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Halophytic flora and vegetation in southern Moravia
and northern Lower Austria: past and present
Jiří Danihelka1,2, Kryštof Chytrý1,3, Martin Harásek1, Petr Hubatka1,4,
Klára Klinkovská1, Filip Kratoš1, Anna Kučerová1, Karolína Slachová1,
Daniel Szokala1, Helena Prokešová4, Eva Šmerdová1, Martin Večeřa1
& Milan Chytrý1*
We dedicate this article to the memory of Vít Grulich (1956–2022), a botanist extraordinaire
1Department of Botany and Zoology, Faculty of Science, Masaryk University, Kotlářská 2, CZ-611 37
Brno, Czech Republic; 2Institute of Botany of the Czech Academy of Sciences, CZ-252 43 Průhonice,
Czech Republic; 3Department of Botany and Biodiversity Research, University of Vienna,
Rennweg 14, A-1030 Vienna, Austria; 4Administration of the Protected Landscape Area Pálava,
Náměstí 32, CZ-692 01 Mikulov, Czech Republic
* corresponding author: chytry@sci.muni.cz
Abstract: Halophytic habitats are distinctive components of the landscape in southern Moravia,
Czech Republic, and the Pulkau valley in northern Lower Austria. We provide a historical over-
view of their flora and vegetation since the early 19th century and describe the current state
assessed in the field at all remaining sites in 2020. We summarized the historical and current dis-
tribution of 17 species with the strongest affinity for saline habitats in the study area: Bupleurum
tenuissimum,Cirsium brachycephalum,Crypsis aculeata,C. schoenoides,Galatella cana,Glaux
maritima,Juncus gerardii,Plantago maritima,Salicornia perennans,Samolus valerandi,
Scorzonera parviflora,Spergularia marina,S. media,Suaeda prostrata,Taraxacum bessarabicum,
Triglochin maritima and Tripolium pannonicum.Ofthese,Galatella cana,Salicornia perennans,
Suaeda prostrata and Triglochin maritima are regionally extinct. We also characterized the
bryoflora typical of saline habitats. We classified historical and newly recorded relevés and
identified 14 halophytic associations belonging to the alliances Chenopodion rubri,Meliloto
dentati-Bolboschoenion maritimi,Cypero-Spergularion salinae,Salicornion prostratae,Pucci-
nellion limosae,Juncion gerardii and Festucion pseudovinae. The vegetation of the alliance
Salicornion prostratae had disappeared in the study area by the 1970s. The alliance Festucion
pseudovinae, with the association Centaureo pannonicae-Festucetum pseudovinae, was recog-
nized as a new vegetation type for the Czech Republic. Using soil pH and electrical conductiv-
ity measurements, we evaluated the relationships of individual species and vegetation types to
soil salinity. This synthetic study shows that the once well-developed halophytic flora and vege-
tation in the study area steadily declined from the early 19th century to the 1980s, initially
mainly due to drainage and after the mid-20th century due to the combination of drainage and
cessation of grazing. The introduction of conservation management in the 1990s and ecological
restoration contributed to stabilizing plant diversity at the last saline sites. The future of halo-
phytic flora and vegetation depends on the continuation of conservation management.
Keywords: Czech Republic, flora, habitat, halophyte, history, inland saltmarsh, Lower Austria,
Moravia, nature conservation, saline, vegetation
Preslia 94: 13–110, 2022
doi: 10.23855/preslia.2022.013 Preslia
Received: 8 Aug 2021, Revised: 18 Nov 2021, Accepted: 22 Nov 2021, Published: 24 Mar 2022
Introduction
Inland halophytic flora and vegetation find their optimum mainly in arid and summer-
warm areas such as southern Ukraine, southern Russia, Middle and Central Asia and the
Middle East (Walter 1974, Eliáš et al. 2020). In such areas, salt steppes and salt meadows
are considered intrazonal vegetation within the steppe zone (Mucina et al. 2016).
Halophytes are favoured there by high soil salinity caused by intense evaporation, which
promotes the upward migration of salts to the upper soil horizons. The high salinity is det-
rimental to many plant species and favours a small number of adapted species, which are
considered obligate halophytes. Macroclimate is a strong driver of the broad-scale distri-
bution of halophytes even in central Europe, where they occur in the dry lowlands. While
the saline habitats and specialized halophytes are widespread and common in the central
parts of the Great Hungarian Plain (Molnár et al. 2008), they become rarer in the precipi-
tation-richer areas in the foothills of the Carpathians and the Alps (Vicherek 1973). In
such marginal areas, including southern Moravia and Lower Austria, other salinity
sources are important besides evaporation in dry macroclimate, especially springs in
mineral-rich Tertiary deposits. These halophytic habitats also depend on regular distur-
bance, especially traditional livestock and poultry grazing on villages’ common land.
Due to their recent decline, they are listed in the European Habitats Directive 92/43/EHS,
and most of their remaining sites are protected.
The halophytic flora and vegetation in southern Moravia are confined to the lowlands,
including the wider environs of the lower stretches of the Dyje and Svratka rivers, extend-
ing slightly into Lower Austria along the Pulkau stream. Their distribution is naturally
bounded by colline landscapes of the Bohemian Massif in the west and north-west, the
Weinviertler Upland in the south and the Western Carpathian foothills in the east. The
halophytic habitats in this area share some common features in vegetation physiognomy
and species composition that differ from those of halophytic habitats in eastern Austria,
southern Slovakia and Hungary. Consequently, southern Moravia with the adjacent
Pulkau valley in Lower Austria can be considered a biogeographically distinct region of
halophytic flora and vegetation.
The unique flora of saline habitats has attracted botanists since the beginnings of
floristic research in central Europe. The first literature records of halophytic species in
the study area date back to the 1820s and mostly refer to the now-vanished natural saline
lakes of Kobylí and Čejč (Hochstetter 1825). In the 19th century, some saline habitats
were drained, which triggered a discussion among botanists about the decline of halo-
phytic flora (Krzisch 1859, Bubela 1882, Spitzner 1894). In the 19th century, numerous
halophytic species were reported in regional floras from different parts of southern
Moravia (Rohrer & Mayer 1835, Makowsky 1863, Oborny 1879, 1886, Formánek 1887,
1892) and Lower Austria (Neilreich 1859, Beck 1890–1893, Halácsy 1896). The first
comprehensive botanical study of saline habitats in southern Moravia (Laus 1907) con-
tained descriptions of the flora and plant communities at the most important sites. Šmarda
(1953) described the state of southern Moravian halophytic flora and vegetation in the
mid-20th century, and Vicherek (1962, 1973) described halophytic plant communities
based on systematic vegetation sampling in the early 1960s.
Halophytic flora and vegetation declined dramatically in the 1960s and 1970s.
Vicherek (1962, 1973) was the last to document with relevés the still well-developed
14 Preslia 94: 13–110, 2022
halophytic plant communities at most southern Moravian sites. The decline of halophytic
species after 1950 was mainly related to the cessation of the traditional grazing manage-
ment in the village’s common pastures (Hutweiden), which led to vegetation succession,
and exacerbated by interventions into the natural water regime. Although most botanists
were aware of this process, the policy-driven intensive agriculture of the 1950s to 1980s
did not provide any opportunity to practice management that would conserve the last
remnants of halophytic habitats. The succulent halophytes Salicornia perennans and
Suaeda prostrata went extinct in southern Moravia and Lower Austria in the 1970s and
1980s. The quality of halophytic habitats was very bad at that time and continued to
decline, as described in the last systematic survey of the southern Moravian halophytic
flora by Grulich (1987).
After the political changes of 1989, the Nature Conservation Agency of the Czech
Republic launched funding programmes in 1993 to support and manage nationally valu-
able natural sites. After 2004, these programmes were continued with the EU support.
Management by grazing and mowing was introduced at the remaining saline sites (e.g.
Dedek 2016). These measures have halted the decline in populations of several endan-
gered halophytic species; however, they have been limited to small protected areas. The
general lack of grazing in today’s landscape and increasing drought events do not allow
for the restoration of halophytic habitats at most of their original sites.
In Austria, land use also changed during the 20th century, although less drastically
than in the former Czechoslovakia (Devátý et al. 2019). Currently, the protection of sites
with halophytic vegetation in Lower Austria is mainly ensured by designation as Natura
2000 sites. However, the management plans for these areas are not legally binding
(Geitzenauer et al. 2016).
This article aims to give a comprehensive overview of the past and present state of
halophytic flora and vegetation in southern Moravia and the nearby Pulkau valley in
northern Lower Austria. Our main objectives are (i) to describe the main sites of
halophytic flora and vegetation, (ii) to review the past and present distribution of plant
species with the greatest affinity for saline habitats and (iii) to provide a classification of
halophytic communities based on historical and current data. We also describe the
bryoflora of saline habitats and evaluate the relationship between individual species and
vegetation types and soil salinity.
Methods
Study sites
The study area (Fig. 1) is in southern Moravia (south-eastern Czech Republic) and the
Pulkau valley in the adjacent northern part of Lower Austria (north-eastern Austria). Its
western part is located in the Western Carpathian foredeep (Dyjsko-svratecký úval),
while the eastern part lies in the Lower Morava Valley (Dolnomoravský úval), which is
part of the Vienna Basin, and in the Trkmanka stream valley in the hilly landscape of the
flysch Carpathians. The halophytic habitats are found in depressions filled with Neogene and
Quaternary sediments (Demek & Mackovčin 2006). The last marine regression took place
there in the Miocene, leaving saline sediments. Mineral springs rich in sodium, magne-
sium and sulphates occur in some of these depressions. The mean annual temperature is
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 15
9–10 °C, and the annual precipitation total is 450–550 mm (Tolasz et al. 2007). Precipita-
tion peaks in summer, but long droughts can occur in late spring or summer in some
years.
Systematic descriptions of the state of halophytic flora and vegetation in southern
Moravia in their time were provided by Laus(1907), Šmarda (1953) and Grulich (1987).
Many publications dealt with the flora of individual sites or localities of selected halo-
phytes. In 1959–1965, Vicherek (1962, 1964, 1973) carried out systematic vegetation
survey and recorded relevés at most sites where halophytic vegetation was still preserved.
We use the regional division of saline sites into districts proposed by Grulich (1987)
(Fig. 1; Supplementary Data S1). Each of these districts is a group of nearby sites with
extant or vanished halophytic flora and vegetation located in the same basin or drained by
the same stream. Grulich (1987) defined eight districts named after the nearby village(s)
and, in one case, after a stream (Daníž). He only dealt with halophytes in southern Moravia;
therefore, we added the Pulkau district in a nearby area in Lower Austria. We also added
another district in Moravia (Damnice-Olbramovice district), which Grulich (1987) did not
recognise because of the small number of halophyte records available to him. However,
we found another previously neglected source of halophyte records (Oborny 1912) that
supported the delimitation of this new district. In addition, we re-assigned the halophyte
16 Preslia 94: 13–110, 2022
Fig. 1. Halophytic districts in southern Moravia and northern Lower Austria.
occurrences west, south-west and south of the town of Mikulov from the Hevlín-Nový
Přerov district to the Sedlec district, as this area is drained by the Včelínek stream, which
flows through the Sedlec district.
We reviewed botanical literature from Hochstetter (1825), the first scientific report on
Moravian flora, to the present, plant records in the Pladias database, which contains data
from botanical literature, herbarium specimens and unpublished observations from the
Czech Republic (Wild et al. 2019, Chytrý et al. 2021), the herbarium database JACQ
(jacq.org) and relevés (vegetation-plot records) in the Czech National Phytosociological
Database (Chytrý & Rafajová 2003). Using these data, we prepared descriptions of the
historical status of halophytic flora and vegetation in individual districts and at the most
important sites within these districts, especially where halophytic flora and vegetation
have survived to the present day.
In many cases, the geographic location of some former halophytic sites was uncertain. To
determine such locations, we compared the information from herbarium labels and litera-
ture with the maps of the 2nd and 3rd Military Survey of the Habsburg Monarchy, differ-
ent versions of which are now digitized and publicly available (Anonymous 2001–2017,
2021a, b, Timár et al. 2011). The original topographic sections (scale 1:25,000) of the 3rd
Military Survey were particularly useful as they contain information on land use, distin-
guishing between arable land, meadows and common pastures. To interpret the geo-
graphic location of the relevés recorded by J. Vicherek 1959–1965, which contained
short and sometimes unclear location information, we used the orthophoto maps based on
aerial photographs from the 1950s available on the INSPIRE geoportal (Anonymous
2021b).
We visited all sites with preserved halophytic vegetation on 9–13 August 2020,
described their current state, recorded vascular plant species with particular attention to
halophytes and subhalophytes, and sampled relevés of halophytic vegetation. We sup-
plemented this primary dataset with records from occasional visits to some sites in sum-
mer 2021.
Flora
We reviewed the historical and current distribution of 16 halophytes considered by
Grulich (1987) as species with the strongest affinity for the saline habitats in the study
area (sometimes referred to as “obligate halophytes”), i.e., Bupleurum tenuissimum,
Cirsium brachycephalum,Crypsis aculeata,C. schoenoides,Galatella cana,Glaux
maritima,Plantago maritima subsp. ciliata,Salicornia perennans,Samolus valerandi,
Scorzonera parviflora,Spergularia marina,S. media,Suaeda prostrata,Taraxacum
bessarabicum,Triglochin maritima and Tripolium pannonicum subsp. pannonicum. We
added Juncus gerardii, which was omitted by Grulich (1987) due to numerous identifica-
tion errors in herbaria and literature. We refer to these 17 species as halophytes. There are
also other species associated with saline habitats in Moravia, but they also occur on non-
saline soil; we refer to them as subhalophytes.
Of the 17 halophytes selected for our study, recently published distribution maps and
lists of reviewed records were prepared within the Pladias project for Crypsis aculeata
and C. schoenoides (both Danihelka & Kaplan in Kaplan et al. 2020), Plantago maritima
(Danihelka & Kaplan in Kaplan et al. 2018), Salicornia perennans (Danihelka & Kaplan
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 17
in Kaplan et al. 2017), Spergularia marina and S. media (both Kúr & Ducháček in Kaplan
et al. 2016), Suaeda prostrata (Danihelka & Kaplan in Kaplan et al. 2017) and Tripolium
pannonicum (Danihelka in Kaplan et al. 2017). A list of verified records and a map for
Cirsium brachycephalum and Triglochin maritima were prepared in collaboration with
P. Bureš, those for Scorzonera parviflora with V. Grulich and those for Taraxacum
bessarabicum with J. Štěpánek and J. Zámečník. For the remaining five species, we revised
and recorded the specimens available in the herbaria BRNM, BRNU, GM, HR, MMI,
MZ, OL, OLM, OP, PR, PRC, VYM and ZMT (acronyms follow Thiers 2021). We found
some additional records in these herbaria of the species already covered by the Pladias
project. To improve spatial and temporal coverage, we extracted additional records from the
literature. For most species, we used numerous literature and unpublished field records
available in the Pladias database (www.pladias.cz). However, for J. gerardii and, to
a large extent, for T. bessarabicum,wemainlyreliedontherecordsdocumentedbyher
-
barium specimens. The records of halophytes from the Pulkau district were mainly taken
from the literature and databases, especially from the JACQ database (www.jacq.org).
The herbarium specimens were revised, their labels photographed and stored in a data-
base together with the records extracted from the literature. They were georeferenced
using digital maps from the portal Mapy.cz and various historical maps mentioned above.
We then imported them into the Pladias database using its consistency-checking proce-
dures (Wild et al. 2019). The records were further managed in the Pladias database.
In several cases, we used information from the literature for herbarium specimens and
information from herbarium labels for records extracted from the literature to make
a record more precise in terms of time and location. For example, it is known from vari-
ous sources that Ch. F. Hochstteter lived in Brno in 1817–1824 and that the Měnín Fish-
pond (Mönitzer See) was drained in 1822. Therefore, his undated specimens can be dated
to 1817–1824 and those collected on the shores of Měnín Fishpond to 1817–1821. Simi-
larly, we assigned all records from the 1950s onwards of Salicornia perennans and
Suaeda prostrata from the vicinity of the village of Terezín in the Čejč district to the
Zápověď site, as at that time, it was demonstrably the only site in the area where these two
succulent halophytes still occurred.
In total, we compiled ~3433 records of halophytes from the nine southern Moravian
districts (all stored in the Pladias database) and ~136 records from the Pulkau district in
Lower Austria (Supplementary material Data S2). Of the 3433 records, ~1946 records
were already available in the Pladiasdatabase. We added ~1487 records, of which ~1321
were supported by herbarium specimens. We collected most of the relevant plant records,
both from herbarium collections and the literature, that have accumulated since the
beginning of botanical research in the area in the early 19th century. Only for the Pulkau
district, some herbarium records may be lacking, but it is still the most representative
dataset on halophyte flora in the area. The recently added specimens from local collec-
tions (e.g. VYM and ZMT) contributed only a few unique records. Based on this experi-
ence, we believe that the spatial and temporal pattern of halophyte distribution described
here will not change significantly, even if more herbarium specimens and literature
records are found in the future.
For all the species and districts, we created graphs showing records of all the 17
halophyte species within each district on the timeline.
18 Preslia 94: 13–110, 2022
The taxonomy and names of vascular plants follow Kaplan et al. (2019) or, for those
not occurring in the Czech Republic, Fischer et al. (2008). The Red List categories given
in the Species chapter follow the latest Red Lists for the Czech Republic (Grulich 2017)
and Austria (Niklfeld 1999).
In addition to vascular plants, we examined the bryoflora of saline habitats. We
reviewed the literature from the 20th century and the current literature on the distribution
of bryophytes in southern Moravia. Taxonomic concepts and nomenclature follow
Kučera et al. (2012).
Vegetation
We extracted all relevés from southern Moravia from the Czech National Phytosocio-
logical Database (CNPD; Chytrý & Rafajová 2003) that contained at least one of the
above 17 halophytes or subhalophytes Lotus tenuis or Trifolium fragiferum, and added all
the other relevés from publications in which at least one relevé contained at least one of
the 17 selected species. We then searched in unpublished sources not digitized in the
CNPD for other relevés of halophytic vegetation and added them to this dataset. These
newly added relevés came mainly from the unpublished manuscripts of J. Vicherek from
the early 1960s and from unpublished reports of protected area surveys conducted in the
last two decades.
During our visit at individual sites on 9–13 August 2020, we sampled at least one
relevé in a relatively homogeneous area of each visually distinguishable type of halo-
phytic vegetation. All the relevés had a standard size of 10 m2. Most of them were squares
of 3.16 × 3.16 m, but if there was no homogeneous vegetation in a square area, we used
a rectangular plot of 10 m2. We measured the geographic coordinates of all four corners
of each plot using a GPS (Topcon HiPer SR with Getac PS336 Data Collector) with
a positioning accuracy of ~5 cm. In each relevé, we recorded all species of vascular plants
and bryophytes and estimated their cover using the nine-grade Braun-Blanquet scale
(Westhoff & van der Maarel 1978). In total, we sampled 67 relevés. This dataset is avail-
able in the Zenodo repository (Danihelka et al. 2021).
For vegetation classification, we used a selection of relevés from the CNPD, added the
relevés newly digitized from the literature, and the new relevés sampled in the field. This
dataset contained 729 relevés. All the newly digitized and newly sampled relevés were
deposited in the CNPD. We corrected errors in species identifications that we had discov-
ered based on our field experience and examination of herbarium specimens. We also
replaced broad species concepts used in the older literature with currently established
concepts of narrower taxa that occur at saline sites. For example, records of Carex
vulpina and Festuca valesiaca at saline sites were changed to C. otrubae and F. pulchra,
respectively. Groups of closely related species often misidentified by botanists were
combined into aggregates. For example, because the herbarium collections showed
a high proportion of Juncus compressus specimens from saline sites to be misidentified
as J. gerardii, we had to combine these two species into the J. compressus agg.
We classified this relevé dataset using the expert system CzechVeg-ESy, developed
for the national vegetation classification of the Czech Republic (Chytrý 2007, 2011). We
replaced the original definitions of marsh associations, as available in version 1-2020-01-
12 of the expert system (Chytrý et al. 2020), with the definitions developed by Landucci
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 19
et al. (2020). The classification was performed using the Juice 7.1 program (Tichý 2002).
We carefully reviewed the classification of each relevé by the expert system and investi-
gated why some relevés were misclassified, not classified at all, or assigned to more than
one association. Based on this experience, we improved the expert system by refining the
logical definitions of associations of specialized halophytic vegetation. For some refined
definitions, we used new features implemented in the expert system language for vegeta-
tion classification (ESy) described by Tichý et al. (2019). This refinement increased the
number of correctly classified relevés.
It was not our aim to define new associations that are not recognized in Vegetation of the
Czech Republic (Šumberová 2007a, b, Šumberová et al. 2007, 2011). However, we
noticed that our data from 2020 and some newly obtained datasets contain several relevés
corresponding to the association Centaureo pannonicae-Festucetum pseudovinae, described
from Slovakia and not recognized in the national vegetation classification of the Czech
Republic (Šumberová et al. 2007). Therefore, we added this association to the expert system.
Most halophytic associations are species-poor vegetation types defined by the domi-
nance of a single highly specialized species. However, this is not the case for the halo-
phytic grasslands of the class Festuco-Puccinellietea. To test whether the associations of
this class defined by the expert system reflect real patterns of species composition, we
compared the expert-system-based classification of this class with an unsupervised divisive
classification (Twinspan;Hill 1979). We applied Twinspan to therelevés assigned by the
expert system to any association of the class Festuco-Puccinellietea. We used the R pack-
age Twinspan (Oksanen 2019) with pseudospeciescut levels corresponding to 0, 2, 5, 10
and 20% cover. To avoid classification bias due to a disproportionate number of similar
relevés from some sites, we performed a stratified resampling of the dataset. We took
only two randomly selected relevés from each group of relevés assigned to the same asso-
ciation and sampled by the same researcher at the same site and in the same year. This
stratification reduced the number of Festuco-Puccinellietea relevés from 293 to 166.
These relevés were classified by Twinspan. Because Twinspan tends to divide groups into
subgroups of similar size, vegetation types represented by a few relevés may not be rec-
ognized at higher division levels. Therefore, we interpreted groups at different hierarchi-
cal levels and accepted those that were most similar to the accepted associations or corre-
sponded to ecologically defined subgroups within the associations. We then compared
the associations defined by the expert system with a set of accepted Twinspan groups
using alluvial diagrams created with the R library ggaluvial v. 0.12.3 (Brunson 2020).
From the classified dataset of 729 relevés, we selected 426 relevés assigned by the
expert system to the associations of specialized halophytic vegetation. We considered the
associations that were either defined by a dominance of one halophytic species or contained
multiple halophytic species. We applied the same stratified resampling as described above,
resulting in 246 relevés. We used this resampled dataset to create a synoptic table and cal-
culate the diagnostic, constant and dominant species of each association.
All the halophytic associations of the classes Bidentetea,Phragmito-Magno-Caricetea,
Crypsietea aculeatae and Thero-Salicornietea strictae were defined through the domi-
nance of a single halophytic species. Consequently, their classification was straightfor-
ward within the concept of dominance-based associations and did not require further test-
ing. However, the classification of halophytic grasslands of the class Festuco-
Puccinellietea was based mainly on the combination of the occurrence of sociological
20 Preslia 94: 13–110, 2022
species groups. Therefore, we tested whether this classification reflects the patterns of
variation in total species composition using two methods. First, we calculated the
Twinspan classification of the stratified subset of 166 relevés assigned to the associations
of the Festuco-Puccinellietea class and compared the Twinspan clusters with these asso-
ciations. Second, we prepared a DCA ordination (Hill & Gauch 1980, calculated using
the vegan v. 2.5-7 library in R; Oksanen et al. 2020) of the same relevés and plotted the
association assignment of each relevé by the expert system onto the ordination diagram.
Diagnostic species were defined based on the frequency of their presence/absence
within and outside the groups of relevés belonging to different associations. To be con-
sidered diagnostic, a species had to meet four conditions simultaneously: (i) to have
a value of the phi coefficient of association with the group of relevés greater than 0.25, (ii)
to have a probability of observed occurrence concentration within that group based on
Fisher’s exact test of less than 0.05, (iii) to have the greatest frequency in the relevés of
the focal association with a constancy ratio greater than 0.25, and (iv) to have more than
one occurrence in the dataset. The phi coefficient was calculated for the virtually equal-
ized sizes of relevé groups according to Tichý & Chytrý (2006). The constancy ratio is
defined as the ratio between the percentage species frequency within the focal group in
which frequency is highest and the next highest frequency in any other group (Dengler
2003). If a species had a phi coefficient greater than 0.50, it was considered highly
diagnostic.
Constant species were defined as those occurring in more than 40% of the relevés
assigned to the focal association. Species occurring in more than 80% of the relevés were
considered highly constant.
Dominant species were defined as those occurring in more than 20% of relevés with
a cover greater than 25% or a lower cover that is greatest in the relevé. If they fulfilled
these conditions in more than 50% of the relevés,they were considered highly dominant.
Relationships to salinity
We collected soil samples from a depth of 2–10 cm in 3–4 places within each relevé sam-
pled in August 2020. We mixed these samples for each relevé and measured the pH and
electrical conductivity of the mixed sample in a suspension with distilled water
(soil:water ratio ~ 2:5) using the Hach HQ40D multimeter (Hach, Loveland, CO, US). To
evaluate the relationships between individual plant species and salinity, we related the
presence/absence data of all species to measured soil pH and conductivity using a gener-
alized linear model with binomial distribution and logit link function. We used the glm()
function from the R-core package stats (R Core Team 2020). We used McFadden’s
pseudo-R2(McFadden 1974) to quantify the proportion of variation explained by the
model. For species with significant (p < 0.05) or marginally significant (p < 0.1) models, we
drew the plots of species occurrence probability using the packages ggplot2 (Wickham
2016), ggtext (Wilke 2020), gtools (Warnes et al. 2020) and gridExtra (Auguie 2017).
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 21
Sites
Měnín-Šaratice district
This district lies between the villages of Blučina, Otmarov, Sokolnice, Šaratice, Otnice,
Těšany and Moutnice in a large and shallow depression of the Litava, Dunávka and Říčka
streams (Grulich 1987). This is the northernmost part of southern Moravia where
halophytes occurred naturally. It includes the area of the former large Měnín Fishpond
(Mönitzer Teich) with some smaller fishponds and mineral water springs between the vil-
lages of Nesvačilka (Neudorf) and Šaratice (Scharatitz).
The former Měnín Fishpond was between the villages of Měnín, Blučina and
Moutnice. Although mostly referred to as a lake in the literature, it was an artificial pond
created in 1492 (Hurt 1960). Soils on its shores were rich in salt, especially calcium and
magnesium sulphate (Laus 1907). The fishpond was drained in 1822 and converted
mainly into arable land (Hurt 1960). The nearby smaller fishponds werealso drained dur-
ing the 19th century, and most of the natural springs were adjusted for collecting water
using a system of tubes and wells during the 20th century (Laus 1907, Grulich 1987).
In the past, this district harboured all the 17 halophytes except Samolus valerandi and
Triglochin maritima. The past occurrence of Glaux maritima is uncertain: the single non-
dated herbarium specimen allegedly collected near Měnín may be mislabelled.
The only records of the halophytic flora at the Měnín Fishpond before its drainage are
from Hochstetter (1825). He explored the area likely between 1817–1821 because his
1825 article indicates he was unaware of the fishpond drainage. He observed vegetation with
Cirsium brachycephalum (Cnicus polyanthemus), Crypsis aculeata,Plantago maritima,
Salicornia perennans,Scorzonera parviflora and Suaeda prostrata on the fishpond
shores. He also collected Galatella cana in a nearby grove and Taraxacum bessarabicum
(Leontodon lividus) in an adjacent wet meadow. He further collected Bupleurum tenuissi-
mum,Scorzonera parviflora,Spergularia media (misidentified as S. salina)andTripolium
pannonicum, but these records were published only later by Rohrer & Mayer (1835).
Makowsky (1856) found Crypsis aculeata and C. schoenoides in saline habitats near
the villages of Měnín, Moutnice and Otmarov and confirmed the occurrence of Sali-
cornia perennans,Suaeda prostrata and Tripolium pannonicum. He returned to the area
several times and collected most of the halophytes once recorded by Hochstetter. How-
ever, the occurrences of Cirsium brachycephalum and Galatella cana have not been con-
firmed since Hochstetter’s times (for the latter, see Danihelka 2008). Scorzonera parvi-
flora was last recorded in 1900. The halophytic sites near Otmarov were repeatedly vis-
ited by Brno botanists from the late 1860s because of their easy accessibility via railway,
and numerous herbarium specimens have been preserved. However, already Laus (1907:
14), who visited the area in 1894, reported that the area had been cultivated, and the
occurrences of halophytes reduced to a few spots, mainly ditches and road verges in vil-
lages’ common pastures. He considered the surroundings of Vladimirov farmstead near
Otmarov, where he collected Crypsis aculeata, as the best-preserved halophytic site.
Halophytic flora gradually disappeared near Otmarov, although some halophytes, e.g.
Bupleurum tenuissimum and Plantago maritima, survived there until the 1930s.
Krist (1935) referred to the site near the village of Šaratice as almost destroyed,
harbouring only Plantago maritima and Tripolium pannonicum. Šmarda (1953) de-
scribed, partly based on information from the algologistJ. Bílý Jr., remnants of halophytic
22 Preslia 94: 13–110, 2022
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 23
Fig. 2. Halophytic localities in the Měnín-Šaratice district: (A) Kalužiny 2020, (B) Zřídla u Nesvačilky 2020,
(C) Císařská obora 2020. Photo credits: K. Chytrý (A), M. Chytrý (B), E. Šmerdová (C).
vegetation at several sites within the district but reported few halophytic species. Frag-
ments of halophytic vegetation were still preserved along the Dunávka stream west of
Sokolnice between the railway and the road. In 1950, J. Šmarda recorded there
Bupleurum tenuissimum,Juncus gerardii,Plantago maritima and Spergularia media.
Salicornia perennans, collected there in 1912, and Suaeda prostrata, found in 1912 and
1922, were no longer present. The last specimens of Salicornia perennans were collected
by S. Staněk and J. Suza around mineral springs near the villages of Nesvačilka and
Šaratice in the early 1920s, and the last specimen of Suaeda prostrata was collected by J.
Šourek near Nesvačilka in 1946.
Vicherek (1973) was still able to record relevés of halophytic vegetation in a shallow
depression between Moutnice and Nesvačilka, probably near the springs of the Nesva-
čilka stream. Between 1962 and 1964, he found there Crypsis aculeata,C. schoenoides,
Spergularia marina,S. media,Taraxacum bessarabicum and Tripolium pannonicum.
This site may be identical with the place recorded on herbarium labels as Luža, where
E. Vítek collected Crypsis aculeata,Salicornia perennans and Suaeda prostrata in 1912.
These last spots were destroyed by drainage probably in the 1960s. In 1982, only
Plantago maritima was known to survive in the district, occurring at two sites. However,
targeted searches after 2000 also yielded a few records of Bupleurum tenuissimum and
Spergularia marina. Nowadays, strongly impoverished fragments of halophytic vegeta-
tion remain mainly at the three small sites described below.
Kalužiny (Fig. 2A). – This site is located ~1.75 km S of the village of Šaratice. In old her-
barium labels, it is referred to as “Šaratica”, “prameny Šaratice”, “prameny hořké vody”
or “gradovna”. Currently, it is covered mainly with meadows dominated by Bromus erec-
tus in drier places and meadows with Festuca arundinacea,Inula britannica and
Serratula tinctoria in wetter places. A meadow with Medicago sativa and Festuca
pratensis was sown along the eastern edge of the site. These meadows are regularly
mown, and the wet places are disturbed by heavy machinery. Only a few small wet
depressions still contain halophytes and subhalophytes, such as Plantago maritima and
Lotus maritimus. The site is protected as a natural water source.
Zřídla u Nesvačilky (Fig. 2B). – The Natura 2000 site Zřídla u Nesvačilky, also protected
as a natural water source, consists of two non-contiguous sites. The larger one, in old her-
barium labels usually referred to as “Wolfovy studánky” or “prameny Šternovky”, is
located ~2 km NE of the village of Nesvačilka. Currently, there are some wet places
between the wells in a fenced area. In 2006, a few individuals of Lactuca saligna were
recorded there by J. Podhorný and J. Komárek (Grulich in Hadinec & Lustyk 2007). In
2020, we observed Plantago maritima in one place outside the fenced area as the last spe-
cies of the 17 halophytes studied here. There were patches of Festuca pulchra grassland
(association Centaureo pannonicae-Festucetum pseudovinae). However, the dominant
vegetation type in this area is dry to mesic non-halophytic grassland. The smaller site is
located ~2 km NE of Nesvačilka. It is covered by a grassland composed of generalistspe-
cies with no halophytes. Both sites are regularly cut and mulched. Although the most
appropriate management for halophytes is grazing with soil surface disturbances, it
would be at odds with using the site for collecting mineral water (Pekárová 2015).
24 Preslia 94: 13–110, 2022
Císařská obora (Fig. 2C). – The Císařská obora (Emperor’s Game Preserve) site is
located ~5 km S of the village of Měnín, south of the former Měnín Fishpond. It is a part
of the Natura 2000 site Rumunská bažantnice. Not much is known about its past. Accord-
ing to the 2nd Military Survey maps (in Moravia dating to 1836–1840), the site was cov-
ered mainly by meadows. The 3rd Military Survey maps (1876–1878) showed it covered
by a forest (Galdhof-Wald), with a small patch of arable land in the middle. The aerial
photographs from the 1950s are difficult to interpret, but the area was mainly covered
by meadows and (possibly) clearings. Currently, the site is located inside a fenced game
preserve. There are ditches and small fishponds with islands created in the late 1990s
(Juřica et al. 2015). Phragmites australis stands occur on fishpond shores and stands
of Bolboschoenus maritimus,B. planiculmis,Carex distans and C. otrubae near ditches.
Islands are covered by unmown ruderal grasslands dominated by Calamagrostis epigejos
and Agrostis gigantea with tall herbs Althaea officinalis,Cirsium canum and Dipsacus
fullonum. Other grasslands in the area are mown. Subhalophytes such as Centaurium
pulchellum,Lotus maritimus,L. tenuis,Melilotus dentatus and Teucrium scordium are
scattered in the area, but we found none of the 17 halophytes studied here.
Velké Němčice district
This district lies in a shallow depression along the Starovický stream between the villages
of Velké Němčice (Gross Niemtschitz), Uherčice (Auerschitz) and Starovice (Steurowitz).
In the north, it borders the Měnín-Šaratice district.
In the past, this district supported all the 17 halophytes except Cirsium brachycepha-
lum,Galatella cana,Samolus valerandi and Triglochin maritima. The past occurrence of
Glaux maritima is uncertain: there are two records from relevés recorded by J. Vicherek
(Vicherek 1962), but no herbarium specimen has been found. There are also historical
records of Triglochin maritima (e.g. Vicherek 1962), but we consider them erroneous
because the only herbarium specimen reported for this district (cf. Bureš 2010) corre-
sponds to T. palustris.
The occurrences of halophytes in this district were discovered by botanists as late as
1921. Fietz et al. (1923) described patches of halophytic vegetation developed around
a small fishpond (Hofteich) at the southern edge of Velké Němčice, now filled with earth
and built up. At this site, they recorded Bupleurum tenuissimum,Crypsis schoenoides,
Plantago maritima,Scorzonera parviflora,Spergularia marina and Tripolium pannoni-
cum. They found the best-developed patches of halophytic vegetation ~1.3 km SSE of the
town. This “Niemtschitzer Salzsteppe” (later referred to as Plácky or Brodač) was a small
remnant of the former extensive saline meadows and pastures in the shallow valleys of
the Starovický and Křepický streams. It had an approximately triangular shape (like the
current Plácky Nature Monument) and an estimated area of 5 ha. Fietz et al. (1923)
described various vegetation types, including patches dominated by annual species of
Atriplex,Juncus gerardii,Tripolium pannonicum and Festuca (probably F. pulchra)and
patches co-dominated by Salicornia perennans and Suaeda prostrata. In addition, they
noted the occurrence of Plantago maritima,Scorzonera parviflora,Spergularia marina
and S. media. Some halophytes, including Salicornia perennans and Suaeda prostrata,
were also abundant on a dirt road close to the site. Non-succulent halophytes occurred in
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 25
a broader area because Fietz et al. (1923) reported patches of halophytic vegetation also
east of the road Velké Němčice–Hustopeče.
The Plácky site was frequently visited by botanists in the 1920s and 1930s, as docu-
mented by numerous herbarium specimens. Bupleurum tenuissimum,Crypsis aculeata,
C. schoenoides,Salicornia perennans,Scorzonera parviflora,Spergularia marina,
Suaeda prostrata and Tripolium pannonicum were collected at the Plácky site in large
quantities for the exsiccate series Herbarium florae Reipublicae bohemicae slovenicae
and distributed worldwide. However, Krist (1935) reported that the halophytic site near
Velké Němčice had been “reduced”, and the surrounding wetlands drained by deep
ditches. Consequently, the site remained dry throughout the year.
From the 1940s onwards, the Plácky site was repeatedly surveyed by V. Vlach,whose
unpublished records were summarized by Husák & Jatiová (1984). In 1942, Vlach
recorded Crypsis aculeata,Juncus gerardii,Salicornia perennans,Spergularia media,
Taraxacum bessarabicum and Tripolium pannonicum. According to his records, local
conditions for halophytic flora, grazed by livestock, were optimal until 1945. In that year,
a weir on the Svratka river was removed, and the drought of 1947 caused a significant
decline in groundwater level. Consequently, halophytes started to decline, and ruderal
species increased. In 1950, Vlach failed to confirm the occurrence of Crypsis aculeata
and Taraxacum bessarabicum, while Salicornia perennans was only represented by
a few individuals. Jan Šmarda studied this site in 1950, recording Bupleurum tenuissi-
mum,Plantago maritima,Salicornia perennans,Suaeda prostrata and Taraxacum
bessarabicum (Šmarda 1953). He considered Plácky as one of the most valuable
halophytic sites in Moravia and recommended its strict protection. Although the site was
put under conservation in 1950, the decline of halophytic vegetation continued. Between
1958 and 1965, the area south of Velké Němčice was drained by subsurface pipe drain-
age, and a system of drainage ditches was established around the perimeter of the Plácky
site a few years later. Still, J. Vichererek was able to record relevés of vegetation domi-
nated by Salicornia perennans and Suaeda prostrata between 1959 and 1965 (Vicherek
1973). This was the last record of the former, while for Suaeda prostrata,thereisaher
-
barium specimen collected by F. Weber allegedly in 1973. In the mid-1970s, only
Plantago maritima,Spergularia media,Scorzonera parviflora and Tripolium pannoni-
cum were observed at the site (Husák & Jatiová 1984). A visit to the site in 2001 by
J. Danihelka yielded no records of halophytes. Conservation management was restored in
2005 when regular mowing was introduced. Since 2017, the site has been grazed by
horses (M. Hájek, pers. comm.). The current halophytic flora of Plácky is extremely
impoverished. J. Danihelka & K. Fajmon collected Spergularia marina next to the south-
western edge of the reserve in 2007, and P. Novák observed a small population of
Bupleurum tenuissimum and probably also Juncus gerardii in 2012. In addition,
Chenopodium chenopodioides was repeatedly recorded in the southern part of the reserve.
Currently, there are no well-developed halophytic plant communities there (Fig. 3A).
Some parts of the reserve, especially its north-western corner, are dry and covered by
subruderal grasslands dominated by Carex praecox,Festuca rupicola and Poa angusti-
folia. Wet depressions are overgrown by Carex riparia.
26 Preslia 94: 13–110, 2022
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 27
Fig. 3. Halophytic localities in the Velké Němčice, Šakvice and Rakvice districts (A) Plácky 2020, (B) Štinkovka
2021, (C) Trkmanec-Rybníčky 2020. Photo credits: K. Chytrý (A), M. Chytrý (B), E. Šmerdová (C).
Starovický Fishpond, the second major halophytic site of the Velké Němčice district,
was located at the northern edge of the village of Starovice northeast of Hustopeče. The
3rd Military Survey maps (1876–1878) show two small fishponds near the northern edge
of the village, surrounded by a common pasture. Aerial photographs from the 1950s show
a similar pattern, with a football playground replacing pasture at the south-eastern corner
of the area. The halophytes occurred around the fishponds and possibly also in a pasture
land strip along the Starovický stream up to 1 km towards the north-west. Probably in the
late 1960s or during the 1970s, the southern part of the halophytic site was built up, and
its northern part was destroyed by enlarging the fishpond towards the village. The west-
ern half of the former site (west of the Starovický stream) is now completely drained and
partly used as a tennis playground, while most of the remaining area is covered by ruderal
grassland (in places with Carex otrubae). The fishpond littoral is largely overgrown by
Phragmites australis.
The occurrences of halophytes near Starovice were discovered by botanists in the
early 1920s. At this site or nearby, they collected Bupleurum tenuissimum,Crypsis
schoenoides,Juncus gerardii,Plantago maritima,Spergularia marina,S. media,Suaeda
prostrata,Taraxacum bessarabicum and Tripolium pannonicum. The first description of
the Starovice site was published by Šmarda (1953), who in 1951 recorded Bupleurum
tenuissimum,Juncus gerardii,Plantago maritima,Scorzonera parviflora,Spergularia
marina,S. media,Suaeda prostrata,Taraxacum bessarabicum and Tripolium pannonicum.
Vicherek (1962, 1973), who studied the Starovice site in 1959–1965, confirmed the
occurrence of Bupleurum tenuissimum,Glaux maritima,Plantago maritima,Salicornia
perennans,Spergularia marina,S. media,Suaeda prostrata,Taraxacum bessarabicum
and Tripolium pannonicum. He was still able to record vegetation dominated by both suc-
culent halophytes. However, the halophytic habitats were destroyed soon after his survey.
The last herbarium specimen of Suaeda prostrata is from 1968. Still, a few halophytes
survived until the 1970s: Tripolium pannonicum was last collected in 1977 and Plantago
maritima in 1980.
In 1961, F. Kühn studied weed flora of patches with saline soils on arable land in the
surroundings of Velké Němčice (Kühn 1963). He found Spergularia marina at about ten
sites, at some of them accompanied by Crypsis schoenoides (in Table 1 of his article
reported as Heleochloa alopecuroides+schoenoides). Of the formerly rich halophytic
flora in the Velké Němčice district, only Bupleurum tenuissimum,Spergularia marina
and probably also Juncus gerardii have survived until recently, all in tiny populations.
Some subhalophytes still occur in the floodplain of the Starovický stream ~1 km N to
1.5 km NNW of Starovice. Shallow pools were created there in 2013–2015 as a part of
a restoration project. In 2021, the edges of the Phragmites australis stands fringing the
pools contained several subhalophytes such as Bolboschoenus maritimus,B. planiculmis,
Carex otrubae,C. secalina,Centaurium pulchellum,Lotus tenuis,Melilotus dentatus,
Puccinellia distans and Trifolium fragiferum (Chytrý & Danihelka 2021). However, the
site is relatively dry and unmown, which does not provide good conditions for
halophytes. None of the 17 halophytes studied in this article was found there.
28 Preslia 94: 13–110, 2022
Šakvice district
This district encompasses the shallow depressions of the Popický and Štinkovka streams
between the village of Pouzdřany (Pausram) in the west, the town of Hustopeče (Auspitz)
in the north and the village of Starovičky (Klein-Steurowitz) in the east. Some halophytes
also occurred in narrow stream valleys north of Popice (Poppitz) and Hustopeče.
The flora of the district included Bupleurum tenuissimum,Crypsis aculeata,C. schoe-
noides,Glaux maritima,Juncus gerardii,Plantago maritima,Salicornia perennans,
Samolus valerandi,Scorzonera parviflora,Spergularia marina,S. media,Suaeda prostrata,
Taraxacum bessarabicum,Triglochin maritima and Tripolium pannonicum. It is likely
that Cirsium brachycephalum also occurred in this district, but no herbarium specimen
has been found.
The most valuable and well-known halophytic site was around the Šakvice railway
station (formerly called Hustopeče station, Bahnhof Auspitz or Nordbahn-Station
Auspitz). It was discovered in the 1860s by J. Wessely, then a parish priest in Hustopeče.
Because of its location on the railway from Vienna to Brno (opened in 1839), botanists
frequently visited this site, and its flora was documented by numerous herbarium speci-
mens. All the species listed in the previous paragraph except Cirsium brachycephalum
occurred there. Triglochin maritima was discovered there as a new species of the
Moravian flora in 1900. There is also a herbarium specimen of Samolus valerandi alleg-
edly collected from this site, but it may be a labelling mistake.
Laus (1907) was the first to describe the halophytic flora around the Šakvice railway
station in detail. At that time, halophytes occurred on both sides of the railway, but the
best-developed halophytic vegetation was found in the place where the local railway to
Hustopeče branched off. In this narrow triangle, Laus observed stands dominated by
Salicornia perennans andprobablyalsobySuaeda prostrata.
Laus (1907) already reported that one part of the saline habitats wasdestroyed in 1894
when the local railway to Hustopeče was built. Another part disappeared when the second
railway track and a flyover were built in the early 1920s (Krist 1935). Triglochin
maritima was last collected in 1923. As reported by J. Bílý (sec. Šmarda 1953), it once
occurred in a small saline meadow north-west of the station. The last specimen of Suaeda
prostrata was collected in 1927 and that of Salicornia perennans in 1931. Krist (1935)
recorded Plantago maritima and Tripolium pannonicum but no other halophytes. Šmarda
(1953) only found Juncus gerardii (last collected in 1952) and Plantago maritima,and
concluded that the once-famous halophytic site near the Šakvice station “belongs to the
past age”. Indeed, these were the last records of halophytes from this site.
Another halophytic site occurred ~1.8 km W of the village of Starovičky on both
banks of the Štinkovka stream. The 2nd Military Survey maps (1836–1840) show there
meadows on the bottom of a drained fishpond (still referred to as a fishpond: Un.
Steurowitzer Teich). Halophytes occurred in these wet saline meadows and along the
Starovičský stream roughly up to the village centre (Šmarda 1953), where Glaux
maritima was recorded in 1952. The flora of this site was rich in halophytes, including
Bupleurum tenuissimum,Crypsis schoenoides,Glaux maritima,Plantago maritima,
Samolus valerandi,Scorzonera parviflora,Spergularia marina,S. media and Tripolium
pannonicum. Šmarda explored the site in 1952 and found Plantago maritima,Scorzonera
parviflora,Taraxacum bessarabicum and Tripolium pannonicum (Šmarda 1953). Also in
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 29
1952, V. Skřivánek (sec. Šmarda 1953) found Scorzonera parviflora and Taraxacum bes-
sarabicum somewhere between this site and the village. Jiří Vicherek still sampled saline
grasslands of the association Scorzonero parviflorae-Juncetum gerardii with Plantago
maritima,Scorzonera parviflora and Tripolium pannonicum in 1962 or 1963 (Vicherek
1973). Plantago maritima and Tripolium pannonicum were last collected in 1971, while
Juncus gerardii was still recorded there in 2021 (Chytrý & Danihelka 2021).
A targeted search in summer 2021 along the Štinkovka stream between the south-east-
ern edge of Hustopeče and a dam of the Lower Nové Mlýny reservoir resulted in the dis-
covery of halophytic habitats on a temporarily abandoned wet arable land and ex-arable
land converted to semi-natural areas as a part of restoration projects (Chytrý & Danihelka
2021). The richest halophytic and subhalophytic flora, including whole halophytic plant
communities, was found along the Štinkovka between Šakvice and Starovičky. Of the
halophytes, Crypsis schoenoides,Juncus gerardii,Samolus valerandi and Spergularia
marina were recorded, each species at two sites.
In the past, some halophytes also occurred further upstream along the Štinkovka at the
north-western outskirts of Hustopeče. The site was referred to as an ice-skating rink
(Eislaufplatz) and later a swimming school (Schwimmschule) on herbarium labels.
Samolus valerandi was collected there repeatedly from 1897 and last observed in 1954 or
1955 (J. Tománek sec. Sedláček & Dvořák 1983). It co-occurred with Plantago maritima,
Taraxacum bessarabicum and Tripolium pannonicum. Šmarda (1953) considered the site
destroyed. Today, there is an open-air swimming pool and a swimming hall. In 2018,
S. valerandi was found (together with a few subhalophytes) on the drained bottom of the
Přední Fishpond ~2 km north of its former site in Hustopeče (Ambrozek & Melichar in
Lustyk & Doležal 2019).
Further well-developed patches of halophytic vegetation once existed in a shallow
depression of the Popický stream near the road ~1.5 km SE of the village of Popice. The
3rd Military Survey map (1876–1878) shows strips of grasslands along the Popický
stream and its left-hand tributary. This site most likely hosted Suaeda prostrata, collected
there or nearby in 1894–1913, as well as Crypsis aculeata,Juncus gerardii,Plantago
maritima,Scorzonera parviflora,Spergularia marina and Tripolium pannonicum.The
record of Cirsium brachycephalum (Šmarda 1953) may also refer to this site. The last
record of Tripolium pannonicum dates back to 1930. In 1984, V. Grulich and R. Řepka
collected Plantago maritima and Spergularia marina between this site and the railway.
However, the site was destroyed in the same year by reclamation (Grulich 1987).
The occurrences of halophytes along the Popický stream above the north-western
edge of Popice were discovered probably by Šmarda (1953), who recorded only Juncus
gerardii. Local vegetation was described by Vicherek (1962). His relevés include
Cirsium brachycephalum,Juncus gerardii,Plantago maritima,Scorzonera parviflora,
Taraxacum bessarabicum and Tripolium pannonicum. These saline habitats were proba-
bly destroyed in the 1960s, as suggested by the absence of newer records of halophytes.
In addition, Šmarda (1953) found Juncus gerardii,Plantago maritima,Spergularia
marina and Tripolium pannonicum in grazed places along the stream in the village and on
the local playground.
Halophytic vegetation also occurred further westwards between the villages of Popice
and Pouzdřany, probably ~1.5 km WNW of Popice, but the exact location is unknown.
Laus (1907) recorded there Bupleurum tenuissimum,Plantago maritima,Spergularia
30 Preslia 94: 13–110, 2022
marina,S. media,Suaeda prostrata and Tripolium pannonicum. Šmarda (1953) observed
Plantago maritima,Spergularia marina and Tripolium pannonicum somewhere in that
area in 1952. Single records of halophytes also exist from other places in the district, e.g.
from the eastern shores of the former Šakvický Fishpond south-east of the village of
Šakvice, now flooded by the Lower Nové Mlýny reservoir.
The first halophyte species to disappear from the Šakvice district was Triglochin
maritima in the 1920s, while both succulent halophytes vanished in the late 1920s or
early 1930s. Plantago maritima was last observed in the district in the mid-1980s. Nowa-
days, Crypsis schoenoides,Juncus gerardii,Samolus valerandi and Spergularia marina
occur in the district, except for J. gerardii in large populations.
Restored halophytic sites along the Štinkovka stream near Šakvice (Fig.3B).–Since
2017, the village of Šakvice has implemented a series of ecological restoration projects
aimed at converting the wet arable land along the Štinkovka stream into areas with semi-
natural vegetation. They created a pond and severalshallow pools, planted forests, sowed
grasslands and left some areas on the former arable land without management. After rains,
some places in these areas are shallowly flooded. In summer 2021, several halophytic
species and plant communities were found in these areas (Chytrý & Danihelka 2021).
One site occurs on the right bank of the Štinkovka between the road Šakvice–Starovičky
and the railway Brno–Břeclav ~2.2–2.8 km ESE of Šakvice. There were large popula-
tions of Crypsis schoenoides and Samolus valerandi in the stands of Bolboschoenus
maritimus and B. planiculmis (association Tripleurospermo inodori-Bolboschoenetum
planiculmis). Several species of subhalophytes also occurred in the stands of Carex
secalina (Agrostio stoloniferae-Juncetum ranarii) and ephemeral wetlands on wet ex-
arable land (Veronico anagalloidis-Lythretum hyssopifoliae). Another site, restored in
2018, occurred near a new fishpond on both sides of Štinkovka 1.5–1.9 km E of Šakvice.
Also here, abandoned wet arable land was overgrown by Bolboschoenus planiculmis
stands containing large populations of Crypsis schoenoides,Samolus valerandi and
Spergularia marina. In places, there were patches with salt efflorescences on the soil sur-
face and species-poor vegetation with Crypsis schoenoides, corresponding to the associa-
tion Heleochloëtum schoenoidis. The deposit of the substrate removed from the pond and
pools were covered by extensive stands of Carex secalina (Agrostio stoloniferae-
Juncetum ranarii) and in trampled places by open grasslands dominated by Puccinellia
distans and Spergularia marina (association Puccinellietum limosae). Narrow reed
stands dominated by Schoenoplectus tabernaemontani developed along the banks of the
new fishpond. Large populations of Samolus valerandi and Spergularia marina were
also found in wet maize fields on the right bank of the Štinkovka adjacent to the north-
eastern edge of this restored site.
Rakvice district
The Rakvice district lies in a shallow depression along the lower Trkmanka stream and its
left-hand tributary Bílovický stream between the southern edge of the town of Velké
Pavlovice, the village of Velké Bílovice, the town of Podivín (Kostel) and the village of
Rakvice (Rakwitz). Towards the north-west, halophytes occurred along the railway as far
as the Zaječí (Saitz) station. The latter site, drained by Zaječí stream (a left-hand tributary
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 31
of the Štinkovka stream), might be considered as a part of the Šakvice district. However,
we prefer to follow the district delimitations proposed by Grulich (1987). In the north,
this district borders the Čejč district. In the south-west, it reaches the Dyje floodplain, but
it is impossible to draw a sharp border. While succulent halophytes never occurred within
reach of floods, Bupleurum tenuissimum,Crypsis schoenoides and Cirsium brachy-
cephalum were repeatedly recorded on pasture land inundated during extreme floods or
even within floodplains.
The earliest known herbarium specimen of a halophyte from southern Moravia was
collected in this district by H. Schott, most likely between 1786 and 1792. The specimen
of Plantago maritima labelled “auf der Huthweide bey Kostel” is now preserved in the
herbarium of the Hungarian Natural History Museum in Budapest (BP; Sutorý 1995).
However, this record remained unknown to Moravian botanists for the next two
centuries.
The halophytic flora of the Rakvice district was rich in species and included all the 17
halophytes except for Galatella cana and Triglochin maritima. Both succulent halo-
phytes only occurred near the Zaječí railway station. There are numerous herbarium
specimens of Salicornia perennans from the site near the Rakvice railway station but
only one, possibly mislabelled, specimen of Suaeda prostrata.
The first halophytic site discovered in this district was near the Zaječí railway station.
Alexander Makowsky collected Crypsis aculeata there in 1874 and Bupleurum
tenuissimum and Salicornia perennans in 1879. Laus (1907) reported halophytes to
occur mainly in the ditches along the southern side of the railway and on arable land
south-west of the railway station. However, he noted that they were strongly declining
due to reclamation. Suaeda prostrata was last collected there in 1921, and Salicornia
perennans (by Laus already considered disappeared) in 1924. Other halophytes vanished
from this site roughly at the same time, except for Tripolium pannonicum, which was last
collected in 1952. Šmarda (1953) observed only Bolboschoenus maritimus agg. and
Lotus maritimus at this site.
Species-rich halophytic sites, discovered as late as 1907 by H. Laus, were around the
village of Rakvice (Laus 1909). North of the railway station, he observed Glaux maritima,
Juncus gerardii,Plantago maritima,Salicornia perennans and Scorzonera parviflora.
South of the railway next to the station, Crypsis aculeata,C. schoenoides and Spergu-
laria media occurred in a marshland grazed by geese. In 1909, he also collected Tara-
xacum bessarabicum, while Samolus valerandi was found there in 1926. Halophytes also
occurred along the southern and south-eastern edge of the village. A small fishpond was
indicated on the 3rd Military Survey map (1876–1878) at the southern edge of the village.
Then it was probably surrounded by halophytic vegetation, but now it is filled with earth
and built up. South and south-east of the village, the map showed large tracts of common
pastures and reed beds. As documented by numerous herbarium specimens of halo-
phytes, these pastures and wetlands were at least partly saline. They extended further east
of the village towards the Trkmanka stream and the town of Podivín. Some halophytes,
e.g. Crypsis schoenoides and Spergularia marina, may have occurred further in the
south-east, near Podivín, but locality information on herbarium labels is vague.
Šmarda (1953) confirmed the occurrence of halophytes at both sites near Rakvice. In
1950, he found Crypsis schoenoides,Glaux maritima,Juncus gerardii,Spergularia
marina and Tripolium pannonicum in saline marshlands south-east of the village. Two
32 Preslia 94: 13–110, 2022
years later, he recorded Cirsium brachycephalum in railway ditches next to the station,
along with Juncus gerardii,Plantago maritima,Scorzonera parviflora and Tripolium
pannonicum. In 1959–1964, the halophytic vegetation around Rakvice was documented
by Vicherek (1962, 1973), who recorded numerous relevés containing most of the
halophytes that occurred at both major sites.
However, both halophytic sites near Rakvice were destroyed. Salicornia perennans
was last collected near the railway station in 1932 and vanished probably in the 1930s.
Suaeda prostrata was allegedly collected there in 1925 by F. Weber; however, this may
be a mislabelled specimen, and the species may have never occurred at that site. The halo-
phytic habitats south and south-east of the village were destroyed probably in the 1960s.
In contrast, halophytes in the ditches next to the railway station survived longer: Scorzonera
parviflora and Tripolium pannonicum were last observed in 1981 (Grulich 1987).
The large saline marshlands northeast of Rakvice and south of the Trkmanský dvůr
settlement (below referred to as Trkmanec-Rybníčky and Trkmanské louky) were dis-
covered by S. Staněk and J. Bílý Jr. in 1922, when they collected Cirsium brachy-
cephalum and Tripolium pannonicum there. However, this site was almost disregarded
by Šmarda (1953), who cited a few records of subhalophytes provided by F. Kühn. In
1962 or 1963, J. Vicherek sampled there a relevé of a saline marshland with Bolbo-
schoenus maritimus agg., Cirsium brachycephalum,Juncus gerardii and Scorzonera
parviflora (Vicherek 1973), providing the only record of the latter species from this site.
Samolus valerandi was discovered there only in 1982, and it has been observed repeat-
edly since then.
The last major halophytic site in the Rakvice district existed around the Fabián Fish-
pond at the northern edge of the town of Velké Bílovice. It was discovered in 1921 by
J. Podpěra, who collected Bupleurum tenuissimum,Glaux maritima and Spergularia
media on the shores of this small fishpond.During the following decades, Crypsis
aculeata,C. schoenoides,Plantago maritima,Spergularia marina and Tripolium panno-
nicum were repeatedly collected at this site. Šmarda visited the site in 1951 and recorded
Crypsis aculeata,Glaux maritima,Spergularia marina and Tripolium pannonicum,and
probably also Juncus gerardii (Šmarda 1953). Halophytic vegetation was still present
there in the early 1960s (Vicherek 1962). Last halophytes, namely Crypsis aculeata and
Spergularia marina, were collected in 1971 and 1978, respectively. The fishpond was
filled with earth probably at the turn of the 1970s, and currently, the area is used as aplay-
ground with no remnants of halophytic vegetation.
The current halophytic flora of the Rakvice district is enormously impoverished. Both
succulent halophytes disappeared already in the 1920s. Crypsis aculeata and Glaux
maritima were last collected in 1964, and Bupleurum tenuissimum in 1969. Plantago
maritima,Scorzonera parviflora and Tripolium pannonicum vanished during the early
1990s. Only Cirsium brachycephalum,Crypsis schoenoides,Juncus gerardii,Samolus
valerandi and Spergularia marina still survive. Spergularia media,nowadaysalsopres
-
ent in the district, was collected only twice in 1909 and 1918, and then again in 2008 and
2014 along the motorway. However, the recent occurrences may not be derived from the
autochthonous populations.
Trkmanec-Rybníčky and Trkmanské louky. – This site is located ~2 km E to NE of the
Rakvice village. The 2nd Military Survey map (1836–1840) shows a fishpond (Rakwitzer
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 33
Teich) occupying a large part of this area, and the 3rd Military Survey map (1876–1878)
shows the fishpond to be drained and converted to meadows and probably reed beds. The
latter map also indicates small areas of meadows and common pastures between the
southern corner of this area and the railway. In the 1970s, when the motorway
Praha–Bratislava was built, the road Velké Pavlovice–Podivín, originally following the
right bank of the already channelized Trkmanka stream, was abandoned. The stream
channel was relocated to the east, and the motorway separated the area into the larger
northern and the smaller southern part. The northern part has been protected since 2008
as the Trkmanec-Rybníčky Nature Monument, while the southern part has been proposed
but not yet designated as the Trkmanské louky Nature Monument. Both sites are pro-
tected as Natura 2000 sites.
The Trkmanec-Rybníčky Nature Monument (Fig. 3C) lies in a triangle between the
road Velké Pavlovice–Velké Bílovice, the Trkmanka stream and its tributary Bílovický
stream. In the 1980s and early 1990s, this area was arable land drained by a channel, from
which water was pumped into the Trkmanka stream. The operation of this drainage sys-
tem ceased and the arable fields were gradually abandoned between 1996 and 2006.
Waterlogging of a large part of the area resulted in the succession of Phragmites australis
stands. In 2006–2008, the area was restored, the reed stands were reduced and shallow
pools were dug, in which Crypsis schoenoides and other halophytes appeared. At present,
Trkmanec-Rybníčky comprises areas with different water regimes. The northern part
encompasses halophytic pastures currently managed by horse grazing and mowing. In
wetter parts, these pastures contain halophytes such as Samolus valerandi. The pastures
occur in a mosaic with halophytic marshes with Bolboschoenus planiculmis,Schoeno-
plectus tabernaemontani (associations Astero pannonici-Bolboschoenetum compacti and
Schoenoplectetum tabernaemontani) and other types of wetland vegetation (e.g. Bidenti
frondosae-Atriplicetum prostratae). The central and southern parts of the site are domi-
nated by marshes, and there are also a few ponds, in which stands of Heleochloëtum
schoenoidis were sampled in 2011 and 2015 by P. Lustyk and P. Dřevojan (Dřevojan et
al. 2017).
The Trkmanské louky site (Fig. 4A) is currently managed mainly as arable land. The
proposed Nature Monument aims to protect the critically endangered Cirsium brachy-
cephalum, which grows in fragmentary stands of the association Agrostio stoloniferae-
Juncetum ranarii. Besides crop cultures, Phragmitetum communis is the most wide-
spread vegetation, accompanied by scrub patches with Cornus sanguinea and Calama-
grostis epigejos indicating ongoing succession. Subhalophytes such as Melilotus
dentatus and Centaurium pulchellum can be found rarely at the edges of these extensive
stands. In more disturbed and wet places on arable land where crops grow weakly, saline
vegetation develops with dominant Bolboschoenus planiculmis, accompanied by
Cirsium brachycephalum (Tripleurospermo inodori-Bolboschoenetum planiculmis).
Čejč district
This district included halophytic sites in the valley of the Trkmanka stream and its tribu-
taries around the former salt lakes of Čejč (Čejčské jezero, Tscheitscher See) and Kobylí
(Kobylské jezero, Kobyler See), and a small lake or a fishpond (Smraďák, Smradaker
Teich) between the villages of Terezín (Theresiendorf) and Čejč (Tscheitsch). Towards
34 Preslia 94: 13–110, 2022
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 35
Fig. 4. Halophytic localities in the Rakvice, Čejč and Sedlec districts: (A) Trkmanské louky 2020, (B) Zápověď
2020, (C) Drained bottom of the Nesyt Fishpond in its southern bay 2020. Photo credits: F. Kratoš (A, B),
M. Chytrý (C).
the north, saline habitats occurred in the valley of the Kašnice stream as far as the former
railway station Klobouky u Brna at the eastern edge of the town and along the Spálený
stream as far as the former Kumstát (or Kunštát) mill north of the village of Krumvíř.
Towards the south, the occurrences of some halophytes once continued along the
Trkmanka roughly to the southern edge of Velké Pavlovice, i.e. to the arbitrary border
with the Rakvice district.
The halophytic flora of the Čejč district was rich in species and included all the 17
halophytes except for Galatella cana and Triglochin maritima. The occurrence of
Cirsium brachycephalum is uncertain because there is only a single record from a relevé
(Vicherek 1962).
The salt lake surrounded by saline marshes, fen meadows and steppe slopes harbour-
ing rare species attracted numerous botanists who praised the local flora in superlatives.
For instance, Hochstetter (1825) described it as a place where the Moravian flora “no
doubt develops its rarest products”. Wiesner (1854) referred to the vicinity of Čejč as
“Flora’s treasury in Moravia”, while Bubela (1882) described it as “Eldorado of
Moravian botanists”.
Čejč Lake was situated ~0.9 km SSE of the village of Čejč. The earliest description of
the lake is found in a book about the healing waters of the Margraviate of Moravia
(Jordán z Klauznburku 1580: 284–293). The author described the lake's water as awfully
bitter, comparing its taste to Centaurium,Gentiana,Artemisia absinthium and the galls of
some animals. He also noticed that no fish could live in the lake, and its water was toxic to
livestock from elsewhere but not to animals from local farms. He further reported that in
summer, the water level was low, and the shores were white [from salt efflorescences].
Much to botanists’ regret, Čejč Lake was drained in 1858 (Krzisch 1859).
Kobylí Lake occupied a shallow depression of the Trkmanka stream between the vil-
lages of Kobylí, Brumovice and Terezín. It was ~5 km long and in its north-eastern part
up to 1.5 km wide. Its water was less saline than that of Čejč Lake (Vlach 1952). It was
reportedly drained as soon as 1834 or 1835 (Spitzner 1894, Vlach 1952), but its northern
part is shown on the 2nd Military Survey map (1836–1840) as still filled with water.
The small lake or a fishpond between the villages of Terezín and Čejč, referred to as
Smraďák (Smradaker Teich) or Rybníček, was located ~0.5 km S of the village of Terezín
(Theresiendorf). It was drained at the same time as Čejč Lake. The bottoms of all three
lakes are now used mainly as arable land. While the shapes of the former Čejč and Kobylí
Lakes may be still traced on current topographic maps, and both place names are still in
use, the former area of the small Smraďák Fishpond has been dissected into three parts by
the local railway Zaječí–Hodonín (opened in 1897) and its branch to Klobouky u Brna.
This district was first floristically explored independently by Ch. F. Hochstetter and
A. Carl, then a county physician in the town of Uherské Hradiště. In 1821, A. Carl col-
lected Scorzonera parviflora and Tripolium pannonicum near Čejč Lake. Next year, he
found Plantago maritima near the villages of Vrbice and Čejč. The first published
floristic records from this district are those from Hochstetter (1825). He reported thatthe
low shores of Kobylí Lake were richly covered with the succulent halophytes Salicornia
perennans and Suaeda prostrata (Chenopodium maritimum)aswellasChenopodium
chenopodioides (C. serotinum)andLepidium coronopus. At Čejč Lake, he found Carex
distans and Scorzonera parviflora. Wilhelm Tkany and Friedrich Kolenati, reporting on
their excursion to the Čejč surroundings with J. Bayer in 1851 (Kolenati 1852), listed
36 Preslia 94: 13–110, 2022
Glaux maritima,Salicornia perennans and Scorzonera parviflora among the species
found.Bayer (Anonymous 1853), in addition, observed Bupleurum tenuissimum,
Crypsis aculeata,Plantago maritima,Spergularia marina,Suaeda prostrata (Schoberia
maritima)andTripolium pannonicum near Čejč Lake. However, already Krzisch (1859),
who found only a few hardly surviving halophytes in the surroundings of Čejč Lake,
regretted the lake’s drainage and envisaged the decline of halophytes.
The drainage of both lakes and the Smraďák Fishpond did not cause immediate
destruction of all halophytic habitats or local extinction of halophytes. For instance,
Steiger (1880) noted Plantago maritima in meadows near Brumovice, probably near the
fishpond at the eastern edge of the village. Bohuňovský (1880) confirmed the occurrence
of Salicornia perennans in a pasture near Terezín and discovered Samolus valerandi at
the site referred to as Na Rybníčku, which corresponds to the remnants of the drained
Smraďák Fishpond. This site was explored by Bubela (1882), who confirmed the occur-
rence of Samolus valerandi and recorded Crypsis aculeata,C. schoenoides,Salicornia
perennans,Suaeda prostrata and Tripolium pannonicum in the saline reed bed and on dry
mud, while Bupleurum tenuissimum,Plantago maritima,Spergularia marina,S. media
and Taraxacum bessarabicum occurred in the adjacent meadow. In the late 1890s, the site
was disturbed during the construction of the railway Zaječí–Hodonín, but halophytes
continued to occur there. The succulent halophytes were last collected at this site in the
mid-1940s, and the habitats must also have been destroyed in the 1940s because Šmarda
(1953) did not mention this site at all.
Botanists continued visiting this district, repeatedly confirming the presence of halo-
phytes at various sites. For instance, Formánek (1884) recorded Scorzonera parviflora
and Tripolium pannonicum near Terezín, and (probably during a second trip) Bupleurum
tenuissimum,Crypsis aculeata (near the sulphur water spring Heliga), C. schoenoides,
Glaux maritima,Plantago maritima,Spergularia marina and Suaeda prostrata near
Čejč. According to Spitzner (1894), who explored the area in 1887, the western shores of
the former Kobylí Lake (east of the village of Brumovice) hosted Salicornia perennans,
Suaeda prostrata and Tripolium pannonicum.Glaux maritima occurred alongside the
stream that once connected Čejč and Kobylí Lakes, i.e. somewhere near the site Na
Rybníčku, while Samolus valerandi was found in cattle tracks on the bottom of the for-
mer Čejč Lake. Spergularia marina grew on the bottom of the former Kobylí Lake, and
Crypsis aculeata and C. schoenoides in ditches probably south-west of the village.
Samolus valerandi was abundant on cattle paths in the former Čejč Lake (Spitzner 1894).
Heinrich Laus visited the district in 1906 and observed only Spergularia marina and
Tripolium pannonicum in the area of the former Čejč Lake (Laus 1907). He did not find
anything of the former halophytic flora between Terezín and Kobylí but still observed
a few sites with halophytes in the district, such asnear the Rovinský dvůr (Rovinka) farm
south of Krumvíř.
Some well-preserved remnants of the halophytic vegetation survived for a long time at
the north-eastern edge of the former Kobylí Lake. Between 1927 and 1929, the surround-
ings of Terezín were thoroughly explored by Gilli (1930). He paid particular attention to the
patches of halophytic vegetation between the railway and the road to Krumvíř ~1.3 km
NNW(–N) of the village, i.e. to the site now referred to as Zápověď. He described the
place as a marshland with a reed stand in the middle. Tripolium pannonicum was found in
large quantities at the edge of the reed stand, Glaux maritima,Juncus gerardii,Plantago
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 37
maritima,Scorzonera parviflora and Spergularia media occurred in the surroundings,
and Salicornia perennans was present at one spot with salt efflorescences. The Zápověď
site, on the 3rd Military Survey maps (1876–1878) indicated as a common pasture, was
frequented by botanists from the 1930s onwards, as documented by numerous herbarium
specimens. This site was spared from draining due to its location in a narrow strip of land
between the road and railway. It was managed as a nature reserve by a non-governmental
nature conservation society from 1944 and designated as a State Nature Reserve in 1952
(Vlach 1952, Smejkalová 1990). In 1949, the site was explored by Šmarda (1953) and
roughly at the same time also by Vlach (1952). The latter author described the site in
detail and noted several vegetation types, including stands with both succulent halo-
phytes. He recorded Glaux maritima,Juncus gerardii,Plantago maritima,Salicornia
perennans,Scorzonera parviflora,Spergularia marina,S. media,Suaeda prostrata,
Taraxacum bessarabicum and Tripolium pannonicum. Šmarda (1953) found the same
species as Vlach and, in addition, collected Bupleurum tenuissimum. Jiří Vicherek stud-
ied halophytic vegetation in the Čejč district in 1959–1964. In the relevés from the
Zápověď site, he recorded Bupleurum tenuissimum,Crypsis aculeata,Juncus gerardii,
Plantago maritima,Salicornia perennans,Scorzonera parviflora,Spergularia media,
Suaeda prostrata,Taraxacum bessarabicum and Tripolium pannonicum (Vicherek
1962, 1973). However, his record of Suaeda prostrata in two relevés from 1959–1961 is
the last record of this species from the site. Salicornia perennans was last collected in
1967. Other halophytes disappeared in Zápověď by the early 1970s, primarily due to the
drainage of the surrounding land (Grulich 1987).
Gilli (1930), apart from describing the Zápověď site in detail, reported the occurrence
of halophytes elsewhere in the northern part of the district. He found Bupleurum
tenuissimum,Plantago maritima and Tripolium pannonicum in the meadows between
the road and the Trkmanka stream north of Zápověď towards Krumvíř. Plantago
maritima dominated vegetation in some places, and Spergularia media occurred on ara-
ble land. He further recorded Spergularia media around a small fishpond in the southern
part of Krumvíř, and Plantago maritima,Scorzonera parviflora and Tripolium panno-
nicum in the meadows along the Spálený stream between the village and the Kumstát
mill. In contrast, he did not confirm the occurrence of Scorzonera parviflora near the rail-
way station Klobouky u Brna, where it was once found by Steiger (1880). In the early
1950s, Marvanová & Marvan (1959) found Spergularia marina along the Spálený stream
south of Krumvíř, Samolus valerandi together with Juncus gerardii in a drained fishpond
south-west of the Rovinský dvůr farmstead and Plantago maritima at two sites near Čejč
and on a path to the Samolus site south of Krumvíř. Vicherek (1962, 1973) recorded
relevés of saline marshlands with Juncus gerardii,Plantago maritima,Samolus valerandi,
Taraxacum bessarabicum and Tripolium pannonicum along the Trkmanka stream north
of the Zápověď site towards Krumvíř, which is the area once explored by Gilli (1930) and
Marvanová & Marvan (1959). He also sampled saline marshlands with Juncus gerardii
and Scorzonera parviflora near the Trkmanka stream between the villages of Terezín and
Brumovice. These were probably the last remaining sites with halophytic vegetation suit-
able for vegetation sampling in the Čejč district.
As mentioned above, occurrences of some halophytes continued along the Trkmanka
stream as far as the southern edge of the town of Velké Pavlovice. There were several
records of Glaux maritima between 1857 and 1949 from saline grasslands next to the
38 Preslia 94: 13–110, 2022
village of Bořetice, probably somewhere close to its northern edge. Glaux maritima also
used to occur at the northern edge of Velké Pavlovice along the Trkmanka stream and east
of the railway station at its southern periphery; the last observation from the latter site
dates back to 1952 (Z. Hrabal sec. Šmarda 1953). Juncus gerardii was collected some-
where near the town in 1931.
In the mid-1980s, no occurrences of halophytes in the Čejč district were known apart
from a single record of Spergularia marina from 1983 at the northern edge of Terezín
(Grulich 1987). In the late 1980s, the protection of the degraded Zápověď site was recom-
mended for abolition, which happened probably in the early 1990s (Buček & Smejkalová
1987; but see Smejkalová 1990). However, in 2003, the Zápověď site was restored by
bulldozing shallow pools of different depths at the formerly saline sites. Halophytic vege-
tation quickly developed around them (Slavík in Hadinec & Lustyk 2009). Beside these
shallow pools, a fishpond was built in 2005, but it is deep and therefore unsuitable for
halophytic vegetation. The upper soil layer of the shallow pools was removed in the
2010s to support the halophytes dependent on early successional stages (P. Slavík, pers.
comm.). Spergularia media emerged there in 2005, Glaux maritima in 2007, Crypsis
aculeata in 2008 and Spergularia marina in 2015, all probably from the soil seed bank.
The site currently hosts halophytes and subhalophytes such as Bolboschoenus maritimus,
Carex secalina,Glaux maritima and Spergularia media (Fig. 4B). Grasslands around the
pools are regularly mown. In drier parts, there are saline grasslands of Loto tenuis-
Potentilletum anserinae, while the places flooded for a longer time but drying out in sum-
mer are occupied by a Puccinellietum limosae grassland with Puccinellia distans and
Spergularia media. Reed stands of Phragmites australis dominate deeper parts of the
pools usually flooded throughout the year. Stands of Bolboschoenus maritimus and
B. planiculmis (Astero pannonici-Bolboschoenetum compacti) occur at the edges of
these reed beds and in other flooded places.
Apart from the new occurrences of halophytes at Zápověď, Samolus valerandi was
found in 2017 in a wet meadow in the Trkmanka valley ~2.5 km south-east of the village
of Krumvíř, i.e. near the sites where it was once recorded by Marvanová & Marvan
(1959) and Vicherek (1962).
Sedlec district
The Sedlec district lies in the shallow valley of the Včelínek (Niklasgraben) stream
roughly between the towns of Mikulov (Nikolsburg) and Lednice (Eisgrub). In the west,
it also includes a small area adjacent to the western edge of Mikulov drained by a name-
less left-hand tributary of the Včelínek and a small area in Lower Austria drained by the
Niklasgraben stream, which is the uppermost reach of the Včelínek. The soil salinity in
the district is caused by sulphates (mainly magnesium sulphate) from Tertiary deposits,
but in the village of Sedlec (Voitelsbrunn, Selce), it was formerly strengthened by a min-
eral spring. In the 15th century, several fishponds were built on the Včelínek stream
between Mikulov and Lednice, just on the historical border between Moravia and Lower
Austria. They included the Porzteich Fishpond (now partly preserved as Nový Fishpond)
west of Sedlec and a series of four fishponds east of Sedlec, namely Nesyt (Nimmer-
satteich or Steindammteich), Hlohovecký (Bischofwarter Teich), Prostřední (Mitter-
teich) and Mlýnský (Mühlteich). The Czech name Nesyt and its German equivalent
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 39
Nimmersatteich mean Sateless pond, which refers to water shortage in this dry and warm
area.
The halophytic flora of the Sedlec district used to be rich in species and included all
the 17 halophytes considered here except Cirsium brachycephalum,Galatella cana and
Triglochin maritima. The presence of Glaux maritima is doubtful. However, the district
was poorly accessible from both Brno and Vienna due to its peripheral location. There-
fore, it was rarely visited by botanists, and the occurrence of halophytes was reported rel-
atively late. The earliest record of a halophyte from the district is that of Tripolium
pannonicum in the Flora of Lower Austria (Neilreich 1859), but the origin of this record
is unknown. Plantago maritima,Scorzonera parviflora,Spergularia media (Lepigonium
marginatum)andTripolium pannonicum were collected by F. R. Müncke, who served as
a pharmacist in the Prussian army and explored the site during the Prussian military cam-
paign against Austria in 1866 (Müncke 1868). His records are labelled Valtice (Felds-
berg) and probably refer to the occurrences of halophytes at the south-eastern edge of the
Nesyt Fishpond. Formánek (1887, 1892) reported Spergularia marina,Taraxacum bes-
sarabicum and Glaux maritima from Sedlec. However, the occurrence of G. maritima has
never been documented by a herbarium specimen, and the records in the relevés from the
1960s are most likely mistakes. The occurrences of the succulent halophytes, Salicornia
perennans and Suaeda prostrata, were discovered in 1914 by F. Zimmermann (Wildt
1915, Zimmermann 1916). In 1922, J. Bílý Jr. found Samolus valerandi on the Nesyt
shores near the Sedlec railway station. At that time, well-developed halophytic vegeta-
tion was found along both banks of the Včelínek stream near the western shore of Nesyt,
an area marked as a common pasture on the 3rd Military Survey maps.
Since the 1920s, the sites around the Nesyt Fishpond have been repeatedly visited by
Moravian botanists. Anton Fröhlich, a secondary school teacher in Mikulov, reported
Bupleurum tenuissimum in 1933 and made numerous observations about local halophytic
flora and vegetation (Fröhlich 1935, 1940, 1943). Zapletálek (1933, 1939) provided
a description of the flora and vegetation in the surroundings of the Nesyt Fishpond.
His description of vegetation developed on the drained Nesyt bottom, summarized in
an unpublished thesis, was partly reproduced by Šmarda (1953). Jan Šmarda visited the
site in 1950 and confirmed the occurrence of Bupleurum tenuissimum,Juncus gerardii,
Plantago maritima,Salicornia perennans,Scorzonera parviflora,Spergularia marina,
S. media and Suaeda prostrata near the Sedlec railway station (Šmarda 1953). At that
time, a concentric zonation of halophytic vegetation was still preserved, with the stands
of Puccinellia distans and Salicornia perennans in the lowest part in the middle of the
site. Fröhlich summarized his observations in a brief report (Fröhlich & Švestka 1956)
accompanied by a distribution map of selected species around the western half of the
Nesyt Fishpond. Many of the records were indicated as old; however, this map shows that
some halophytes, including Suaeda prostrata, once occurred quite far in the south-west,
south of the railway. The aerial photographs from the 1950s show a sizable treeless area
on both sides of the Včelínek stream between the western tip of the Nesyt Fishpond and
the village of Sedlec. The area was crossed by walking paths and contained trampled
patches with sparse vegetation. It was probably still used as a common pasture and
provided suitable conditions for halophytic vegetation.
In the early 1960s, the vegetation on the western shores of the Nesyt Fishpond was
explored by Vicherek (1962, 1973). His relevés are not dated, but he collected herbarium
40 Preslia 94: 13–110, 2022
specimens at this site in 1960–1962. He still recorded stands with Salicornia perennans
and Suaeda prostrata, patches with annual grasses Crypsis aculeata and C. schoenoides,
and saline marshlands with Scorzonera parviflora. Most of these relevés were probably
sampled in the current nature reserve and on the drained Nesyt bottom. However, he also
recorded stands with Crypsis aculeata at the northern margin of the fishpond and saline
grassland with Plantago maritima,Puccinellia distans,Spergularia media,Taraxacum
bessarabicum and Tripolium pannonicum at the southern and south-eastern shores. This
suggests that the halophytic vegetation was still well preserved and relatively widespread
at that time.
Since the 1960s, the occurrences of halophytes were gradually restricted to the area of
the current Slanisko u Nesytu National Nature Reserve. This reserve was officially estab-
lished in 1961, but there was some kind of protection earlier, probably from 1953
(Grulich 1987), as Fröhlich & Švestka (1956) refer to the site as a halophytic reserve. The
common pasture between the Včelínek stream and the southern edge of the village of
Sedlec was later converted into a football playground. The frequent mowing and trampling
by football players turned out to support halophytes. Spergularia marina and Taraxacum
bessarabicum survived on the playground until1999; however, we found none of them at
this site in 2020. During the 1960s, grazing at the Slanisko u Nesytu site gradually ceased,
perhaps partly due to its legal protection. No goose grazing was observed there in the
1970s (V. Grulich, pers. comm.). Consequently, the habitat quality deteriorated, and both
succulent halophytes, Salicornia perennans and Suaeda prostrata, went locally, hence
also nationally, extinct there in 1976 and 1986, respectively (Grulich 1987).
In the past, halophytes also occurred scattered elsewhere around the Nesyt Fishpond.
For instance, Scorzonera parviflora was collected in the southern bay of the Nesyt Fish-
pond in 1930 by J. Zapletálek. Fröhlich & Švestka (1956) reported Crypsis aculeata and
C. schoenoides from that place. Vicherek (1973) recorded a relevé with Juncus gerardii,
Orchis palustris and Tripolium pannonicum, probably in the same place, in the early
1960s. Small patches of halophytic vegetation with Bupleurum tenuissimum,Plantago
maritima,Spergularia marina and some subhalophytes once existed along a walking
path on the southern shores of the Výtopa Fishpond (adjacent to the south-eastern corner
of Nesyt) and in a shallow depression next to the north-eastern corner of Výtopa
(Danihelka & Hanušová 1995). Both sites were indicated as common pastureson the 3rd
Military Survey maps. The early records of halophytes by Müncke (1868) may also refer
to this part of the district. To our knowledge, Bupleurum tenuissimum was last seen there
in 2004, while Plantago maritima and Juncus gerardii were still present in 2020 and 2021.
Further eastwards, some halophytes, namely Crypsis schoenoides,Samolus valerandi
and Tripolium pannonicum, were scattered on the shores and drained bottoms in the
Lednice Fishpond system. Crypsis schoenoides was collected on the drained bottom of
the Hlohovecký Fishpond in 2021. The easternmost halophytic site in this district was at the
western edge of the Dyje floodplain below the dam of the Mlýnský Fishpond (Mühlteich)
south-east of Lednice. In the early 1960s, Vicherek (1973) recorded there a relevé of
a saline marshland with Juncus gerardii,Plantago maritima and Tripolium pannonicum.
One herbarium specimen of Juncus gerardii from 1960 and a few specimens of
Taraxacum bessarabicum from the 1920s–1960s were probably collected at the same
site. However, no halophytes have been found in this part of the district since the 1960s.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 41
Although concentrated to the western shore of the Nesyt Fishpond, some halophytes
also occurred along the Včelínek stream westwards as far as south and west of Mikulov.
There was a herbarium specimen from the mid-19th century and a literature record from
1949 (J. Bílý sec. Šmarda 1953) of Plantago maritima from that area. Scorzonera parvi-
flora once occurred near the Tiergartenhof north-west of the village of Kleinschwein-
barth in Lower Austria (S. Domas sec. Makowsky 1863). Also, the herbarium records of
Crypsis schoenoides from the Seeäcker saline meadows along the border between
Moravia and Lower Austria may be attributed to the Sedlec district. In 2007, a large pop-
ulation of Samolus valerandi developed temporarily in a saline marshland with Bolbo-
schoenus planiculmis in a shallow wet depression on arable land ~1.6 km W of Mikulov,
north of the railway to Znojmo (Paukertová & Sedláček in Hadinec & Lustyk 2007:326).
Several subhalophytes (e.g. Centaurium pulchellum,Lythrum hyssopifolia and Schoeno-
plectus tabernaemontani) were also found there.
Currently, the Sedlec district has the best-preserved halophytic flora of all the ten dis-
tricts considered in this study. Although both succulent halophytes have disappeared,
Bupleurum tenuissimum,Crypsis aculeata,C. schoenoides,Juncus gerardii,Plantago
maritima,Scorzonera parviflora,Spergularia marina,S. media and Tripolium panno-
nicum still occur at the Slanisko u Nesytu site. Moreover, Crypsis schoenoides and Samolus
valerandi have been repeatedly recorded on the drained bottoms of the Nesyt and
Hlohovecký fishponds. A few individuals of Crypsis aculeata and Tripolium panno-
nicum were observed in 2020 on the drained bottom of the Nesyt Fishpond in its southern
bay (Fig. 4C). In addition, Juncus gerardii and Plantago maritima have survived at
another site south of the small Výtopa Fishpond.
Slanisko u Nesytu (Fig. 5A). – The Slanisko u Nesytu National Nature Reserve, the most
species-rich halophytic site in the study area, is located ~0.4 km S of the village of Sedlec
along the western margin of the Nesyt Fishpond. The reserve is divided into the western
and eastern parts by a shallow ditch. Most of its area is currently covered by sub-
halophytic grasslands. The occurrences of halophytes are confined mainly to shallow
depressions in both parts of the reserve, being slightly more common in the eastern part.
Until the 1990s, targeted conservation management was limited. In the mid-1970s, shal-
low ditches were dug in the western part of the reserve to create a habitat for annual
halophytes (Grulich 1987). The ditches had a positive effect for some years until they
were overgrown by Agrostis stolonifera. In 1986, a square-shaped pit was dug in the east-
ern part of the reserve to re-establish Salicornia perennans (Grulich 1987, in litt. 2021).
This attempt failed, and the pit was ultimately overgrown by Phragmites australis
(Danihelka 2005). In 1991, topsoil was removed in a strip about 100 m long and 3 m wide
in the western part of the reserve (Danihelka & Hanušová 1995). This shallow ditch now
harbours small populations of Crypsis aculeata and C. schoenoides, particularly in rainy
years (Dřevojan et al. 2017), and sometimes also several individuals of Samolus
valerandi (population derived from seeds collected from the bottom of Nesyt) and
Scorzonera parviflora. Regular mowing was re-established in 1993, and sheep grazing
was introduced in 2000. The intensity and spatial extent of these management measures
have increased since then. Nowadays, the halophytic vegetation is grazed by horses,
which seems to be a particularly suitable management practice for saline habitats (Kmet
et al. 2018). The best developed halophytic community (association Puccinellietum
42 Preslia 94: 13–110, 2022
limosae) with Bupleurum tenuissimum,Plantago maritima,Taraxacum bessarabicum
and Tripolium pannonicum is preserved in the eastern part of the reserve. Slightly wet
to mesic parts of the site are covered by the grasslands of Loto tenuis-Potentilletum
anserinae and drier parts with Festuca pulchra by Centaureo pannonicae-Festucetum
pseudovinae. Wet halophytic grasslands of Scorzonero parviflorae-Juncetum gerardii,
once common at the site, were developed in the 1990s in a ditch created for conservation
purposes in the western part (Hanušová 1995) and last recorded there by J. Danihelka in
2004.
Slanisko v trojúhelníku. – This recently discovered small halophytic site is located ~1.6 km
W of the town of Mikulov in a shallow wet depression on arable land north of the railway
Mikulov–Znojmo. In 2006, a part of the depression was filled with earth. In 2007, a popu-
lation of Samolus valerandi, counting several thousands of individuals, was discovered
there (Paukertová & Sedláček in Hadinec & Lustyk 2007: 326).Later, a part of the earth
was removed to support the occurrence of halophytes. In 2020, the site was relatively dry,
covered with a mosaic of bare soil and annual ruderal vegetation. Small plants of
S. valerandi occurred in large numbers in pits dug in the previous year as a part of conser-
vation management. However, the prospects of this site are uncertain because it is located
in the route of a planned motorway.
Slanisko u Výtopy. – Small remnants of halophytic vegetation are preserved along the
walking path south of the Výtopa Fishpond ~1.9 km N of the Valtice railway station.
A small population of Plantago maritima, counting about 100 individuals, occurs in the
Centaureo pannonicae-Festucetum pseudovinae grassland dominated by Festuca pulchra.
Juncus gerardii occurs on wet arable land next to the south-eastern bay of the fishpond.
Damnice-Olbramovice district
The Damnice-Olbramovice district includes the former halophytic sites in a shallow val-
ley of the Miroslavka stream between the Miroslavský Fishpond south-west of the village
of Suchohrdly u Miroslavi in the west and the village of Vlasatice in the east, as well as
a shallow valley of the Olbramovický stream between the village of Olbramovice in the
north-west and Vlasatice in the south-east. This district is proposed here in addition to the
eight districts recognized in the Moravian part of the study area by Grulich (1987).
The flora of the Damnice-Olbramovice district has always been poor in halophytes for
its peripheral location and likely a low degree of soil salinity. As botanists rarely visited
the district, some halophyte occurrences may have vanished without being recorded. Of
the 17 halophytes, the district used to harbour Glaux maritima,Plantago maritima,Spergu-
laria marina,Tripolium pannonicum, and probably also Juncus gerardii (Šmarda 1953).
The first botanist to discover halophytes in this district was H. Zimmermann. In 1883, he
collected Plantago maritima in a meadow north-west of the village of Damnice (Damitz)
and noted Spergularia marina and Tripolium pannonicum as co-occurring species on the
herbarium label of this Plantago specimen (see also Oborny 1886). Andreas Ripper
explored the area a few years later and recorded Glaux maritima in the villages of Damnice,
Jiřice u Miroslavi (Iritz) and Troskotovice (Treßkowitz; Oborny 1912). He also confirmed
theoccurrenceofPlantago maritima at the site discovered earlier by Zimmermann
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 43
44 Preslia 94: 13–110, 2022
Fig. 5. Halophytic localities in the Sedlec and Hevlín-Nový Přerov districts: (A) Slanisko u Nesytu 2016, (B)
Dobré Pole 2020, (C) Novosedly 2020. Photo credits: M. Chytrý (A), K. Chytrý (B, C).
(“between Damnice and the Miroslav railway station”) and added another record near
Troskotovice. The halophytic site north-west of Damnice survived at least until 1952 when
J. Šmarda and M. Součková collected Plantago maritima and Tripolium pannonicum and
observed Juncus gerardii there. They also found some subhalophytes in a wet meadow
upstream below the Suchohrdelský Fishpond (Šmarda 1953).
The occurrences of halophytes along the Olbramovický stream were discovered even
later. Johann Hruby collected Plantago maritima at the margin of the village of Malé
Želovice (Klein-Seelowitz, currently part of Olbramovice) in 1927. In 1948, J. Horňanský
Sen. collected this species near the road between Olbramovice and Branišovice, i.e.
roughly in the same place. In 1951, F. Kühn added a record of Tripolium pannonicum
from a ditch north-east of Branišovice (Šmarda 1953). Spergularia marina was collected
near Branišovice by J. Vicherek in 1962 and for the second time by L. Bravencová in
2007, which is the latest record of any halophyte from our list in this district. To our
knowledge, no halophytic vegetation has been preserved in the district.
Hevlín-Nový Přerov district
The Hevlín-Nový Přerov district includes halophytic sites adjacent to the left side of the
Dyje river floodplain between the villages of Dyjákovice (Tajax), Hevlín (Höflein),
Hrabětice (Grafendorf) and Jevišovka (Fröllersdorf, Frélichov), halophytic sites adjacent
to the right side of the Dyje floodplain between Nový Přerov (Neu-Prerau) and
Novosedly (Neusiedl, Nové Sídlo), as well as those in a shallow basin drained by the
Mikulovka/Polní stream east of the town of Mikulov (Nikolsburg). Most sites were in
southern Moravia, but a few halophytes also occurred in the adjacent part of Lower Aus-
tria east of the settlement of Alt-Prerau. Grulich (1987) also included in this district the
area with halophytes west of the town of Mikulov, but we consider it part of the Sedlec
district because it is drained by the Včelínek stream.
The halophytic flora of the Hevlín-Nový Přerov district was originally rich in species
and included all the 17 halophytes except Galatella cana and Triglochin maritima.The
first records of halophytes from this area date back to the early 1840s. Siegfried Reissek
(Reisseck 1841) found Salicornia perennans,Spergularia marina and Suaeda prostrata
on the way from the town of Drnholec (Dürnholz) to the village of Dobré Pole (Gutten-
feld), probably somewhere near Novosedly. However, the exact location of the site is
unknown as the description in the original source does not match the locations of halo-
phytic sites as currently known.
The halophytic site in Nový Přerov was discovered by A. Oborny in 1883. He col-
lected there Bupleurum tenuissimum,Crypsis aculeata,C. schoenoides,Salicornia
perennans,Samolus valerandi,Spergularia media,Suaeda prostrata and Taraxacum
bessarabicum. Schierl (1896) reported the occurrence of Glaux maritima. All these
records refer to the surroundings of a small fishpond in the northern part of the village and
saline meadows between the northern part of the village and the railway from Novosedly
to the town of Laa an der Thaya in Lower Austria, abandoned in the 1920s.
In 1920 the site was explored by H. Iltis, a secondary school teacher in Brno. He col-
lected most of the species found there once by A. Oborny, including both succulent halo-
phytes and, in addition, Glaux maritima. In 1922, J. Bílý Jr. and S. Staněk collected Bupleu-
rum tenuissimum,Plantago maritima,Salicornia perennans,Scorzonera parviflora,
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 45
Spergularia media and Suaeda prostrata in saline meadows between Nový Přerov and
Jevišovka. It is unclear whether these finds are from the same site as those collected by A.
Oborny and H. Iltis, as the site is referred to as “beyond the railway”, i.e. possibly
between the abandoned railway and the Dyje floodplain.
For the subsequent decades, the Nový Přerov halophytic site was almost neglected.
Šmarda (1953) did not visit it. Jiří Vicherek studied its vegetation probably in 1962. His
relevés (Vicherek 1973) contained Bupleurum tenuissimum,Crypsis aculeata,Glaux
maritima,Juncus gerardii,Plantago maritima,Spergularia marina,S. media,Taraxa-
cum bessarabicum and Tripolium pannonicum. In 1980–1984, V. Grulich collected there
herbarium specimens of Bupleurum tenuissimum,Juncus gerardii,Plantago maritima,
Spergularia marina,Taraxacum bessarabicum and Tripolium pannonicum. At that time,
the halophytes were confined to the bottom and shores of a small fishpond at the village's
northern edge. Vicherek returned to the site in 1994 and collected Bupleurum tenuis-
simum and Plantago maritima. The last record of Tripolium pannonicum dates back to
1992. The fragments of saline habitats south-west of the fishpond were destroyed by
afforestation in 1998. However, a small population of Plantago maritima still survives in
trampled places ~250 m NNE of the church.
The occurrences of halophytes in Novosedly or nearby, possibly observed already by
S. Reissek, were rediscovered in 1883 by A. Ripper, a theology student, and A. Oborny,
who found Crypsis aculeata and C. schoenoides somewhere between Jevišovka and
Novosedly (Oborny 1883). In 1884, H. Zimmermann collected Crypsis aculeata in the
village or nearby. Probably at the same time, A. Ripper (sec. Oborny 1912) recorded
Glaux maritima directly in the village (“auf Dorfanger”). Šuk (1956) described the halo-
phytic vegetation in the surroundings of Novosedly as nearly destroyed by intensive
grazing. He listed only a few subhalophytes. Jiří Vicherek studied the vegetation of this
site probably in 1962. His relevés contained Bupleurum tenuissimum,Crypsis aculeata,
Juncus gerardii,Plantago maritima,Spergularia marina,S. media,Taraxacum bessarabi-
cum and Tripolium pannonicum (Vicherek 1973). Vít Grulich visited the site in 1982 and
1983 and collected Crypsis aculeata,C. schoenoides,Juncus gerardii,Spergularia
marina,S. media and Taraxacum bessarabicum. He considered the site relatively well
preserved (Grulich 1987). Since the 1980s, the site has been repeatedly visited by bota-
nists. In 1993, a nature reserve was established there to protect the remaining occurrences
of halophytes. However, most of them disappeared due to the lack of management, and
Crypsis schoenoides is the only halophyte still found there, though irregularly.
There were other halophytic sites near Novosedly in the past. For instance, H. Zim-
mermann collected Salicornia perennans and Crypsis schoenoides near Jevišovka in
1884. This site may have been located either in a common pasture ~1.5 km W of the vil-
lage or, less likely, on the right bank of the Dyje river towards Nový Přerov. Roughly at
the same time, A. Ripper found Bupleurum tenuissimum and Tripolium pannonicum in
common pastures between Jevišovka and the town of Hrušovany nad Jevišovkou
(Grußbach), possibly in the same place where H. Zimmermann found Crypsis and
Salicornia,andGlaux maritima even further towards the east near Hrušovany nad
Jevišovkou. A record of Plantago maritima and Tripolium pannonicum from 1949 (or
already from the 1920s?) in a common pasture near Jevišovka (J. Bílý sec. Šmarda 1953)
may also refer to this site. However, all these occurrences were destroyed a long time ago.
46 Preslia 94: 13–110, 2022
The halophytic site at the village of Dobré Pole was also discovered by A. Ripper, who
recorded abundant populations of Glaux maritima and Bupleurum tenuissimum;healso
found the former species near the village of Březí (Brattlesbrunn). In 1934, O. Thenius
and F. Teuber, hobby botanists from Brno, collected Glaux maritima,Spergularia
marina and S. media near a small fishpond in Dobré Pole, which is the halophytic site
preserved until now. Anton Fröhlich collected Glaux maritima in Dobré Pole repeatedly
between 1924 and 1954, Tripolium pannonicum in 1941 and a non-dated specimen of
Spergularia marina. All of these 20th-century herbarium records certainly originated
from the area of the current nature reserve at the southern edge of the village. Šmarda
(1953) did not visit this site. He only reported the occurrence of Glaux maritima (“in
large quantities”) based on the information from J. Bílý Jr. The undocumented record of
Cirsium brachycephalum, reportedly based on the information provided to him by F.
Švestka, is likely erroneous. Vratislav Šuk, in theearly 1950s working as a pharmacist in
Mikulov, listed Glaux maritima,Plantago maritima,Spergularia media,Taraxacum
bessarabicum and Tripolium pannonicum from the Dobré Pole site (Šuk 1956). Jiří
Vicherek recorded no relevés at this site (see Vicherek 1973). Vít Grulich visited the site
repeatedly during the 1980s and documented the occurrence of Glaux maritima,
Plantago maritima,Spergularia marina,S. media,Taraxacum bessarabicum and
Tripolium pannonicum. He considered the site relatively well preserved (Grulich 1987)
and recommended it for conservation. In the early 1990s, Bupleurum tenuissimum was
rediscovered at the Dobré Pole site after more than a century; the latest record of this
species is from 2008.
The occurrences of halophytes in the western part of the district were also discovered
by A. Ripper. Probably in 1884, he found Bupleurum tenuissimum and Tripolium
pannonicum near Hevlín and B. tenuissimum near the Travní dvůr (Trabinghof) farm-
stead west of Hrabětice. The site north of Hevlín was rediscovered half a century later by
Tomaschek (1933). In a saline meadow located ~1.1 km NNE of the village (currently,
there is a brick factory), Tomaschek confirmed the presence of Tripolium pannonicum
and added Plantago maritima and Taraxacum bessarabicum as new records. In addition,
he found Plantago maritima south-west of Travní Dvůr and discovered the westernmost
halophytic site of this district ~1 km NW of Dyjákovice. At the latter site, described as
a small saline meadow with ditches, he found Plantago maritima,Scorzonera parviflora,
Spergularia marina,S. media and Tripolium pannonicum, accompanied by numerous
subhalophytes. Though no exact information is available, all these occurrences must have
vanished probably a long time ago. The only surviving halophyte in this part of the dis-
trict is Spergularia marina, collected near Hevlín between 1992 and 2012 (Němec et al.
2014). The find of Cirsium brachycephalum in an abandoned loam pit ~1.2 km ENE of
Hevlín (Dřevojan 2017; still present there in 2020) is the only reliable record of this spe-
cies from the Hevlín-Nový Přerov district.
The current halophytic flora in this district is enormously impoverished. The two suc-
culent halophytes, Salicornia perennans and Suaeda prostrata, were last collected in
1922. Samolus valerandi was last collected even a year earlier. Scorzonera parviflora
was last seen in the early 1930s. Crypsis aculeata was last seen at the Novosedly site in
1994. Glaux maritima,Plantago maritima,Spergularia marina,S. media,Taraxacum
bessarabicum and Tripolium pannonicum still survive in this district, but their occur-
rences have mostly been reduced to a single site.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 47
48 Preslia 94: 13–110, 2022
Fig. 6. Halophytic localities in the Hevlín-Nový Přerov and Pulkau districts: (A) Nový Přerov 2020, (B) Hevlín
2020, (C) Hintausacker 2020. Photo credits: E. Šmerdová (A, B), M. Chytrý (C).
Nový Přerov (Fig. 6A). – A remnant patch of halophytic and subhalophytic habitats is
located at the north-eastern edge of the village of Nový Přerov ~250 m NNE of the
church. Nowadays, Plantago maritima occurs there as the only halophyte. In 2020, the
area was grazed with a mixed herd of sheep, goats and one horse and covered by
a subhalophytic pasture with Achillea collina,Althaea officinalis,Dipsacus laciniatus,
Inula britannica,Lotus tenuis,Pulicaria dysenterica and Senecio erraticus. Subhalo-
phytic trampled grasslands with Lotus maritimus,L. tenuis,Pulicaria dysenterica and
Trifolium fragiferum (Loto tenuis-Potentilletum anserinae) also occur in the village
along pedestrian paths.
Dobré Pole (Fig. 5B). – The Dobré Pole saline site is located at the southern edge of the
village of Dobré Pole. A part of the area has been used as a football playground for
decades. In the 1980s, a pond at the saline site was deepened to collect waste and sewage
water from the village (Lysák 2016a). This intervention disrupted the water regime and,
together with some other measures, caused desalination of its soils. Since the disruption
of the water regime, the saline site has been drying out, and the herb layer cover is
increasing (Lysák 2016a). After a wastewater treatment plant was constructed in the vil-
lage, the pond sediment was removed and nowadays, it is only used for collecting the
wastewater. The football playground and its surroundings (~4 ha in total) were desig-
nated as the Slanisko Dobré Pole Nature Reserve in 1993. Trampling on the playground
and in the surroundings supports some competitively weak halophytes and subhalo-
phytes. The playground is covered mainly by the Loto tenuis-Potentilletum anserinae
association, and the heavily trampled spots, partly on the access road, are covered by the
Centaureo pannonicae-Festucetum pseudovinae and in places also by Puccinellietum
limosae grasslands. Glaux maritima is restricted to a depression north of the fishpond.
The area between the pond and the core of the saline site is covered by a subhalophytic
tall-forb community of Althaea officinalis,Dipsacus laciniatus and Pulicaria
dysenterica. This community also occurs in several slightly wet spots in the broader sur-
roundings of the saline site. Currently, the site is regularly mown. In addition, the areas
outside the playground are grazed as part of conservation management.
Novosedly (Fig. 5C). – This saline site is located at the north-eastern edge of Novosedly.
Its area was used by local people for hay making and poultry grazing. However, it was
gradually abandoned and changed into an illegal dumping place for waste and building
rubble. In the late 1980s, some digging and earth moving disrupted the water regime,
which may have caused desalination and supported the spread of Phragmites australis
(V. Grulich, in litt. 2021). Although the site was put under conservation in 1993 (Slanisko
Novosedly Nature Reserve), most of the remaining halophytes disappeared in the subse-
quent years due to the lack of management and succession of Phragmites australis and
other competitive grasses. In 2008, P. Šmarda recorded only subhalophytes Carex distans,
Chenopodium chenopodioides,Melilotus dentatus,Pulicaria dysenterica,Schoenoplectus
tabernaemontani and Trifolium fragiferum (Šmarda 2008), but none of the 17 halophytes
analyzed in the current study. Still, Taraxacum bessarabicum was collected there a year
later. In 2011, dense reed stands were removed together with the topsoil, waste and build-
ing rubble. Since then, the site has been regularly grazed for conservation purposes
(Lysák 2016b), which resulted in the re-appearance of Crypsis schoenoides. In 2020, the
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 49
area was covered mainly by subhalophytic grassland of the Loto tenuis-Potentilletum
anserinae association dominated by Agrostis stolonifera,Carex otrubae,Deschampsia
cespitosa,Festuca pratensis agg. and Potentilla anserina, with the occurrence of Carex
distans,Lotus maritimus,L. tenuis,Melilotus dentatus,Pulicaria dysenterica and
Trifolium fragiferum. An artificial depression was created in the central part of the site to
support halophytes. In favourable years, its bottom is inhabited by Atriplex prostrata,
Carex secalina,Chenopodium chenopodioides and Crypsis schoenoides (Lysák 2016b).
The peripheral parts of the reserve arecoveredbysemi-drygrasslandswithAchillea
collina,Agrimonia eupatoria,Brachypodium pinnatum,Centaurea jacea,Festuca rubra,
F. rupicola,Origanum vulgare and Petrorhagia prolifera.
Hevlín (Fig. 6B). – This site, an abandoned and flooded sand or loam pit ~1.2 km ENE of
Hevlín, is indicated as such already on the 3rd Military Survey map (1876–1878). No
detailed information about its flora had been available until recently. In 2014, reed beds
were partly removed from its northern and southern parts. In 2016, P. Dřevojan found
there a population of Cirsium brachycephalum (Dřevojan 2017). Although this species
has never been reported from this area, it may have been present there for a long time.
Conservation management was introduced in 2020, aimed at reducing reed stands at the
C. brachycephalum sites.
Daníž district
The Daníž district includes the halophytic sites once found along the Daníž stream south-
east of Znojmo (Znaim) between the town of Šatov (Schattau) in the west and the village
of Jaroslavice (Joslowitz) in the east, as well as along the Vrbovecký stream, a tributary
of Daníž. Although Plantago maritima was found at the northern edge of the district near
the village of Vrbovec (Urbau) as early as 1871 (Oborny 1872), the halophyte-rich sites
along the Daníž had remained unknown. They were discovered in the late 1920s or early
1930s by O. Tomaschek, a primary school teacher in Jaroslavice. His report (Tomaschek
1933) comprehensively summarized the distribution of halophytes in the Znojmo admin-
istrative district, representing the westernmost occurrence of halophytes in Moravia. He
provided detailed descriptions of the four most valuable halophytic sites near the villages
of Jaroslavice, Strachotice (Rausenbruck), Dyjákovičky (Klein-Tajax) and Šatov. How-
ever, halophytes were more widespread within the district (for instance, he recorded
Plantago maritima at 14 additional sites). František Švestka collected Glaux maritima in
Šatov in 1946 and near the village of Chvalovice in 1947. However, no other information
is available about the halophytic flora and vegetation at these sites. Of the 17 halophytes
dealt with in this study, Glaux maritima,Juncus gerardii,Plantago maritima,Scorzonera
parviflora,Spergularia marina,Taraxacum bessarabicum,andTriglochin maritima
used to occur in the Daníž district.
The largest halophytic grassland once existed ~3 km WSW to 2 km WNW of
Jaroslavice, in a valley of a right-side tributary of the Daníž stream, lined by steep hill-
sides in the south-east. Tomaschek (1933) recorded there Glaux maritima,Plantago
maritima,Scorzonera parviflora,Taraxacum bessarabicum and Triglochin maritima,as
well as numerous subhalophytes. In addition, Juncus gerardii was collected there
in 1947. The occurrences of all the species found by Tomaschek (1933) were allegedly
50 Preslia 94: 13–110, 2022
confirmed by Švestka (1947a); however, his brief report published in a natural history
journal for teachers may have been entirely based on the records published by Tomaschek
(1933).
The next halophytic site upstream along the Daníž was located south-west of Stra-
chotice (Rausenbruck) near the road to the hamlet of Hnízdo (Gnast). Glaux maritima,
Scorzonera parviflora and Plantago maritima, accompanied by some subhalophytes,
occurred in this saline grassland surrounded by arable fields (Tomaschek 1933).
The third halophytic site was located near a brick factory south of the village of
Dyjákovičky. Tomaschek (1933) observed Plantago maritima and Taraxacum bessarabi-
cum there, the latter in large numbers, and five subhalophytes. Halophytic vegetation also
occurred in grazed places in the village. In 1963 or 1964, J. Vicherek recorded there five
relevés containing Glaux maritima,Spergularia marina and Taraxacum bessarabicum
(Vicherek 1973).
The westernmost halophytic site, located between the villages of Šatov and Hnanice
(Gnadlersdorf), hosted fewer halophytes than the previous sites, and the only halophyte,
noted by Tomaschek (1933) and Švestka (1947b), was Glaux maritima, accompanied by
some subhalophytes.
The halophytic vegetation in the Daníž district disappeared probably during the 1950s
and 1960s. In the early 1960s, Vicherek found halophytic vegetation worth sampling
only in the village of Dyjákovičky (Vicherek 1973). Grulich (1987) reported that all the
halophytes that formerly occurred in the Daníž district had vanished. However, at least
Juncus gerardii still survives at a handful of sites, as documented by the records from the
eastern edge of the village of Šatov (2012), the southern edge of the village of Vrbovec
(2011) and the former halophytic site south-west of Strachotice (2011; Němec et al. 2014).
Pulkau district
This district lies in the shallow valley of the Pulkau stream in northern Lower Austria. It is
a long and narrow area between the village of Zellerndorf (south of the town of Retz) and
the town of Laa an der Thaya, as well as in the shallow depression of the Entersgraben
stream between Laa an der Thaya and the town of Staatz. The best-developed halophytic
vegetation occurred in the past between the town of Seefeld-Kadolz and the village of
Wulzeshofen, particularly around the village of Zwingendorf.
The 3rd Military Survey maps from the early 1870s show numerous large tracts of
meadows and common pastures along the Pulkau stream and its tributaries. Some of these
meadows and especially common pastures, particularly in the eastern part of the district,
were likely saline and hosted halophytic species, including both succulent halophytes.
However, the stream channelization to prevent floods started already in the 1830s
(Kalbrunner 1855). Since the 1930s, large areas have been drained. The groundwater
level dropped by 0.5 m or more, and large tracts of meadows and pastureswere converted
to arable fields (Oberleitner et al. 2006). Consequently, many saline habitats and occur-
rences of halophytes were probably destroyed before being described by botanists. In
addition, the occurrences of halophytes in the Pulkau valley have always been considered
less attractive and less visited by Austrian botanists than those around Lake Neusiedl.
Therefore, they received less attention, and the available information is scarce.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 51
The flora of the Pulkau district used to be rich in halophytes. It included Bupleurum
tenuissimum,Crypsis aculeata,C. schoenoides,Glaux maritima,Juncus gerardii,
Plantago maritima,Salicornia perennans,Samolus valerandi,Scorzonera parviflora,
Taraxacum bessarabicum,Spergularia marina,S. media and Suaeda prostrata.Ithas
always been the only area in Austria with the occurrence of Glaux maritima.
Based on the opinion of Neilreich (1859), the earliest record of a halophyte species in
the Pulkau district may be found in Schultes (1814), who reported the occurrence of
Salicornia perennans (S. herbacea) near the town of Ernstbrunn. Although Ernstbrunn is
located ~20 km S of the Pulkau valley, this is a reasonable interpretation of Schultes’
record. Dolliner (1842) found Glaux maritima in wet places around the village of Staatz.
The occurrences of most halophytes in the district were discovered by H. Kalbrunner,
a pharmacist and the mayor of the town of Langenlois (Kalbrunner 1855). In the village
of Mailberg, he observed Glaux maritima, covering the shores of the Schlossteich Fish-
pond, and near the village, he also found Scorzonera parviflora and Spergularia marina.
Glaux maritima was also very common along the Pulkau between the town of Seefeld-
Kadolz and the village of Wulzeshofen. Kalbruner further found Plantago maritima
around the village of Zwingendorf, and Suaeda prostrata (Schoberia maritima) around
the village of Hadres. Based on information from A. Unger, Neilreich (1859) added the
records of Crypsis aculeata from the area between Staatz and Laa an der Thaya and the
surroundings of Zwingendorf. Based on a record by M. Matz, he reported Salicornia
perennans to occur between the villages of Platt and Watzelsdorf south of the town of
Retz and between the village of Haugsdorf and Laa an der Thaya, in places very abun-
dant. He added the records of Suaeda prostrata from near Seefeld, Zwingendorf and Laa
an der Thaya, a record of Spergularia marina near Zwingendorf, and records of S. media
along the Pulkau stream from Mailberg as far as Laa an der Thaya. According to the same
source, Plantago maritima occurred all over the district.
The eastern part of the Pulkau district became easily accessible by train from Vienna
in 1870 when a new railway line Vienna–Brno was put in operation. In July 1871, the
halophytic flora between Laa an der Thaya and Seefeld was explored by AugustL. Reuss,
his brother and their friends (Reuss 1873). They observed halophytes in ditches and field
margins along the road from Laa an der Thaya to Wulzeshofen. Further westwards,
between Wulzeshofen and Zwingendorf, they reported large tracts of infertile ground, in
many places with a thick layer of salt efflorescences. Halophytes were becoming less fre-
quent towards Seefeld. They collected Cirsium brachycephalum in road ditches and in
a reed bed north of the road Laa an der Thaya–Wulzeshofen, Crypsis schoenoides and
Juncus gerardii between Wulzeshofen and Zwingendorf, Salicornia perennans in ditches
between Hanfthal and Wulzeshofen, Samolus valerandi in a ditch, Scorzonera parviflora
and Tripolium pannonicum in a dried-out wetland between Wulzeshofen and Zwingen-
dorf, and Spergularia marina and S. media between Laa an der Thaya and Wulzeshofen.
Spergularia media occurred as far as Zwingendorf. Plantago maritima was common,
while Glaux maritima occurred in large quantities in the villages and along road ditches
from Wulzeshofen as far as Zwingendorf and beyond. Reuss (1873) concluded that there
was no other area in Lower Austria with such well-developed halophytic flora.
For the next 100 years, a small number of plant records are available from the Pulkau
district. In the late 1870s or early 1880s, Scorzonera parviflora was recorded near Laa
and der Thaya by H. Braun, and near Kadolz by A. Oborny (Halácsy & Braun 1882).
52 Preslia 94: 13–110, 2022
Roughly at the same time, K. Rechinger collected Glaux maritima in Zwingendorf for the
exsiccate series Flora exsiccata austro-hungarica. In 1897, J. Haring collected Tripolium
pannonicum in a meadow near the Zellerndorf railway station, roughly in the same place
where M. Matz recorded Salicornia perennans in the 1850s. Probably in 1904, A. Teyber
found Bupleurum tenuissimum in Zwingendorf, which was the first record of this species
in the district. In 1953, H. Melzer recorded Glaux maritima,Salicornia perennans,
Scorzonera parviflora,Suaeda prostrata and Tripolium pannonicum on the bottom of
a former fishpond, probably at the Hintausacker site near Zwingendorf, and also found
Cirsium brachycephalum somewhere nearby (Melzer 1955). In 1960, he collected
Salicornia perennans and Suaeda prostrata at the site known as Saliterweide (also
Saliterwiese or Saliterheide) ~2.4 km ESE of Zwingendorf.
The Hintausacker site at the NW edge of Zwingendorf was still used as a goose pasture
in the 1970s. However, the number of grazing animals dropped, and some parts of the site
were used as a waste place. Waste was repeatedly removed from the area. In 1979, the
Hintausacker site was purchased by the Land Niederösterreich and, together with
Saliterweide, included in the newly designated Zwingendorfer Glaubersalzböden Nature
Reserve (Naturschutzgebiet). In the mid-1990s, the site was cleaned from waste again,
bulldozed, and some earth was removed. Reed stands were reduced by repeated mowing,
and some spontaneously established trees and shrubs were cut. In contrast, Saliterweide
was left almost unmanaged, except for some interventions into the water regime, such as
removing subsurface drainage pipes and enlarging the pond (Holzer et al. 2002).
From the mid-1920s to the 1970s, halophytic flora in the area was documented by
Jurasky (1980). At the Hintausacker site, he recorded Glaux maritima,Plantago
maritima,Salicornia perennans,Scorzonera parviflora,Spergularia marina,S. media,
Suaeda prostrata,Taraxacum bessarabicum and Tripolium pannonicum.Awetdepres-
sion ~1 km E of Zwingendorf, north of the road to Wulzeshofen, probably a former ice
pond, harboured both the succulent halophytes and Scorzonera parviflora,Spergularia
media and Tripolium pannonicum. This habitat was later destroyed by filling with
earth (Oberleitner et al. 2006). In contrast, at the Saliterweide site, Jurasky only found
Plantago maritima, but he reported the occurrence of salt efflorescences at the margins of
adjacent arable fields. Hütterer & Albert (1993) sampled Glaux maritima,Juncus
gerardii,Plantago maritima,Spergularia media,Taraxacum bessarabicum and Tripolium
pannonicum for an ecophysiological study at the Hintausacker site in 1989. They also
sketched a vegetation map of the site, which roughly corresponds to the situation as
observed in 2020. A shallow pond in the north-eastern corner of the site was surrounded
by Phragmites australis stands except for the southern side, which was probably
disturbed by horses, poultry or people and fringed by the Astero pannonici-Bolbo-
schoenetum compacti vegetation. The next vegetation zone in slightly drier places was
Loto tenuis-Potentilletum anserinae with Melilotus dentatus and Tripolium pannonicum
and trampled (perhaps also grazed) Potentilla anserina grassland, in places with Glaux
maritima and Inula britannica.
Holzer et al. (2002) recorded Glaux maritima,Juncus gerardii,Plantago maritima,
Spergularia media and Tripolium pannonicum at the Hintausacker site in 2001. They
reported a mosaic of various vegetation types, including shallow ponds either with
Bolboschoenus marshes or completely occupied with Phragmites reed beds, open stands
with Glaux maritima, wet to intermittently wet saline meadows, monodominant stands of
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 53
Calamagrostis epigejos and a young self-established stand of Fraxinus excelsior and
Acer negundo. Some halophytes were also present on an abandoned arable field south-
west of the reserve. Cirsium brachycephalum,Juncus gerardii,Plantago maritima and
Tripolium pannonicum still occurred at Saliterweide. Holzer et al. (2002) discovered an
additional halophytic site between the farmstead Alicenhof and the western edge of
Zwingendorf, including a large reed bed with Phragmites australis and a saline meadow.
Of the halophytes, Glaux maritima,Plantago maritima and Tripolium pannonicum were
present at this site. In summer 2021, we found none of these species there.
The current halophytic flora of the Pulkau district is strongly impoverished and
includes Cirsium brachycephalum (last record from 2016), Glaux maritima,Plantago
maritima,Scorzonera parviflora,Spergularia marina,S. media and Tripolium panno-
nicum. Locally disappeared halophytes include Crypsis aculeata and C. schoenoides
(last recorded in the late 19th century), Salicornia perennans and Suaeda prostrata (per-
sisting until the late 1970s or early 1980s), Taraxacum bessarabicum (last observed
about ten years later) and Samolus valerandi (last record in 1988). All the remaining
occurrences of halophytes in the Pulkau district are confined to both Zwingendorf sites,
especially Hintausacker. However, subhalophytes still occur at other sites in the Pulkau
district. In July 2021, we observed Bolboschoenus maritimus,Centaurium pulchellum,
Lotus tenuis and Melilotus dentatus in wet places on arable land 1.5 km E of Obritz,
Bolboschoenus maritimus,Carex otrubae,Lotus tenuis,Melilotus dentatus,Puccinellia
distans,Rumex stenophyllus and Trifolium fragiferum in the sports grounds and around
ponds at the southern edge of Seefeld-Kadolz, and Achillea asplenifolia,Bolboschoenus
maritimus,Carex distans,C. otrubae,Lotus maritimus,L. tenuis,Melilotus dentatus and
Rumex stenophyllus in reed beds and meadows between the western edge of Zwingendorf
and the Alicenhof farmstead.
Hintausacker (Fig. 6C). – This last well-preserved site of halophytic vegetation in the Pulkau
district is located at the north-western edge of the village of Zwingendorf north to north-east
of the church. It is one of the two parts of the protected area Zwingendorfer Glaubersalzböden
with an area of 5.38 ha. Nowadays, it is the only Austrian locality of Glaux maritima.The
southern part of the site is covered by a mesic meadow dominated by Festuca arundinacea
and Arrhenatherum elatius. The most widespread halophytic vegetation at this site corre-
sponds to the association Loto tenuis-Potentilletum anserinae. The meadows and sub-
halophytic grasslands are left unmown or mown in longer than annualintervals, which allows
tall grasses and herbs to dominate and suppress competitively weak halophytes. Most halo-
phytes and subhalophytes, including Carex distans,C. secalina,Centaurium pulchel-
lum,Glaux maritima,Juncus gerardii,Plantago maritima,Puccinellia distans,Scorzo-
nera parviflora,Spergularia media and Tripolium pannonicum occur in the Puccinellietum
limosae vegetation in shallow depressions with relatively sparse vegetation, including the
places where topsoil was removed. At wetter places, there are also patches of saline marshes
with Bolboschoenus maritimus (Astero pannonici-Bolboschoenetum compacti). Two
shallow ponds in the northern part of the site are surrounded by stands of Phragmites aus-
tralis,Bolboschoenus maritimus agg. and Carex riparia.
Saliterweide. – This abandoned pasture is located between arable fields ~2.4 km ESE of
the church in Zwingendorf, south of the road to Wulzeshofen. It is the second of the two
54 Preslia 94: 13–110, 2022
parts of the protected area Zwingendorfer Glaubersalzböden with an area of 10.33 ha.
The largest part of the site is covered with an unmanaged grassland with Agrostis
gigantea,A. stolonifera,Dactylis glomerata,Elytrigia repens,Festuca rupicola and Poa
angustifolia, here and there with individuals of Plantago maritima. Trampled places and
grassy roads are dominated by Festuca pulchra with abundant Plantago maritima (asso-
ciation Centaureo pannonicae-Festucetum pseudovinae).Stands of Phragmites australis
and rich populations of Bolboschoenus maritimus,Lotus tenuis and Plantago maritima
occur around a small pond in the south-western part of the site. Cirsium brachycephalum
is reported from this site, but we did not find it in July 2021.
Other occurrences of halophytes
Some occurrences of halophytes were also recorded at a few sites outside the above-
described districts. Plantago maritima,Taraxacum bessarabicum,Tripolium pannonicum,
and probably also Juncus gerardii were recorded in a shallow depression of the Jevišovka
stream near or in the village of Stošíkovice na Louce north-east of the town of Znojmo
in the early 1950s (Šmarda 1953). Juncus gerardii and Tripolium pannonicum once
occurred around a fishpond at the south-eastern edge of the village of Hodonice east of
Znojmo (Šmarda 1953). In the 1880s, Taraxacum bessarabicum and Tripolium panno-
nicum were found on common pastures somewhere between the towns of Břeclav and
Lanžhot and the village of Moravská Nová Ves (A. Ripper in Oborny 1912). The exact
location of this site is unknown. While J. gerardii may be still present at the Hodonice site
(last collected in 2013), all the remaining occurrences likely disappeared a long time ago.
Species
Here we review the historical and current distribution and ecology of 17 halophytes that
are (or were) significantly related to saline habitats in Moravia. Their Red List classifica-
tion in the Czech Republic and Austria is given in brackets after the species names, fol-
lowing Grulich (2017) and Niklfeld (1999), respectively: RE – regionally extinct; CR –
critically endangered, EN – endangered; VU – vulnerable.
The presence of these species in the halophytic districts of southern Moravia and the
Pulkau valley since the 1820s is summarized in Figs. 7 and 8. Individual occurrence
records mentioned in the text and shown in these figures are summarized in Supplemen-
tary Data S2. Although we prepared the occurrence dataset with extraordinary care, it is
most likely not free of mistakes. Labelling mistakes in herbaria are probably more fre-
quent in halophytes than in other vascular plants. As halophytes have never been wide-
spread, they were often collected for exchange or used as a present to other botanists. The
new owners sometimes replaced the original locality information with new labels, which
may contain imprecise information. Also, specimens collected during excursions may
have been labelled without knowledge of local topography and reliable maps. There are
also errors in the collection date. In addition, some botanists collecting in large quantities
and selling their specimens to museums, e.g. F. Weber, may have deliberately misdated
specimens to reduce the number of duplicates. For these reasons, if a single record or
specimen of a species exists from a particular site, the occurrence of this species should
be considered uncertain. The same holds for temporal outliers.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 55
56 Preslia 94: 13–110, 2022
1969
1969
1977
1981
1963
1942
1967
1971
1994
1981
1821
1997
1923
1965
1931
1932
1967
1976
1922
1980
1976
1963
1980
1970
1933
1933
1978
1976
1930
1934
1970
1965
1960
1963
1966
1963
1989
1973
1977
1971
1980
1967
1952
1953
1824
1960
1960
1963
1959
1964
1890
1960
1952
1964
1963
1980
1984
1981
1977
1952
1947
1921
1988
1946
1973
1927
1921
1960
1983
1922
1980
1923
1947
Tripolium pannonicum
Taraxacum bessarabicum
Triglochin maritima
Spergularia media
Suaeda prostrata
Scorzonera parviflora Spergularia marina
Salicornia perennans Samolus valerandi
Juncus gerardii Plantago maritima
Galatella cana Glaux maritima
Crypsis aculeata Crypsis schoenoides
Bupleurum tenuissimum Cirsium brachycephalum
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020 1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
Fig. 7. Recorded occurrences of 17 halophytic species in individual halophytic districts. Red crosses with
a date indicate the last record for locally extirpated species, defined as those not recorded in the district after
2000. Many records are not shown because they are not dated. This applies particularly to herbarium specimens
from the 19th century.
Bupleurum tenuissimum (Red Lists: CZ: CR; AT: EN)
Bupleurum tenuissimum is an annual herb occurring in the western Mediterranean, in the
Atlantic parts of Europe from Spain to Denmark, including Great Britain, the western
Baltic area, central Germany and the Pannonian region, with scattered occurrences
towards the east reaching as far as southern Ukraine (Meusel et al. 1978).
This species is a competitively weak halophyte usually growing in open, often dis-
turbed vegetation. In the study area, it was recorded in nearly all the types of saline grass-
lands of the class Festuco-Puccinellietea.
Bupleurum tenuissimum was present in all the halophytic districts except Damnice-
Olbramovice and Daníž. The last records in the Rakvice and Šakvice districts date back to
1951 and 1953, respectively. In the Čejč district, it was last collected in 1977. In the
Pulkau district, it was discovered near the village of Zwingendorf in 1904 by A. Teyber
(Anonymous 1905) and re-recorded roughly at the same site in 1981. Currently, the larg-
est population of this species, comprising thousands of individuals, occurs at the Slanisko
u Nesytu site in the Sedlec district. There are also recent records from the Měnín-Šaratice,
Velké Němčice and Hevlín-Nový Přerov districts, in each of them from a single site, but the
populations are small. Due to its inconspicuous appearance, this plant can be easily over-
looked in the field, and the size of its populations fluctuates depending on the weather of
a particular year. Therefore, it may have survived unnoticed at some additional sites.
Apart from the occurrences in the halophytic districts, Bupleurum tenuissimum was
once found near the village of Komárov at the southern periphery of the city of Brno
(A. Makowsky sec. Oborny 1886) and at two sites near the village of Popovice east of the
town of Uherské Hradiště (Staněk 1926). However, both occurrences have not been con-
firmed since then.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 57
12
1
1
7
13
7
1
5
1
1
7
12
5
3
3
9
4
1
2
9
14
9
13
5
13
3
1
10
8
15
12
1
14
13
11
9
8
2
16
11
11
15
15
14
13
1
12
5
16
11
13
10
11
15
13
2
4
5
6
16
9
9
6
11
13
13
1
8
2
10
15
2
1
2
5
2
12
10
10
15
5
3
4
5
5
12
1
10
1
8
14
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
Study area
1820 1840 1860 1880 1900 1920 1940 1960 1980 2000 2020
Fig. 8. Number of species of the 17 halophytes dealt in this study recorded per two-decade periods in individual
halophytic districts of southern Moravia and northern Lower Austria and across all the districts (Study area).
This Figure summarizes the records for individual species shown in Fig. 7. The period 2001–2020 also includes
several records from the year 2021.
Cirsium brachycephalum (Red Lists: CZ: CR, AT: EN; EU Habitats Directive 92/43/EEC)
Cirsium brachycephalum is a biennial to short-lived monocarpic species, seldom repro-
ducing vegetatively (Prokešová 2013). It is endemic to the Carpathian Basin, having most
localities in Hungary and southern Slovakia and the westernmost distribution limit in
southern Moravia and Lower Austria (Meusel & Jäger 1992).
Cirsium brachycephalum is a competitively weak and moisture-demanding species. It
prefers heavy, saline soils that are at least occasionally flooded. It is usually considered
a halophyte (Grulich 1987), although its affinity for saline habitats is not as strong as gen-
erally believed (Zlinská 2004, Prokešová 2013). It also grows in subhalophytic marshes
and meadows, margins of reed beds, wet fallows, wetlands on arable land and at other
occasionally disturbed sites (Bureš 2004). In the study area, it was recorded mainly in
saline marshes of the alliance Meliloto dentati-Bolboschoenion maritimi and saline
grasslands of the association Agrostio stoloniferae-Juncetum ranarii.
Unlike most other halophytes, Cirsium brachycephalum hasalwaysbeenrareinthe
study area. It was first collected on the shores of the Měnín Fishpond between 1817 and
1821 (Hochstetter 1825, see also Danihelka in Hadinec & Lustyk 2007). Reisseck (1841)
reported this species to occur somewhere north-east of the town of Lednice. Albin Wildt
found it near the village of Rakvice in 1909 and 1911. Currently, C. brachycephalum
occurs in a few places north-east of this village at the Trkmanec-Rybníčky and Trkmanské
louky sites. In the past, the species was also recorded in the ditches along the railway south-
east and north-west of Rakvice and in the Dyje floodplain south of the village (herbarium
specimens and records from 1950–1969). There is also a herbarium specimen allegedly
collected near the village of Terezín (F. Weber 1930 PR) and a record from a relevé sam-
pled north of Terezín in 1958–1961 (Vicherek 1962), both in the Čejč district. In addition,
C. brachycephalum was recorded in a saline marsh north-west of the village of Popice in
the Šakvice district in 1958–1961 (Vicherek 1962). A single record from the village of
Dobré Pole (F. Švestka sec. Šmarda 1953) in the Hevlín-Nový Přerov district is probably
erroneous. The recent find of C. brachycephalum near the village of Hevlín (Dřevojan
2017) is the first reliable record of this species in that district. Currently, it is the only site of
this species in the Czech Republic besides the sites near Rakvice. This site is close to the
occurrences in the Pulkau district in Lower Austria. In this district, the species was reported
from the village of Unternalb south of the town of Retz (Adler et al. 1996) and the area
between the village of Zwingendorf and the town of Laa an der Thaya. A small population
of C. brachycephalum was observed in 2016 at the Saliterweide site near Zwingendorf
(N. Sauberer sec. Dřevojan 2017), being the last population of this species in northern
Lower Austria. However, we did not find this species at Saliterweide in July 2021.
In addition to the occurrences described above, Cirsium brachycephalum was col-
lected in a fen meadow at the edge of the Morava river floodplain south of the village of
Moravský Písek in 1946. This is the only record from that site, and the occurrence proba-
bly vanished soon after being discovered.
Crypsis aculeata (Red Lists: CZ: CR; AT: VU)
Crypsis aculeata is an annual grass distributed in the submeridional and meridional zones
in Europe, Asia and northern Africa. It also occurs in some areas of the temperate zone
with continental climates. Its occurrences in Lower Austria, southern Moravia and south-
58 Preslia 94: 13–110, 2022
ern Slovakia are located at the northern limit of its range (Conert 1998, Danihelka &
Kaplan in Kaplan et al. 2020).
Typical habitats of this halophyte in Moravia are periodically flooded places, such as
temporary pools on pastures and arable land, shores and bottoms of drained fishponds
with saline water, and margins of saline reed beds. Crypsis aculeata is a dominant species
of the association Crypsietum aculeatae. It occurs on heavy soils rich in nitrogen and sol-
uble sulphates and chlorides.
Crypsis aculeata was once present in all the districts but Damnice-Olbramovice and
Daníž. However, it started to decline with the drainage of saline lakes and fishpondsin the
19th century. Its decline accelerated in the early 20th century with the drainage of saline
pastures followed by their conversion to arable land. Its occurrence at the Plácky site, the
only one in the Velké Němčice district, was last documented in 1942. The occurrence
near the village of Moutnice in the Měnín-Šaratice district (Vicherek 1973) vanished
probably soon after 1963. The last herbarium record from the Šakvice district dates back
to 1967. In the Rakvice district, C. aculeata was last collected in 1971 at the saline site
surrounding the former Fabián Fishpond, now filled with earth, in the town of Velké
Bílovice. In the Čejč district, it was recorded frequently until 1966, e.g. at the Zápověď
site near the village of Terezín, but was not found for the next few decades. In 2008, it was
rediscovered at Zápověď in a shallow pool dug in 2003 as a part of an ecological restora-
tion project (Hadinec & Lustyk 2009). The last record of this species at the Novosedly
site in the Hevlín-Nový Přerov district dates back to 1994 (J. Vicherek sec. Hanušová
1995). Only one small population, once established from seeds collected from another
site at the Nesyt Fishpond (Danihelka & Hanušová 1995), still survives at the Slanisko
u Nesytu site near the village of Sedlec. In addition, a few plants were seen on the drained
bottom near the southern bay of the Nesyt Fishpond in 2020. For the Pulkau district, only
two 19th-century records and one record from 1981 exist for C. aculeata.
Crypsis schoenoides (Red Lists: CZ: CR; AT: CR)
Crypsis schoenoides is an annual grass with a similar life history and distribution as
C. aculeata. It occurs in the submeridional and meridional zones and the continental part
of the temperate zone in Europe, western and Central Asia, Siberia, northern and subtrop-
ical Africa and Madagascar. The northern range limit of this species is in south-eastern
Austria, southern Moravia and southern Slovakia (Conert 1998, Danihelka & Kaplan in
Kaplan et al. 2020).
Crypsis schoenoides grows at periodically flooded sites such as shores and bottoms of
regularly drained fishponds and oxbows and temporary pools in saline grasslands and
arable fields. It is a dominant species of the association Heleochloëtum schoenoidis.It
grows on heavy soils rich in nitrogen, phosphorus, sulphates and chlorides. However, its
affinity for saline habitats is weaker than in C. aculeata (Šumberová 2007a).
Crypsis schoenoides was once present in all the districts but Damnice-Olbramovice
and Daníž. It declined similarly and for the same reasons as C. aculeata but slower
because of its weaker affinity for saline habitats. The last herbarium specimen in the
Velké Němčice district was collected in 1959, probably at the Plácky site. The last
records from the Měnín-Šaratice and Čejč districts date back to the 1960s. Only two 19th-
century records of C. schoenoides exist for the Pulkau district (Beck 1890–1893). After
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 59
2000, C. schoenoides was observed in saline habitats in four districts: In the Šakvice dis-
trict, where the last records were from 1977, two populations of this species were found
in summer 2021 on abandoned wet arable land along the Štinkovka stream between the
villages of Šakvice and Starovičky (Chytrý & Danihelka 2021). In the Rakvice district, it
still occurs at the Trkmanec-Rybníčky site. In the Hevlín-Nový Přerov district, it was
found at the Slanisko Novosedly site, and in the Sedlec district, it still occurs at the
Slanisko u Nesytu site and on the drained bottom of the Nesyt Fishpond, locally in large
quantities (Dřevojan et al. 2017).
In addition to the occurrences at sites described above, Crypsis schoenoides was
repeatedly recorded in 1994–2012 on gravel bars and in an oxbow of theDyje river south-
west of the town of Lanžhot.
Galatella cana (Red Lists: CZ: RE; AT: EN)
Galatella cana is a perennial clonal herb endemic to the western and southern margin of
the Pannonian Lowland. It was recorded in southern Moravia, eastern Austria, Hungary,
northern Serbia and south-western Romania (Wagenitz 1979, Grulich & Feráková 1999).
Galatella cana was found only once on the territory of the current Czech Republic
between 1817 and 1821 by Ch. F. Hochstetter, somewhere near the eastern shores of the
former Měnín Fishpond in the Měnín-Šaratice district (Hochstetter 1825). Hochstetter
described its habitat as follows: “In einem kleinen Gehölze seitwärts vom See steht eine
neue Pflanze für Deutschlands Flora, nämlich Aster canus W. et K., ebendaselbst Senecio
tenuifolius [i.e. S. erucifolius]...!”. This corresponds to the circumstances known from
the closest sites of this species near the village of Baumgarten an der March in Lower
Austria and at the Šúr site near Bratislava in south-western Slovakia (Grulich & Feráková
1999). For a long time, no herbarium specimen of this species from the Měnín site had
been known (Grulich 1987), and its former occurrence in Moravia was sometimes ques-
tioned. However, a specimen collected by Hochstetter near Měnín was found in the her-
barium of the University of Vienna in 2007 (Danihelka 2008). Other specimens from the
same site were later discovered in the State Museum of Natural History Stuttgart
(Danihelka 2009) and recently in the Silesian Museum in Opava, the latter with a tran-
scribed label. No records of G. cana exist from the Pulkau district.
Glaux maritima (Red Lists: CZ: CR; AT: CR)
Glaux maritima is a low-growing perennial herb with a circumpolar distribution. It
occurs mainly along the coasts of western and northern Europe, eastern Asia, and western
and eastern North America, as well as in inland saline habitats in areas with arid climates
in eastern Europe, Siberia and North America. It also occurs scattered in saline habitats
across central and western Europe (Meusel et al. 1978).
In the study area, Glaux maritima prefers wet saline sites, in the past grazed mainly by
geese and ducks, often in villages along ditches and streams and around fishponds
(Grulich 1987). It occurs in halophytic grasslands of the associations Puccinellietum
limosae,Scorzonero parviflorae-Juncetum gerardii and Loto tenuis-Potentilletum
anserinae.
Glaux maritima used to occur in most Moravian districts of halophytic flora. How-
ever, some of the records are uncertain. Only one herbarium specimen is known from the
60 Preslia 94: 13–110, 2022
Měnín-Šaratice district, allegedly collected near the village of Měnín in the late 19th cen-
tury; however, it could be a labelling mistake. Only three records exist from the Velké
Němčice district, all from relevés recorded by Vicherek (1962, 1973) at the southern edge
of the village of Velké Němčice and near the village of Starovice probably in 1960 and
1965. In the Šakvice district, G. maritima occurred near the Šakvice railway station and
in the village of Starovičky; at the latter site, it was last collected in 1952. From the
Rakvice district, G. maritima disappeared during the 1960s. The species was also abun-
dant in the Čejč district, but some occurrences disappeared already in the 19th century
after the lakes of Kobylí and Čejč were drained. At the Zápověď site near the village of
Terezín, G. maritima used to be collected frequently until 1965; thereafter, it disappeared
but was found again in 2007 after shallow pools were created in the former nature
reserve. The species was also present in at least three villages of the Damnice-
Olbramovice district; however, these occurrences, recorded most likely in the 1880s
(A. Ripper sec. Oborny 1912), disappeared probably in the early 20th century. Glaux
maritima still survives at the Dobré Pole site in the Hevlín-Nový Přerov district. In the
Daníž district, it was recorded in four villages, with the last record in the village of
Dyjákovičky dating back to 1962 or 1963 (Vicherek 1973). Itwas once widespread in the
Pulkau district; a large population still survives at the Hintausacker site near
Zwingendorf. There are also three undocumented records from the Sedlec district, but we
consider them erroneous. The earliest one (Formánek 1887) is probably based on a mis-
identification. One of the records in two different relevés by Vicherek (1962) is
a misprint, and the other one may be a location mistake.
In addition to the occurrences described above, there is an undocumented record of
Glaux maritima from the Hrubá louka site north of the town of Hodonín (J. Bílý sec.
Šmarda 1953) and an unclear record from the surroundings of Retz (Janchen 1977).
Juncus gerardii (Red Lists: CZ: CR; AT: VU)
Juncus gerardii is a perennial graminoid distributed along the coasts of Europe and the
eastern coasts of North America, as well as at inland saline sites of Europe, western tem-
perate Asia, Siberia and North America (Hultén & Fries 1986).
The species occurs in wet saline meadows and marshes, at mineral springs and rarely
in wetlands on arable land. It is also found on non-saline but mineral-rich clayey soil. In
southern Moravia, it frequently occurs in most types of saline vegetation, often dominat-
ing or co-dominating the saline grasslands of the Juncion gerardii alliance.
Juncus gerardii is very similar to the frequently co-occurring J. compressus, and both
species have often been misidentified. For instance, there are ~100 Juncus specimens origi-
nally identified as J. gerardii from the Czech Republic at BRNU, of which ~50 were
reidentified as J. compressus in our revision. Consequently, records of J. gerardii not
documented by a herbarium specimen, especially records from relevés, are unreliable.
For this reason, we accept only a few records of this species that are not supported by
a herbarium specimen, thus certainly underestimating the species’ frequency. In vegeta-
tion analyses, we merged J. compressus and J. gerardii into the J. compressus agg.
Among the 17 halophytes dealt with in this study, Juncus gerardii is the most wide-
spread species, both in southern Moravia and Lower Austria. It has several isolated
occurrences outside the ranges of other halophytes. Also, it is usually the last surviving
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 61
halophyte at former halophytic sites. Based on the inspected herbarium specimens and
field experience, we assume that in the past, it was present in all the districts, though there
is no herbarium specimen from the Damnice-Olbramovice district. There are also recent
records from all the other districts, most of them supported by herbarium specimens
collected after 1990.
Plantago maritima (Red Lists: CZ: CR; AT: VU)
Plantago maritima is a perennial clonal herb with a cosmopolitan but fragmented distri-
bution. In Europe, it occurs mainly along its western Atlantic and northern coasts, in the
mountains of its southern part and inland saline habitats in the areas with continental cli-
mates (Meusel et al. 1978). Three to six subspecies are recognised in Europe (Chater &
Cartier 1976, Marhold 2011). Plants occurring in the steppe zone of Eurasia are tradition-
ally separated as P. maritima subsp. ciliata (Chrtek 2000), which is the taxon found in the
study area.
Plantago maritima naturally occurs in grasslands on alkaline to neutral, heavy soils. In
the study area, it occurs in most types of halophytic vegetation, ranging from salt marshes
of the Meliloto dentati-Bolboschoenion maritimi alliance through annual saline vegetation
of the alliances Cypero-Spergularion salinae and Thero-Salicornion to the saline grass-
lands of the alliances Puccinellion limosae,Juncion gerardii and Festucion pseudovinae.
It is most frequent in the association Centaureo pannonicae-Festucetum pseudovinae.
Plantago maritima once occurred in all the districts of halophytic flora in the study
area. However, the last record from the Daníž district dates back to 1947 and that from the
Damnice-Olbramovice district to 1952. In the late 1970s and during the 1980s, its popula-
tions disappeared from the Velké Němčice, Šakvice, Rakvice and Čejč districts. The spe-
cies still occurs in the Měnín-Šaratice, Sedlec, Hevlín-Nový Přerov and Pulkau districts,
usually at one or two sites in each district. However, most populations consist of a small
number of individuals. The largest population survives at the Slanisko u Nesytu site.
In the study area, Plantago maritima occurred far beyond the ranges of other halo-
phytes except for Juncus gerardii, usually on heavy, saline to slightly saline or mineral-
rich soils. In the west, its range extended to the village of Podmolí west of the town of
Znojmo, in the north to the city of Brno, and in the eastthe towns of Kyjov and Hodonín.
However, none of the populations at the limits of its historical range has survived until
now. In addition, this species was recorded at two isolated sites in the Bílé Karpaty Mts;
the occurrence in an abandoned loam pit west of the town of Uherský Brod may still exist.
Recently, P. maritima is spreading along the roads where salt is used for winter de-icing
(Danihelka & Kaplan in Kaplan et al. 2018). However, no such records are available from
the study area.
Salicornia perennans (Red Lists: CZ: RE; AT: VU)
The taxonomy of the genus Salicornia is still insufficiently resolved (Kadereit et al.
2007). The populations once found in southern Moravia and Lower Austria are now
assigned to S. perennans (syn. S. prostrata), but records under the name S. herbacea in
earlier literature also refer to this species. It is an annual diploid succulent herb occurring
at inland saline sites from the Carpathian Basin in the west to the Aral Sea in the east
(Kadereit et al. 2007). The southern Moravian sites were at the north-western limit of its
range (Danihelka & Kaplan in Kaplan et al. 2017).
62 Preslia 94: 13–110, 2022
The southern Moravian stands of Salicornia perennans were confined to the parts of
saline sites with strongly saline soils that were flooded in the spring but usually turned
very dry in the mid-summer. The dominance of this species defined the association
Salicornietum prostratae, but it also co-occurred in stands dominated by the other annual
succulent halophyte species, Suaeda prostrata.
Salicornia perennans was once present in all the districts but Daníž. It was likely
abundant in pastures on the flat shores of the former Měnín Fishpond and Kobylí and
Čejč lakes (Hochstetter 1825, Krzisch 1859). Some populations survived at other sites,
but their decline accelerated in the early 20th century. After World War II, S. perennans
was only observed in four districts: two of the occurrences (both in nature reserves) van-
ished during the 1960s because of the drainage of the surrounding agricultural land. At
the Slanisko u Nesytu site in the Sedlec district, this species survived for another decade,
but the population size gradually diminished during the early 1970s (Grulich 1987). The
likely causes of population decline include abandonment (especially cessation of poultry
grazing), changes in water regime and desalination. In 1976, when the fate of the last sur-
viving Salicornia population became clear, the remaining five dwarfish specimens were
collected and later deposited at the BRNU herbarium. The last records from two sites near
the village of Zwingendorf in the Pulkau district date back to the early 1970s or 1980s
(Jurasky 1980).
In 1986, an unsuccessful attempt was made by P. Tomšovic from the Institute of Bot-
any in Průhonice to re-introduce Salicornia perennans at the Slanisko u Nesytu site. The
seeds were collected at Lake Neusiedl. However, the seeds sown at the site did not germi-
nate, while young plants grown in a greenhouse and transplanted to the site failed to fruc-
tify (Grulich 1987, in litt. 2021, Danihelka 2005).
Samolus valerandi (Red Lists: CZ: CR; AT: EN)
Samolus valerandi is an annual halophyte distributed mainly in the Mediterranean, west-
ern Europe, along the Baltic coast and in the steppe zone of western Asia, towards the east
reaching Pakistan and north-western India. It is also found in southern Africa. Its distri-
bution in central Europe is fragmented, confined to saline marshes (Meusel et al. 1978).
The populations of the New World are now separated at the species level as S. parviflorus
(Choleva 2009).
In the study area, Samolus valerandi is found in saline marshes of the alliance Meliloto
dentati-Bolboschoenion maritimi and wet saline grasslands of the alliance Juncion
gerardii, in ditches, on the bottoms of drained fishponds and in temporary pools on arable
land. It never reaches high cover.
Although being native to the study area, Samolus valerandi was first collected as late
as 1871 near the village of Zwingendorf in the Pulkau district (Reuss 1873) and 1880
between the villages of Terezín and Čejč in the Čejč district (Bohuňovský 1880). It has
been recorded in the Šakvice, Rakvice, Čejč, Sedlec, Hevlín-Nový Přerov and Pulkau
districts. In the Hevlín-Nový Přerov district, S. valerandi was last collected in 1921. In
the Čejč district, it was last collected in 1962 and,after more than 50 years, again in 2017
in a wet saline meadow north of the village of Terezín. In the Šakvice district, it once
occurred at the northern edge of the town of Hustopeče, with the last record dating back to
1955. In 2017, a small population was discovered on the drained bottom of the Přední
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 63
Fishpond ~1.5 km N of the former site. In 2021, large populations were discovered on
abandoned wet arable land along the Štinkovka stream between the villages of Šakvice
and Starovičky. There are numerous recent records from the Rakvice and Sedlec districts.
In the Rakvice district, an abundant population of S. valerandi was recently found mainly
at the Trkmanec-Rybníčky site. In the Sedlec district, this species has been repeatedly
observed since the 2000s, mainly on the drained bottom of the Nesyt Fishpond and once
also on the drained bottom of the Hlohovecký Fishpond. A new occurrence was discov-
ered in 2007 in a wet part of an arable field ~1.6 km W of the town of Mikulov. In the
Pulkau district, S. valerandi was only found in the surroundings of the village of
Zwingendorf in 1871 and for the second time in 1988.
In addition to the occurrences associated with halophytic sites, Samolus valerandi was
recorded in a fen meadow in the Dyje floodplain west of the village of Brod nad Dyjí in
1922 and a reed bed west of the village of Pavlov in 2002.
Scorzonera parviflora (Red Lists: CZ: CR; AT: EN)
Scorzonera parviflora is a perennial herb with a large continuous distribution range in the
steppe zone in south-eastern European Russia, Kazakhstan, adjacent parts of western
Siberia and Kyrgyzstan. It also occurs in Transcaucasia, central Anatolia, the Carpathian
Basin, central Germany and southern France, with scattered occurrences mainly in south-
ern Ukraine, Romania and northern Bohemia (Meusel & Jäger 1992).
In the study area, Scorzonera parviflora was recorded mainly in periodically wet saline
meadows of the association Scorzonero parviflorae-Juncetum gerardii on heavy, clayey
soils. However, it also occurs in saline marshes and other types of halophytic vegetation.
The species was once present in all the districts except Damnice-Olbramovice. In the
Měnín-Šaratice district, it occurred near the village of Měnín (last collected in 1900) and
in saline meadows at mineral springs south of the village of Šaratice, where it was last
seen in 1944 and then again in 2017. In the Velké Němčice district, this species was
repeatedly collected at the Plácky site, with the last record from 1975 or 1976 (Husák &
Jatiová 1984). There is also one record from the southern edge of the village of Velké
Němčice approximately from 1963 (Vicherek 1973) and another one from the village
of Starovice from 1950 (Šmarda 1953), but at least the latter is uncertain. In the Šakvice
district, the S. parviflora population near the railway station was destroyed in the early
1920s. However, another population survived at the saline site west of the village
of Starovičky until about 1963 (Vicherek 1973). In the Čejč district, this species was
particularly abundant, but the last record from the Terezín site dates back to 1970. In the
Rakvice district, S. parviflora once occurred at several sites; the population in wet rail-
way ditches next to the Rakvice railway station survived until the early 1980s (Grulich
1987). The species was once abundant on the western shores of the Nesyt Fishpond, but it
also occurred along its southern shores and in the railway ditches west of the village of
Sedlec (Fröhlich 1940). One of the two last extant populations of S. parviflora in the
study area occurs at the Slanisko u Nesytu site. Its size fluctuates depending on moisture,
declining in dry years. Only small numbers of individuals were observed during the
2010s. However, numerous plants were observed in 2021, supported by a humid spring
of that year. The species was also recorded north of the village of Kleinschweinbarth in
Lower Austria (S. Domas sec. Makowsky 1863). In the Hevlín-Nový Přerov district,
64 Preslia 94: 13–110, 2022
S. parviflora was collected near the village of Nový Přerov in 1922, and there has been no
other reliable record since then. In the Daníž district, it was found by Tomaschek (1933)
at three sites, but these populations were probably destroyed in the 1950s. In the Pulkau
district, S. parviflora once occurred at several sites between the villages of Großkadolz
and Mailberg and the town of Laa an der Thaya (Halácsy 1896); however, it has only sur-
vived until the present at the Zwingendorf site.
There are old literature records of Scorzonera parviflora from the southern margin
of the city of Brno (Oborny 1886) and the surroundings of the town of Rajhrad
(F. Mitrowsky sec. Oborny 1886), both relatively close to the occurrences in the Měnín-
Šaratice district. The undocumented records from the surroundings of the towns of
Uherské Hradiště (L. Schlögl sec. Oborny 1886) and Hodonín (C. Theimer sec. Oborny
1886) are almost certainly erroneous as no suitable sites have ever existed in those areas.
Spergularia marina (Red Lists: CZ: CR; AT: CR)
Spergularia marina is an annual or short-lived perennial prostrate to ascending herb.
It has a nearly cosmopolitan distribution, but it is unclear which parts of its distribution
range are primary and where it has been introduced. In the Old World, it occurs mainly
along the European and Mediterranean coasts and in inland saline habitats of the
Carpathian Basin, steppe zone of eastern Europe, western Siberia and Central Asia
(Meusel et al. 1965, Kúr & Ducháček in Kaplan et al. 2016).
This competitively weak species grows in sparse vegetation, in which it can be locally
one of the most abundant species. In the study area, it has been most often recorded on
wet shores and drained bottoms of ponds, especially in annual vegetation of the associa-
tions Chenopodietum rubri,Crypsietum aculeatae,Heleochloëtum schoenoidis and
Spergulario marginatae-Suaedetum prostratae. It also occurs in halophytic grasslands
and on wet arable land but seems to avoid saline marshes.
Spergularia marina was once present in all the districts, though there is no herbarium
specimen known to us from the Daníž district. Records after the year 2000 are available
from all the districts except Daníž. In the Šakvice district, the last herbarium specimen
was collected in 1984, and only in 2021, large stands of this species were found at restored
sites and abandoned wet arable land east of the village of Šakvice. This demonstrates that
S. marina can survive in wet saline places on arable land unnoticed for decades.
While declining in its original habitats, Spergularia marina is now spreading along
roads salted for winter de-icing and in cities (Kúr & Ducháček in Kaplan et al. 2016).
Spergularia media (Red Lists: CZ: CR; AT: VU)
Spergularia media is a prostrate to ascending perennial herb. It is native to coastal areas
and inland saline sites in Europe, northern Africa, Kazakhstan and northern Iran (Meusel
et al. 1965, Kúr & Ducháček in Kaplan et al. 2016).
Spergularia media has similar habitat requirements as S. marina but is less frequent on
exposed wet bottoms and more frequent on spring-wet and summer-dry soils with high
salt concentration, especially in the Puccinellietum limosae grasslands. In the past, it was
constantly present in the stands of succulent halophytes (alliance Thero-Salicornion). It
also occurs in the stands of annual halophytic grasses (alliance Cypero-Spergularion
salinae). Unlike S. marina, it is less tolerant to mechanical disturbances and is mainly
found in natural habitats.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 65
Spergularia media was once present in all the districts but Damnice-Olbramovice.
There is only one literature record (not supported by a herbarium specimen) from the
Daníž district, namely from the saline site north-west of the village of Dyjákovice
(Tomaschek 1933). Spergularia media declined together with other halophytes, mainly
due to drainage. It first disappeared from the Šakvice district, where it was last collected
in 1929. The last herbarium specimen from the Velké Němčice district is from 1963,
when it was collected at the Starovice site. In the Měnín-Šaratice district, the species was
last collected in 1978 at the Zřídla u Nesvačilky site. In the Čejč district, S. media was
repeatedly collected from the 1820s until 1967, when it was last found at the Zápověď
site. It was rediscovered there in 2005 after the creation of shallow pools. In the remain-
ing districts, this species still survives at one or a few sites.
Recently, Spergularia media has also been recorded along roads treated by de-icing
salt. However, the number of such records is much smaller than for S. marina (Kúr &
Ducháček in Kaplan et al. 2016).
Suaeda prostrata (Red Lists: CZ: RE; AT: EN)
Suaeda prostrata is a prostrate to ascending annual succulent herb. It is a member of the
taxonomically difficult S. maritima aggregate, which in Europe comprises four species
(Uotila 2011). Suaeda prostrata is a continental species distributed from the Carpathian
Basin to south-western Siberia (Freitag et al. 1996, Freitag & Lomonosova 2006). It
reaches the north-western limit of its range in southern Moravia, where it is the only spe-
cies of Suaeda. The records of S. maritima and S. pannonica in earlier sources from the
study area also belong to this species (Danihelka & Kaplan in Kaplan et al. 2017).
The stands dominated by Suaeda prostrata (association Spergulario marginatae-
Suaedetum prostratae) developed on strongly saline soils that were at least shortly
flooded in the spring but dried out in summer. This species was more tolerant to the high
content of nitrogen than Salicornia perennans (Šumberová 2007b).
Like Salicornia perennans,Suaeda prostrata was once present in all the districts but
Damnice-Olbramovice and Daníž. Both species usually co-occurred and also declined
simultaneously. The occurrences in the Šakvice, Rakvice and Nový Přerov districts van-
ished already in the 1920s. After World War II, Suaeda prostrata was only observed in
five districts: the occurrence in the Měnín-Šaratice district vanished probably in the
1940s, the last remaining population in the Čejč district at the Zápověď site disappeared
in the late 1960s, and two populations in the Velké Němčice district vanished in the late
1960s or early 1970s. The population decline at the Slanisko u Nesytu site in the Sedlec
district started around the year 1980 (Grulich 1987). In the Pulkau district, S. prostrata
was last recorded in the late 1970s at two sites (Jurasky 1980), namely Hintausacker next
to the village of Zwingendorf and in a shallow wet depression ~1 km E of the village, now
filled with earth.
Taraxacum bessarabicum (Red Lists: CZ: CR; AT: EN)
Taraxacum bessarabicum is a small perennial herb. It is one of the few easily identifiable
Taraxacum species, a member of the Taraxacum sect. Piesis, which comprises late-flow-
ering (summer to autumn), sexually reproducing halophytic species. It has an extensive
distribution range from western Europe to Central Asia. Unlike apomictic species,
66 Preslia 94: 13–110, 2022
T. bessarabicum is morphologically variable but this variation exhibits no spatial pattern
across its geographic range (Meusel & Jäger 1992, Kirschner et al. 1994, 2011).
Taraxacum bessarabicum is confined to nutrient-rich saline soils (Kirschner et al.
1994, Dudáš et al. 2016). In the study area, it was recorded most frequently in theassocia-
tion Puccinellietum limosae but also occurred in other types of halophytic grasslands and
annual halophytic vegetation.
In the past, Taraxacum bessarabicum occurred in all the halophytic districts of the
study area. However, there is only one record from the Damnice-Olbramovice district, by
A. Ripper from the village of Troskotovice, which may be dated to the 1880s (Oborny
1912). In the Šakvice district, this species was last collected in 1922. There are only two
records of T. bessarabicum from the Daníž district undocumented by herbarium speci-
mens, near the villages of Dyjákovičky and Slup (Tomaschek 1933); these occurrences
probably disappeared soon after World War II. The last populations in the Velké
Němčice, Čejč and Rakvice districts disappeared probably in the 1960s. The last herbar-
ium specimen collected in the Měnín-Šaratice district near the village of Otmarov dates
back to 1970; this species disappeared from this district in the early 1970s. In the Pulkau
district, T. bessarabicum was last observed in 1989 at the Hintausacker site near the vil-
lage of Zwingendorf. Recent records have only been known from the Hevlín-Nový
Přerov and Sedlec districts. In the former, this species still may survive at the Novosedly
and Dobré Pole sites, whereas a small population at the Nový Přerov site vanished in the
1990s. The occurrences in the Sedlec district, once surrounding the whole Nesyt Fish-
pond, have been reduced to the Slanisko u Nesytu site, while the occurrence in slightly
saline meadows south-east of the town of Lednice was documented by herbarium
specimens in 1921–1962.
In addition to the occurrences described above, there is a herbarium specimen of
Taraxacum bessarabicum collected in 1860 by A. Makowsky in Královo Pole, now a part
of the city of Brno; however, this record is dubious as it is not listed in his flora of the
Brno county published three years later (Makowsky 1863). Based on the record by
A. Ripper (sec. Oborny 1912) from the 1880s, this species also occurred in slightly saline
pastures between the town of Břeclav and the village of Tvrdonice.
Triglochin maritima (Red Lists: CZ: RE; AT: VU)
Triglochin maritima is a perennial herb with a wide distribution range in Europe, Asia
and the Americas. In Europe, it is most common in the western and northern European
coastal areas and on the Baltic coast. It is also scattered along the Mediterranean coast.
Further, it occurs around the Black Sea and in the Pontic and Pannonian regions (Hultén
& Fries 1986).
Triglochin maritima was one of the rarest halophytes in the study area. It naturally
occurred only in two districts. In 1900 it was discovered near the Šakvice railway station.
However, this occurrence was destroyed in the early 1920s when the second track was
built (cf. Krist 1935). Another population existed in the Daníž district in a saline meadow
west of the village of Jaroslavice. It was first observed there in 1926 by Tomaschek
(1933) and for the last time collected by F. Šmarda in 1947. The species was considered
extinct until 2000 when two individuals were found in a shallow ditch at the Slanisko
u Nesytu site in the Sedlec district (Danihelka & Šmarda 2001). The last plant was
observed at that site in 2005. The species was likely planted there.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 67
In addition to the occurrences at the sites with halophytic vegetation, Triglochin
maritima was collected in 1947–1949 east of the village of Ratíškovice between the
towns of Bzenec and Hodonín. However, this occurrence probably disappeared shortly
after being discovered. Although Bureš (2010) accepted some additional undocumented
records from southern Moravia, we consider them erroneous because of numerous identi-
fication mistakes we have seen in herbaria. The specimen cited therein from the Starovice
site (J. Dvořák 1959 OP) is T. palustris.
Tripolium pannonicum (Red Lists: CZ: CR; AT: VU)
Tripolium pannonicum is a short-lived perennial or annual herb. It has a discontinuous
distribution range from the British Isles through the temperate zone in eastern Europe,
southern Siberia and Central Asia to China and Japan. This speciesis very variable, and its
populations distributed along the European coast are sometimes separated as T. panno-
nicum subsp. tripolium. The inland populations, assigned to the type subspecies, are scat-
tered from north-eastern Spain over central Germany, the Carpathian Basin, south-east-
ern and eastern Europe to the temperate zone of Asia (Meusel & Jäger 1992).
Tripolium pannonicum grows on heavy clayey or loamy saline soils well supplied
with nutrients. Being a weak competitor, it requires disturbance by grazing cattle or poul-
try (Danihelka in Kaplan et al. 2017). In southern Moravia and the Pulkau valley, it
occurs or previously occurred in most types of halophytic vegetation, being most fre-
quent in the halophytic grasslands of the associations Puccinellietum limosae and
Scorzonero parviflorae-Juncetum gerardii and the stands of succulent halophytes of the
alliance Thero-Salicornion.
Tripolium pannonicum was once present in all the districts. During the 1950s, it disap-
peared from the Šakvice, Damnice-Olbramovice and Daníž districts. From each of the
latter two districts, only one herbarium specimen and one field record exist. In the Měnín-
Šaratice district, T. pannonicum survived until the early 1970s near the village of Otnice,
while the last record from the frequently visited Zřídla u Nesvačilky site dates back to the
early 1960s (Vicherek 1973). In the Velké Němčice district, T. pannonicum was last col-
lected in 1973 at the Starovice site. In the Čejč district, this species survived until the late
1960s at the Zápověď site, while in the Rakvicedistrict, it was last collected in 1980 in the
railway ditches near the Rakvice railway station. At present, T. pannonicum occurs in
a large population at the Slanisko u Nesytu site in the Sedlec district. A few plants were
also found on the drained bottom near the southern bay of the Nesyt Fishpond in 2018 and
2020. Populations found at the Dobré Pole site in the Hevlín-Nový Přerov district and at
the Hintausacker site near Zwingendorf in the Pulkau district are smaller.
In addition to the occurrences described above, Tripolium pannonicum was observed
in slightly saline pastures between the town of Břeclav and the village of Tvrdonice. This
record (A. Ripper sec. Oborny 1912) dates back to the 1880s.
Bryophytes
Most records of bryophytes from saline sites were provided by J. Podpěra in the series
Results of the bryological research in Moravia (Podpěra 1906, 1913, 1923). He de-
scribed, as new to science, Chrysohypnum helodes var. salinum (current name Campylia-
delphus elodes) from the saline habitats along the railway near the Šakvice station
68 Preslia 94: 13–110, 2022
(Podpěra 1906) and recorded it repeatedly there (Podpěra 1913, 1923). This taxon was
also found by Fietz et al. (1923) at the Plácky site near Velké Němčice. However, accord-
ing to Bradáčová et al. (2015), populations of Campyliadelphus elodes do not differ
between saline and non-saline habitats. Furthermore, these authors found a mixture of
C. elodes and the halophytic bryophyte Conardia compacta in Podpěra’s syntype stored
in the herbarium PR. In 1912, J. Podpěra discovered Hennediella heimii, a moss species
characteristic of saline habitats, near the railway between the village of Rakvice and the
town of Podivín (Podpěra 1912, 1913). In the 1990s, it was considered as critically
endangered (Váňa 1995) and afterwards as missing (Kučera & Váňa 2005) until it was
rediscovered in 2017 at the Slanisko u Nesytu site near Sedlec (Kučera et al. 2017). Along
with H. heimii, Kučera et al. (2017) found other endangered or critically endangered
halophytic bryophytes at this site, namely Conardia compacta,Didymodon tophaceus
subsp. sicculus and Pterygoneurum cf. kozlovii.
The specialized halophytic mosses co-occur at halophytic sites with species character-
ized by broad distribution and broad ecological niches. At the sites with Hennediella
heimii, Podpěra (1912) also recorded Trichostomum crispulum. Other non-specialist species,
such as Leptodictyum riparium,Physcomitrium pyriforme,Pterygoneurum subsessile,
P. ovatum,Tortula protobryoides and T. truncata were observed at saline sites by Šmarda
(1953). Bryological surveys of the Slanisko u Nesytu site in 2004 (Kubešová & Novotný
2004) and 2017 (Kučera et al. 2017) yielded ~30 species, including Amblystegium
serpens,Barbula unguiculata,Brachythecium rutabulum,Drepanocladus aduncus,
Homalothecium lutescens,Hygroamblystegium humile,Leptodictyum riparium,Physco-
mitrium pyriforme,Syntrichia papillosa and Tortula acaulon.
In August 2020, we collected bryophytes in relevés at the sites with preserved
halophytic vegetation in southern Moravia and the Pulkau district. Among altogether 86
bryophyte specimens collected, we identified 29 terricolous species. However, none of
them was a specialist of saline habitats. The most frequent species were Brachythecium
rutabulum,Drepanocladus aduncus,Homalothecium lutescens,Hygroamblystegium
humile,Leptodictyum riparium,Oxyrrhynchium hians and various Bryum species, e.g.
B. caespiticium. All the identified bryophytes are classified as least concern in the Czech
Republic except for Drepanocladus polygamus, which is considered vulnerable (Kučera
et al. 2012). Although this species regularly occurs in some saline habitat types, these
habitats are endangered in general, and individual populations are small. A complete list
of bryophytes may be extracted from the table of our relevés (Danihelka et al. 2021).
Vegetation
Vegetation classification: overall patterns
The modified expert system CzechVeg-ESy classified the dataset of 729 relevés to the
associations. Of these, 426 relevés were classified to 14 associations dominated by
halophytes or defined through their occurrence (hereafter called “halophytic associations”;
Supplementary Data S3). Other relevés were classified to non-halophytic associations,
remained unclassified or were transitional (i.e. classified to more than one association).
A comparison of the Festuco-Puccinellietea associations classified by the expert sys-
tem with Twinspan classification is shown in an alluvial diagram (Fig. 9). Some sister
clusters in the Twinspan dendrogram were merged to match the classification by the
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 69
expert system as much as possible. Still, the match of both classifications was not perfect.
The associations TCA01 Puccinellietum limosae and Centaureo pannonicae-Festucetum
pseudovinae were almost entirely contained within one Twinspan cluster each. However,
each of these two clusters also contained a significant proportion of the relevés of another
association. The associations of the alliance Juncion gerardii corresponded to 2–4
clusters each.
In the DCA ordination, the first axis corresponded to soil moisture increasing from left
to right (Fig. 10). The second axis corresponded to increasing salinity and the proportion
of halophytes from the bottom to the top. The Puccinellietum limosae (TCA01) and
Centaureo pannonicae-Festucetum pseudovinae (TCC01) relevés were located mainly at
the edges of the triangular cloud of points in the ordination diagram, which corresponded
to the classification of these two associations to separate alliances. However, the three
associations of the Juncion gerardii were overlapping, with Loto tenuis-Potentilletum
anserinae (TCB02) encompassing the largest variation in species composition, spanning
from the mesophytic types on the bottom left to the wet types from drained fishpond
bottoms on the right.
These two comparisons indicate a mismatch between the traditional classification of
the Juncion gerardii to the associations (here formalized in the expert system) and the
pattern of total species composition (here represented by the Twinspan classification and
DCA ordination). While the traditional classification defines associations based on the
occurrence of a few halophytes that reflect specific habitat conditions within the com-
plexes of halophytic vegetation, the variation in total species composition reflects mainly
the gradients of moisture availability and salinity, i.e. the factors that indicate the
decrease in the habitat quality over the past decades.
Vegetation types
The 14 associations dominated or defined through the occurrence of halophytes were
classified according to the national vegetation classification system of the Czech Repub-
lic (Chytrý 2007, 2011) as follows:
MB Bidentetea Tüxen et al. ex von Rochow 1951
MBB Chenopodion rubri (Tüxen 1960) Hilbig et Jage 1972
MBB01 Chenopodietum rubri Tímár 1950
MBB02 Bidenti frondosae-Atriplicetum prostratae Poli et J. Tüxen 1960
corr. Gutermann et Mucina 1993
MBB04 Chenopodio chenopodioidis-Atriplicetum prostratae Slavnić 1948
corr. Gutermann et Mucina in Mucina et al. 1993
MC Phragmito-Magno-Caricetea Klika in Klika et Novák 1941
MCB Meliloto dentati-Bolboschoenion maritimi Hroudová et al. 2009
MCB01 Astero pannonici-Bolboschoenetum compacti Hejný et Vicherek
ex Oťaheľová et Valachovič in Valachovič 2001
MCB02 Schoenoplectetum tabernaemontani De Soó 1947
TA Crypsietea aculeatae Vicherek 1973
TAA Cypero-Spergularion salinae Slavnić 1948
TAA01 Crypsietum aculeatae Wenzl 1934
70 Preslia 94: 13–110, 2022
TAA02 Heleochloëtum schoenoidis Ţopa 1939
TB Thero-Salicornietea strictae Tüxen in Tüxen et Oberdorfer 1958
TBA Salicornion prostratae Géhu 1992
TBA01 Salicornietum strictae Soó 1964
TBA02 Spergulario marinae-Suadetum prostratae Vicherek in Moravec et al. 1995
TC Festuco-Puccinellietea Soó ex Vicherek 1974
TCA Puccinellion limosae Soó 1933
TCA01 Puccinellietum limosae Soó 1933
TCB Juncion gerardii Wendelberger 1943
TCB01 Scorzonero parviflorae-Juncetum gerardii (Wenzl 1934) Wendelberger 1943
TCB02 Loto tenuis-Potentilletum anserinae Vicherek 1973
TCB03 Agrostio stoloniferae-Juncetum ranarii Vicherek 1962
TCC Festucion pseudovinae Soó 1933
TCC01 Centaureo pannonicae-Festucetum pseudovinae Klika et Vlach 1937
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 71
8
9
20
21
11
3
Expert System Twinspan
Accepted clusters 4th 3rd 2nd 1st
TCB01
TCA01
TCB03
TCB02
TCC01
Fig. 9. A comparison of the associations of the Festuco-Puccinellietea class classified by the expert system
with clusters based on the Twinspan classification. TCA01 = Puccinellietum limosae,TCB01=Scorzonero
parviflorae-Juncetum gerardii,TCB02=Loto tenuis-Potentilletum anserinae,TCB03=Agrostio
stoloniferae-Juncetum ranarii, TCC01 = Centaureo pannonicae-Festucetum pseudovinae.
72 Preslia 94: 13–110, 2022
-2
-1
0
1
2
-2 -1
012
DCA2
TCA01
TCB01
TCB02
TCB03
TCC01
Ach.asp
Ach.mil
Agr.eup
Arr.ela
Bro.ere
Bup.ten
Car.acu
Car.acu
Car.hir
Car.hor
Car.otr
Car.sec
Cir.bra
Cyn.dac
Dac.glo
Dau.car
Ely.rep
Fes.pra
Fes.pul
Gla.mar
Lol.per
Lyc.flo
Lys.num
Lyt.vir
Orc.pal
Pla.lan
Pla.mar
Poa.ann
Puc.dis
Pul.dys
Ran.rep
Ror.syl
Rum.cri
Sco.can
Sco.par
Spe.med
Sym.off
Tar.bes
Teu.sco
Tri.dub
Tri.pra
Tri.pan
Ver.off
-2
0
2
-2 -1
012
DCA1
DCA2
a
a
> 10%
< 10%
Fig. 10. Ordination (detrended correspondence analysis, DCA) of the relevés assigned to the Festuco-
Puccinellietea associations by the expert system. Different colours and symbols indicate the associations (see
Fig. 9 for the code legend). The lower plot shows only diagnostic species of the associations (see Table 1):
black and grey letters indicate species with a fit to the first two axes higher and lower than 10%, respectively.
Tabl e 1. Synoptic table of 14 halophytic associations based on the relevés classified by the expert system. The
numbers are the percentage occurrence frequencies (constancies). In the associations defined by the dominance
of a single species, the dominant species are indicated by dark shading. Light shading indicates diagnostic spe-
cies of individual associations sorted by decreasing values of the phi coefficient of association. Frequencies of
bryophytes were calculated only from our relevés sampled in 2020. Therefore, the associations not sampled in
that year do not contain any bryophyte records. A full version of this table is available as Supplementary Table S1.
Associationnumber 1234567891011121314
Number of relevés 10 5 1 17 13 9 10 9 6 38 29 62 28 10
Number of relevés with bryophytes .1.7.....7.1945
Diagnostic species
1–MBB01Chenopodietum rubri
Chenopodium glaucum 100 . . 6 . 22 40 . 17 11 . 2 11 .
Gnaphalium uliginosum 30.........42..
2–MBB02Bidenti-Atriplicetum prostratae
Atriplex prostrata subsp. latifolia 10 100 100 53 . 33 80 11 17 45 25 13 21 10
3–MBB04Chenopodio chenopodioidis-Atriplicetum prostratae
Chenopodium chenopodioides ..100...30.......
4–MCB01Astero pannonici-Bolboschoenetum compacti
Bolboschoenus maritimus agg. 10 60 100 100 92 11 70 . . 13 32 18 68 .
Mentha aquatica ...24.......34.
5–MCB02Schoenoplectetum tabernaemontani
Schoenoplectus tabernaemontani 10 . . 47 100 . 10 . . . 7 5 14 .
Persicaria amphibia ...1869.10....3..
Eleocharis palustris agg. ...677.....29214.
Hippuris vulgaris ....31.........
Equisetum fluviatile ....23.........
Carex disticha ....15.........
Lythrum salicaria . . . 24 38 . 10 . . 3 4 8 14 .
6–TAA01Crypsietum aculeatae
Crypsis aculeata 40 . . 6 . 100 20 . . 3 4 . . .
7–TAA02Heleochloetum schoenoidis
Crypsis schoenoides .....33100..543..
Chenopodium album agg. .20....40....3..
Solanum tuberosum ......20.......
Matricaria chamomilla ......20..3....
Chenopodium rubrum ......20..3.3..
Anagallis arvensis 10.....20....3..
8–TBA01Salicornietum prostratae
Salicornia perennans .....11.100675....
9–TBA02Spergulario marginatae-Suaedetum prostratae
Suaeda prostrata .......331008....
10–TCA01Puccinellietum limosae
Taraxacum bessarabicum ......101117581113.10
Glaux maritima .........21410..
11–TCB01Scorzonero parviflorae-Juncetum gerardii
Scorzonera parviflora ....23..33.5683.10
Orchis palustris ....15.....25...
Carex acuta ..........112..
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 73
Associationnumber 1234567891011121314
Achillea asplenifolia .........32164.
Teucrium scordium ...6......145..
Lychnis flos-cuculi ..........7...
12–TCB02Loto tenuis-Potentilletum anserinae
Poa annua .........31119..
Cerastium holosteoides .........5.19410
Verbena officinalis ...........8..
Pulicaria dysenterica ......10..814247.
Arenaria serpyllifolia .........3.10..
13–TCB03Agrostio stoloniferae-Juncetum ranarii
Carex otrubae 1020.1846.....32248920
Ranunculus repens ...1231....3323968.
Carex hirta ..........181636.
Cirsium brachycephalum ...1215.....11.36.
Rumex crispus . . . 24 15 . 10 . . . 11 24 50 10
Carex secalina .20.12.....5.232.
Lythrum virgatum .........37.18.
Rorippa sylvestris ..........71321.
Carex hordeistichos ........17.4.2510
Symphytum officinale agg. ...6.......314.
Lysimachia nummularia ...........211.
14–TCC01Centaureo pannonicae-Festucetum pseudovinae
Festuca pulchra .........11.5.100
Achillea millefolium agg. ...6.....114341490
Cynodon dactylon .............30
Scorzonera cana .........114..40
Plantago lanceolata ..........726450
Daucus carota .20.......16423.60
Dactylis glomerata ...........18440
Centaurea jacea 10........82924460
Bupleurum tenuissimum .........1176.40
Plantago maritima . . . 6 38 . 10 44 33 47 29 11 . 90
Arrhenatherum elatius ...........6430
Bromus erectus .............20
Agrimonia eupatoria ...........2.20
Festuca pratensis agg. .........2436342560
Trifolium dubium ..........72420
Trifolium pratense ..........14131130
Species diagnostic for more than one association
Elymus repens 10 80 . 29 8 11 10 11 . 24 25 31 36 80
Tripolium pannonicum . 40 . 24 15 33 20 100 50 84 46 10 25 10
Spergularia media . . . 6 . 22 30 89 83 71 4 . . 10
Puccinellia distans 20 . . 24 . 22 40 89 100 87 32 16 36 .
Lolium perenne .........111139.40
Poa pratensis agg. .........11737750
Other species with > 5% frequency across the whole table
Lotus tenuis 10 20 100 29 8 . 20 11 . 68 96 100 68 90
Juncus compressus agg. 10 60 . 53 77 . 30 56 50 47 100 45 79 40
Agrostis stolonifera agg. 1040 .65773330 . .3979569630
Potentilla anserina 20 20 . 59 46 . 20 . . 47 93 68 82 10
Trifolium fragiferum .........1650696440
74 Preslia 94: 13–110, 2022
Associationnumber 1234567891011121314
Phragmites australis 1020 .59541150 . .3461243920
Melilotus dentatus . 20 100 29 8 . 10 . . 29 46 47 61 30
Odontites vernus subsp. serotinus ...6.....2136583940
Taraxacum sect. Taraxac um ......10..1332505740
Plantago uliginosa 1020 .2415 .50 . . 3363264 .
Inula britannica 10 20 . 29 15 . 20 . . 18 25 35 25 30
Carex distans ...6.....843293650
Polygonum aviculare agg. . 20 . 18 . 22 40 . . 32 7 26 14 .
Senecio erraticus . . . 12 . . 10 . . 13 32 27 25 30
Spergularia marina 70 20 . . . 56 40 11 67 13 7 13 4 .
Mentha pulegium ..........432436.
Centaurium pulchellum 40..18.....1114213210
Potentilla reptans ....8....332292110
Tripleurospermum inodorum . 40 100 29 23 . 30 . . 6 18 21 8 .
Cirsium canum 10..12.....3252125.
Lotus maritimus .........1129132130
Plantago major 104010018.....572314.
Medicago lupulina ..100......3.32440
Juncus bufonius agg. 60 . 100 6 . . 10 . . 3 . 11 36 .
Scorzoneroides autumnalis .........311261810
Poa trivialis ....38.....211129.
Cirsium arvense . .100 6 8 .20 . . 8 4102520
Pulicaria vulgaris .........3182311.
Pastinaca sativa .20.......5423710
Bidens tripartitus 20..6......1121..
Trifolium hybridum ..10012......14821.
Rumex maritimus 10 . 100 6 . . 20 . . . 7 16 4 .
Sonchus arvensis .20.6..20..114811.
Trifolium repens .........3719.20
Potentilla supina . . 100 12 . . 50 . . . 4 6 7 .
Calamagrostis epigejos .........3.1318.
Sonchus asper ...12..10..518214.
Echinochloa crus-galli .20100 6 . .50 . . . 4 6 4 .
Althaea officinalis 10..6.11...311104.
Bryophytes: diagnostic species
4–MCB01Astero pannonici-Bolboschoenetum compacti
Physcomitrium pyriforme – .–29––––– .– . . .
14–TCC01Centaureo pannonicae-Festucetum pseudovinae
Brachythecium albicans – .– .––––– .– . .40
Bryophytes: other species with > 5% frequency across the whole table
Drepanocladus aduncus – .–14–––––14–11 .40
Drepanocladus polygamus – .–14––––– .–21 .20
Brachythecium rutabulum – .– .––––– .–2125 .
Hygroamblystegium humile – .– .–––––14–11 .20
Bryum caespiticium – .–14––––– .–525 .
Leptodictyum riparium – .– .––––– .–16 . .
Brachythecium salebrosum – .– .––––– .–52520
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 75
A synoptic table summarizing the species composition of these 14 halophytic associa-
tions was prepared based on the stratified subset of relevés assigned to these associations
by the expert system (Table 1). The time axes showing the occurrence of each association
and the total number of associations in halophytic districts of the study area are shown in
Figs. 11 and 12. We provide the descriptions of individual associations in the following
text. The highly diagnostic, highly constant or highly dominant species (see Methods) are
shown in bold.
MBB01 Chenopodietum rubri (Fig. 13A)
Diagnostic species: Chenopodium glaucum,Gnaphalium uliginosum,Juncus bufonius agg., Spergulariamarina
Constant species: Chenopodium glaucum,Juncus bufonius agg., Spergularia marina
Dominant species: Chenopodium glaucum
The species-poor annual vegetation dominated by Chenopodium glaucum was described by
Vicherek (1973) from several halophytic sites in southern Moravia. It occurred on fish-
pond shores or in shallow depressions in the complexes of saline grasslands. The soils
were bare, strongly saline and nutrient-rich, wet in spring but drying out in summer.
Vicherek (1973) classified this vegetation to the subassociation Crypsietum aculeatae
chenopodietosum glauci, despite the rare occurrence of Crypsis aculeata in these stands.
He considered this vegetation transitional between the association Crypsietum aculeatae
and ephemeral wetland vegetation of the class Isoëto-Nano-Juncetea. Following the con-
cept accepted in the Vegetation of the Czech Republic (Šumberová & Lososová 2011),
this vegetation belongs to the association Chenopodietum rubri, which comprises stands
of both Chenopodium glaucum and C. rubrum, occurring in both saline and non-saline,
mainly anthropogenic habitats. Vicherek (1973) documented this vegetation by relevés
sampled in the early 1960s in the Velké Němčice, Rakvice, Sedlec and Hevlín-Nový
Přerov districts. There are no newer records from the study area except for a relevé domi-
nated by Chenopodium glaucum recorded during the Natura 2000 mapping in 2001 on
a bottom of an artificial pool in the Císařská obora site in the Měnín-Šaratice district.
However, this relevé is much richer in species than the relevés recorded by Vicherek
(1973). Although this vegetation seems to be absent from halophytic sites in the study
area today, it can temporarily develop on exposed bottoms of fishponds (e.g. Nesyt) and
pools.
MBB02 Bidenti frondosae-Atriplicetum prostratae (Fig. 13B)
Diagnostic species: Atriplex prostrata subsp. latifolia,Atriplex sagittata,Elymus repens,Euphorbiaplatyphyllos
Constant species: Atriplex prostrata subsp. latifolia,Bolboschoenus maritimus agg., Elymus repens,Juncus
compressus agg.
Dominant species: Atriplex prostrata subsp. latifolia,Bolboschoenus maritimus agg., Juncus compressus
agg., Phragmites australis,Spergularia marina,Tripolium pannonicum
This species-poor community is dominated by Atriplex prostrata subsp. latifolia,
a therophyte with an affinity for saline soils, though often occurring also in non-saline
anthropogenic habitats. It develops in places with the fluctuating water level, especially
at the bottoms of summer-drained fishponds and pools. The decay of phytoplankton and
aquatic macrophytes increases nutrient availability, which supports the high productivity
of this vegetation and the occurrence of some species of marshes (e.g. Bolboschoenus
76 Preslia 94: 13–110, 2022
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 77
TCB03. Agrostio stoloniferae-Juncetum ranarii TCC01. Centaureo pannonicae-Festucetum pseudovinae
TCB01. Scorzonero parviflorae-Juncetum gerardii TCB02. Loto tenuis-Potentilletum anserinae
TBA02. Spergulario marginatae-Suadetum prostratae TCA01. Puccinellietum limosae
TAA02. Heleochloëtum schoenoidis TBA01. Salicornietum prostratae
MCB02. Schoenoplectetum tabernaemontani TAA01. Crypsietum aculeatae
MBB04. Chenopodio chenopodioidis-Atriplicetum prostratae MCB01. Astero pannonici-Bolboschoenetum compacti
MBB01. Chenopodietum rubri MBB02. Bidenti frondosae-Atriplicetum prostratae
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1940 1950 1960 1970 1980 1990 2000 2010 2020 1940 1950 1960 1970 1980 1990 2000 2010 2020
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
Fig. 11. Records of halophytic associations in individual halophytic districts of southern Moravia and northern
Lower Austria. Each cross indicates a relevé classified to the association by the expert system. The gap
between 1965 and 1990 is due to the absence of research.
maritimus agg. and Phragmites australis) and ruderal species (e.g. Atriplex sagittata and
Elymus repens). There are also competitively weak halophytes typical of the early
successional stages, especially Spergularia marina. Different authors repeatedly sam-
pled this vegetation on the shores or exposed bottom of the Nesyt Fishpond in the Sedlec
district. In 2020, we also recorded it at the Trkmanec-Rybníčky site in the Rakvice district.
MBB04 Chenopodio chenopodioidis-Atriplicetum prostratae (Fig. 13C)
This association comprises natural vegetation of saline, periodically flooded soils domi-
nated or co-dominated by the halophyte Chenopodium chenopodioides, accompanied by
different species of ephemeral wetlands and some halophytes. The soils are rich in nitro-
gen and show various degrees of salinity (Šumberová & Lososová 2011). In the modified
expert system, we re-defined this association through dominance or co-dominance of
Chenopodium chenopodioides to make it consistent with the definitions of other associa-
tions. The modified definition is more restrictive than the definition in the Vegetation of
the Czech Republic (Šumberová & Lososová 2011), which required >5% cover but not
the dominance of this species. As a result, only one relevé recorded by K. Šumberová on
the bottom of Hlohovecký Fishpond in the Sedlec district in 1997 was classified to this
association. Nevertheless, Chenopodium chenopodioides is repeatedly found on bare wet
soil at other halophytic sites of southern Moravia. Vegetation with significant participa-
tion of this species was documented by relevés, for example, at Zápověď site in the Čejč
district (by K. Šumberová in 2007) and the Trkmanec-Rybníčky site in the Rakvice dis-
trict (here partly in the annual vegetation of the association Heleochloëtum schoenoidis,by
P. Lustyk in 2000 and 2011, and P. Dřevojan in 2015). However, the species has also been
78 Preslia 94: 13–110, 2022
2
1
7
7
5
2
5
5
8
9
2
4
2
7
4
8
1
6
1
12
1
1
7
4
8
2
3
7
4
1
10
2
4
4
4
4
4
5
9
1. Měnín-Šaratice
2. Velké Němčice
3. Šakvice
4. Rakvice
5. Čejč
6. Sedlec
7. Damnice-Olbramovice
8. Hevlín-Nový Přerov
9. Daníž
10. Pulkau
Study area
1940 1950 1960 1970 1980 1990 2000 2010 2020
Fig. 12. Numbers of halophytic associations recorded per two-decade periods in individual halophytic districts
of southern Moravia and northern Lower Austria and across all the districts (Study area). This figure summa-
rizes the records for individual associations shown in Fig. 14. The gap in the 1970s and 1980s indicated the
absence of research between 1965 and 1990. The period 2001–2020 also includes records from the Šakvice and
Pulkau districts from 2021.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 79
Fig. 13. Associations of halophytic vegetation in southern Moravia and northern Lower Austria: (A) MBB01
Chenopodietum rubri, Sedlec, early 1960s; (B) MBB02 Bidenti-Atriplicetum prostratae, Slanisko u Nesytu
2016; (C) MBB04 Fragmentary stand of Chenopodio chenopodioidis-Atriplicetum prostratae, Zápověď 2020.
Photo credits: J. Vicherek (A), M. Chytrý (B), K. Chytrý (C).
repeatedly recorded at various southern Moravian sites that lack well-developed halophytic
vegetation, especially in fishponds and other wet habitats (Dřevojan & Šumberová in
Kaplan et al. 2018). Therefore, vegetation corresponding to this association may also be
found at other sites not reported here.
MCB01 Astero pannonici-Bolboschoenetum compacti (Fig. 14A)
Diagnostic species: Bolboschoenus maritimus agg., Lycopus europaeus,Mentha aquatica,Schoenoplectus
tabernaemontani; Physcomitrium pyriforme
Constant species: Agrostis stolonifera agg., Atriplex prostrata subsp. latifolia,Bolboschoenus maritimus agg.,
Juncus compressus agg., Phragmites australis,Potentilla anserina,Schoenoplectus tabernaemontani
Dominant species: Bolboschoenus maritimus agg.
This relatively species-poor halophytic marsh vegetation occurs in the littoral zones of
water bodies, near mineral springs and in wet depressions in the complexes of halophytic
vegetation (Hroudová et al. 2009). The dominant species are Bolboschoenus maritimus
and B. planiculmis, which can occur either separately or in mixed stands. These species
are accompanied by other halophytes, including Juncus gerardii,Lotus tenuis,Melilotus
dentatus,Puccinellia distans and Tripolium pannonicum. This association is one of the
most common types of (sub)halophytic vegetation in the study area, repeatedly docu-
mented by relevés from several halophytic districts (e.g. by Vicherek 1973 under the
name Bolboschoenetum maritimi continentale). We also recorded relevés belonging to
this association in four districts in 2020. Similar stands develop on temporarily flooded
arable land. They are usually dominated by Bolboschoenus planiculmis, contain some
arable weed species, and are classified as the association Tripleurospermo inodori-
Bolboschoenetum planiculmis Hroudová et al. 2009. Halophytes can also occur in this
association, as documented from wet arable land near Šakvice by Chytrý & Danihelka
(2021).
MCB02 Schoenoplectetum tabernaemontani (Fig. 14B)
Diagnostic species: Bolboschoenus maritimus agg., Carex disticha,Carex riparia,Eleocharis palustris agg.,
Equisetum fluviatile,Hippuris vulgaris,Lythrum salicaria,Persicaria amphibia,Poa trivialis,
Schoenoplectus tabernaemontani
Constant species: Agrostis stolonifera agg., Bolboschoenus maritimus agg., Carex otrubae,Eleocharis
palustris agg., Juncus compressus agg., Persicaria amphibia,Phragmites australis,Potentilla anserina,
Schoenoplectus tabernaemontani
Dominant species: Schoenoplectus tabernaemontani
This vegetation develops on saline or calcium-rich clayey soils at periodically water-
logged sites with changing dry and wet periods (Vicherek 1962, Šumberová et al. 2011).
The dominant species Schoenoplectus tabernaemontani forms dense stands, which may
contain ruderal, subhalophytic and halophytic species such as Carex otrubae,Juncus
gerardii,Plantago maritima and Potentilla anserina. Vicherek (1973) documented
Schoenoplectetum tabernaemontani near the villages of Popice (Šakvice district), Kobylí
(Čejč district), Lednice and at the Nesyt Fishpond (both Sedlec district). In 2020, we
observed (but not sampled) this vegetation at the Trkmanec-Rybníčky site (Rakvice dis-
trict), Nesyt Fishpond and Novosedly site (Hevlín-Nový Přerov district). It also occurs at
the shore of a new fishpond at restored sites near Šakvice (Šakvice district; Chytrý &
Danihelka 2021).
80 Preslia 94: 13–110, 2022
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 81
Fig. 14. Associations of halophytic vegetation in southern Moravia and northern Lower Austria: (A) MCB01
Astero pannonici-Bolboschoenetum compacti, Zápověď 2020; (B) MCB02 Schoenoplectetum tabernaemontani,
Rakvice 2020; (C) TAA01 Crypsietum aculeatae, Slanisko u Nesytu 2016; (D) TAA02 Heleochloetum
schoenoidis, eastern shore of the Nesyt Fishpond 2020; (E) TBA01 Salicornietum prostratae, Terezín 1961.
Photo credits: M. Chytrý (A, C), K. Šumberová (B), K. Chytrý (D), J. Vicherek (E).
TAA01 Crypsietum aculeatae (Fig. 14C)
Diagnostic species: Crypsis aculeata
Constant species: Crypsis aculeata,Spergularia marina
Dominant species: Crypsis aculeata,Spergularia media
This species-poor vegetation is characterized by a high cover or dominance of Crypsis
aculeata, often accompanied by Spergularia marina or S. media. It occurs on exposed
fishpond shores and in periodically inundated depressions that dry out in summer. The
extent of individual stands rarely exceeds 1 m2. In the study area, Crypsietum aculeatae
was only recorded near Rakvice, at the Nesyt Fishpond (Sedlec district) and near
Novosedly (Hevlín-Nový Přerov district; Vicherek 1973). The populations were reduced
dramatically in the 1990s (Danihelka & Hanušová 1995), and later on, this association
was considered to have disappeared from the Czech Republic (Šumberová 2007a). In
2008, a population of C. aculeata was found on the wet bottom of a pool dug at the
Zápověď site in the Čejč district as a part of an ecological restoration project (Hadinec &
Lustyk 2009). However, it was not documented by a relevé. In 2016, this vegetation
developed near the western shore of the Nesyt Fishpond. However, the population of
C. aculeata has rapidly declined at this site since then, and we found only one individual
in 2020. This fluctuation in population size is natural, and the association can re-appear in
the future once bare wet soil is exposed.
TAA02 Heleochloëtum schoenoidis (Fig. 14D)
Diagnostic species: Anagallis arvensis,Chenopodium album agg., Chenopodium rubrum,Crypsis schoenoides,
Echinochloa crus-galli,Matricaria chamomilla,Potentilla supina,Solanum tuberosum
Constant species: Atriplex prostrata subsp. latifolia,Bolboschoenus maritimus agg., Crypsis schoenoides,
Echinochloa crus-galli,Phragmites australis,Plantago uliginosa,Potentilla supina
Dominant species: Crypsis schoenoides,Spergularia marina
These low-growing, open annual grasslands are dominated by Crypsis schoenoides.
Other species, usually occurring in low abundances, include halophytes such as Spergu-
laria marina, species of drained fishpond bottoms such as Chenopodium rubrum and
Potentilla supina, and ruderal plants such as Echinochloa crus-galli or Chenopodium
album. The association occurs on bottoms of drained fishponds or poolsand in seasonally
wet depressions, sometimes on arable land. The soils are wet in spring but dry out in sum-
mer when cracks appear on the surface (Šumberová 2007a). Crypsis schoenoides sur-
vives in a seed bank and can appear on the bare ground after several years since the last
occurrence. Although Vicherek (1973) recorded this vegetation only at one site near the
village of Moutnice, it used to be probably more common in the study area, as inferred
from floristic records of C. schoenoides. In 1993–1994, this vegetation was documented
by relevés at the Nesyt Fishpond (Sedlec district) and near the village of Novosedly
(Hevlín-Nový Přerov district; Danihelka & Hanušová 1995). In 2004, M. Chytrý and
J. Danihelka sampled large stands of Heleochloëtum schoenoidis near the Výtopa Fish-
pond close to the eastern shore of the Nesyt Fishpond. At this site, it occurred in a spring-
flooded field depression in which mud removed from the Nesyt bottom was deposited.
This observation indicates that the species can survive long periods in a seed bank on
pond bottoms. The association was also observed at the Trkmanec-Rybníčky site near
Rakvice and documented by relevés in 2011 by P. Lustyk and in 2015 by P. Dřevojan
82 Preslia 94: 13–110, 2022
(Lustyk in Hadinec & Lustyk 2013, Dřevojan et al. 2017). We observed this vegetation
on the exposed bottom of the eastern shore of the Nesyt Fishpond in 2020. We also sam-
pled this association on abandoned wet arable land along the Štinkovka stream east of
Šakvice in 2021 (Chytrý & Danihelka 2021), which is the first record of its occurrence in
the Šakvice district. It is likely that in the future, Heleochloëtum schoenoidis will also re-
appear at other sites by regeneration of the dominant species from the seed bank follow-
ing mud dredging and depositing or the emergence of wet bare soil.
TBA01 Salicornietum strictae (Fig. 14E)
Diagnostic species: Puccinellia distans,Salicornia perennans,Spergularia media,Tripolium pannonicum
Constant species: Juncus compressus agg., Plantago maritima,Puccinellia distans,Salicornia perennans,
Spergularia media,Tripolium pannonicum
Dominant species: Salicornia perennans
This species-poor vegetation once occurred in highly saline and moist spots dominated
by Salicornia perennans, an annual succulent plant forming sparse and low-growing
stands. In places, this species was accompanied by other halophytes such as Plantago mari-
tima,Puccinellia distans,Spergularia media,Suaeda prostrata and Tripolium panno-
nicum.Salicornietum strictae used to occur in depressions at halophytic sites, where salt
precipitated on the soil surface and, at the same time, the surface was relatively wet
(Vicherek 1973). This vegetation also occurred on dirt roads crossing the complexes of
halophytic vegetation, where it was supported by disturbance. For example, V. Grulich
(pers. comm.) observed it in the tracks of horse-drawn carriages still in the early 1970s. In
the 19th century, this vegetation may have been common along the shores of the former
Měnín Fishpond and salt lakes in the Čejč district. Vicherek (1973) documented Sali-
cornietum strictae by relevés from the early 1960s recorded at the Plácky site (Velké
Němčice district), Zápověď site (Čejč district) and the Nesyt Fishpond (Sedlec district).
However, the populations of the dominant species declined rapidly during the 1960s and
entirely disappeared in the 1970s. This process was related to the cessation of the tradi-
tional management of saline vegetation such as goose and livestock grazing and soil de-
salination.
TBA02 Spergulario marinae-Suadetum prostratae (Fig. 15A)
Diagnostic species: Puccinellia distans,Salicornia perennans,Spergularia marina,Spergularia media,
Suaeda prostrata
Constant species: Juncus compressus agg., Puccinellia distans,Salicornia perennans,Spergularia marina,
Spergularia media,Suaeda prostrata,Tripolium pannonicum
Dominant species: Suaeda prostrata
This association included open, species-poor stands dominated by Suaeda prostrata.
Other species, occurring in low abundances, included Juncus gerardii,Puccinellia
distans,Salicornia perennans,Spergularia marina,S. media and Tripolium pannoni-
cum. The community used to occur in similar habitats as the previous association: shal-
low depressions with an extremely high concentration of salt and nitrates in the soil, often
on livestock and poultry pastures (Vicherek 1973). This vegetation may have been once
common on the shores of the former Měnín Fishpond, Kobylí Lake and Čejč Lake. In the
early 1960s, relevés of this association were recorded by Vicherek (1962, 1973) at the
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 83
Plácky site and near the village of Starovice (both in the Velké Němčice district) and at
the Nesyt Fishpond near Sedlec. These occurrences vanished in the 1970s and 1980s
when Suaeda prostrata went extinct in the Czech Republic.
TCA01 Puccinellietum limosae (Fig. 15B)
Diagnostic species: Glaux maritima,Puccinellia distans,Spergularia media,Taraxacum bessarabicum,
Tripolium pannonicum
Constant species: Atriplex prostrata subsp. latifolia,Juncus compressus agg., Lotus tenuis,Plantago mari-
tima,Potentilla anserina,Puccinellia distans,Spergularia media,Taraxacum bessarabicum,Tripolium
pannonicum
Dominant species: Puccinellia distans
This short open grassland is characterized by the occurrence, and in places also high
cover, of Puccinellia distans. This species can be accompanied by Glaux maritima,
Plantago maritima,Spergularia media,Taraxacum bessarabicum and Tripolium
pannonicum. The community further contains species otherwise typical of subhalophytic
meadows such as Lotus tenuis and Potentilla anserina.Puccinellietum limosae develops
in places that are wet in spring but dry out during summer. Most species of this commu-
nity tolerate trampling, hence the community also develops on footpaths. Puccinellietum
limosae was once common at most halophytic sites in southern Moravia (Vicherek 1962,
1973) but declined in the 1960s. However, as it is less dependent on the water regime, its
decline was less pronounced than the decline of some other associations. During our sur-
vey in 2020, we documented this association by relevésin the Čejč, Sedlec, Hevlín-Nový
Přerov and Pulkau districts. In 2021, we also sampled it near Štinkovka stream in the
Šakvice district (Chytrý & Danihelka 2021).
TCB01 Scorzonero parviflorae-Juncetum gerardii (Fig. 15C)
Diagnostic species: Achillea asplenifolia,Carex acuta,Carex distans,Juncus compressus agg., Lotus tenuis,
Lychnis flos-cuculi,Mentha pulegium,Orchis palustris,Potentilla anserina,Potentilla reptans,Scorzo-
nera parviflora,Teucrium scordium
Constant species: Agrostis stolonifera agg., Carex distans,Juncus compressus agg., Lotus tenuis,Melilotus
dentatus,Mentha pulegium,Phragmites australis,Potentilla anserina,Scorzonera parviflora,Trifolium
fragiferum,Tripolium pannonicum
Dominant species: Carex distans,Juncus compressus agg., Scorzonera parviflora
This association of wet saline grasslands occurs in shallow depressions at halophytic
sites. It is physiognomically similar to wet meadows. The stands are usually dominated
by graminoids, in particular, Juncus gerardii,J. compressus or Carex distans.Grami
-
noids are accompanied by various halophytic dicot herbs, most notably Scorzonera
parviflora and Tripolium pannonicum, which can both attain a high cover. In the early
1960s, this vegetation occurred at most of the southern Moravian halophytic sites
(Vicherek 1962, 1973). However, with further draining and desalination, it gradually
changed to the subhalophytic grasslands with fewer specialist species (Loto tenuis-
Potentilletum anserinae) or non-halophytic grasslands. Later on, this vegetation was
documented by relevés only from the Slanisko u Nesytu sitein the early 1990s (Hanušová
1995) and in 2004 (by J. Danihelka). During our vegetation survey in 2020, we did not
record any relevé corresponding to this association.
84 Preslia 94: 13–110, 2022
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 85
Fig. 15. Associations of halophytic vegetation in southern Moravia and northern Lower Austria: (A) TBA02
Spergulario marginatae-Suaedetum prostratae, Starovice 1962; (B) TCA01 Puccinellietum limosae, Zápověď
2020; (C) TCB01 Scorzonero parviflorae-Juncetum gerardii, Slanisko u Nesytu 2016. Photo credits:
J. Vicherek (A), M. Chytrý (B, C).
TCB02 Loto tenuis-Potentilletum anserinae (Fig. 16A)
Diagnostic species: Arenaria serpyllifolia,Cerastium holosteoides,Lolium perenne,Lotus tenuis,Odontites
vernus subsp. serotinus,Poa annua,Poa pratensis agg., Pulicaria dysenterica,Pulicaria vulgaris,
Scorzoneroides autumnalis,Taraxacum sect. Taraxacum ,Trifolium fragiferum,Verbena officinalis
Constant species: Agrostis stolonifera agg., Juncus compressus agg., Lotus tenuis,Melilotus dentatus,
Odontites vernus subsp. serotinus,Potentilla anserina,Taraxacum sect. Taraxacum,Trifolium fragiferum
Dominant species: Potentilla anserina
This association is currently the most common type of saline grassland in the study area.
It consists mainly of the graminoids Agrostis stolonifera,Juncus compressus and J.
gerardii, and subhalophytic herbs Lotus tenuis,Melilotus dentatus,Potentilla anserina,
Pulicaria dysenterica and Trifolium fragiferum. Besides halophytes and subhalophytes,
various species of mesophilous grasslands, wet soils and ruderal sites are also present,
including Cerastium holosteoides,Lolium perenne,Poa annua,P. pratensis agg. and
Taraxacum sect. Taraxacum. Two subtypes of this association can be distinguished,
mesic and wet. The mesic subtype, characterized by species of mesic meadows and
ruderal vegetation, occurs in trampled areas such as village lawns, paths and extensively
managed football playgrounds. In the past, this subtype was typical of pastures around
villages, and also today, it occurs in places where grazing was reintroduced for conserva-
tion purposes. The wet subtype, characterized by Melilotus dentatus and annual species
of ephemeral wetlands, occurs in wet depressions in the complexes of halophytic vegeta-
tion and on exposed fishpond shores and bottoms. This association represents a transition
between halophytic and non-halophytic vegetation. It was probably always common at
most halophytic sites in the study area. As the only type of halophytic grasslands, it also
occurred outside the natural complexes of halophytic vegetation, e.g. within some vil-
lages. Following the decrease in groundwater level and subsequent soil desalination, var-
ious types of halophytic vegetation lost the most specialized species of saline habitats,
resulting in a change to the grasslands with subhalophytic species of the association Loto
tenuis-Potentilletum anserinae.
TCB03 Agrostio stoloniferae-Juncetum ranarii (Fig. 16B)
Diagnostic species: Agrostis stolonifera agg., Calamagrostis epigejos,Carex hirta,Carex hordeistichos,
Carex otrubae,Carex secalina,Cirsium brachycephalum,Lysimachia nummularia,Lythrum virgatum,
Mentha pulegium,Plantago uliginosa,Potentilla anserina,Ranunculus repens,Rorippa sylvestris,Rumex
crispus,Symphytum officinale agg., Taraxacum sect. Taraxacum ,Trifolium fragiferum
Constant species: Agrostis stolonifera agg., Bolboschoenus maritimus agg., Carex otrubae,Juncus
compressus agg., Lotus tenuis,Melilotus dentatus,Plantago uliginosa,Potentilla anserina,Ranunculus
repens,Rumex crispus,Taraxacum sect. Taraxac um ,Trifolium fragiferum
Dominant species: Agrostis stolonifera agg., Carex otrubae
This type of halophytic grassland is typical of the sites with the repeatedly disturbed soil
surface. It is dominated by graminoids such as Agrostis stolonifera,Carex hordeistichos,
C. otrubae and C. secalina. Ruderal species such as Calamagrostis epigejos,Carex hirta,
Ranunculus repens,Rumex crispus and Taraxacum sect. Taraxacum are also common.
The rare halophyte Cirsium brachycephalum also occurs or once occurred in this vegeta-
tion at some sites. Vicherek (1973) divided this vegetation into three associations based
on the prominence of different sedge species, Agrostio-Caricetum distantis Soó 1939,
Agrostio-Caricetum secalinae Vicherek 1973 and Meliloto-Caricetum otrubae Vicherek
86 Preslia 94: 13–110, 2022
1973. However, the differences in species composition and ecology of these associations
are small, with various transitions; therefore, Šumberová et al. (2007) merged them into
a single association, and our results support this broader concept. Vicherek (1962, 1973)
recorded this association from the Velké Němčice, Šakvice, Rakvice, Čejč and Sedlec
districts. In the latter district, at Slanisko u Nesytu site, relevés were made by J. Danihelka
in 2004. In our vegetation survey in 2020, we recorded this vegetation at the Trkmanské
louky site (Rakvice district), Zápověď site (Čejč district) and near the village of Hevlín
(Hevlín-Nový Přerov district). In 2021, we also recorded it along the Štinkovka stream
near Šakvice (Šakvice district).
TCC01 Centaureo pannonicae-Festucetum pseudovinae (Fig. 16C)
Diagnostic species: Achillea millefolium agg., Agrimonia eupatoria,Arrhenatherum elatius,Bromus erectus,
Bupleurum tenuissimum,Carex distans,Centaurea jacea,Cynodon dactylon,Dactylis glomerata,Daucus
carota,Elymus repens,Festuca pratensis agg., Festuca pulchra,Lolium perenne,Plantago lanceolata,
Plantago maritima,Poa pratensis agg., Scorzonera cana,Trifolium dubium,Trifolium pratense;
Brachythecium albicans
Constant species: Achillea millefolium agg., Carex distans,Centaurea jacea,Daucus carota,Elymus repens,
Festuca pratensis agg., Festuca pulchra,Lotus tenuis,Plantago lanceolata,Plantago maritima,Poa
pratensis agg.
Dominant species: Festuca pulchra
This association comprises grasslands dominated by salt-tolerant narrow-leaved tussock-
forming grass Festuca pulchra, which forms dense stands. Subhalophytic and halophytic
species such as Bupleurum tenuissimum,Lotus tenuis and Plantago maritima frequently
occur in this vegetation. The stands are often adjacent to non-halophytic mesic grasslands
and contain their typical species, e.g. Achillea millefolium agg. (usually A. collina),
Arrhenatherum elatius,Centaurea jacea,Dactylis glomerata,Festuca arundinacea,
F. pratensis,Plantago lanceolata and Poa pratensis agg. This association occurs at the
margins of saline sites where the soil is not inundated, being mesic in spring and dry in
summer. Most stands are trampled, such as at the edges of dirt roads and in pastures. This
association, first described from south-eastern Slovakia (Klika & Vlach 1937), was not
included in the national vegetation classification of the Czech Republic (Šumberová et al.
2007) because of the lack of relevés at that time. Still, Vicherek (1973) published one
relevé from the Nový Přerov site, which he classified to the association Achilleo-
Festucetum pseudovinae Soó (1933) 1947, although all the other relevés of this associa-
tion in his study were from southern Slovakia. A relevé corresponding to this vegetation
was recorded already by Šmarda (1953) at the Nesyt Fishpond (with the dominant species
misidentified as Festuca valesiaca), and other such relevés were recorded by J. Dani-
helka in 2004 at the same site, and by P. Šmarda in 2008 at the Dobré Pole site. In 2020,
we sampled this vegetation at the Zřídla u Nesvačilky site (Měnín-Šaratice district), near
both western and eastern edge of the Nesyt Fishpond (Sedlec district) and at Dobré Pole
site (Hevlín-Nový Přerov district), and observed it at the Plácky site (Velké Němčice dis-
trict). In 2021, we also observed it at the Saliterweide site near Zwingendorf (Pulkau dis-
trict). The higher number of relevés from recent years can be due to avoidance of this veg-
etation type by earlier researchers, who may have considered it degraded vegetation tran-
sitional to other vegetation types. An alternative explanation would be the relatively
recent spread of this vegetation, which developed on solonchak soils after the drop of the
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 87
88 Preslia 94: 13–110, 2022
Fig. 16. Associations of halophytic vegetation in southern Moravia and northern Lower Austria: (A) TCB02
Loto tenuis-Potentilletum anserinae, Zápověď 2020; (B) TCB03 Agrostio stoloniferae-Juncetum ranarii,
Hevlín 2020; (C) TCC01 Centaureo pannonicae-Festucetum pseudovinae, Dobré Pole 2020. Photo credits:
M. Chytrý (A), E. Šmerdová (B), K. Chytrý (C).
water table. This association may represent a degradation phase of the former saline
grasslands of the alliances Puccinellion limosae or Juncion gerardii.
The association Centaureo pannonicae-Festucetum pseudovinae belongs to the alli-
ance Festucion pseudovinae Soó 1933. This alliance, which comprises dry grasslands of
the salt steppe, is distributed mainly in Hungary and southern Slovakia but is also found
at Lake Neusiedl in Austria, in Vojvodina and the Pannonian part of Romania (Eliáš et al.
2013, Dítě et al. 2014). Vicherek (1973) reported Festucion pseudovinae for the Czech
Republic based on the above-mentioned single relevé from Nový Přerov. However, this
alliance was not accepted in the subsequent overviews of Czech vegetation (Vicherek &
Řehořek in Moravec et al. 1995, Šumberová et al. 2007) due to the lack of other records.
Here we add this alliance to the vegetation classification system of the Czech Republic.
During our fieldwork in 2020 and 2021, we sampled this vegetation in three halophytic
districts and observed it in the other two. Compared to the other halophytic associations,
it is characterized by low conductivity and significantly lower pH, indicating possible
recent degradation of the former solonchak soils.
Non-halophytic vegetation types in the complexes of halophytic vegetation
Besides the above-mentioned associations dominated or characterized by the occurrence
of halophytes, complexes of halophytic vegetation in the study area also include some
associations defined by non-halophytes but containing some halophytes as a minor com-
ponent of their species composition. The most common of such associations are
Phragmitetum australis Savič 1926 and Caricetum distichae Nowiński 1927.
Phragmitetum australis includes species-poor stands dominated by Phragmites aus-
tralis. Its halophytic subtype often contains Bolboschoenus maritimus,Juncus compres-
sus,J. gerardii and Potentilla anserina. Some more specialized or rare halophytes such
as Cirsium brachycephalum,Samolus valerandi,Scorzonera parviflora and Tripolium
pannonicum can also occur in this vegetation.
Caricetum distichae is dominated by Carex disticha, which forms dense and extensive
stands through its clonal growth. Other moisture-demanding species such as Carex
otrubae and Iris pseudacorus frequently occur in this vegetation. At the halophytic sites,
Caricetum distichae can be enriched by halophytes such as Juncus gerardii,Plantago
maritima,Scorzonera parviflora and Tripolium pannonicum. Vicherek (1962) described
such vegetation as the subassociation Caricetum distichae juncetosum gerardii Vicherek
1962.
Further vegetation types found in the complexesof halophytic vegetation and contain-
ing some halophytic species include annual wetland vegetation of drained fishpond bot-
toms and temporary pools on arable land. This vegetation appears in irregular intervals,
depending on the water level fluctuations, particularly on the fishponds Nesyt, Výtopa
and Hlohovecký in the Sedlec district. The associations found there include Cyperetum
micheliani Horvatić 1931 (alliance Eleocharition ovatae), Veronico anagalloidis-
Lythretum hyssopifoliae Wagner ex Holzner 1973 (Verbenion supinae)andRumici
maritimi-Ranunculetum scelerati Oberdorfer 1957 (Bidention tripartitae).
There are also weed and ruderal plant communities, especially where halophytic sites
border on arable land or where saline soil was ploughed. These include especially the asso-
ciations Setario pumilae-Echinochloëtum cruris-galli Felföldy 1942 corr. Mucina in
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 89
Mucina et al. 1993 (alliance Spergulo arvensis-Erodion cicutariae)andConvolvulo
arvensis-Elytrigietum repentis Felföldy 1943 (Convolvulo arvensis-Elytrigion repentis),
which were particularly common at the Trkmanské louky and Trkmanec-Rybníčky sites
during our survey in 2020, but also occurred at other halophytic sites. If wet arable land is
abandoned, it can be quickly (within 1–2 years) overgrown by Bolboschoenus planiculmis
stands with weed species. These stands belong to the association Tripleurospermo inodori-
Bolboschoenetum planiculmis Hroudová et al. 2009 (Eleocharito palustris-Sagittarion
sagittifoliae). They often contain various subhalophytes and, in some places, also more
demanding halophytes. For example, Crypsis schoenoides,Samolus valerandi and
Spergularia marina occurred in the stands of this association on abandoned wet arable land
along the Štinkovka stream in the Šakvice district in 2021 (Chytrý & Danihelka 2021).
Relationships to salinity
The pH of soil sampled in the relevés of halophyticvegetation in 2020 ranged from 6.8 to
8.5 (mean 7.8) and conductivity ranged from 238 to 5010 (mean 1633) μS/cm.
The generalized linear models identified 39 species with significant (P < 0.05) or mar-
ginally significant (P < 0.1) relationship to pH or conductivity (Table 2 and graphs in
Supplementary Fig. S1). Contrary to the generally accepted view, Festuca pulchra and
Plantago maritima tended to be significantly more frequent on soils with lower pH and
lower conductivity. In contrast, Agrostis stolonifera and Phragmites australis, species
with a broad ecological range, tended to be more common on high-pH, saline soils. It is
important to interpret these results strictly within the context of the halophytic dataset.
Both Agrostis and Phragmites are species occurring across a broad range of pH and salin-
ity, including non-saline habitats. At the halophytic sites, they occur in wetter places,
which are also more saline. Therefore, their correlation with saline, high-pH soils in the
current dataset is likely mediated through an indirect effect of moisture.
A comparison of vegetation types in terms of pH and conductivity of their soils was
possible for five associations with at least three soil samples (Fig. 17). Loto tenuis-
Potentilletum anserinae (TCB02) and Centaureo pannonicae-Festucetum pseudovinae
(TCC01) had the lowest mean for both the variables, which corresponds to their status of
“degraded” halophytic vegetation occurring on desalinated soils or transitional vegeta-
tion to other grassland types in the marginal zones of halophytic vegetation complexes.
However, the only significant difference was for pH between Centaureo-Festucetum and
the other associations.
90 Preslia 94: 13–110, 2022
Tabl e 2. Results of the generalized linear models linking species occurrence to soil pH and conductivity. Spe-
cies occurring more frequently on higher-pH vs lower-pH soils are called basiphilous vs neutrophilous, respec-
tively. Species occurring more frequently on soils with higher vs lower electric conductivity are called
halophytes vs glycophytes, respectively. Only species with significant (P < 0.05) and, in brackets, marginally
significant (P < 0.1) models are shown.
Species pH Soil conductivity
Achillea collina neutrophilous glycophyte
Agrimonia eupatoria (neutrophilous) -
Agrostis stolonifera (basiphilous) halophyte
Arenaria serpyllifolia - glycophyte
Bolboschoenus maritimus basiphilous -
Bolboschoenus planiculmis - (halophyte)
Bromus erectus neutrophilous -
Carduus acanthoides neutrophilous -
Carex secalina basiphilous (halophyte)
Centaurea jacea neutrophilous (glycophyte)
Cerastium holosteoides - glycophyte
Cirsium brachycephalum basiphilous -
Cirsium canum neutrophilous -
Conyza canadensis - glycophyte
Dactylis glomerata (neutrophilous) glycophyte
Daucus carota neutrophilous -
Dispacus fullonum neutrophilous -
Elymus repens neutrophilous -
Festuca pratensis neutrophilous -
Festuca pulchra neutrophilous (glycophyte)
Glaux maritima (basiphilous) halophyte
Inula britanica (neutrophilous) -
Juncus compressus agg. basiphilous halophyte
Linaria vulgaris neutrophilous -
Lolium perenne - (glycophyte)
Medicago lupulina - glycophyte
Myosotis arvensis neutrophilous -
Phragmites australis basiphilous halophyte
Plantago lanceolata (neutrophilous) glycophyte
Plantago maritima - glycophyte
Plantago uliginosa - (halophyte)
Poa angustifolia (neutrophilous) glycophyte
Potentilla reptans (neutrophilous) -
Puccinellia distans basiphilous halophyte
Serratula tinctoria neutrophilous -
Sonchus arvensis (basiphilous) -
Trifolium pratense (neutrophilous) -
Trifolium repens neutrophilous (glycophyte)
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 91
Discussion
Broader biogeographic context
The study area of southern Moravia and the nearby Pulkau valley in northern Lower Aus-
tria is located on the north-western edge of the Carpathian (= Pannonian) Basin, the larg-
est inland area of well-developed halophytic flora and vegetation in Europe west of the
steppe zone (Wendelberger 1950, Molnár & Borhidi 2003, Eliáš et al. 2013). The similar-
ities and differences between the halophytic flora of the Carpathian Basin and the inland
halophytic flora of Bohemia, Germany, Poland, eastern Romania, the steppe zone of
Eastern Europe and Central Asia, as well as the halophytic flora of the European coasts
have been discussed, for example, by Wendelberger (1950) and Vicherek (1973).
In the north-west of the Carpathian Basin, halophytic vegetation is best developed around
Lake Neusiedl (e.g. Wendelberger 1950, Vicherek 1973, Eliáš et al. 2013, 2020). This area
comprises various saline habitats with different water regimes and degrees of salinity
92 Preslia 94: 13–110, 2022
ns ns ns ns
*
7.0
7.5
8.0
8.5
MCB01
TCA01 TCB02 TCB03
TCC01
pH
ns ns ns ns ns
0
1000
2000
3000
4000
5000
MCB01
TCA01 TCB02 TCB03
TCC01
Conductivity (μS/cm)
Fig. 17. A comparison of soil pH and conductivity among halophytic associations with at least three soil mea-
surements. Data are based on soil samples from the relevés we recorded in 2020. Non-significant differences
are indicated by “ns”; a significant difference at P < 0.05 is indicated by an asterisk. See the syntaxonomic syn-
opsis for the association codes.
including salt pans (Boros et al. 2013). The extent of saline habitats there is large, which
makes local extinctions less likely and colonizations easier than in the more peripheral
parts of the Carpathian Basin with small patches of halophytic habitats, such as in south-
ern Moravia. Consequently, the halophytic flora at Lake Neusiedl includes some species
that have never been recorded in the study area, e.g. the Pontic-Pannonian Suaeda
pannonica (Freitag et al. 1996), Puccinellia limosa and the Pannonian endemic P. peisonis
(Fischer et al. 2008). In contrast, Glaux maritima, which occurs in southern Moravia and
the Pulkau valley, is not found in the other parts of the Carpathian Basin today.
Traditionally, the halophytic flora and vegetation of southern Moravia and the Pulkau
valley have been compared with the halophytic flora and vegetation of south-western and
southern Slovakia, which lies on the northern edge of the Carpathian Basin (Krist 1940,
Vicherek 1973). In general, both areas have similar flora and vegetation, but there are
some notable differences. For example, the two succulent halophytes Salicornia
perennans and Suaeda prostrata have never been found in southern Slovakia; conse-
quently, the vegetation of the alliance Thero-Salicornion has never occurred there either.
Crypsis aculeata and Glaux maritima have always been very rare there; G. maritima dis-
appeared already in the 19th century. In contrast, several species typical of salt steppes of
the alliances Festucion pseudovinae and Puccinellion limosae occurring in southern
Slovakia have never occurred in the study area. Such species include Artemisia
santonicum,Camphorosma annua,Carex divisa,Hordeum geniculatum,Lepidium
cartilagineum,Limonium gmelinii,Pholiurus pannonicus and Plantago tenuiflora.This
can be explained by different ecological conditions, especially by the prevalence of
solonetz, a drier, three-horizon type of saline soils at the Slovak halophytic sites. Such
soils favour the development of salt steppes, which are widespread in Hungary (Molnár et
al. 2008), and also occur in southern Slovakia, where they are represented by several
associations (Vicherek 1973, Dítě et al. 2014). In southern Moravia and the Pulkau val-
ley, salt steppe is only represented by two associations, Puccinellietum limosae and
Centaureo pannonicae-Festucetum pseudovinae, the latter reported here as a new associ-
ation for this area.Higher precipitation and lower summer temperatures in this area do
not support the formation of solonetz. The local halophytic flora and vegetation occur on
solonchak, a wetter, two-horizon type of saline soil that often develops near mineral-
water springs (Vicherek 1973). The prevalence of different salts (mainly sulphates in the
study area vs sodium carbonate in southern Slovakia and on the Hungarian Plain;
Vicherek 1973, Boros et al. 2013) may also contribute to regional differences in halo-
phytic flora. In addition, differences in land use and stochastic processes, such as local
extinction events, may have had an influence. In contrast to the extensive plains of Hun-
gary, Vojvodina, north-western Romania, southern Slovakia and Burgenland, the study
area is much smaller, and the distribution of saline habitats is restricted to small halo-
phytic districts in lowland areas surrounded by hilly landscapes. Small populations of
halophytes in restricted areas may be at greater risk of extinction.
The halophytic flora and vegetation of the study area also show some similarities with
those occurring in isolated inland halophytic areas in more north-western and northern
parts of central Europe, namely north-western Bohemia (Toman 1976, 1988, Šumberová
et al. 2007), Saxony-Anhalt and Thuringia in central Germany (Altehage & Rossmann
1939, Schubert 2001, Piernik 2012, Dítě et al. 2022) and the Kujawy region in Poland
(Wilkoń-Michalska 1963, Piernik 2012). The Bohemian sites have always had a limited
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 93
number of halophytic species, probably due to their isolated location and a small area of
saline habitats. Several continental halophytes occurring in southern Moravia (e.g.
Cirsium brachycephalum,Crypsis aculeata,C. schoenoides,Salicornia perennans,
Suaeda prostrata and Tripolium pannonicum) havenever been found there (Kaplan et al.
2019). Accordingly, halophytic vegetation is poorly developed in Bohemiaand only rep-
resented by the alliances Chenopodion rubri,Meliloto dentati-Bolboschoenion maritimi
and Juncion gerardii (Šumberová et al. 2007). In contrast, German and Polish inland
saline habitats are richer in specialist species, including the succulent halophytes
Salicornia europaea and Suaeda maritima. Traditionally, halophytic vegetation in these
inland areas has been classified in the coastal saltmarsh alliances described from the
North Sea or the Baltic Sea (Schubert 2001, Matuszkiewicz 2007). However, Dítě et al.
(2022) compared the species composition of German and Polish inland halophytic grass-
lands with corresponding plant communities on the Baltic coast and in the Carpathian
Basin. They found these grasslands to be more similar to inland Pannonian than to coastal
halophytic vegetation. Consequently, they proposed reclassifying the German and Polish
inland halophytic grasslands to the continental inland alliances Puccinellion limosae and
Juncion gerardii.
Saline habitats: partly natural, partly man-made
The halophytic flora and vegetation in southern Moravia and northern Lower Austria are
restricted to small areas, mostly shallow depressions with clayey subsoil over Tertiary
deposits rich in magnesium and sodium sulphates. In the dry and summer-warm climate
in this area, the salt was pushed up in the soil and locally formed efflorescences on the
surface. Unique conditions occurred in the salt pan of the former Čejč Lake, which had no
outlet (Jordán z Klauznburku 1580: 284–293). In places, the salt content in the soil was
increased by water from mineral springs, such as near the villages of Šaratice,
Nesvačilka, Čejč and Sedlec. At least in some places, soil salinity was high enough to
allow the occurrence of the Salicornion prostratae vegetation with low-competitive suc-
culent halophytes Salicornia perennans and Suaeda prostrata. For millennia, halophytes
were supported by livestock grazing (Vicherek 1973, Eliáš et al. 2020), which protected
the competitively weak but stress-tolerant halophytes from competing non-halophytes on
moderately saline soils. Since the Middle Ages, new saline habitats had probably devel-
oped around newly created fishponds (Hurt 1960). These fishponds were surrounded by
pastures and meadows, which were wet in the spring but dried out in the summer. Many
fishponds were later drained, but some halophytes survived in grasslands developed on
their drained bottoms.
Additional habitats were created when the Vienna–Brno railway was built in the late
1830s. Halophytes readily colonized the ditches along the railway near the villages of
Rakvice and Šakvice. Halophytes also occurred directly in settlements. The common
land in the villages was freely accessible for grazing livestock, domestic pigs and poultry.
There was hardly any paving, and each village had at least one small pond with unpaved
shallow banks used for horse bathing and domestic geese and ducks. Such sites were suit-
able for halophytes. Records from the 19th and early 20th century show that Glaux
maritima occurred in many villages in the Damnice-Olbramovice, Hevlín-Nový Přerov,
Daníž and Pulkau districts (e.g. Reuss 1873, Oborny 1912). In halophytic districts, Glaux
94 Preslia 94: 13–110, 2022
maritima,Plantago maritima and Spergularia media occurred almost continuously (e.g.
Kalbrunner 1855, Fietz et al. 1923, Gilli 1930). Overall, historical records prove that
humans have long promoted the spread of halophytic flora and vegetation in the study
area.
Not only man-made but also destroyed by man
Halophytic habitats, confined to small areas with a specific combination of topography,
soil properties, moisture, climate and management, were fragile and vulnerable to disrup-
tions, especially changes in the water regime. Such interventions began in the early 19th
century when sugar beet was introduced as a new crop (Tomšovic 1990) to replace cane
sugar, which became unavailable due to the naval blockade of the European continent
during the Napoleonic wars. Sugar beet is salt-tolerant, and its cultivation on drained,
albeit somewhat saline, arable land was much more profitable than grazing or haymaking
on saline grassland. For this reason, the Měnín Fishpond, Kobylí Lake and Čejč Lake
were drained and converted to arable land between 1822 and 1855 (Krzisch 1859,
Spitzner 1894, Hurt 1960).
In some areas, draining of agricultural land was made possible only by river
channelization. For instance, the lower section of the Pulkau stream and the Dyje river
section between its confluence with the Pulkau and the village of Nový Přerov were
channelized already in 1832 (Kalbrunner 1855). The Dyje section between the villages of
Nový Přerov and Mušov was channelized in 1888–1902 (Fiala & Štěpánek 1992). In the
Pulkau district, the once channelized bed of the Pulkau was further deepened downstream
the village of Hadres in 1958, and the channelization continued further upstream to the
village of Zellerndorf (Lindermayer 2008). These measures caused a drop in the ground-
water level in the floodplain, which enabled large-scale drainage and subsequent conver-
sion into arable land of wet habitats adjacent to the stream corridor. The large-scale drain-
ing of agricultural land started in the late 1950s, and 105,000 ha of land were drained in
the former South Moravian region by 1973 (Března et al. 1974).
Already the 19th-century botanists saw the drainage as a cause of the decline of
halophytes (e.g. Krzisch 1859). Still, these interventions were not fatal in the short term
because small patches of halophytic habitats remained preserved at less affected sites.
For instance, in the Čejč district, even the succulent halophytes survived at the Zápověď
and Na Rybníčku sites for more than a century after the drainage of both saline lakes.
However, the halophytic habitats were continuously reduced in size and fragmented, and
the population size of individual halophytes shrank. The decline of well-known
halophytic sites in southern Moravia was briefly discussed by Krist (1935). He also
underlined the drop in groundwater level, caused by drainage, as the main threat to
halophytes, pointing to the drainage of the Plácky site near Velké Němčice. As reported
by Husák & Jatiová (1984), the surroundings of the Plácky site, put under conservation in
1950, was repeatedly drained in 1958–1965 and again in 1970. In 1968–1972, drainage
ditches were dug out around the whole nature reserve, and the groundwater level dropped
to depths of 120–150 cm. Consequently, most halophytesdisappeared by the mid-1970s.
Another factor that strongly accelerated the decline of halophytes was the land-use
change. Grazing livestock and poultry were almost ubiquitous in and around villages
until the 1950s. Common pastureland (Hutweide) indicated in the 3rd Military Survey
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 95
maps, usually in wet parts of the landscape, often corresponds with the localities of
halophytes reported in the literature and on herbarium labels. Grazing animals trans-
ported diaspores of halophytes (e.g. Plantago maritima) along droveways to new suitable
sites and made their occurrence possible on moderately saline soils through disturbances.
However, livestock grazing almost disappeared from southern Moravia with the intro-
duction of collective farming during the 1950s. The surroundings of small fishponds and
other places in villages and their peripheries continued to be grazed by domestic geese
and ducks until the 1960s. However, free-running poultry also gradually disappeared
from rural areas. For instance, the Slanisko u Nesytu site was still grazed by a large num-
ber of domestic geese in the 1960s, but there was no longer any grazing in the 1970s
(Danihelka 2005). Husák & Jatiová (1984) reported that the livestock grazing at the
Plácky site ceased in the mid-1960s. In contrast, the Hintausacker site near Zwingendorf
in Lower Austria was grazed by geese until the 1980s, and one of the small ponds there
was used for horse bathing (Holzer et al. 2002). This spontaneous management was able
to keep at least some parts of the site without tall-growing vegetation, i.e. suitable for less
competitive halophytes. However, many abandoned halophytic grasslands outside the
villages were overgrown by dense stands of Phragmites australis.
The changes in land use were paralleled by the changes in rural settlements. The pub-
lic space in the villages was gradually changed to resemble towns. Along with the elimi-
nation of small dumping places and free-roaming domestic animals, roads were paved,
streams deepened, straightened, channelized or hidden into subsurface pipes, and small
village ponds, initially with flat shores and fluctuating water level, were either filled with
earth or changed into water reservoirs with a vertical embankment of stone and concrete.
Unpaved public spaces were converted to regularly mown lawns or small city-like parks,
often with plantations of ornamental trees and shrubs. As a result, habitats suitable for
halophytes completely disappeared from most villages or were reduced to small and iso-
lated patches. Vegetation types with specialized halophytes (e.g. the alliance Salicornion
prostratae or associations Puccinellietum limosae,Scorzonero parviflorae-Juncetum
gerardii and Agrostio stoloniferae-Juncetum ranarii) were converted into species-poor
grasslands dominated by generalist grasses such as Festuca arundinacea and Lolium
perenne. In a better case, they were transformed to the Loto tenuis-Potentilletum anseri-
nae grasslands containing a few less demanding subhalophytes. In southern Moravia,
these changes were almost finished by the late 1970s, while this process was somewhat
delayed in Lower Austria. As a result, Glaux maritima, once recorded directly in village
centres, completely disappeared from these sites during the 1960s. In 1963, this species
was recorded from grazed places in the village of Dyjákovičky in the Daníž district,
which was the last such record in southern Moravia.
In summary, most of the halophytic habitats in southern Moravia and the Pulkau valley
were destroyed by draining and subsequent conversion into arable land. Small patches of
halophytic habitats in villages were also destroyed directly through improvements of
public spaces. The remaining halophytic habitats at the village peripheries and those out-
side villages, usually too small for intensive farming, were abandoned and, consequently,
disappeared due to a secondary succession of reed stands and woody vegetation.
These patterns and causes of the decline of halophytic flora and vegetation were simi-
lar to those observed in the whole Carpathian Basin (e.g. Eliáš et al. 2020), southern
Slovakia (Vicherek 1973, Dítě et al. 2014) and elsewhere in eastern Austria (Oberleitner
96 Preslia 94: 13–110, 2022
et al. 2006). However, the timing and relative importance of human interventions and
succession after abandonment were site-specific.
From the non-intervention conservation to targeted management and restoration
The fast decline of saline habitats during the 20th century stimulated interest in their con-
servation and led to designating several sites as protected areas, namely Zápověď
u Terezína (designated as a private reserve in 1944, as a Nature Reserve in 1952 and abol-
ished in the early 1990s), Plácky (designated in 1950, now Nature Monument), Slanisko
u Nesytu (1961, now National Nature Reserve), Zwingendorfer Glaubersalzböden (1979,
Naturschutzgebiet consisting of two parts, Hintausacker and Saliterweide), Slanisko
Dobré Pole and Slanisko Novosedly (both 1993, now Nature Reserves) and Trkmanec-
Rybníčky (2008, Nature Monument). Other localities without a status of (National)
Nature Reserve or (National) Nature Monument were designated as the Sites of Commu-
nity Importance (SCI) within the Natura 2000 network: Zřídla u Nesvačilky, Rumunská
bažantnice (including Císařská obora) and Trkmanské louky. Consequently, the most
valuable of the remaining halophytic sites in southern Moravia and the Pulkau valley are
currently under protection except for Zápověď, the site with Cirsium brachycephalum
near Hevlín and the newly discovered halophytic sites along the Štinkovka stream near
Šakvice.
Around the mid-20th century, nature conservationists supported non-intervention as
the primary conservation approach because disturbances were considered detrimental.
For instance, Krist (1935) described the remaining halophytic patches at the Plácky site
as “trampled by cattle” and referred to the vegetation as “destroyed”. He considered graz-
ing pressure as one of the causes of halophyte decline, in addition to drainage. Fröhlich &
Švestka (1956), describing the Slanisko u Nesytu site near Sedlec, complained about the
“spread of weeds” and disturbances of “natural plant communities” caused by grazing
and mowing. However, referring to the attempts to convert a part of this site to arable land
in 1932, they already recognized the positive effect of disturbances on halophytes: “luck-
ily, former attempts to use saline soils for economic purposes failed […]. The cultivation
of soil supported the occurrence of halophytes, while the crops suffered”.
Until the mid-1970s, there was no conservation management in any of the protected
halophytic sites. Only the sites at village peripheries were managed spontaneously by
local people who used them for grazing by free-roaming chickens, occasional haymaking
(e.g. Dobré Pole), and sometimes waste removal (e.g. Hintausacker near Zwingendorf;
Holzer et al. 2002). The Dobré Pole site, for some decades the only site of Glaux
maritima in southern Moravia, was saved by local footballers, who mowed and trampled
the playground and the adjacent area. In contrast, the legally protected Zápověď and
Plácky sites, both located far from the nearest villages, were entirely abandoned and lost
their halophytic flora and vegetation.
Nature conservation authorities realized the importance of management in halophytic
reserves in the 1970s. In 1976, they proposed a project to raise the groundwater level at
the Plácky site, but it was not implemented (Husák & Jatiová 1984). At the Slanisko
u Nesytu reserve, shallow ditches were dug out in the mid-1970s to create suitable habi-
tats for halophytes (Grulich 1987, in litt. 2021). However, these ditches were left
unmanaged and overgrew by perennial grasses, mainly Agrostis stolonifera (Danihelka
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 97
2005, Čtyroký et al. 2007). Husák & Jatiová (1984) recommended regular disturbances
of soil surface at the Plácky site to reduce vegetation cover and create suitable conditions
for competitively weak halophytes. These proposals failed partly due to the political and
economic situation in Czechoslovakia at that time: nature conservation authorities had
limited power and almost no financial resources for conservation management. Even if
some money was allocated for conservation purposes, finding a company or a coopera-
tive farm with required machinery and free working capacities was challenging. Never-
theless, the situation of the Hintausacker site in Zwingendorf was similar despite the
different political and socioeconomic conditions in Austria. Although the site has been
formally protected since 1979, conservation management by bulldozing, scrub removal
and mowing started as late as 1995 (Holzer et al. 2002).
The Slanisko u Nesytu site was probably the first halophytic reserve in the study area
with regular conservation management. In 1991, a shallow ditch was created in the west-
ern part of the reserve, in which halophytic grasslands of Scorzonero parviflorae-
Juncetum gerardii developed spontaneously (Danihelka & Hanušová 1995, Hanušová
1995). Since 1993, the reserve has been mown regularly. Grazing was restored in 2000,
first with a small flock of sheep and goats and in two-year intervals (Danihelka 2005,
Čtyroký et al. 2007). All these early management measures were limited by the lack of
money and logistic difficulties. There were neither small farmers with necessary machin-
ery nor private persons specialized in the management of protected areas. The local coop-
erative farm was also unable to work in small areas and perform fine-scale conservation
management. The traditional shepherding skills were forgotten, and no animals were
available. However, funding was slowly increasing, and nature conservation authorities
gradually improved their skills and experience in active management.
Regular grazing was established at some sites after 2010. Among livestock, horses
proved to be the most effective agents in restoring saline habitats for their high mobility
and weight (Kmet et al. 2018). Apart from the plant biomass removal, they disturb the
soil surface, and by compressing the soil, they enhance evaporation, hence salt precipita-
tion at the soil surface. The establishment of grazing resulted in pronounced improve-
ment of conditions at some saline sites. Some protected areas are also affected by
eutrophication due to their direct contact with surrounding arable fields and, in the
absence of management, by expanding competitively strong species, both native such as
Phragmites australis andaliensuchasSolidago gigantea.
Recent evidence also shows that restoration of saline habitats is possible at some sites
where they have disappeared. Two notable examples of successful (yet unintentional)
restoration of halophytic vegetation are the Zápověď site near Terezín and saline habitats
along the Štinkovka stream near Šakvice. At both sites, no halophytes were recorded for
several decades. However, after creating shallow pools and depositing removed earth in
the surroundings, several halophytic species appeared spontaneously in large popula-
tions, most probably by germinating from the soil seed bank. For example, Crypsis
aculeata,Glaux maritima,Spergularia marina and S. media appeared at Zápověď after
its restoration in 2003 (Slavík in Hadinec & Lustyk 2009) while Crypsis schoenoides,
Samolus valerandi and Spergularia marina appeared near Šakvice after wet arable land
was transformed into a semi-natural area in 2018 (Chytrý & Danihelka 2021). Moreover,
the whole halophytic plant communities corresponding to well-defined phytosocio-
logical associations soon developed at these sites. Similar developments occurred after
98 Preslia 94: 13–110, 2022
the restoration of the Trkmanec-Rybníčky site in 2006-2008. These cases of successful
restoration of halophytic habitats and their specialized flora and vegetation are important
examples for future restoration projects, demonstrating that partial return of halophytes to
the landscape is possible. However, the long-term preservation of halophytic flora and vege-
tation at restored sites requires continued management, including the removal of reeds and
other tall plants through grazing, mowing and periodical disturbance of the soil surface.
Our database of exactly located relevés sampled in 2020 (Danihelka et al. 2021) pro-
vides the basis for the future monitoring of vegetation and habitat changes, which can
support proper management planning. However, the disrupted water regime at most
halophytic sites and their surroundings limits the full recovery of halophytic flora and
vegetation. Restoration of the water regime in protected areas is currently the main chal-
lenge for in situ conservation of halophytic flora and vegetation and restoration of its
habitats.
Halophytes: those lost and those to be saved
Halophytic sites in the study area lost a considerable proportion of their former floristic
diversity. The pattern shown in Fig. 8 resulted from two different processes. From the
beginning until the 1930s, the graph mainly showsprogress in floristic research, whereas
in later years, it mainly reflects the destruction of individual sites and species. The recent
halophytic flora of the study area is strongly impoverished. Galatella cana,Salicornia
perennans,Suaeda prostrata and Triglochin maritima went regionally extinct. These
losses are hardly replaceable, at least in the short time, as shown by the failed attempt to
re-establish the population of Salicornia perennans at the Slanisko u Nesytu site in the
mid-1980s (V. Grulich, in litt. 2021). At least the two succulent halophytes require
a much higher salinity degree than currently encountered in the study area. At present,
conservation efforts should thus focus on protecting the remaining 13 halophytes still
occurring in the area and their habitats.
Of the remaining species, the least endangered ones are Juncus gerardii,Plantago
maritima and Spergularia marina.Juncus gerardii cansurviveevenindegradedsaline
habitats as long as they are humid in the spring. Although it is often neglected and con-
fused with J. compressus, it seems to be the most widespread halophyte of the study area.
Plantago maritima is still present at several sites, and at least the populations at the
Slanisko u Nesytu, Dobré Pole, Hintausacker and Saliterweide sites are still large enough
not to be in immediate danger of local extinction.Its survival may beexplained by its rel-
atively loose relations to salinity: in our dataset, it tends to grow preferably at sites with
lower salinity (Table 2). Spergularia marina occurs in most halophytic districts, but most
of its populations are small. It is more often found on wet arable land next to the
halophytic sites than directly within these sites. Conservation management of this species
includes continued cultivation of periodically flooded arable land. The species appears
there in wet years when the crop fails to develop or when a wet patch is left uncultivated.
Plantago maritima,Spergularia marina and S. media have recently spread along roads
salted for winter de-icing and in cities (Ducháček & Kúr in Kaplan et al. 2016, Danihelka
& Kaplan in Kaplan et al. 2018). However, nothing is known about the genetic relation-
ship between the autochthonous populations from saline habitats and the recently spread-
ing populations.
Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 99
Glaux maritima and Tripolium pannonicum currently occur at three sites. Both of
them have two populations that are large enough to escape the immediate risk of extinc-
tion. These species readily respond to mowing and grazing applied as a part of conserva-
tion management. The current status of Spergularia media is similar to the previous two
species. Cirsium brachycephalum is currently present at two or three sites; the population
at the Trkmanec-Rybníčky and Trkmanské louky sites seems to be stabilized and large
enough, while the populations at the Hevlín and Saliterweide sites are small and require
monitoring and conservation management. We failed to confirm the occurrence at
Saliterweide in July 2021. Only two of the populations of Samolus valerandi known to us
are found in protected areas, both being large enough and monitored by nature conserva-
tion authorities. In addition, there are recent records from further five sites, and other
occurrences may be discovered in the future on drained fishpond bottoms somewhere
else in the area. At the Nesyt Fishpond, S. valerandi co-occurs with Crypsis schoenoides.
The latter species forms locally large stands there on exposed parts of the bottom in dry
years and does not seem to be in danger. However, this may change in the future if non-
saline water from the Dyje river was pumped into the Lednice Fishpond system, as cur-
rently discussed. This happened already in the early 1980s and may have been one of the
factors causing the desalination at the Slanisko u Nesytu site (Danihelka & Hanušová
1995).
The populations of four remaining species, Bupleurum tenuissimum,Crypsis aculeata,
Scorzonera parviflora and Taraxacum bessarabicum, require monitoring and targeted
management.Of them, only Bupleurum tenuissimum is not in immediate danger because
the population at the Slanisko u Nesytu site is rich in individuals and well responding to
management. Taraxacum bessarabicum may have survived only at two or three sites, in
populations counting less than several hundred or several dozen plants. Crypsis aculeata
and possibly also Scorzonera parviflora are on the brink of extinction in the study area.
Populations of both species have suffered from a series of dry years since 2015. However,
S. parviflora profited from the humid spring of 2021, and an abundant population
appeared at the Slanisko u Nesytu site. Relatively rich population occurs at the Hintau-
sacker site near Zwingendorf, and two plants were observed at the Kalužiny site in the
Měnín-Šaratice district in 2017. Crypsis aculeata has survived at one to three sites: it was
last recorded at the Zápověď site in 2008, while at the Slanisko u Nesytu site, it is con-
fined to a few square meters in an artificial ditch, and the population size fluctuates from
a few to several hundred plants. A few plants were observed on the drained bottom of the
Nesyt Fishpond in 2020.
Although the decline of saline vegetation in the last two centuries was caused mainly
by human activities, some halophytes were able to take advantage of those changes. Dur-
ing the last decades, some halophytes spread massively along motorways. The critically
endangered species Puccinellia distans and Spergularia marina became common ele-
ments of salted roadsides while being at extinction risk in natural habitats. The former
species dominated ruderal grasslands along salted roads in the cities already in the late
1970s (Hadač et al. 1983). Limonium gmelinii, a species of continental salt steppes, was
found as a new non-native species of Moravian flora on a motorway edge near Brno in
2009 (Kocián et al. 2016). Since the roadsides can be a partial substitute for some drier
types of natural saline habitats, their colonization by other halophytes is possible.
100 Preslia 94: 13–110, 2022
Despite the spread of some halophytes in anthropogenic habitats, natural and semi-
natural occurrences of halophytes and their plant communities are in general decline. In
the long term, most halophyte species are at risk of regional extinction due to the extinc-
tion debt phenomenon (Figueiredo et al. 2019). The reasons are obvious: many popula-
tions are small in numbers, some survive in suboptimal conditions, and there are no suit-
able habitats to colonize. Therefore, targeted management of the remaining halophytic
habitats and restoration at selected sites where they have disappeared seems to be the only
way to conserve their populations.
Supplementary materials
Fig. S1. – Results of the generalized linear models (binomial distribution, logit link function) linking the proba-
bility of species occurrence to soil pH and conductivity
Table S1. – Full version of the synoptic table (Table 1) of 14halophytic associations based on the relevés classi-
fied by the expert system
Data S1. – Shape files with the delimitation of ten halophytic districts of southern Moravia and northern Lower
Austria
Data S2. – Floristic records of the 17 halophytic species analyzed in this study
Data S3. – Metadata on relevés classified by the expert system
Supplementary materials are available at www.preslia.cz
Acknowledgements
This study is a result of the Geobotanical Project course at Masaryk University. It was prepared by Botany stu-
dents KC, MH, PH, KK, FK, AK, KS and DS supervised by JD and MC, with contributions from HP, EŠ and
MV. JD, MV and MC were funded by the Czech Science Foundation (project no. 19-28491X), JD also by long-
term research development project no. RVO 67985939 of the Czech Academy of Sciences and the other authors
by the Vegetation Science Group, Department of Botany and Zoology,Faculty of Science, Mas aryk University.
We thank the late Vít Grulich for valuable comments, Harald Niklfeld and Christian Gilli for some halophyte
records from northern Lower Austria, Nature Conservation Agency of the Czech Republic for providing us
unpublished survey reports and Židlochovice Forest Enterprise for research permit for the Císařská obora site.
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Danihelka et al.: Halophytic flora and vegetation in southern Moravia and northern Lower Austria 109
Halofytní flora a vegetace jižní Moravy a severních Dolních Rakous:
minulost a současnost
Slaniska představují významnou součást jihomoravské krajiny v území mezi Znojmem, Brnem a Břeclaví,
jakož i v údolí Pulkavy v příhraniční části Dolních Rakous. V této studii shrnujeme historií slanomilné flóry
a vegetace tohoto území od počátku19. století a na základě terénního výzkumu v létě 2020 popisujeme souč asný
stav zbylých halofytních lokalit. Lokality slanisk dělíme do deseti okrsků, a to měnínsko-šaratického, velko-
němčického, šakvického, rakvického, čejčského, sedleckého, damnicko-olbramovického, hevlínsko-novopře-
rovského, danížského a pulkavského. Nejlépe vyvinutá slaniska se v minulosti nacházela ve velkoněmčickém,
šakvickém, rakvickém, čejčském a sedleckém okrsku. V současnosti se nejlépe zachovalé halofytní biotopy
nacházejí v sedleckém, hevlínsko-novopřerovském a pulkavském okrsku. Dále jsme zpracovali přehled sou-
časného a historického rozšíření 17 druhů, které se ve studovaném území vzhledem k silné vazbě na zasolené
půdy považují za halofilní, a to Bupleurum tenuissimum,Cirsium brachycephalum,Crypsis aculeata,C. scho-
enoides,Galatella cana,Glaux maritima,Juncus gerardii,Plantago maritima,Salicornia perennans,Samolus
valerandi,Scorzonera parviflora,Spergularia marina,S. media,Suaeda prostrata,Taraxacum bessarabicum,
Triglochin maritima aTripolium pannonicum.ZnichGalatella cana,Salicornia perennans,Suaeda prostrata
aTriglochin maritima už na jižní Moravě i v údolí Pulkavy vyhynuly. Fytocenologické snímkyhalofytní vege-
tace jsme klasifikovali do 14 asociací, které patří do svazů Chenopodion rubri,Meliloto dentati-Bolboschoeni-
on maritimi,Cypero-Spergularion salinae,Salicornion prostratae,Puccinellion limosae,Juncion gerardii a
Festucion pseudovinae. Na základě půdní reakce a konduktivity změřené z půdních vzorků jsme analyzovali
vztah jednotlivých druhů a vegetačních typů k míře zasolení půdy. Naše syntéza ukazuje, že kdysi dobře vyvi-
nutá halofytní flóra a vegetace ve studovaném území trvale ustupovala od počátku 19. století až do 80. let minu-
lého století, zpočátku většinou v důsledku odvodňování a od poloviny 20. století také vlivem ukončení pastvy.
Ochranářská péče od 90. let zastavila úbytek halofytů na posledních několika lokalitách. Budoucnost halofyt-
ních biotopů závisí na dlouhodobé a cílené ochranářské péči a ekologické obnově.
How to cite: Danihelka J., Chytrý K., Harásek M., Hubatka P., Klinkovská K., Kratoš F., Kučerová A., Slachová
K., Szokala D., Prokešová H., Šmerdová E., Večeřa M. & Chytrý M. (2022) Halophytic flora and vegetation in
southern Moravia and northern Lower Austria: past and present. – Preslia 94: 13–110.
Preslia, a journal of the Czech Botanical Society
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www.preslia.cz
This is an open access article under the terms of the Creative Commons Attribution License, which permits use,
distribution and reproduction in any medium, provided the original work is properly cited.
110 Preslia 94: 13–110, 2022
