional limiting factor that is supposed to affect the distribution and occurrence
of E. hungarica is change in land-use. The species was often recorded in rice fields, which
provided an adequate depth of water for a sufficientlength of time for it to complete its life
cycle. The Pannonian Basin is situated at the northern limit of rice production. From the
16th century onwards sporadic attempts at rice cultivation were made in different coun-
tries. The practice of cultivation was established and the main production areas in Hungary
determined before World War II. Rice production had a strong political priority in the
1950s and the area under rice cultivation rapidly increased (from ~ 20,000 hectares in
1949 to 87,500 hectares in 1955; Hajdú 2006). Fortunately, the weed flora of rice fields
was thoroughly investigated during this period (Soó 1948, Ubrizsy 1948, 1961, Csapody
1953) and E. hungarica was recorded at numerous locations. Rice was also cultivated in
Serbia (Vojvodina) and Slovakia in the past (Hejný 1960, Tošić 1970) but to a much less
extent than in Hungary. Although the presence of rice paddy fields had a similar effect on
the flora of Slovakia, e.g. the increase in the number of localities of Beckmannia
eruciformis was related to experimental rice cultivation (Dítě et al. 2011), E. hungarica
was not recorded in rice paddy fields in Slovakia or Serbia. The area under rice was
Takács et al.: Elatine hungarica in the Pannonian Basin 201
quickly reduced in Hungary after 1970. Nevertheless, the decrease in the number of
records of E. hungarica cannot be explained only by the decline in rice cultivation. In
Hungary there is no documented record of this species in 1960–1998 (Molnár V. et al.
1998, 1999). In this period the intensity of floristic research in Hungary, especially in the
potential habitats of E. hungarica (e.g. inundated arable fields) was probably too low.
Similarly, apparent rarity is recorded for other ephemeral wetland species in the Czech
Republic (Šumberová 2003, Šumberová et al. 2012) and Slovakia (Zlacká et al. 2006,
Eliáš et al. 2011). After 1998, as a consequence of surveys that focused on inundated
areas, unprecedented numbers of localities were recorded.
In Hungary, E. hungarica was recorded early mostly in rice paddy fields and in the last
decade most frequently in arable fields, which are, at least in certain years, subject to
inland inundation and become temporary wetlands (Lukács et al. 2013). Most of these
areas are used for agricultural crop production. The presence of this species in wet arable
fields can be explained at least in three ways: (i) the seeds are transferred from natural hab-
itats to agricultural land by waterfowl (epi- and/or endozoochory), (ii) by human activities
or (iii) the species is a relict of the former natural wetland flora, whose seeds survive in the
soil in unfavourable dry years. Transmission by waterfowl is documented for some Isoëto-
Nanojuncetea species. For instance, seeds of Carex bohemica in mud attached to the feet
of mallards (Hohensee & Frey 2001) and propagules of 21 plant species in mud attached to
the body of waterfowl (Kerner 1868, 1895). In addition, recent studies emphasize the
importance of epizoochorous (Vivian-Smith & Stiles 1994) and endozoochorous
(Charalambidou & Santamaría 2002, Green et al. 2002, Mueller & van der Valk 2002,
Charalambidou et al. 2003, Chang et al. 2005, Wongsriphuek et al. 2008, Brochet et al.
2009, 2010, Figuerola et al. 2010) transport of marsh plants and aquatic macrophytes.
Nevertheless, some ecologists (e.g. Clausen et al. 2002) are sceptical about the real impor-
tance of ornithochory in long-distance dispersal of seed. In a similar vein, the conspicuous
fact that E. hungarica has not been recorded in the floodplains of the Danube, Sava and
Drava rivers argues against effective dispersal by water birds. If small seeded plants like
Elatine have conspicuous gaps in their distributions it is much more probable that they are
habitat- rather than dispersal limited. This is also clear from the huge but usually scattered
distribution range of many wetland annuals, e.g. Tillaea aquatica,Coleanthus subtilis and
some Elatine species (Hultén & Fries 1986). However, this phenomenon may be
explained by other ecological factors (such as soil characteristics), which needs to be
addressed in future investigations.
There is evidence that human vehicles (including soles of boots) and other technical
equipment greatly contribute to the dispersal of plant propagules (e.g. Clifford 1959,
Bakker et al. 1996). Seed dispersal between isolated ponds carried on vehicles and boots is
reported for the Czech Republic for some Elatine species (Šumberová et al. 2012). It is
probable that vehicles used in farming might also disperse large numbers of propagules.
However, this mode of dispersal is limited to local or regional scales.
On the other hand, seed of the genus Elatine retain the ability to germinate for a long
period (e.g. Margittai 1939, Deil 2005), e.g. E. triandra for more than 50 years (Kasahara
et al. 1967). Relevant also is that there had been a relatively large area of permanently
inundated land in the Pannonian Basin. Until the middle of the 19th century, when the reg-
ulation of rivers started in this region, 20.5% of area had been permanently and an addi-
tional 17.8% periodically covered by water (Hydrographic Institute 1938). Nonetheless,
202 Preslia 85: 193–207, 2013
Pálfai (2003) reports a potential excess of temporary water on 60% of the plains in Hun-
gary even after the regulation of the rivers was completed.
Conservation outlook
The relative abundance of E. hungarica is connected with the availability of suitable tem-
porary wet habitats, especially in agricultural fields. Intensive agricultural cultivation (e.g.
ploughing, treading) and inundation creates hundreds of hectares of optimally wet,
exposed surfaces. However, local and regional scale drainage of inundated arable fields,
which is in the interest of farmers, causes this habitat to dry out before this species can ger-
minate. In addition, E. hungarica (and other dwarf mud dwellers) is threatened by the
abandonment of traditional extensive cultivation methods, and the general application of
advanced agrotechnology and herbicides. According to Timár & Ubrizsy (1957) 2,4-
dichlorophenoxyacetic acid and MCP (2-methyl-4-chlorophenoxyacetic acid), which are
widely used chemicals in agriculture are known to be harmful to this species.
Elatine hungarica is evaluated as missing (possibly extinct) in the Slovak Republic
(Holub et al. 1999: 414, Feráková et al. 2001, Ťavoda & Goliašová 2008). However, it was
recently recorded at two localities in Slovakia (Molnár V. et al. 1999, Király & Eliáš 2011),
hence the species must be considered as critically endangered (CR). The same happened
in Hungary, where Bagi (1998) presumed that the species was extinct, even though pro-
tected in this country since 1993 but without specified level of threat (Király 2007).
Overall, this study has a clear conservation message. Because of the general rarity of
this species, and the strong correlation of its occurrences with annual rainfall and the area
of land inundated, the best strategy for conservation is the hydrographical rehabilitation of
selected sites or creation of suitable habitats and inclusion of the species in a vulnerable
legislation category. The success of the former strategy is ensured by the ability of
E. hungarica to create a long lasting seed bank. Because of the latter we propose for
E. hungarica the category Least Concern (LC) on the global scale and Vulnerable (VU) on
a regional scale according to Bilz et al. (2011). This may result in E. hungarica being
given a preferred role in nature conservation projects.
See www.preslia.cz for Electronic Appendix 1
Acknowledgements
The authors thank János Podani for his comments and linguistic corrections, Zoltán Túri for georeferencing the
map used as the basis for Fig. 3., Judit Kapocsi, Róbert Vidéki, Norbert Pfeiffer, Gergely Gulyás, Lajos Felföldy,
Antal Széll, László Tóth, Viktor Virók for participation in the field work, László Barta (Cluj-Napoca, Romania)
for providing herbarium data and János Csiky and András Mészáros for personal communications. We are very
grateful to three anonymous reviewers for their appropriate and constructive suggestions and for their proposed
corrections that improved the earlier version of this paper. The work was supported by the TÁMOP-4.2.2/B-10/1-
2010-0024 and TÁMOP 4.2.4.A/2-11-1-2012-0001 grants, both co-financed by the European Union and Euro-
pean Social Fund. The work of G. Sramkó was funded by the EU Marie Curie Actions (EU7KP) co-funded,
NKTH and OTKA provided Mobilitas grant (no. MB08-A 80332). The work of G. Király was supported by
OTKA 67666 and the field research of P. Eliáš jun. was funded by grant VEGA no. 2/0003/11. Tony Dixon and
Petr Pyšek kindly improved our English.
Takács et al.: Elatine hungarica in the Pannonian Basin 203
Souhrn
Elatine hungarica Moesz, drobný mokřadní efemerofyt s centrem výskytu ve střední a východní Evropě, je ve
většině zemí Panonského ekoregionu považován za taxon vyhynulý (EX) nebo velmi vzácný a ohrožený (EN),
nebo je hodnocen jako druh s nedostatečnými údaji výskytu (DD). Stanoviště tohoto druhu patří mezi biotopy ev-
ropského významu (tj. jsou uvedeny v Příloze I. směrniceo stanovištích EU). V předloženém příspěvku předsta-
vujeme historické i aktuální rozšíření tohoto druhu v Panonském ekoregionu zpracované na základě dat získa-
ných studiem literatury a herbářových údajů, doplněných o 160 terénních záznamů získaných v letech 1998
a 2010. V rámci ekoregionu se druh vyskytuje roztroušeně ve velké části Maďarska, zatímco v Srbsku a na Slo-
vensku byl zaznamenán pouze ojediněle. Časové rozložení floristických záznamů je velmi nerovnoměrné. Elati-
ne hungarica byla zaznamenána v průběhu posledních 213 let (mezi lety 1798–2011) pouze v 27 případech/le-
tech. Analýzou dat jsme zjistili pozitivní korelaci mezi přítomností druhu a množstvím srážek, jakož i mezi pří-
tomností druhu a velikostí oblasti zaplavení. Vrchol jeho výskytu ve čtyřicátých a padesátých létech minulého
století souvisí s tehdejším velkoplošným pěstováním rýže v Maďarsku. Dnes je většina nálezů soustředěna v ze-
mědělských oblastech, které jsou v letech s vysokým úhrnem srážek vystaveny periodickým záplavám vytvářejí-
cím dočasné mokřady. Historické i současné rozšíření E. hungarica vykazuje rovněž nápadné regionální rozdíly:
ačkoliv druh bývá v blízkosti řeky Tisy a její přítoků nalezen dosti často, na obdobnýchlokalitách v blízkosti Du-
naje se vyskytuje velmi roztroušeně až vzácně.
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Received 19 July 2012
Revision received 12 December 2012
Accepted 10 January 2013
Takács et al.: Elatine hungarica in the Pannonian Basin 207
