20/1 • 2021, 217–224
DOI: 10.2478/hacq-2021-0001
Syntaxonomical analysis of sandy grassland
vegetation dominated by Festuca vaginata
and F. pseudovaginata in the Pannonian basin
Károly Penksza1 , Péter Csontos2, Gergely Pápay1,*
Key words: coenotaxon, DanubeTisza Interfluve, degraded sandy
grassland, diagnostic species.
Ključne besede: cenotakson,
ozemlje med Donavo in Tiso,
degradirana peščena travišča,
diagnostične vrste.
Received: 29. 01. 2019
Revision received: 27. 10. 2020
Accepted: 03. 12. 2020
Co-ordinating Editor:
Orsolya Valkó
Abstract
Festucetum vaginatae Rapaics ex Soó 1929 em. Borhidi 1996 is a characteristic
association of the calcareous sandy areas of the Pannonian basin; its dominant
grass species is Festuca vaginata. Another typical species of these sandy areas is
the newly discovered F. pseudovaginata. The question is whether F. pseudovaginata
forms an independent coenotaxa? Our study proved that F. vaginata and
F. pseudovaginata populations grow separately and compose different associations.
Stands dominated by F. pseudovaginata had a higher species richness and
harboured twice as many Festuco-Brometea species compared to the Festucetum
vaginatae stands. Diagnostic species of the Festucetum pseudovaginatae association
are Festuca pseudovaginata, Colchicum arenarium, Ephedra distachya, Koeleria
majoriflora, and Astragalus onobrychis. The number of species, the density of the
individuals, and the variability and diversity of the vegetation separated it from
the Festucetum vaginatae association; thus, it can be considered an independent
endemic association. Festucetum pseudovaginatae has its own differentiating and
dominant species: Carex stenophylla, Cynodon dactylon, Eryngium campestre,
Kochia laniflora.
Izvleček
Festucetum vaginatae Rapaics ex Soó 1929 em. Borhidi 1996 je značilna
asociacija na apneniških peščenih območjih v Panonski nižini; dominantna
trava je Festuca vaginata. Druga značilna vrsta teh peščenih območij je novo
odkrita vrsta F. pseudovaginata, za katero se pojavlja vprašanje, ali gradi tudi
samostojne sintaksone. Z našo raziskavo smo dokazali, da populacije vrst F.
vaginata in F. pseudovaginata uspevajo ločeno in gradijo različne asociacije.
Sestoji, v katerih prevladuje F. pseudovaginata so vrstno bogatejši in v njih se
pojavlja dvakrat več vrst razreda Festuco-Brometea v primerjavi s sestoji Festucetum
vaginatae. Diagnostične vrste asociacije Festucetum pseudovaginatae so: Festuca
pseudovaginata, Colchicum arenarium, Ephedra distachya, Koeleria majoriflora in
Astragalus onobrychis. Število vrst, gostota osebkov, variabilnost in raznolikost
vegetacije jo jasno loči od asociacije Festucetum vaginatae, zato jo lahko
obravnavamo kot samostojno endemično asociacijo. Festucetum pseudovaginatae
označujejo lastne razlikovalne in dominantne vrste: Carex stenophylla, Cynodon
dactylon, Eryngium campestre, Kochia laniflora.
1 Szent István University, Institute of Crop Production Sciences, H–2100 Gödöllő, Páter K. u. 1, Hungary
2 Institute for Soil Science and Agricultural Chemistry, Centre for Agricultural Research H–1022 Budapest, Herman O. út 15, Hungary
* Corresponding author: E-mail: geri.papay@gmail.com
217
20/1 • 2021, 217–224
Introduction
Pannonic sandy grasslands are important habitats harbouring several endemic and rare plant species, included
in the Natura 2000 network as ‘Pannonic sand steppes’
(habitat code: 6260). Festuca vaginata forms psammophytic communities on calcareous areas in the Pannonian basin. The alliance of Festucion vaginatae Soó 1929
occurs from the western shores of the Black Sea to the
Pannonian Basin along the Danube valley, encompassing
the Pontic and Pannonian regions. Festucetum vaginatae
Rapaics ex Soó 1929 em. Borhidi 1996 is the most widespread indigenous association in this alliance.
Three associations of sandy grasslands occur in acidic
sandy areas of the Pannonian region, of which Festuco
dominii-Corynephoretum Borhidi (1958) 1996 (Borhidi
et al. 2012) is the most widespread. Festuca dominii is
considered to be the dominant species of this association, which is why it is named after this species. Festuco
vaginatae-Corynephoretum Soó in Aszód 1935 is characteristic in the eastern parts of the Pannonian flora region
(Nyírség), where Festuca vaginata is the dominant grass
species. Achilleo ochroleucae – Corynephoretum (Hargitai
1940) Borhidi 1996 is found mainly in the middle sand
dune range in the area between the Danube and Tisza rivers. The stands are dominated by the greyish tussocks of
Corinephorus canescens forming and open and loose mat
with 40–50% vegetation cover (Borhidi et al. 2012).
There are several taxonomic debates around Festuca
species occurring in the Pannonian sandy grasslands. The
currently valid classification of Festuca dominii Krajina is
Festuca psammophila subsp. dominii (Krajina) P. Šmarda,
which was clarified by Šmarda et al. (2007). Even though
the basic species is missing from the Pannonian region
(Penksza 2019), according to both the collected and the
herbarium examples. Many of the examined Festuca vaginata individuals were found to have short (0.1)0.2–0.4
mm spikes on the glume, though this can be considered
a variety of the F. vaginata species (Šmarda et al. 2007),
which taxa do not have their own range.
Festuca pseudovaginata is a recently discovered fescue species of the Pannonian sandy grasslands (Penksza
2003). The distinctive features of F. pseudovaginata are
the following (Figure 1). The stem is 20–35 cm high and
hairless. The basal leaves are hairless, with a light green
colour, they are not glaucous. The surface of base leaves is
smooth. The base leaves have a continuous sclerenchyma
ring. The sclerenchyma ring is extraordinarily thick, and
not uniform in the side of the mesophyllum. In young
leaves ragged sclerenchyma rings can also be observed.
The awn on the outer lemma is 1.2–1.8 mm long. The
locus classicus of this species is Kis-tece legelő (Kis-tece
218
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
pasture) situated near Vácrátót, Hungary (Penksza 2003).
Our preliminary examinations on the distribution and
habitat characteristics of F. pseudovaginata performed
by Szabó et al. (2017) revealed that the species prefers
degraded habitats, which was also confirmed by the soil
parameters of the studied populations. After the discovery of the species, our aim was to search systematically
the Pannonian sandy regions to reveal new occurrences
and to examine the plant community composition of the
sandy grasslands where F. pseudovaginata occurs.
Materials and Methods
During the systematic sampling of sandy grasslands, Festuca specimens were collected at 20 sites in the Carpathian Basin. In this paper we present data from two sites:
the Tece pasture near Vácrátót and from Szigetmonostor
on Szentendre Island where both F. vaginata and F. pseudovaginata occurred.
The two examined areas are located in the northern
part of the Danube-Tisza Interfluve, under similar abiotic
conditions. The mean temperature in the coldest month
(January) is 3 °C, in the warmest month (July) 20.5 °C,
the mean annual temperature is 9.5 °C. Precipitation is
550–600 mm (Péczely 2006). Due to the low precipitation and poor water–holding capacity of the soil, sites
are characterized by an extremely dry and warm microclimate (Péczely 2006). The close surface geology of the
Danube–Tisza Interfluve region is determined by mainly
carbonatic eolian sediments of the Danube River. At the
higher, dry landscape positions carbonatic shifting sands
and humic sandy soils (Arenosols) can be found, which
are characterized by unfavourable physical and chemical
properties (high permeability and low water and nutrient
storage capacity), thus have low fertility (Dövényi 2010,
Stefanovits 2010).
The main difference between the two areas lies in
landscape use and history. The pasture at Tece has been
affected by continuous landscape transformation activities, including the cutting of the woody component of
the forest steppe (Fekete et al. 1976). The area at Szigetmonostor on the Szentendre Island on the Danube has
been being used similarly for centuries, i.e. pasturing on
natural grasslands. The mosaic structure of the foreststeppe complexes were not affected, as pasturing has
been typical on the grassland patches without deforestation (Böhm 2015).
In both sites, a series of F. vaginata and a series of F. pseudovaginata dominated plots were sampled. Coenological
sampling was carried out in May-June and September
2018, we recorded the percentage cover of vascular plant
species in quadrats of 2 m × 2 m. In total, 15 relevés
20/1 • 2021, 217–224
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
Figure 1: Appearance of the two fescue taxa. A: Tussocks of F. pseudovaginata (left) and F. vaginata (right); B: inflorescence of F. pseudovaginata;
C: inflorescence of F. vaginata). Photos: Dr. Károly Penksza
Slika 1: Izgled dveh vrst bilnic. A: Šopa F. pseudovaginata (levo) in F. vaginata (desno); B: socvetje F. pseudovaginata; C: socvetje F. vaginata).
Fotografije: Dr. Károly Penksza.
219
Results
A total of 69 vascular plants were recorded in the coenological samples in the Festuca pseudovaginata dominated grasslands. There were 25 species in grasslands
where Festuca vaginata was dominant. As a result, there
were significant differences in the species richness of
the two grassland types. Analysis of variance followed
by Tukey-Kramer test supported significant differences
of average species numbers between all possible pairs
of grassland types and study sites, except for the pair of
F. vaginata (Tece) versus F. vaginata (Szigetmonostor).
Comparison of the two grassland types’ species numbers
(regardless of study sites) showed a strong significant
difference (nF. vag.= 15, nF. pseudovag.= 19, meanF. vag.= 13.9,
meanF. pseudovag = 29.4, t = 8.789, p < 0.0001).
The description of the plant communities dominated
by F. pseudovaginata are given below.
Association Festucetum pseudovaginatae ass. nova
hoc. loco (Table 2) holotype SzFp1
Diagnostic species for the association: Festuca pseudovaginata, Colchicum arenarium, Ephedra distachya,
Koeleria majoriflora, Astragalus onobrychis
Constant species: Carex stenophylla, Cynodon dactylon,
Eryngium campestre, Kochia laniflora
The species occurring both in F. vaginata and F. pseudovaginata grasslands were the ones typical in natural and
semi-natural sandy grasslands. Of these, the species of
Pannonian psammophytic grasslands (Festucion vaginatae
Soó 1929): Arenaria serpyllifolia, Bromus squarrosus, Cen220
taurea arenaria, Erysimum diffusum, Fumana procumbens
and Koeleria glauca. The various species of Atlantic sandy
grasslands (Corynephoretalia Klika 1934) were present in
both types: Cerastium semidecandrum, Rumex acetosella,
Veronica dillenii.
The species typical of arid and semi-arid rocky and
steppe grasslands (Festuco-Brometea), were mostly found
only in quadrats with F. psedovaginata (e.g. Alyssum alyssoides, Erophila verna, Hypericum perforatum, Phleum
phleoides, Poa angustifolia and P. bulbosa). Carex liparicarpos, C. stenophylla (continental sand steppes – Festucetalia vaginatae & rupicolae species), Viola kitaibeliana
and Thymus praecox (Festucetalia vaginatae & valesiacae)
were present only in Festuca pseudovaginata grasslands. In
the more disturbed F. pseudovaginata grasslands in Tece,
there was a large number of various ruderal plant species
characteristic of Chenopodietea, Chenopodietea & Secalietea, Secalietea, Aphanion, such as Ambrosia artemisiifolia,
Anchusa officinalis, Anthemis austriaca, Apera spica-venti,
Conyza canadensis, and with the dominance of Cynodon
dactylon. These are well-separated in the classification
(Figure 2). There is a clear separation between the two
associations. The F. vaginata group is more uniform, with
F. pseudovaginata diverging at two higher levels of difference. This is also apparent in the DCA analysis (Figure 3).
were recorded in the centre of the patches dominated by
F. vaginata and 19 relevés in the F. pseudovaginata stands,
resulting in a total of 34 relevés. Species names follow
the nomenclature of Király (2009), phytosociological nomenclature follows Borhidi et al. (2012).
We used an agglomerative cluster analysis technique
(SYN–TAX program package): a fusion algorithm was a
combinatorial method (minimizing increase of variance)
and the correlation was used as comparative function
(Podani 1997). Calculations were performed in the SYN–
TAX IV program (Podani 2001). For comparing the species numbers of relevés in the different grassland types
and sites, one way ANOVA and Tukey-Kramer Multiple
Comparisons test as a post hoc test were used, since the
raw data showed Gaussian distribution and fulfilled homoscedasticity test. Species numbers of the two grassland
types was also compared by an unpaired t-test based on
data merged according to the two sites. Differences were
considered significant at p < 0.01 level. DCA ordination was used to visualize the species composition of the
F. vaginata and F. pseudovaginata series in the two sites.
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
20/1 • 2021, 217–224
Figure 2: Classification of the sample areas. (FpTe: Festuca
pseudovaginata series at Tece; FpSz: F. pseudovaginata series at
Szigetmonostor; FvTe: F. vaginata series at Tece; FvSz: F. vaginata series
at Szigetmonostor).
Slika 2: Klasifikacija vzorčenih območij (FpTe: serija Festuca
pseudovaginata pri kraju Tece; FpSz: serija F. pseudovaginata pri kraju
Szigetmonostor; FvTe: serija F. vaginata pri kraju Tece; FvSz: serija
F. vaginata pri kraju Szigetmonostor)
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
20/1 • 2021, 217–224
Figure 3: DCA analysis of the samples. The abbreviations of the
species names are given in Appendix 1.
Slika 3: DCA analiza vzorcev. Okrajšave imen vrst so v Prilogi 1.
Discussion
Based on our results, we can refine the earlier conclusion of Borhidi et al. (2012) according to which the only
dominant fescue species in Hungarian open sandy grasslands is F. vaginata. Our study shows that the species
composition of grasslands dominated by Festuca vaginata
and F. pseudovaginata is different. We have shown that
both grassland types contain numerous species from the
typical elements of the Festucion vaginatae, Festucetalia
vaginatae & valesiacae and rupicolae coenotaxa. The high
abundance of Cynodon dactylon and the appearance of
numerous species typical of ruderal vegetation types are
a clear indication of the anthropogenic disturbances in
areas colonised by F. pseudovaginata. Literature dealing
with the effects of grazing confirms that dominance of
Cynodon dactylon is often a consequence of high grazing pressure and is therefore can be used as an indicator
of intensive grazing and trampling (Szentes et al. 2011,
2012, Deák et al. 2014).
We found no species indicating anthropogenic degradation in the F. vaginata type. In addition, in the F. vaginata stands a well-developed moss-lichen synusium occurred, which is also an indication of the good habitat
quality. Contrary, in F. pseudovaginata grasslands we
detected several disturbance-tolerant and weed species.
Based on these results, it is likely that F. pseudovaginata
grasslands appear as part of a regenerative process that
starts after severe disturbances in the place of former for-
ests; however, due to the ongoing smaller disturbances,
they do not reach the state of equilibrium typical of
natural habitats and this species composition retains its
disturbed nature. Similarly, Borhidi et al. (2012) consider the Cynodonti-Festucetum pseudovinae Soó (in Aszód
1935) association description to be valid and confirm the
role of grazing in the development of that association.
This is another example for the formation of an association as a result of grazing that can be expected in the
mosaic of complex forested habitats and treeless habitats
prone to salinisation. These observations confirm our
current results, namely that new associations may develop in the mosaic of woody and barren habitats as a result
of grazing and disturbances of other origin, and that the
appearance of brown forest soil can play a significant role
in the separation of the two vegetation types (see also
Szabó et al. 2017).
We assume that the two studied Festuca species are
good indicators of the various soil types and of many
properties of the topsoil (see also Bartha et al. 2004,
2008). Bajor et al. (2016) examined the effect of removing woody vegetation on the vegetation of open sandy
grasslands. Their results show that after the removal of
woody species, a large number of ephemeral species appears, similarly to old fallow land (Prach et al. 2007,
Valkó et al. 2016) and spontaneously regenerating grasslands (Deák et al. 2015, Valkó et al. 2017). These species
are replaced by perennials in the later phases of the successional progress (Török et al. 2009; Csecserits et al.
2011, Albert et al. 2014), primarily because perennials
are probably better able to compete (Prach et al. 1997).
According to Bajor et al. (2016), the most common perennial grass species after cutting of woody vegetation was
F. pseudovaginata.
Conclusions
Festuca vaginata is the only dominant fescue species in
both calcareous areas and acidic areas in the Pannonian region. However, we showed that the indigenous
Festuca pseudovaginata also forms populations in the
central sandy areas of the Carpathian Basin. The Festuca
vaginata vegetation types hold many natural species and
also show aspects of a composition containing spring and
autumn annual or ephemeral plant species. The grasslands dominated by Festuca pseudovaginata show greater
species richness and can be considered as an independent
endemic biocenosis based on the number of species, the
density of individuals, and the variability and diversity
of the vegetation.
221
20/1 • 2021, 217–224
Acknowledgements
The work was supported by OTKA K-125423. We acknowledge the general support of the Kiskunság National Park,
the Duna-Ipoly National Park, the Fertő-Hanság National
Park, Budapest Waterworks, The Mayor’s Office, Budapest
and 20430-3/2018/FEKUTSTRAT project. For P. C. the
research was funded by the Széchenyi 2020 programme,
the EU European Regional Development Fund and the
Hungarian Government (GINOP-2.3.2-15-2016-00056).
Gergely Pápay https://orcid.org/0000-0002-0321-3859
References
Albert, Á. J., Kelemen, A., Valkó, O., Miglécz, T., Csecserits, A., Rédei,
T., Deák, B., Tóthmérész, B. & Török, P. 2014: Trait–based analysis of
spontaneous grassland recovery in sandy old–fields. Applied Vegetation
Science 17: 214–224. DOI: https://doi.org/10.1111/avsc.12068
Bajor, Z., Zimmermann, Z., Szabó, G., Fehér, Z., Járdi, I., Lampert,
R., Kerényi-Nagy, V., Penksza, P., L Szabó, Z., Székely, Z., Wichmann,
B. & Penksza, K. 2016: Effect of conservation management practices
on sand grassland vegetation in Budapest, Hungary. Applied Ecology
and Environmental Research 14 (3): 233–247. DOI: http://dx.doi.
org/10.15666/aeer/1403_233247
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
Király, G. 2009: Új magyar füvészkönyv. Magyarország hajtásos
növényei (New Hungarian Herbal. The Vascular Plants of Hungary.
Identification Key) [in Hungarian]. Aggtelek National Park
Directorate, Aggtelek, 616 pp.
Péczely, G. 2006: Éghajlattan (Climatology) [in Hungarian]. Nemzeti
Tankönyvkiadó, Budapest, 336 pp.
Penksza, K. 2003: Festuca pseudovaginata, a new species from sandy
areas of the Carpathian basin. Acta Botanica Hungarica 45: 356–372.
Penksza, K. 2019: Additions to the knowledge of Hungarian Festuca
taxa I. Taxa of Festuca psammophila series in the Festuca vaginata species
complex. Botanikai Közlemények 106(1): 65–70.
DOI: https://doi.org/10.17716/BotKozlem.2019.106.1.65
Podani, J. 1997: Syn–Tax 5.1: New version for PC and Macintosh
computers. Coenoses 12: 149–152.
Podani, J. 2001: SYN–TAX IV. Computer programs for data analysis
in ecology and systematics. in: Feoli, E. & Orlóci, L. (eds.): Computer
assisted vegetation analysis, Chapter 39: 437–452.
Prach, K., Lepš, J. & Rejmánek, M. 2007: Old Field Succession in
Central Europe: Local and Regional Patterns. In: Cramer, V. A.,
Hobbs, R. J. (eds.): Old fields: dynamics and restoration of abandoned
farmland. Island Press, Washington, DC, pp. 180–202.
Prach, K., Pyšek, P. & Šmilauer, P. 1997: Changes in species traits
during succession: a search for pattern. Oikos 79: 201–205.
Bartha, S., Campetella, G., Canullo, R., Bódis, J. & Mucina,
L. 2004: On the importance of fine–scale spatial complexity in
vegetation restoration studies. International Journal of Ecology and
Environmental Sciences 30: 101–116.
Šmarda, P., Šmerda, J., Knoll, A., Bureš, P. & Danihelka, J. 2007:
Revision of Central European taxa of Festuca ser. psammophilae Pawlus:
morphometrical, karyological and AFLP analysis. Plant Systematics
and Evolution 266: 197–232. DOI: https://doi.org/10.1007/s00606007-0532-3
Bartha, S., Campetella, G., Ruprecht, E., Kun, A., Házi, J., Horváth,
A., Virágh, K. & Molnár, Z. 2008: Will interannual variability in sand
grassland communities increase with climate change? Community Ecology 9(Suppl): 13–21. DOI: https://doi.org/10.1556/ComEc.9.2008.S.4
Stefanovits, P. 2010: A tájak talajviszonyai. In: Stefanovits P., Filep Gy.
& Füleki Gy. Talajtan, Mezőgazda Kiadó, Budapest, 470 p.
Böhm, É. I. 2015: A Szentendrei-sziget tájtörténete. (Landscape history of the Szentendre Island) [in Hungarian]. Private edition, 104 pp.
Borhidi, A., Kevey, B. & Lendvai, G. 2012: Plant communities of
Hungary. Akadémiai Kiadó, Budapest, 544 pp.
Csecserits, A. Czúcz, B., Halassy, M, Kröel-Dulay, G., Rédei, T.,
Szabó, R., Szitár, K. & Török, K. 2011: Regeneration of sandy old–
fields in the forest steppe region of Hungary. Plant Biosystems 145:
715–729. DOI: https://doi.org/10.1080/11263504.2011.601340
Deák, B., Valkó, O., Alexander, C., Mücke, W., Kania, A., Tamás,
J. & Heilmeier, H. 2014: Fine–scale vertical position as an indicator
of vegetation in alkali grasslands Case study based on remotely
sensed data. Flora 209: 693–697. DOI: https://doi.org/10.1016/j.
flora.2014.09.005
Deák, B., Valkó, O., Török, P., Kelemen, A., Miglécz, T., Szabó, S.,
Szabó, G. & Tóthmérész, B. 2015: Micro–topographic heterogeneity
increases plant diversity in old stages of restored grasslands. Basic
and Applied Ecology 16: 291–299. DOI: https://doi.org/10.1016/J.
BAAE.2015.02.008
Dövényi Z. (ed.) 2010: Magyarország kistájainak katasztere. (Inventory
of Microregions in Hungary) [in Hungarian] MTA Földrajztudományi
Kutatóintézet, Budapest, 876 pp.
Fekete, G., Précsényi, I., Molnár, E. & Melkó, E. 1976: Niche studies
on some plant species of a grassland community. I. Comparison of
various measurements. Acta Botanica Hungarica 22: 321–354.
222
Szabó, G., Zimmermann, Z., Catorci, A., Csontos, P., Wichmann,
B., Szentes, S., Barczi, A. & Penksza, K. 2017: Comparative study on
grasslands dominated by Festuca vaginata and F. pseudovaginata in the
Carpathian Basin. Tuexenia 37: 415–429.
DOI: https://doi.org/10.14471/2017.37.018
Szentes, S., Dannhauser, C., Coetzee, R. & Penksza, K. 2011:
Biomass productivity, nutrition content and botanical investigation
of Hungarian Grey cattle pasture in Tapolca basin. Animal welfare,
Etológia és Tartástechnológia 7(2): 180–198.
Szentes, S., Sutyinszki, Z., Szabó, G., Zimmermann, Z., Házi,
J., Wichmann, B., Hufnágel, L., Penksza, K. & Bartha, S. 2012:
Grazed Pannonian grassland beta–diversity changes due to C4 yellow
bluestem. Central European Journal of Biology. 7(6): 1055–1065.
DOI: https://doi.org/10.2478/s11535-012-0101-9
Török, P., Matus, G. & Papp, M. 2009: Seed bank and vegetation
development of sandy grasslands after goose breeding. Folia Geobotanica 44: 31–46. DOI: https://doi.org/10.1007/s12224-009-9027-z
Valkó, O., Deák, B., Török, P., Kelemen, A., Miglécz, T. &
Tóthmérész, B. 2017: Filling up the gaps – Passive restoration does
work on linear landscape scars. Ecological Engineering 102: 501–508.
Valkó, O., Deák, B., Török, P., Kelemen, A., Miglécz, T., Tóth, K. &
Tóthmérész, B. 2016: Abandonment of croplands: problem or chance
for grassland restoration? Case studies from Hungary. Ecosystem
Health and Sustainability 2(2): e01208.
DOI: https://doi.org/10.1002/ehs2.1208
Appendix 1: Cover of the vascular plant species recorded in sandy grasslands dominated by Festuca vaginata and F. pseudovaginata in Tece and Szigetmonostor. The abbreviation
of species names used in Figure 3 are also given here.
Dodatek 1: Pokorovnost cevnic zabeleženih na peščenih traviščih z vrstama Festuca vaginata in F. psudovaginata pri krajih Tece in Szigetmonostor. Prikazane so tudi okrajšave
vrstnih imen v sliki 3.
FpSz11
2 2 2 2 1
0 0 1 0 0
0 0 0 0 0
0 0 0 2 2
0 0 0 0 0
0 0 0 0 0
0 0 0 0 0
0 0 0 0 0
2 2 2 1 1
0 0 0 0 0
2 0 2 2 2
1 2 2 2 2
0 0 0 0 0
2 2 2 2 2
2 2 2 2 2
0 0 0 0 0
0 0 0 0 0
0 0 0 0 0
0 0 2 0 0
4 0 0 0 0
0 0 0 0 0
0 0 0 0 0
4 2 2 4 2
0 0 0 0 0
0 0 0 0 0
0 0 0 0 0
0 0 0 0 0
0 0 0 0 0
5 2 2 2 2
2 2 2 1 1
1 2 2 2 2
4 2 2 2 0
10 20 15 20 15
0 0 0 0 0
0 2 8 6 5
0 0 0 0 0
FpSz10
2
2
0
0
0
0
0
0
2
0
2
1
0
2
2
0
0
0
2
0
0
0
4
0
0
0
0
0
2
2
2
2
5
0
0
0
FpSz9
FpSz8
FpSz7
FpSz6
FpSz5
FpSz4
FpSz1
FpSz3
FpSz2
FpTe8
FpTe7
FpTe6
FpTe5
FpTe4
FpTe3
FpTe2
FpTe1
FvSz9
FvSz8
FvSz7
FvSz6
FvSz5
FvSz4
FvSz3
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2
0 0 0 0 20 10 8 10 10 4 5 8 5 10 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 2 0 2 0 2 0 2 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 2 2 2 2 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 2 0 2 2 2 0 2 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 1 0 0 0 0
2 2 2 1 1 2 2 2 4 2 2 2 5 2 2 2 2 2 2 2 2 2 1 1
0 2 2 2 2 0 0 0 0 0 5 15 15 10 5 10 10 4 5 2 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 1 1 0 0 2
0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 2 2 2 2 2 4 0 0
0 0 0 0 0 0 0 2 0 2 0 2 2 2 0 0 0 0 0 0 0 0 2 2
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 2 2 2 2 2 5 0 2
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 0 4 0 2 0 0 0
0 2 2 2 4 2 4 2 4 2 2 4 2 2 2 4 2 2 2 4 4 2 0 0
0 1 0 2 0 2 4 2 2 2 2 4 2 4 2 2 2 2 2 2 2 2 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 10 2 0 2 2 2 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 2
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 5 5 4 5 5 5 2 2
0 0 0 0 0 0 0 0 0 0 0 0 0 0 2 1 2 2 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 2 0 1 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 1 0 1 0 1 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 5 10 10 0 4 4 0 0 2 2
0 0 0 0 0 1 1 1 2 2 2 2 2 2 1 2 2 1 2 2 2 0 1 1
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 4 2
0 0 0 2 2 2 5 5 5 2 2 4 8 5 0 0 2 3 0 0 2 0 10 8
0 0 0 0 0 0 0 0 0 0 0 0 0 0 35 2 20 35 30 20 45 45 25 20
15 25 30 25 30 20 15 20 15 35 30 15 10 15 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
0 0 0 0 0 5 4 5 4 5 4 0 0 2 0 0 0 0 0 0 0 0 0 0
FvSz2
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
20
0
0
FvSz1
FvTe
FvTe5
FvTe1
FvTe4
Tece
1
0
0
2
0
0
0
0
2
0
2
2
0
2
2
0
0
0
0
0
0
0
1
0
0
0
0
0
2
1
2
2
8
0
0
0
2
0
0
4
0
0
0
0
2
0
2
1
0
2
0
0
0
0
0
0
2
0
2
0
2
0
0
0
4
1
4
10
10
0
0
0
2
0
0
0
0
0
0
0
0
0
4
1
0
2
0
0
0
0
0
0
1
0
2
0
2
0
0
0
4
0
4
10
15
0
0
0
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
Aca
Acp
Alt
Alto
Ama
Ano
Anr
Aps
Ars
Arc
Aso
Brs
Brt
Cal
Cas
Cei
Cea
Ces
Chj
Chg
Coa
Con
Cyd
Elr
Epd
Eqr
Erc
Erc
Ery
Erd
Euc
Eus
Fp
Fv
Fvv
Fup
Festuca pseudovaginata
Szigetmonostor
20/1 • 2021, 217–224
223
Achillea collina
Achillea pannonica
Alkanna tinctoria
Alyssum tortuosum
Ambrosia artemisiifolia
Anchusa officinalis
Anthemis ruthenica
Apera spica-venti
Arenaria serpyllifolia
Artemisia campestris
Astragalus onobrychis
Bromus squarrosus
Bromus tectorum
Carex liparicarpos
Carex stenophylla
Cenchrus incertus
Centaurea arenaria
Cerastium semidecandrum
Chondrilla juncea
Chrysopogon gryllus
Colchicum arenarium
Corispermum nitidum
Cynodon dactylon
Elymus repens
Ephedra distachya
Equisetum ramosissimum
Erigeron canadensis
Erodium cicutarium
Eryngium campestre
Erysimum diffusum
Euphorbia cyparissias
Euphorbia seguierana
Festuca pseudovaginata
Festuca vaginata
Festuca vaginata x valesiaca
Fumana procumbens
FvTe3
Tece
FvTe6
Festuva vaginata
Szigetmonostor
FpSz3
FpSz4
FpSz5
FpSz6
FpSz7
FpSz8
FpSz9
FpSz10
FpSz11
0 0
0 0
0 0
0 0
0 0
0 0
0 10
0 0
2 2
4 5
0 2
0 0
0 0
0 0
0 0
5 5
0 0
0 0
1 0
5 5
0 0
0 0
0 0
2 2
0 0
0 0
2 1
0 0
0 0
2 0
0 0
0 0
5 10
1 0
0 0
0 0
0 0
0 0
0 0
0 0
FpSz2
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
2 2 1 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 10 5 10 5 10
0 0 0 0 0 0
2 4 2 2 2 2
4 5 3 4 10 5
0 0 2 0 0 2
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 10 5 10 5 10
0 0 0 0 0 0
0 0 0 0 0 0
1 2 1 0 1 0
0 4 5 5 5 5
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
2 2 1 2 2 2
0 0 0 0 0 0
0 0 0 0 0 0
0 2 0 2 0 2
0 0 0 0 1 1
0 0 1 0 0 0
0 2 0 2 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 5 10 5 5 10
0 0 0 1 1 1
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
0 0 0 0 0 0
FpSz1
0 0 0
2 2 0
0 0 0
0 0 0
2 2 2
0 0 0
0 0 0
0 0 0
2 5 8
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
5 10 5
4 4 4
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
0 0 0
2 2 2
10 8 10
0 0 0
15 10 5
0 0 0
0 0 0
0 0 0
0 0 0
1 0 0
0 0 0
0 0 0
0 0 0
FpTe8
0
0
0
0
2
0
0
0
2
0
0
0
0
0
0
0
0
5
0
0
0
0
0
0
0
0
0
0
2
5
0
2
0
0
0
0
0
0
0
0
FpTe7
0
0
0
0
2
0
0
0
2
0
0
0
0
0
0
0
0
4
2
0
0
0
0
0
0
0
0
0
2
4
0
4
0
0
0
0
0
0
0
0
FpTe6
FvSz7
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
5
2
0
0
0
0
0
0
0
0
0
2
5
0
0
0
0
0
0
0
0
0
0
FpTe5
FvSz6
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
5
10
0
0
0
0
8
0
0
0
0
2
4
0
0
0
0
0
0
1
0
0
0
FpTe3
FvSz5
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
8
2
0
0
0
0
5
0
0
0
0
2
4
0
0
0
0
0
0
0
0
0
0
FpTe2
FvSz4
0
0
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
5
1
0
0
0
0
0
0
0
0
0
2
2
0
0
0
0
0
0
0
0
0
0
FpTe1
FvSz3
0
0
0
0
2
0
0
0
2
0
0
0
0
0
0
1
0
8
1
0
4
0
0
1
0
0
0
1
0
4
0
2
0
0
0
0
0
0
0
0
FvSz9
FvSz2
0
0
0
0
1
0
0
0
2
0
0
0
0
0
0
1
0
4
5
0
0
0
0
0
0
0
0
0
0
0
0
2
0
0
0
0
0
0
0
0
FvSz8
FvSz1
0 0
0 0
0 0
0 0
1 1
0 0
0 0
0 0
2 2
0 0
0 0
0 0
0 0
0 0
0 0
0 1
0 0
10 15
2 5
0 0
2 0
0 0
0 0
0 1
0 0
0 0
0 0
0 0
0 0
0 2
0 0
5 8
0 0
0 0
0 0
0 0
0 0
0 0
0 0
0 0
FvTe6
FvTe3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
8
0
0
0
0
0
0
0
0
0
0
1
4
0
4
0
0
0
0
0
0
0
0
FvTe5
FvTe
FvTe4
FvTe1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
4
5
0
0
0
0
0
0
0
0
0
0
2
0
2
0
0
0
0
0
0
0
0
2
0
2
0
0
2
0
4
2
0
0
1
0
0
0
2
2
2
0
0
2
0
2
0
0
1
0
0
1
0
2
0
0
0
2
2
0
2
0
1
2
0
5
0
0
1
2
4
2
0
0
1
0
2
2
5
2
2
0
0
2
0
2
0
0
2
0
0
1
0
2
2
0
0
2
2
0
4
1
1
2
0
4
0
0
1
5
4
2
0
0
0
2
0
2
2
2
2
0
0
2
0
2
0
1
0
0
0
2
0
1
5
0
0
2
0
0
4
1
4
2
0
2
0
0
1
4
3
2
10
0
0
0
2
2
4
2
4
1
0
4
0
4
0
2
1
0
0
2
0
0
4
0
0
2
0
0
4
0
4
2
0
2
0
0
1
0
4
2
5
0
0
2
0
2
5
2
2
0
0
2
0
2
0
2
2
0
1
2
0
1
5
0
0
0
0
0
0
1
1
2
0
2
0
0
1
4
4
2
0
0
0
2
4
2
5
2
2
0
0
2
1
2
0
0
0
0
0
2
0
0
6
0
0
2
0
0
0
1
1
2
0
1
0
0
1
0
2
2
5
0
0
0
2
2
4
4
2
0
0
2
1
1
0
2
0
0
1
1
0
0
6
0
0
2
0
0
0
0
1
2
0
4
0
0
2
4
4
4
0
0
0
0
2
4
2
2
2
0
0
4
1
1
0
0
2
0
1
2
0
1
5
0
0
2
0
0
0
0
1
2
0
2
0
0
1
0
2
4
5
0
0
0
2
4
5
2
4
1
0
4
0
2
0
0
0
0
1
2
0
0
5
0
0
2
0
0
2
0
1
2
0
2
0
0
1
0
4
4
10
0
0
0
2
4
2
2
4
1
0
4
0
2
0
0
2
0
1
2
0
1
4
0
0
2
0
0
4
1
1
2
0
2
0
0
1
5
2
2
5
0
0
0
0
2
2
2
2
1
0
4
0
0
0
0
0
0
1
2
0
0
5
0
0
1
0
0
2
1
2
Károly Penksza, Péter Csontos & Gergely Pápay
Syntaxonomical analysis of sandy grassland vegetation dominated by
Festuca vaginata and F. pseudovaginata in the Pannonian basin
Gav
Gya
Heo
Kib
Hou
Hyp
Kol
Kom
Mem
BL
Oeb
Orm
Pea
Peo
Php
Pla
Poa
Pob
Pol
Poo
Pot
Rua
San
Ses
Seh
Ses
Sev
Sic
Sio
Stb
Tha
Ths
Trr
Trt
Tra
Vec
Vep
Ver
Via
Vil
20/1 • 2021, 217–224
Galium verum
Gypsophila arenaria
Helianthemum ovatum
Hieracium bauhinii
Holosteum umbellatum
Hypericum perforatum
Kochia laniflora
Koeleria majoriflora
Medicago minima
moss-lychen synusium
Oenothera biennis
Orchis morio
Peucedanum arenarium
Peucedanum oreoselinum
Phleum phleoides
Plantago arenaria
Poa angustifolia
Poa bulbosa
Polygonum arenarium
Portulaca oleracea
Potentilla arenaria
Rumex acetosella
Salvia nemorosa
Secale sylvestre
Sedum hillebrandtii
Sedum sexangulare
Setaria viridis
Silene conica
Silene otites
Stipa borysthenica
Thesium arvense
Thymus serpyllum
Tragus racemosus
Tribulus terrestris
Trifolium arvense
Veronica chamaedrys
Veronica praecox
Veronica prostrata
Vicia angustifolia
Vicia lathyroides
Festuca pseudovaginata
Szigetmonostor
Tece
FpTe4
224
Festuva vaginata
Szigetmonostor
Tece
Károly Penksza