HACQUETIA 7/1 • 2008, 5–20
DOI: 10.2478/v10028-00¸8-0001-8
Distribution and communities of
Crypsis aculeata and Heleochloa schoenoides
in Slovakia
Pavol Eliáš jun.*, Daniel Dítě**, Vít Grulich*** & Marek Sádovský****
Abstract
The distribution and communities of two annual grasses Crypsis aculeata and Heleochloa schoenoides were
studied in Slovakia during 2003–2007. Herbarium and field data were used to reconstruct the occurrence of
both taxa. A strong decline in C. aculeata localities was observed. This was mainly due to the destruction of
periodically flooded saline habitats. By contrast, the occurrence of H. schenoides was changed only slightly.
This species survives in secondary habitats (rural roads, field margins, field depressions). Point maps of
historical and recent distribution are presented. Halophile vegetation of Crypsis aculeata and Heleochloa schoenoides has been sampled with the adapted Braun-Blanquet method. All collected relevés have been classified
using the JUICE software. The results showed that the Crypsidetum aculeate Wenzl 1934 community is missing
now in Slovakia, but Heleochloetum schoenoidis (Soó 1933) Ţopa 1939 was still recorded at a few sites. However,
the community contained many ruderal plant species.
Key words: Crypsis aculeata, Heleochloa schoenoides, halophytes, saline soils, distribution, vegetation, Slovakia
Izvleček
Raziskave razširjenosti dveh enoletnih trav Crypsis aculeata in Heleochloa schoenoides smo raziskovali na Slovaškem med letoma 2003 in 2007. Za rekonstrukcijo pojavljanja obeh vrst smo uporabili herbarijske in terenske
podatke. Opazili smo močno upadanje števila nahajališč, kjer se pojavlja C. aculeata. Razlog je predvsem
uničenje periodično poplavljenih slanih rastišč. Nasprotno pa se je pojavljanje vrste H. schenoides le malo
spremenilo. Vrsta lahko uspeva tudi na sekundarnih rastiščih (kolovozi, robovi njiv, uleknine na njivah).
Historično in recentno razširjenost smo prikazali na točkovnih kartah. Halofilno vegetacijo vrst Crypsis aculeata in Heleochloa schoenoides smo vzorčili s prilagojeno Braun-Blanquetovo metodo. Vse popisno gradivo smo
klasificirali s programom JUICE. Rezultati so pokazali, da asociacije Crypsidetum aculeate Wenzl 1934 na Slovaškem ne najdemo več, medtem ko je bila asociacija Heleochloetum schoenoidis (Soó 1933) Ţopa 1939 najdena
na nekaj rastiščih, vendar se v njej pojavljajo številne ruderalne rastlinske vrste.
Ključne besede: Crypsis aculeata, Heleochloa schoenoides, halofiti, slana tla, razširjenost, vegetacija, Slovaška
1. INTRODUCTION
the most important factor in their formation. The
saline soils contain toxic concentrations of soluble
salts in the root zone. The soluble salts consist of
chlorides and sulphates of sodium, calcium and
magnesium (Buol et al. 1997, Boros 2003). Areas
Saline soils belong to hydromorphic soils which
are strongly influenced by intensive water evaporation, and the salt dynamics of ground water is
* Department of Botany, Slovak University of Agriculture, Tr. A. Hlinku 2, SK-949 76 Nitra, Slovakia, pelias@afnet.
uniag.sk
** Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 14, SK-84523, Bratislava, Slovakia, daniel.dite@
savba.sk
*** Department of Botany and Zoology, Masaryk University, Kotlářská 2, CZ-611 37 Brno, Czech Republic, grulich@
sci.muni.cz
**** Nábrežná 8, SK-941 03 Úľany nad Žitavou, Slovakia, msadovsky@gmail.sk
Hacquetia 7/1 • 2008, 5–20
Typical dominants of periodically flooded saline soils are annual grasses Crypsis aculeata (L.)
Aiton and Heleochloa schoenoides (L.) Host ex Roem.
Crypsis aculeata occupies a wide area from North
Africa, Spain and Portugal to northern China and
Mongolia. Distribution of Heleochloa schoenoides is
very similar. However, the area of the species gets
larger towards Tibet and northwestern India; in Africa from Senegal to Mozambique, Malawi, Tanzania, and Madagascar (Conert 1983). The northern
distribution border of both species passes through
Central Europe (Holub & Grulich 1999a, b), occurrence of these species in this region has always
been rare. The situation of historical and present
occurence in the Czech Republic was described
by Grulich (1987). Both species are evaluated as
critically endangered in Slovakia (Feráková & al.
2001) and the Czech Republic (Holub & Procházka 2000). In Austria, Crypsis acuelata is evaluated as
endangered, while Heleochloa schoenoides as strongly endangered (Niklfeld & Schratt-Ehrendorfer
1999).
Crypsis acuelata and Heleochloa schoenoides plant
communities are usually included in classis Crypsidetea aculeatae Vicherek 1973, ordo Crypsidetalia
aculeatae Vicherek 1973 and alliance Cypero-Spergularion salinae Slavnić 1948. Within the frame of
this alliance both taxa are components of some associations and subassociations (see Vicherek 1973,
Mucina & Maglocký 1985, Mucina 1993, Borhidi
1996, Pop 2002 etc.).
of saline soils are widely distributed over the world
and constitute approximately 10 % of the terrestrial surface (Szabolcz 1991, O’Leary & Glenn 1994).
In Central Europaean countries with an arid continental climate, saline soils are widely spread
mainly throughout Hungary, where alkali habitats
cover nearly 400 000 ha (Molnár &Vajda 2000).
In Slovakia, saline soils covered by halophytic
plant communities are distributed mainly in warm
and dry lowland regions. The largest areas of saline soils are concentrated in the Danube Lowland – they are distributed from the surroundings
of towns of Komárno and Štúrovo to the town of
Nitra (Krist 1940, Krippelová 1965). The second
area of saline soils is in the East Slovakian Lowland
in the neighbourhood of the villages of Malčice,
Zemplínske Kopčany, Malé Raškovce, and Veľké
Raškovce (Vicherek 1964). The last small area
near the town of Malacky in the Záhorská nížina
Lowland (Krist 1940) was destroyed in the sixties.
Saline soils created by the mineral springs in basins of northern Slovakian mountains represent
special habitats (Šmarda 1961, Vicherek 1973, Dítě
& al. 2004).
The areas of saline soils have all been markedly
reduced during the last few decades due to human
activities. For instance, Osvačilová & Svobodová
(1961) mentioned approximately 8300 ha of soil
with saline vegetation in the Danube Lowland, but
today only 500 ha have been re-discovered there
(Sádovský & al. 2004b). It is this reduction that
strongly affects these plant communities distributed on soils with high salinity and a high level of
underground water level during spring months.
For example, we confirmed that Camphorosmetum
annuae now covers only a few square metres in Slovakia, but the typical vegetation was found only at
some of the last square decimetres. Initial stages of
periodically flooded saline soils are now rare and
are usually developed in secondary biotopes (fields
and field borders, depressions in rural roads etc.).
Plant communities of periodically flooded
saline habitats represent unique initial stages of
plant succession. They cover river banks as well as
terrain depressions. The formation of these plant
communities is closely associated with annual water level decrease and we can usually find it in the
second half of the vegetation period. This type of
biotope is rare in Slovakia nowadays; it has predominantly developed in wet depressions of intensively farmed fields. Vegetation of saline flooded
banks is even more seldom – only two micro-localities still exist (Sádovský & al. 2004a).
The aims of our paper were:
a) to describe the historical and recent distribution of Crypsis aculeata and Heleochloa schoenoides
in Slovakia,
b) to assess the recent stage of initial Crypsis aculeata and Heleochloa schoenoides saline communities
in Slovakia,
c) to compare present vegetation data with older
works published by Krist (1940) and Vicherek
(1973).
2. Material and Methods
The study was carried out during the years 2003–
2007. The data concerning the distribution of the
species were obtained from herbariums BP, BRA,
BRNU, BRNM, NI, NIM, KO, MMI, PMK, PR, PRC,
SAV, SLO and ZV. Herbarium specimens collected during field research are stored in herbarium
P. Eliáš jun., D. Dítě, V. Grulich, M. Sádovský: Distribution and Communities of Crypsis Aculeata and Heleochloa Schoenoides …
Figure 1: Area of study.
Slika 1: Območje raziskav.
3. Results and Discussion
NI. Herbarium abbreviations are used according
to Holmgren & al. (1990) and Vozárová & Sutorý
(2001). The active field research was carried out to
confirm recent localities of these species. Results
of this study are presented on the point maps constructed using DMAP software. The grid on the
maps (5’×3’) follows one that was described by
Niklfeld (1971).
The vegetation relevés were sampled according
to the Zürich-Montpellier approach using the adapted Braun-Blanquet scale (Barkman & al. 1964). All
relevés were stored using the TURBOVEG database
software (Hennekens & Schaminée 2001) and data
analyzed in JUICE software (Tichý 2002).
The nomenclature of ferns and flowering plants
follows Marhold & Hindák (1998), the names of
syntaxa are used in accordance with Molnár &
Borhidi (2003). The abbreviations of works published before 1956 follows Futák & Domin (1960).
Phytogeographical divisions of Futák (1980) are
also used (Fig. 1).
3.1 Distribution of Crypsis aculeata (L.)
Aiton in Slovakia
The species is distributed only on the Danube Lowland in Slovakia. In the course of our study we found
all together 30 localities of Crypsis aculeata. Half of
them were documented by herbarium specimens,
the other part was mentioned only in the literature
(see Fig. 2 and data below). However, we would
here like to refer to frequent determination mistakes – young individuals of Heleochloa schoenoides
were often determined as Crypsis aculeata. Hence
we take it that approximately a third of all the literature data are probably erroneous and therefore
they need to be used with care. Our deduction is
based on the fact that C. aculeata is an obligate halophyte and it grows only on strongly salinizated
soils of saline lakefronts (Krist 1940). On the other
hand, Heleochloa schoenoides, although included in
Hacquetia 7/1 • 2008, 5–20
because no saline habitats exist in this region (neither presently nor historically) and Kupčok did not
include this species in his review of Pukanec flora
(Kupčok 1956).
the group of obligate halophytes (Krist 1940) colonizes even more dry sub-saline and ruderal habitats
(field depressions, rural roads, fallows etc.). Sádovský & al. (2004a) interpreted it as an effect of a
wider ecological amplitude of H. schoenoides.
Novák (1954) also mentioned the species from
saline biotopes of the Záhorská nížina lowland, but
he did not point out any exact site. We consider
this record doubtful.
We established a marked reduction of Crypsis
aculeata distribution (Fig. 2.) with respect to strong
changes in Slovak saline areas (see Sádovský & al.
2004b). The species occurs now only at two microsites in the village of Tvrdošovce. We ascertained
during our field research that all other localities
were destroyed – predominantly by land reclamation (Búč, Dolný Jatov, Hájske, Močenok, Nový
Život, Palárikovo, Šurany, Trnovec nad Váhom,
Veľká Dolina, Zlatná na Ostrove), but also by building development (Komárno, Malá Maňa) and forestation by hybrid poplars (Veľké Kosihy).
A specimen of Crypsis aculeata collected by Samuel Kupčok was found in the herbarium BP. The
specimen’s label mentioned the village of Pukanec
(the Štiavnické vrchy Mts. in the Carpaticum phytogeographical region). We assumed that this specimen probably had its label accidentally exchanged,
3.2 Distribution of Heleochloa schoenoides
(L.) Host ex Roemer in Slovakia
The occurrence of Heleochloa schoenoides is concentrated in the southwestern and southeastern part of
Slovakia. Most of the 30 localities are known from
the territory of the Danube Lowland, but one locality was reported from a lower reach of the river of
Ipeľ (Zalaba) and circa 10 locations were found in
the East Slovakian Lowland (Fig. 3).
Novák (1954) mentioned the occurrence of this
species in the Záhorská nížina Lowland. However,
the origin of this reference is not clear. Similarly
as in the case of Crypsis aculeata, we found no other literature reference or herbarium specimen of
Heleochloa schoenoides from this region. However, a
recent occurrence of Heleochloa schoenoides on the
sand bank of the Dyje River (near the junction with
the Morava River) was published in South Moravia (Šumberová et al. 2000). A recent occurence
is also known from the Austrian part of Marchfeld
Figure 2: Historical and recent distribution of Crypsis aculeata in Slovakia: * – data from literature, ● – before 1980, ○ – 1980
onwards.
Slika 2: Historična in recentna razširjenost vrste Crypsis aculeata na Slovaškem: * – literaturni podatki, ● – pred 1980,
○ – po 1980.
P. Eliáš jun., D. Dítě, V. Grulich, M. Sádovský: Distribution and Communities of Crypsis Aculeata and Heleochloa Schoenoides …
Figure 3: Historical and recent distribution of Heleochloa schoenoides in Slovakia: ○ – before 1980, ● – 1980 onwards.
Slika 3: Historična in recentna razširjenost vrste Heleochloa schoenoides na Slovaškem: ○ – pred 1980, ● – po 1980.
(Schratt-Ehrendorfer 1999). These findings evoke
an idea of the species possibly occurring on the
Záhorská Lowland in Slovakia as well.
Heleochloa schoenoides hybridizes with its sibling
species H. alopecuroides. Both species grow in similar ecological conditions and are often found together. The hybrid – Heleochloa × bernatskyana (Degen) Boros – was reported from the surroundings
of the Mužľa Village. The specimen is deposited in
the herbarium in Budapest (Boros 1917 BP). This
hybrid needs further study: it is not clear if the cited specimen represents a real hybrid combination
or only an extreme variability of one of the supposed parents.
Recent distribution of the species more or less
corresponds to pre-existent localities. Contrary to
previous species, Heleochloa schoenoides is adapted to
the strong disturbance of saline habitats (amelioration, ploughing) and it also frequently colonizes
secondary sites. We found the species at 18 localities on the Danube Lowland. We did not confirm
its occurrence on the East Slovakian Lowland. On
the other hand, Zlacká (2006) confirmed one locality in East Slovakia in 2005 and many localities were
found here earlier in the past (Vicherek 1964).
On the basis of these data, we now assume a more
frequent occurrence of H. schoenoides in the area.
Many plant species of denuded beds (e. g. Carex bohemica, Dichostylis micheliana, Schoenoplectus supinus
etc.) occur irregularly – in years with favourable climatic conditions they create large populations but
in years of unfavourable climatic conditions they
are missing (Hejný 1960, Holub 1999).
3.3 Communities with dominance of
Crypsis aculeata in Slovakia
We found a biotope of periodically flooded saline
banks with the presence of Crypsis aculeata only at
two microlocalities during our study. Both were
situated in the urban area of Tvrdošovce village
(Fig. 4).
Vegetation on flooded lakesides fluctuated distinctively from year to year depending on the water
levels of the two lakes mentioned above. A high
abundance of vascular plants was observed during
the relatively dry and warm years of 2003 and 2004.
By contrast in years 2005–2006 the water level was
relatively stable and no contiguous plant cover
was discovered. Abundance of Crypsis aculeata was
always rather low in those two habitats – the species covered only 5–50 % area of the relevés, usually 10–25 %. The number of all species recorded
Hacquetia 7/1 • 2008, 5–20
Figure 4: Village of Tvrdošovce: one of two last microsites of Crypsis aculeata in Slovakia. Denuded waterbanks of the small
pool are in optimal conditions covered by an ephemeral community with presence of this species. Photo: D. Dítě, 21. 9. 2004.
Slika 4: Pri vasi Tvrdošovce: eno od dveh zadnjih mikrorastišč vrste Crypsis aculeata na Slovaškem. Goli bregovi manjših
mlak so v najugodnejših razmerah porasli z efemerno združbo te vrste. Foto: D. Dítě, 21. 9. 2004.
per plot was also relatively low. The species richness varied from 6 to 16 species per relevé (average
9.8). On the basis of these data we classified both
communities as association Heleochloetum schoenoidis
(Soó 1933) Ţopa 1939 [syn. Cyripsidetum schoenoidis
(Soó 1938) Ţopa 1939]. It must be noted that this
vegetation was also strongly ruderalized (Table 1,
relevés no. 3–10).
We found that the recent conditions are not
optimal for the existence of Crypsidetum aculeatae
at the two above mentioned microlocalities in the
area of Tvrdošovce village (and in the whole of
Slovakia respectively). We therefore suppose that
this plant community is now missing in Slovakia.
The last occurrence of Crypsidetum aculeatae in Slovakia was thus confirmed by Valachovič (1995) in
the same village in the first half of the nineties.
It is interesting that the author did not mention
Heleochloa schoenoides in his relevés, since we found
Crypsis aculeata always together with H. schoenoides
there. This change could be a result of desalination
of the soil.
By comparing our data with the work of Vicherek (1973) from the former Czechoslovakia,
the correctness of our conclusions that the studied
vegetation is now missing in Slovakia can be clearly
confirmed. The author differentiated two subassociations within the scope of Crypsidetum aculeatae
using data from South Moravia and south-west Slovakia. Subassociation typicum Vicherek 1973 is characterised by strong dominance of Crypsis aculeata
species (over 75 %) and the very low number of
other taxa per relevé (3–6 species). Subassociation
chenopodietosum glauci Bojko ex Vicherek 1973 was
distinguished based on the dominance of Chenopodium glaucum (abundance 75–100 %) and Spergularia salina (up to 50 %). By contrast Crypsis aculeata
was present only sparsely, the author mentioned
maximal abundance up to 25 %. The number of
species per relevé was low again (3–7 species).
10
P. Eliáš jun., D. Dítě, V. Grulich, M. Sádovský: Distribution and Communities of Crypsis Aculeata and Heleochloa Schoenoides …
Figure 5: The Kelemen-Szék Lake in Hungary, typical association of Crypsidetum aculeatae on denuded saline bank. Photo:
D. Dítě, 11. 9. 2007.
Slika 5: Jezero Kelemen-Szék Lake na Madžarskem, tipična asociacija Crypsidetum aculeatae na golih slanih brežinah. Foto:
D. Dítě, 11. 9. 2007.
Vicherek (1973) regarded Crypsidetum aculeatae as a Pontic-pannonian community. Outside
Slovakia, the association was reported from northeastern Austria (Bojko 1931, 1932, Wendelberger
1959), southeastern part of the Czech Republic (Vicherek 1962, 1973, Šumberová 2007), central and
eastern Hungary (Soó 1934, Timár 1954, Borhidi
1996, Molnár & Borhidi 2003), northern part of
Serbia (Slavnić 1948, Kojić et al. 1998), Romania
(Ţopa 1939, Sanda et al. 1999, Popescu 2005a) and
Ukraine (Bilik 1937, 1963). Hilbig (2000) mentioned the community from Mongolia which was
the reason why Šumberová (2007) assumed occurrence of the association on salt steppes in southern Russia. On the other hand, Mucina (1993)
mentioned the community only from Austria, the
Czech Republic, Slovakia, Hungary, and Serbia. In
addition to that, Micevski (1965) described a Macedonian endemic community Crypsidetum aculeatae
balcanicum, which was developed in small depres-
sions and channels in Ovče Pole. The author described this community on the basis of the occurrence of species Puccinelia convoluta and Spergularia
marginata which are not present in Crypsidetum aculeatae described from Central Europe. Revision of
his data is needed in the future.
A survey of the saline lakes in Central Hungary
(surroundings of the villages of Akasztó, Fülopszálás, Harta and Solt) was made in 2006 and 2007
to obtain comparative data of vegetation with Crypsis aculeata. We found a typical cover of the species
comparable to data given by Timár (1954), Wendelberger (1959) and Vicherek (1973) (Tab. 2). We
recorded a wide area of dry saline lakes overgrown
by thousands of individuals of the species (Fig. 5).
On the other hand, it seems that this community is
abundant only locally in Hungary, because Varga
& Várgáné (1999a) included it in the Red book of
Hungarian vegetation. Similarly, the community is
sporadic in Austria – it occurs only in the area of
11
Hacquetia 7/1 • 2008, 5–20
3.4 Communities with dominance of
Heleochloa schoenoides in Slovakia
the Neusiedler See (Mucina 1993, Körner 2006).
In Serbia Crypsidetum aculeatae is rare and occurs
mainly in northern Serbia (Vojvodina: Potamišje,
Derečka bara). Only a few isolated sites are found
in southern parts of the country in the surroundings of the Bujanovac settlement (Vučković 1983,
Knežević 1994, Zlatković et al. 2005). According to
Popescu (2005a) recent distribution of Crypsidetum
aculeatae in Romania is limited; the community still
occurs in regions of Muntenia, Moldova, Dobrogea
and in the Danube Delta. The author supposes that
the community presently covers an area of approximately 5–10 hectares only and that conservation of
the association is inadequate. Similarly, Micevski’s
Crypsidetum aculeatae balcanicum has recently been
threatened by limited distribution and is under
direct destruction (Matevski et al. 2003). Apart
from Slovakia, the community is also missing in the
Czech Republic (Danihelka & Hanušová 1995, Dítě
2006 ined., Šumberová 2007) and moreover the
species survive there thanks only to human support
(Holub & Grulich 1999a).
Occurrence of vegetation with dominance of Heleochloa schoenoides was recorded at several localities
during recent times. In 2004 we found typical vegetation mainly at the bottom of a small periodical
pool near the Tvrdošovce railway station, (Tab. 1,
relevés 1, 2, Fig. 6). It was distinguished by great
dominance of the species (nearly 100 %) and low
richness and abundance of other species (5 species
in relevés with abundance to 5 %). Similar vegetation was found at a periodically flooded habitat of a
field depression in the surroundings of the village
of Močenok in 2006 (Tab. no. 1, relevés no. 12, 16–
20). However, also many synanthropic species grew
there. Other habitats of Heleochloa schoenoides were
found on field margins and at open places strongly
affected by man (field depressions, field roads, wet
fallows, Fig. 7). The presence of ruderal species was
caused by lower soil salinity and direct contact with
intensively managed fields. Therefore this vegeta-
Figure 6: Heleochloetum schoenoidis on drying lake bank in Tvrdošovce. Photo: P. Eliáš jun., 30. 8. 2004.
Slika 6: Heleochloetum schoenoidis na izsušenem jezerskem bregu pri vasi Tvrdošovce. Foto: P. Eliáš jun., 30. 8. 2004.
12
P. Eliáš jun., D. Dítě, V. Grulich, M. Sádovský: Distribution and Communities of Crypsis Aculeata and Heleochloa Schoenoides …
tion was modified and the presence of many ruderal plant species was recorded (e. g. Amaranthus
retroflexus, Atriplex sagittata, A. tatarica, Echinochloa
crus-galli, Lepidium ruderale, Polygonum rurivagum
etc.). In some extreme cases the cover of ruderal
species was close to 50 % per plot. Despite this fact,
all samples in Slovakia can be assigned to the association Heleochloetum schoenoidis (Soó 1933) Ţopa
1939 [syn. Crypsidetum schoenoidis (Soó 1938) Topa
1939] even though they are more or less ruderalized. In some cases, when the soil salinity was low,
we found a transition to communities of the class
Isoëto-Nanojuncetea (e.g. on a dry bed of a small pool
in Komárno – Ďulov dvor farmstead). The same pattern was recorded by Šumberová (2007) in degraded saline pools of Southern Moravia. We suppose
that this change is probably irreversible and will
result in full extinction of the community.
The community is distributed in the Ponticpannonian region (Vicherek 1973). Reliable data
about its occurrence were given from the Czech
Republic (Vicherek l. c., Šumberová 2007), Slovakia (Krist 1940, Vicherek 1964, 1973, Sádovský &
al. 2004b), Hungary (e. g. Soó 1940, 1947, Bodrogközy 1962, Borhidi 1996, Molnár & Borhidi 2003),
Romania (Ţopa 1939, Pop 2002, Popescu 2005b),
and Mongolia (Hilbig 2000). Following Hilbig’s data Šumberová (2007) suggested the occurrence of
the community in Ukraine and in steppe areas of
southern Russia. Nevertheless, the community was
not mentioned in published surveys of saline vegetation of Ukraine and Russia (Solomakha 1996,
Freitag et al. 2001, Voityuk 2005). This stage could
also be caused by different syntaxonomical treatment of the Heleochloa schoenoides communities.
Vicherek (1973) mentioned that Heleochloetum
schoenoidis included two subassociations on the territory of the former Czechoslovakia: subass. typicum
and subass. spergularietosum marginatae. Similarly as
in the case of Crypsidetum aculeatae association the
presence of 3–5 plant species was typical for subass.
typicum. The second subassociation was characterised by slightly higher species richness (7–9 plant
taxa per relevé). In our relevés it was approximately two times greater due to penetration (invasion?) of ruderal species (8–16 plant species per
relevé, Tab. 1). Therefore it is now impossible to
distinguish the above mentioned subassociations.
Heleochloetum schoenoidis sites were mainly on the
area of agricultural crops and they were usually destroyed in years with dry weather. The same stage
was pointed out by Šumberová (2007); the author
reported ca. 10 species per plot.
Figure 7: Rural roadside – a secondary habitat of Heleochloa
schoenoides. Photo: P. Eliáš jun., 16. 7. 2005
Slika 7: Kolovoz – nadomestni habitat vrste Heleochloa schoenoides. Foto: P. Eliáš jun., 16. 7. 2005
The comparative findings were made near the
villages of Solt (Table 1, relevés no. 13, 14) and
Balmazújváros in Hungary (Table 1, relevé no. 11).
Only ruderalized Heleochloetum schoenoidis vegetation was found and the community was established
just on secondary habitats (field depressions, field
margins). Despite our findings, the community is
the most common in Hungary; the occurrence was
mentioned mainly from the large region of the
Great Hungarian Plain (Soó 1939, 1964). In this
region approximately 1200 ha of saline lakes and
13,000 ha of salt meadows and marshes were mentioned (Molnár & Varga 2000). However, Varga &
Várgáné (1999b) have recently included the association into the Red book of Hungarian vegetation.
Distribution in Romania is very scattered, Popescu
13
Hacquetia 7/1 • 2008, 5–20
(2005b) mentioned the community mainly from
districts of Transylvania, Banat, Muntenia, and
Moldova. The author estimated that the total area covered by Heleochloetum schoenoidis is no larger
than 35–40 hectares, and the community needs
active protection. The association is infrequent in
Serbia as well (Knežević 1990, Kojić et al. 1998). It
is centralised in the Vojvodina region and some
localities are situated in the central and southern
part of the country. The community is also very
rare in the Czech Republic, recent occurrence was
confirmed in south-eastern Moravia (Danihelka &
Hanušová 1995, Šubmerová 2007). These data suggest that the community is presently uncommon in
the whole of Central and Eastern Europe.
We are indebted to Rozália and Zoltán Orosi
(Harta, Hungary) and Alexander Fehér (Nitra,
Slovakia) for help with field research in Hungary. This paper was particularly supported by the
grants of the Grant Agency for Science “VEGA”,
Bratislava, Slovakia (grants No. 1/3446/06 and
No. 1/0672/08) and of the Ministry of Education
of the Czech Republic (grant no. 0021622416).
6. References
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4. Conclusions
A stronger decrease of saline areas was recorded
in the last 30–40 years in the Czech Republic and
Slovakia. Most of the habitats were fully destroyed
by agriculture and construction activities (Sádovský
et al. 2004b). The small fragments were conserved
to the present time, but the habitats are situated
among crop fields where the water regime is being
changed. Habitat degradation caused changes of
vegetation cover. Soil salinity has decreased and the
attack of weedy species has been recorded. Because
Crypsis aculeata is unable to adapt to these changes
the number of localities markedly decreased and
the species is threatened with extinction. Typical
communities with dominance of the species were
not found at two still existing micro-sites. It survived
scattered in ruderalized vegetation. On the other
hand, Heleochloa schoenoides is partly adapted to
the ecologically changed conditions and thus
historical and present distribution of the species is
practically identical. But the Heleochloa schoenoides
communities were modified dramatically – they
occurred usually at secondary habitats and the
ruderal species were very abundant. Finally, our
data pointed out that Crypsis aculeata and Heleochloa
schoenoides communities are now endangered in
the whole Pannonian region.
5. Acknowledgements
We are grateful to Milan Valachovič (Bratislava,
Slovakia) for valuable comments and contributions, Viktor Kerényi-Nagy (Budapest, Hungary)
for his help with the herbarium data excerption.
14
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1951 BRNU; Šourek 1951 PR). – Hájske, Mešterik
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Kolník & Sádovský 2002 NI; Grulich 2002 BRNU
– near the railway station). – Tvrdošovce, small fishpond on northern margin of the village (Grulich
1985 MMI; Pluhař 1986 BRNU; Grulich & Kochjarová 1988 SLO). – Palárikovo, near railway station
(Weber 1936 BRA). – Palárikovo, Veľké Čiky farmstead (Weber 1933 BRNM; Smejkal 1965 BRNU).
– Palárikovo, Malé Čiky [= Čiastka] farmstead
(Weber 1935 BRA; Valenta 1937 BRA; Krist 1938
BRNU, Krist; 1940). – Palárikovo, Bačala farmstead
(Weber 1935 BRA). – Búč (Dvořák 1965 BRA; Unar
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b) Literature references without confirmation
of herbarium specimens:
4. The Záhorská nížina lowland (Novák 1954)
[not mapped]. 6. Tôň, wetland site in the village
surrounding (Futák 1949 ined.). – Opatovský
Sokolec, southern margin of the village (Vicherek
1973). – Okoč, southern margin of the village (Vicherek 1973). – Kľúčovec, bank of small pond in
the village (1975) (Svobodová1988: 11). – Malé Kosihy, in the village (Vicherek 1973). – Žihárec, near
the railway station (Vicherek 1973). – Palárikovo,
Horný Jur farmstead (Futák 1949 ined.). – Chotín,
bank of small pond in the village (1974) (Svobodová1988: 11). – Pribeta, saline site near railway
station (Futák 1949 ined.). – Mužľa (Feichtinger
1899: 392; Domin 1933: 247). – Nána (Feichtinger
1899: 392; Domin 1933: 247). – Kamenín (Feichtinger 1899: 392; Domin 1933: 247).
List of localities of Heleochloa schoenoides:
2. Zalaba (Hejný 1951 SLO). 4. Záhorská nížina
lowland, saline soils (Novák 1954) [not mapped].
6. Bratislava, wetlands near Istrochem factory
(Feráková 1972 SLO). – Bratislava-Vajnory, Čierna
Voda farmstead (Ondrášek 1986 MMI). – Svätý
Jur, Pálffyovský dvor farmstead (Holuby 1913
PRC). – Svätý Jur, site Šúr (Zigmundík 1913 BRA;
Holub & Grulich 1999b) [data probably refer to
the previous]. – Hájske (Krist 1936 BRNU, 1937
BRNU; Weber 1931 BRNM, 1937 BRA; Krist 1940;
Vicherek – SW margin of the village; Sádovský &
al. 2004a). – Hájske, Mešterik farmstead (Weber
7. Appendix
List of localities of Crypsis aculeata:
a) Herbarium specimens and confirmed literature references
6. Nový Život, part Eliášovce (Mergl 1902 SAV).
– Veľké Kosihy, salt pasture near pri skladoch (Májovský 1963 SLO; Májovský & al., Acta Fac. Rer.
Natur. Univ. Comen., Bot. XXIII: 7, 1974). – Zlatná na Ostrove (Weber 1936 BRA; 1949 BRNM; Vicherek 1973). – Komárno, near the railway station
(Futák 1949 ined., Šmarda 1951 BRNM; Smejkal
17
Hacquetia 7/1 • 2008, 5–20
1935 BRA, 1936 BRNM; Krist 1937 BRNU; Klokner 1955 SLO; Weber 1970 PR, 1977 PR; Eliáš jun. &
Dítě 2005 NI). – Močenok, Síky farmstead (Weber
1935 BRA, 1936 BRNM; Krist 1937 BRNU; Krist
1940; Klokner 1955 SLO; Unar 1965 BRNU ut “salt
meadow Močenok”; Weber 1970 PR, 1977 PR; Eliáš
jun. & Dítě 2005 NI). – Cabaj-Čapor, Szík farmstead
[locality probably refers to the previous] (Vlach
1935 PRC). – Šaľa, Andelek farmstead (Weber 1936
BRNM). – Trnovec nad Váhom, near the railway
station (Weber 1935 BRA, 1936 BRNM). – Horný
Jatov (Krist 1936 BRNU; Weber 1936 BRNM; Sádovský 2002 NI). – Dolný Jatov, Čierny vršok farmstead (Osvačilová 1953 PRC, Dostál 1955 PRC). –
Dolný Jatov (Weber 1935 BRA, 1936 BRNM; Holub
& Grulich 1999b; Grulich 1988 MMI – wet place 1
km SSW from the village). – Polný Kesov (Dostál
1955 PRC). – Komjatice, Ružový dvor farmstead
(Vlach 1934 PRC; Krist 1940; Sádovský & al. 2004a).
– Tvrdošovce (Krist 1924 BRNU, 1936 BRNU;
Krist 1940; Weber 1935 BRA, 1936 BRA, BRNM;
Pospíšil 1952 BRNM; Pokluda 1961 BRNM; Dostál
1966 PR; Vicherek 1973 – small fishpond on N part
of the village; Weber 1977 PR; Grulich 1985 MMI
– small fishpond on N part of the village; Grulich
1985 MMI – wet pastures on NW part of the village;
Grulich 1986 MMI – 1.5 km NW from the railway
station; Pluhař 1986 BRNU; Grulich & Kochjarová
1988 SLO; Vozárová 1994 BRA; Svobodová 1994
NI; Eliáš jun., Dít, Kolník & Sádovský 2002 NI).
– Palárikovo (Krist 1924 BRNU; Krist 1940; Scheffer 1923 BRNU – in agris; Weber 1935 BRA, 1936
BRNM; Skřivánek 1948 BRNM; Futák 1949 SLO;
Dvořák 1952 BRNM; Pospíšil 1952 BRNM; Šourek
1954 PR – NW from the railway station; Pokluda
1961 BRNM; Vicherek 1973 – NE margin of the
village; Grulich 1985 MMI – NW from the railway station; Holub & Grulich 1999b). – Palárikovo, Berchtold farmstead (Weber 1936 BRNM).
– Palárikovo, Bačala farmstead (Weber 1935 BRA,
1936 BRNM). – Palárikovo, Veľké Čiky farmstead
(Weber 1936 BRNM; Klokner 1955 SLO; Smejkal 1965 BRNU; Unar 1965 BRNU ut Heleochloa
alopecuroides). – Palárikovo, Malé Čiky farmstead
[= Čiastka] (Weber 1933 BRNM, 1935 BRA, 1936
BRNM; Krist 1937 BRNU; Krist 1940; Weber 1977
PR; Grulich 1988 MMI; Eliáš jun. & Sádovský 2003
NI). – Šurany, Akomáň farmstead (Weber 1936
BRNM; Weber 1977 PR; Grulich 1987 MMI; Holub
& Grulich 1999b). – Malá Maňa, small fishpond in
the village (Vicherek 1973). – Žihárec, small fishpond near the railway station (Vicherek 1973). –
Tôň (Futák 1949 SAV; Vicherek 1973). – Okoč (Krist
1940; Vicherek 1973). – Veľké Kosihy (Májovský
1963 SLO; Holub & Grulich 1999b). – Okánikovo,
Mostové Nature Reserve [= Dérhidia] (Čvančara &
Šourková 1972 BRNM, BRNU, PR; Grulich 1986
MMI; Eliáš jun. & Dítě 2004 NI). – Okoličná na
Ostrove, near the railway station (Májovský 1963
SLO; 1966 SLO, 1969 SLO). – Okoličná na Ostrove, Ekelský dvor farmstead (Krippelová & Zahradníková 1962 SAV). – Zlatná na Ostrove (Valenta
1936 BRA; Weber 1936 BRA, BRNM, 1937 BRNM;
Šourek 1954 PR; Hajdúk & Krippelová 1962 SAV;
Holub & Grulich 1999b; Grulich 1987 MMI – 0.5
km E from the railway station). – Kameničná (Feráková 1971 SLO; Řehořek & Svobodová 1971 NI ut
Crypsis aculeata). – Komárno, osada Pavel (Klokner
1966 PMK). – Komárno, osada Nová Stráž (Krist
1940; Magic 2002 BRA). – Komárno, near railway
station (Nábělek 1936 BRA; Hrabětová 1947 BRNU
ut Crypsis aculeata; Hrabětová 1951 BRNU – field
margin; Dvořák 1952 BRA – dno jamy neďaleko
mostu; Smejkal 1951 BRNU; Šmarda 1951 BRNM;
Hejný 1953 PR; Šourek 1954 PR). – Komárno,
Ďulov dvor farmstead (Klokner 1967 PMK; Eliáš
jun. 2003 NI). – Chotín (Futák 1949 SLO; Šourek
1954 PR – W from elevation point 109 m). – Iža,
Bokroš hamlet (Svobodová 1991 NI; Eliáš jun., Dítě
& Sádovský 2006 NI). – Pribeta (Klika 1937 PR).
– Búč (Feichtinger 1899: 392; Májovský 1963 SLO;
Vicherek 1963 BRNU, Biológia, 19: 463, 1964;
Dvořák 1965 BRA; Čvančara 1970 BRNM, 1971 PR;
Nevrlý 1971 BRNM; Dorotovičová 1994 PMK; Eliáš
jun., Dítě & Sádovský 2005 NI). – Gbelce (Krist
1936 BRNU; Vicherek 1973). – Mužľa (Feichtinger
1899: 392; Domin 1933v: 247; Šmarda 1949 BRNM;
Májovský 1963 SLO; Eliáš jun. &. Sádovský 2006
NI). – Štúrovo, Boží kopec hill, waterbank of the
Danube (Hejný 1953 PR). – Nána (Feichtinger
1899: 392; Domin 1933v: 247). – Kamenný Most,
Čistiny Nature Reserve [= Irtoványi rétek] (Šourek
1954 PR). – Kamenín (Feichtinger 1899: 392;
Domin 1933v: 247; Klika 1931 NI, 1936 NI; Nábělek
1936 SAV; Weber 1936 BRNM; Futák 1948 SLO;
Osvačilová 1956 NI ut Crypsis aculeata; Černoch,
Biológia, 15/11: 818, 1960; Májovský 1965 SLO;
Řehořek 1981 NI; Murín, Feráková & Činčura 1982
SLO; Grulich 1985 MMI; Svobodová 1982 NI; Eliáš
jun. 2005 NI). 8. Kačanov, S obce (Vicherek, Biológia 19: 556, 1964). – Malčice, 2 km SW from the
village (Vicherek, Biológia 19: 556, 1964). – Beša
(Zlacká 2005 NI). – Somotor, under northern
slope of Tarbucka hill (Vicherek, Biológia 19: 556,
1964). – Viničky, river branch of the Bodrog River
(Májovský 1964 SLO). – Veľký Kamenec, S from
18
P. Eliáš jun., D. Dítě, V. Grulich, M. Sádovský: Distribution and Communities of Crypsis Aculeata and Heleochloa Schoenoides …
the village (Vicherek 1962 BRNU; Vicherek, Biológia 19: 556, 1964). – Velký Kamenec, hamlet Bukov
[= Ružový dvor farmstead?], river branch Velká
Krčava (Hejný 1953 PR). – Strážne, waterbank of
Krčava canal 3 km W from the village (Vicherek,
Biológia 19: 556, 1964). – Malý Horeš, SW from the
village (Vicherek, Biológia 19: 556, 1964). – Leles,
3 km N from the village (Vicherek Biológia 19:
556, 1964). – Leles, Kapoňa hamlet, sandy field
south from the village (Májovský & Záborský 1961
SLO). – Ruská, in the village (Vicherek, Biológia
19: 556, 1964; Vicherek 1973). – Morčany, W from
the village (Vicherek, Biológia 19: 556, 1964) [not
mapped; unidentified locality].
Table 1: Analytical table of recent Crypsis aculeata and Heleochloa schoenoides plant communities in Slovakia
and Hungary.
Tabela 1: Analitična tabela recentnih rastlinskih združb s Crypsis aculeata in Heleochloa schoenoides na Slovaškem
in Madžarskem.
Relevé number:
Relevé surface (m2)
Coverage E1 %
Coverage E0 %
Number of species per relevé
Potentilla anserina
Xanthium strumarium
Persicaria lapathifolia
Chenopodium glaucum
Crypsis aculeata
Agrostis stolonifera
Carex secalina
Plantago major ssp. intermedia
Lotus tenuis
Bidens tripartitus
Spergularia salina
Tripleurospermum perforatum
Polygonum rurivagum
Echinochloa crus-galli
Puccinellia distans
Matricaria chamomilla
Plantago major
Potentilla supina
Elytrigium repens
Atriplex tatarica
Heleochloa schoenoides
Atriplex prostrata
Juncus compressus
Aster tripolium ssp. pannonicus
1
16
80
0
5
r
1
r
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
+
.
.
2
16
95
0
5
1
r
r
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
5
.
r
.
3
10
45
0
7
+
+
.
3
b
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
1
1
+
.
4
8
35
0
7
+
r
.
.
b
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
b
+
+
.
5
4
75
0
8
1
+
.
1
a
1
.
.
.
.
.
.
.
.
.
.
.
.
.
.
a
1
a
.
6
2
35
0
6
+
.
.
1
a
.
.
.
.
.
.
.
.
.
.
.
.
.
.
.
b
1
.
1
7
10
70
0
16
+
+
.
+
3
a
r
+
1
a
.
.
.
.
.
.
.
.
.
.
3
1
a
+
8
9
60
0
12
.
+
.
+
1
3
+
+
+
+
+
.
.
.
.
.
.
.
.
.
b
.
a
a
9
8
60
0
9
.
.
.
.
b
1
r
+
+
.
.
.
.
.
.
.
.
.
.
.
a
+
b
a
10
16
70
0
14
1
r
r
.
b
1
.
.
1
+
1
.
.
.
a
.
.
.
.
.
3
1
a
1
11
12
25
0
8
.
+
.
.
.
.
.
.
.
.
+
.
.
.
+
+
.
.
.
.
b
.
.
+
12
16
95
0
15
.
.
.
.
.
.
.
1
+
.
a
+
+
1
.
.
.
.
+
.
5
a
.
.
13
8
55
0
13
.
.
.
+
.
.
.
+
.
.
a
+
+
+
1
.
.
.
.
.
3
1
.
14
16
60
0
11
.
.
.
.
.
.
.
.
.
.
1
+
+
.
+
.
.
.
.
.
4
+
+
.
15
16
45
0
10
.
.
.
.
.
.
.
.
.
.
.
+
a
a
1
.
.
.
1
.
3
1
.
.
16
16
85
0
15
.
.
.
+
.
.
.
+
.
.
.
.
1
1
.
+
.
.
.
+
4
+
.
.
17
16
80
2
15
.
.
.
.
.
.
.
.
.
.
.
+
a
1
.
.
1
r
.
.
5
+
1
.
18
16
45
0
8
.
.
.
.
.
.
.
.
.
.
.
+
b
a
.
r
+
.
.
.
3
.
+
.
19
9
60
0
12
.
.
.
.
.
.
.
.
.
.
.
+
1
a
1
.
+
+
.
.
3
+
.
+
20
8
70
0
11
.
.
.
.
.
.
.
.
.
.
.
.
3
r
+
+
r
.
1
+
a
.
.
.
21
16
45
0
16
.
.
.
.
.
.
.
.
1
.
.
.
3
+
1
+
.
+
1
.
a
+
+
.
22
10
45
0
8
.
.
.
.
.
.
.
.
.
.
.
.
3
r
+
.
.
.
+
+
a
.
.
.
* Characteristic species of Cypero-Spergularion salinae Slavnić 1948 are marked in bold.
Species recorded in two relevés only: Artemisia
monogyna 1 (20), + (21), Atriplex littoralis + (22), r
(23), Bolboschoenus maritimus 2a (4), 1 (13), Cirsium
arvense + (12), + (19), Cynodon dactylon + (13), r
(21), Hibiscus trionum r (16), r (17), Chenopodium
hybridum r (14), r (16), Kochia scoparia 1 (15), +
(23), Lepidium ruderale 1 (11), r (15), Plantago tenuiflora 1 (11), 2a (22), Rumex stenophyllus + (18, 19),
Taraxacum sect. Ruderalia + (12), + (19), Triticum
aestivum r (12), 1 (16).
19
Hacquetia 7/1 • 2008, 5–20
Species recorded in one relevé only: Alisma
plantago-aquatica r (13), Amaranthus retroflexus r
(16), Atriplex sagittata 1 (23), Brassica napus + (16),
Capsella bursa-pastoris + (16), Datura stramonium +
(12), Eleocharis palustris 1 (17), Epilobium tetragonum
1 (17), Hordeum geniculatum + (11), Chenopodium
rubrum 1 (13), Chenopodium urbicum + (12), Inula
britanica + (12), Juncus bufonius + (14), Lythrum hyssopifolia 1 (17), Pulicaria vulgaris r (23), Persicaria
amphibia 1 (7), Persicaria hydropiper r (17), Rumex
crispus + (14), Sclerochloa dura r (22), Sonchus arvensis + (7), Sonchus asper + (20), Trifolium fragiferum
ssp. bonannii r (17), Trifolium repens + (23).
Localities of relevés (number of relevé, locality,
exposition, elevation, altitude, sampling date):
1, 2: Tvrdošovce, denuded bottom of lake near
the railway station, 110 m n. m., 21. 9. 2004, 3–6:
Tvrdošovce, denuded waterside of lake, 115 m n.
m., 21. 9. 2004, 3: South, 5°, 4: South-eastern, 3°,
5: East, 7°, 6: East, 2°. 7–10: 7: Tvrdošovce, denud-
ed waterside of lake near the railway station, East,
3°, 8. 10. 2003, 8: South-eastern, 3°, 8. 10. 2003, 9:
East, 6°, 115 m n. m., 21. 9. 2004. 10: East, 2°, 8. 10.
2003, 11: Hungary, Hajddúbőszórmény, Kispród
farmstead, saline pasture near road, 100 m a. s. l.,
11. 7. 2007, 12: Močenok, Siky farmstead, depression in cereal field, 130 m a. s. l., 5. 10. 2006, 13,
14: Hungary, Solt, dry depression in maize field, 98
m n. m., 18. 9. 2006. 16: Močenok, Siky farmstead,
depression at rural road through a cereal field, 130
m a. s. l., 5. 10. 2006., 15. Veľké Kosihy, Mostové Nature Reserve (=Dérhidja), slopes on ploughed salt
meadow, 115 m n. m., 17. 9. 2006, 17, 19: Močenok,
Siky farmstead, depression in lucerne field, 13. 9.
2005, 18: Močenok, Siky farmstead, dry depression
in lucerne field, 13. 9. 2005, 20, 22: Veľké Kosihy,
Mostové Nature Reserve (=Dérhidja), rural road,
115 m a. s. l., 21. 9. 2004, 21: Veľké Kosihy, rural
road in ploughed part of Mostové Nature Reserve,
115 m a. s. l., 21. 9. 2004.
Tab. 2: Analytic table of the Crypsidetum aculeatae Wenzl 1934 recorded in Hungary
Tabela 2: Analitična tabela asociacije Crypsidetum aculeatae Wenzl 1934 na Madžarskem.
Relevé number:
Relevé surface m2
Coverage E1 (%)
Coverage E0 (%)
Number of species per relevé
Crypsis aculeata
Puccinellia distans
Bolboschoenus maritimus
1
16
35
0
1
3
.
.
2
16
40
0
1
3
.
.
3
16
70
0
2
4
+
.
4
16
60
0
2
4
1
.
5
16
85
0
2
5
.
+
6
16
80
0
1
5
.
.
7
16
80
0
1
5
.
.
Localities of relevés (number of relevé, locality, exposition, elevation, altitude, sampling date):
1–7: Kiskunság National Park, denuded waterside of the Kelemén-Szék Lake, 92 m a. s. l., 11. 9. 2007.
Recieved 14. 1. 2008
Revision recieved 31. 3. 2008
Accepted 3. 4. 2008
20
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