HACQUETIA 11/1 • 2012, 5-15
DOI: 10.2478/v10028-012-0001-6
succession model with CORmEPHORUS
CANESCENS in abandoned sandy fields
(w Slovakia)
Milan VALACHOVIC
Abstract
The succession of vegetation on acidic sandy fields after abandonment was studied and obtained results were compared with similar data from eastern part of Europe. The vegetation changes head towards oligotrophic Corynephorus-grasslands, because nutrients from the upper layer of soils are gradually washed out. Dry, occasionally blowing sand enables the growth of psammophytes very well. The paper compares the succession models on abandoned sandy fields, and semi-natural acidic sandy dunes in western part of Slovakia, southern Moravia, Hungary and Croatia. In spite of amazing similarities, some differences are displayed using spectra of life forms and presentation of weeds.
Keywords: arable land, inland sand dunes, weed, Corynephorion, Borska nižina Lowland. Izvleček:
Proučevali smo sukcesijo vegetacije na opuščenih kislih peščenih poljih in primerjali rezultate s podobnimi podatki iz vzhodne Evrope. Spremembe vegetacije so usmerjene proti oligotrofnim traviščem z vrsto Co-rynephorus, saj se hranila postopomo izperejo iz zgornjih talnih plasti. Suh pesek, ki ga občasno prinese veter omogoča uspevanje psamofitom. V članku smo primerjali sukcesijske modele na opuščenih peščenih poljih in polnaravnih kislih peščenih Dinah v zahodnem delu Slovaške, južne Moravske, Madžarske in Hrvaške. Kljub veliki podobnosti smo izpostavili določene razlike s spektrom življenskih oblik in prisotnostjo plevelnih vrst. Ključne besede: obdelana tla, celinske peščene dine, pleveli, Corynephorion, Borska nižina.
1. INTRODUCTION
The Corynephorus-grassland represents widespread pioneer vegetation on active mobile dunes along the exposed Atlantic coasts from Portugal to Denmark. It occur regularly along the southern Baltic Sea (Hulten & Fries 1986: map 342) and decline as inland sand dune grasslands in the Central and Eastern European lowlands (Rych-novska 1963, Hrsak 2004, Lajer 2004). The natural Corynephorus-grassland prefers acidic sands, but the particular species Corynephorus canescens occurs also on neutral or slightly basic dunes. For a growth of C. canescens is much more important open sandy substratum which is unstable, than accurate pH value, because this grass can auton-
omously regulate the soil pH by its root system (Rychnovska 1963).
Beside dominant Corynephorus, another psammophytes with constant frequency are present, such as Acetosella vulgaris, Agrostis tenuis, A. vinea-lis, Hypochoeris radicata, Jasione montana, Logfia minima, Pilosella officinarum, Spergula morisonii, Teesdalia nudicaulis. This species assemblage well defines the dry acidophilous and nutrient-poor plant communities of the alliance Corynephorion canescentis Klika 1931 in larger European region (cf. Korneck 1978, p. 32, Dengler 2004). Some taxa with atlantic and subatlantic distribution, e.g. Am-mophila arenaria or Carex arenaria differ west European stands from those occurred more eastern, but generally, the species composition of alliance
i Institute of Botany, Slovak Academy of Sciences, Dubravska cesta 9, SK-845 23 Bratislava, Slovak Republic, e-mail: milan.valachovic@savba.sk
is conspicuously uniform. This fact is recently well known and achieved by analyses based on large data from whole Europe (cf. Dorsing 2008).
Succession in abandoned fields and sandy areas has been investigated in numerous older studies (e.g. Prach 1985, Tilman 1987, Falinski et al. 1993) but is still very attractive for recent investigations (e.g. Putten et al. 2000, Csecserits & Rédei 2001, Deng 2001, Ejrnœs et al. 2003, 2008, Koubkovâ 2007, Torok et al. 2008). The secondary succession on sandy soils obviously heads towards pine forest communities (Prach et al. 1993), but in special situation gives a chance for possible renaturalization of abandoned fields towards sandy semi-natural grasslands. A lower attention was given to the stands, where the sandy vegetation was fully destroyed, and for longer time managed as crop fields.
The aim of the paper is (i) to describe processes of succession on sandy soils, partly on natural dunes, but predominately on secondary stands; and (ii) to compare some succession models, known from eastern part of Europe with situation in abandoned fields in Slovakia.
2. MATERIAL AND METHODS
A total of 21 phytosociological relevés were carried out in the Borskâ nizina Lowland, equally from abandoned fields (10 rels.) as well as near natural dunes (11 rels.) in accordance of the Braun-Blanquet approach. The relevés were entered to the Turboveg database (Hennekens & Schaminée 2001) and classified using modified TWINSPAN classification (Hill 1979, Rolecek et al. 2009) and carried out by using JUICE software (Tichy 2002). All imput data are stored in Central database at the Institute of Botany (http://ibot. sav.sk/cdf). The final table depicts result of comparison of 52 relevés with similar stands found in East European regions (Hrsak 2004, Lâjer 2004, Koubkovâ 2007), and with earlier stadium from the similar localities (Mâjekovâ 2005). The sorting of taxa to group of psammophytes and weeds in large extends follows index prepared by Jurko (1990). The spectra of life forms were calculated according this source as well. The abbreviations - T (therophytes = annuals and bienniales), H (hemicryptophytes), G (geophytes), C (chamae-phytes), P (phanerophytes), M (mosses), and L (lichens). The nomenclature of plant species follows Marhold & Hindâk (1998).
3. RESULTS
The TWINSPAN classification separates in the first step the relevés from abandoned crop fields and group of those stands, which were under human impact but without agronomical and agro-chemical procedures. While the first group (column A-B in Table 1) represents the development after abandonment of crop fields, the next three columns depict the succession on mechanically disturbed sandy areas. The fertilization and cultivation of fields in the past is manifested by presence of numerous weeds, e.g. Anthemis ruthenica, Conyza canadensis, Digitaria sanguinalis and many others. Especially in the earlier phases after abandonment, the fields are covered by nitrogen demanding plants, such as Agrostis gigantea, Chenopo-dium album, Crepis tectorum, Elytrigia repens or Echi-um vulgare, later the nutrients are rather quickly and easily wash out and the abundance of weeds changed in favour of more oligotrophic weeds, such as Agrostis capillaris, Digitaria sanguinalis, Trifo-lium arvense etc. (see column B in Table 1). The dry sand on the upper layer of soil prefers psammo-phytes. The Corynephorus canescens, together with shallow rooted therophytes, such as Logfia minima, Spergula morisonii, Teesdalia nudicaulis, Veronica dile-nii, occupied the washed sand, but the nutrients in the deeper layer of soils give a sufficient conditions for growth of some weeds, like Hypochoeris radicata. The percentage of therophytes in column A is conspicuously high (59%, see Figure 1, 2).
70
60 50 40 30 20 10 0
									
E									
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A	B
Therophytes Hemicryptophytes
Geophytes 0 Chamaephytes D Lichens
Figure 1: Percentage spectra of selected life forms in columns A-E.
Slika 1: Spekter izbranih življenskih oblik v odstotkih v stolpcih A-E.
C
D
E
Figure 2: Weed Echium vulgare is replaced by oligotrophic Jasione montana and Logfia minima.
Slika 2: Plevel Echium vulgare nadomestita oligotrofni vrsti Jasione montana in Logfia minima.
The succession on open sandy dunes started obviously with acidophilous mosses - Polytri-chum piliferum and lichens of the genus Cladonia. This model on the abandoned fields was not observed. Only sporadic were found lichens, often the Cladonia fimbriata. The result is partially misrepresented, because in some published data the cryptogamic flora has been overlooked (cf. Layer 2004, column C, in Table 1).
In the column D are associated data from military grounds and ranges in Moravia and Slovakia, where traditionally absent agronomical interventions, but large areas (several hectares) are under extensive disturbance due military activities. The number of psammophytes here is much
Figure 3: Corynephorus-grasslands in military range near Malacky, Križnica.
Slika 3: Travišča z vrsto Corynephorus na vojaškem vadbišču pri naselju Malacky, Križnica.
higher and positive correlate with size of area and availability of propagules from neighbouring habitats (sandy grasslands and pine forests of Di-crano-Pinion alliance). Here is also high percentage of lichens in the relevés, 17% (Figure 1), while the role of therophytes is suppressed (Figure 3).
A last column (E) differentiates species like Artemisia campestris, Carex praecox, Tithymalus cy-parissias, and Festuca vaginata, all plants typical for sandy soils with neutral or slightly basiphil-ous reaction.
4. DISCUSSION
Natural (near-natural) stands on sandy dunes
The natural pioneer communities from the alliance Corynephorion canescentis are generally poor in vascular plants but rich in lichens, primarily of the genus Cladonia (Table 1, Figure 4). The floristic composition of herb layer is conspicuously uniform in whole area of its distribution. Among initial components of natural succession belongs mostly Spergula morisonii, Teesdalia nudi-caulis, Veronica dilenii (cf. Juskiewicz-Swaczyna 2009); in Eastern Europe also Thymus serpyllum (Chytry 2007). Stands of Corynephorus-grasslands are obviously in this region named like separate association Thymo angustifolii-Corynephoretum canescentis Krippel 1954 and more dense grasslands than as the Festuco dominii-Corynephoretum Borhidi (1958) 1996. These are differentiated by Pannonian or continental species such as Carex ericetorum, Dianthus armeriastrum, D. serotinus,
Figure 4: Corynephorus canescens growing in lichen carpet. Figure 4: Vrsta Corynephorus canescens uspeva na preprogi lišajev.
Festuca vaginata subsp. dominii, Thymus serpyllum subsp. angustifolius, Veronica dillenii, Viola kitaibe-liana and some others taxa (cf. Hrsak 2004, Lajer 2004).
Vicherek (1972) documented the easternmost occurrence of such sand dunes steppes near Kiev (Ukraine), where the taxon Spergula morisonii is very rare; therefore these stands were associated to the Veronico dillenii-Corynephoretum Passarge 1960, which Vicherek considered as subcontinental vicariate association.
In recent studies (Chytry 2007, Dorsing 2008, Juskiewicz-Swaczyna 2009) the authors unified all stands, formerly described under the various names, into one association, the Cornicular-io aculeatae-Corynephoretum canescentis Steffen 1931 (Syn.: Spergulo morisonii-Corynephoretum canescentis R. Tx. (1928) 1955; Spergulo vernalis-Corynephoretum canescentis R. Tx. (1928) Libbert 1933) and certain differentiation lies only on level of subassociations (Dorsing l.c.), variants or geographical races (Juskiewicz-Swaczyna l.c.).
Secondary stands on sandy fields
The secondary grasslands with dominance of Corynephorus, sporadically developed on abandoned crop-fields after several years, showed differences against the above discussed stands of the Corniculario aculeatae-Corynephoretum canescentis association.
There are noticeable differences between habitats, which were ploughed, and then after 2 or 3 years afforested again by pine trees, or which were deforested, and for longer time exploited as crop-fields. As an example of succession on young pine plantings can be emphasized study by Somsak (1976). He observed fallows on sandy soils in Borska nizina lowland and described than typical plant community Setario viridis-Eri-geronetum Somsak 1976. These stands are characterized by dominance of many weeds, like Conyza canadensis, Agrostis tenuis, Setaria viridis, Carex hirta, Apera spica-venti combined with lower presence by psammophytes such as Corynephorus canescens, Festuca vaginata subsp. dominii, and Logfia arvensis.
22 years long succession on abandoned sandy fields in Denmark studied Deng (2001). According his observation the annual weeds fade out as first (during 3-5 years) and Jasione montana and Acetosella vulgaris became dominant, slowly supplement with persistent perennials such as Pilosel-la officinarum. The Corynephorus canescens grows
Figure 5: Abandoned field near Studienka with psammo-phyte vegetation.
Slika 5: Opuščena polja pri naselju Studienka s psamofitsko vegetacijo.
generally in this field very sparsely and after 12 years completely disappeared and whole field returned to Calluna vulgaris heathland. Falinski et al. (1993) studied vegetation dynamic on the abandoned sandy farmlands during secondary succession in Poland. The pioneer phase of vegetation after 2-3 years of the abandonment represent weed plants such as Digitaria ischaemum, Spergula arvensis accompanied by herbaceous perennials, among which psammophytes prevail - Jasione montana, Helichrysum arenarium, Hiera-cium pilosella, Hypericum perforatum, Hypochoeris radicata, Acetosella vulgaris, Solidago virgaurea, etc. The occurrence of psammophilous grass Corynephorus canescens as well as therophytes like Logfia minima is also characteristic (Figure 5).
The floristic correspondence with abandoned fields in Borska nižina lowland here is remarkable. The phase with Juniperus communis (cf. Falinski et al. 1993) is not common in our study site, but final phase to stable pine forests after more than 100 years after the field abandonment should by one alternative also in Slovakia, but required to stop human impact to the fields (artificial forestation, repeated transformation into crop-fields, urbanization, etc.) for so long time.
Succession models
Typical succession model on natural acidic sandy dunes is generally described as (a) phase with open, shifting sand (dominate mosses and lichens), practically without vascular plants except the first tuft of grass Corynephorus canescens, (b) phase of dry grasslands with psammophytes
such as Pilosella officinarum Jasione montana, etc., (c) phase of closed grasslands with dominate Festuca sp. div., and (d) development towards Callu-na vulgaris-heathland or direct to various oak pine forests. The primary succession is relatively slow. Small-scale disturbances (anthills, molehills, and holes) keep the process of succession towards sand grasslands in equilibrium for longer time (Jentsch et al. 2002, Tschope & Tielborger 2010).
Different situation is on sandy fields. Accumulation of organic matter gives gradually the opportunity for more demanding plants and onset of next phases with various nitrophilous and mesophilous plants (Ejrn^s et al. 2003). Very frequent situation is that fields are attack by rather specialized invasive aliens, in first stage annuals, herbaceous weeds and neophytes already present in the seed bank, latter also by persistent perennials (Van der Putten et al. 2000) and finally by alien woody species, such as Acer negundo and Robinia pseudoacacia (Pysek et al. 1998). Hrsak (2004) found direct relation between accumulation of organic matter and nutrients and changes in floristic pattern on sandy dunes in Croatia. Starting plants, like Corynephorus canescens and Festuca vaginata are replaced by Calamagrostis epigeios, Conyza canadensis, Oenothera biennis, Soli-dago gigantea, and shrubby Sambucus nigra. Very similar succession model is common also in our study area.
In southern Moravia Koubkova (2007) investigated floristic variability and secondary succession in sandy vegetation on disturbed open sands after three decades, when J. Vicherek started study this vegetation. The results showed that soil pH did not change markedly, but the number of species with affinity to nutrients, soil moisture and neutral reaction (in sense to Ellenberg indication values) increased.
Secondary succession on abandoned fields with tendency to return toward more or less natural or near-natural vegetation is not typical phenomenon. Nevertheless, on sandy fields, where the low amount of nutrients eliminate demanding ruderal species during 10 years after abandonment (Csecserits & Redei 2001) the succession model head towards higher abundance of psammophytes. The nutrient poor soils support sand specialists and pioneer species of the open dunes, such as Spergula morissonii and others. The abandonment of fields with nutrient poor soils may lead to the formation of semi-natural grasslands, especially in situation, when prop-
agules in seed bank or from neighbouring habitats are present (cf. Ejrn^s et al. 2008, Sefferova Stanova et al. 2011). Appropriate management on similar stands give a chance to create conditions for recover of rare plant populations and Natura 2000 habitats typical for acidophilous open sandy dunes, such as 2340 - Pannonic inland dunes, and 6260 - Pannonic sand steppes (Stanova & Valachovic 2002).
5. ACKNOWLEDGEMENT
The author acknowledges to specialist for kindly determination of some hardly distinguishable taxa, P. Smarda (Festuca), M. Peniastekova (Scle-ranthus), I. Pisut and A. Dingova (Cladonia). The research was supported from the Grant Agency of Slovak Academy of Sciences VEGA No. 0181 (Dynamics and succesion of grassland vegetation) and VEGA No. 0672 (Diversity changes in neighbouring zones with agrocoenoses).
6. REFERENCES
Csecserits, A. & Redei, T. 2001: Secondary succession on sandy old-fields in Hungary. Applied Vegetation Science 4: 63-74. Deng, H.J. 2001: Succession from farmland to heathland: a case for conservation of nature and historic farming methods. Biological Conservation 97: 319-330. Dengler, J. 2004: 21. Klasse: Koelerio-Corynepho-retea Klika in Klika et V. Novak 1941 -Sandtrockenrasen und Felsgrusfluren von der submeridionalen bis zur borealen Zone. In: Berg, C., Dengler, J., Abdank, A. & Isermann, M. (eds), Die Pflanzengesellschaften Mecklenburg-Vorpommerns und ihre Gefährdung - Textband. Landesamt f. Umwelt, Naturschutz und Geologie Mecklenburg-Vorpommern. Weissdorn-Verlag Jena, pp. 301-326 Dörsing, M. 2008: Classification of Corynephorion vegetation. Abstracts and excursion guides. 17th International Workshop European Vegetation Survey, Brno, 1-5 May 2008. Ejrn^s, R., Hansen, D.N. & Aude, E. 2003: Changing course of secondary succession in abandoned sandy fields. Biological Conservation 109: 343-350. Ejrn^s, R., Liira, J., Poulsen, R.S. & Nygaard B. 2008: When has an abandoned field become a
semi-natural grassland or heathland? Environmental Management 42: 707-716.
Falinski, J.B., Cieslinski, S. & Czyzewska, K. 1993. Dynamic-floristic atlas of Jelonka Reserve and adjacent areas. Distribution of vascular plant species, bryophytes and lichens on the abandoned farmlands during secondary succession [Atlas dynamiczno-florystyczny Rezerwatu Jelonka i przyleglych obszarów. Rozmieszczenie gatunkow roslin naczynio-wych, mszakow i porostów na nieuzytkach po-rolnych podlegajacych sukcesji wtórnej]. Phy-tocoenosis, Vol. 5 (N.S.), Suppl. Cartogr. Ge-obot. 3, Bialowieza Geobotanical Station, Warszawa-Bialowieza. 139 pp.
Hill, M.O., 1979: TWINSPAN - A Fortran program for arranging multivariate data in an ordered two-way table by classification of the individuals and attributes. Cornell Univ. Ithaca. NY.
Hrsak, V., 2004: Vegetation succession and soil gradients on inland sand dunes. Ekológia, Bratislava 23: 24-39.
Hultén, E. & Fries, M., 1986: Atlas of North European vascular plants: north of the Tropic of Cancer, Vol. 1., map. 342. Koeltz Scientific Books, Königstein.
Jentsch, A., Friedrich, S., Beyschlag, W. & Nezad-al, W., 2002: Significance of ant and rabbit disturbance for seedling establishment in dry acidic grasslands dominated by Corynephorus canescens. Phytocoenologia 32: 553-580.
Jurko, A., 1990: Ekologické a socioekonomické hodnotenie vegetácie. Príroda. 195 pp.
Juskiewicz-Swaczyna, B. 2009: The psammo-philous grassland community Corniculario ac-uleatae-Corynephoretum canescentis in the Masurian Lake District (NE Poland). Tuexenia 29: 391-408.
Korneck, D. 1978: Corynephoretalia. In: Oberdorfer, E. (ed.), Süddeutsche Pflanzengesellschaften. Teil II. 2. Aufl. Gustav Fischer Verlag. Stuttgart - New York, pp. 31-45.
Koubková T., 2007: Nelesní vegetace písku na Hodonínsku a její zmeny po tíech desetiletích. [Non-forest vegetation of the sand area near Hodonín, Czech Republic and its changes after three decades]. Diplom. Theses, Masaryk univ. Brno. 72 pp.
Lájer, K. 2004: A rejtöke (Teesdalia nudicaulis) elöfordulasa és a Bolsö-somogyi ezüstperjés gyepek mai állapota. [The occurrence of Shepherd's Cress (Teesdalia nudicaulis) and the
recent status of Grey Hair-grasslands in Bolso-Somogy]. Somogyi Muzeumok Kozlemenyei 16: 257-264.
Majekova, J. 2005: Flora a vegetacia na poliach a uhoroch Borskej niziny. Diplom. Theses, Pri-FUK Bratislava.
Marhold, K. & Hindak, F. 1998: Checklist of nonvascular and vascular plants of Slovakia. Veda, Bratislava, 687 pp.
Prach, K. 1985: Succession of vegetation in abandoned fields in Finland. Annales Botanici Fennnici 22: 307-314.
Prach, K., Fanta, J., Lukesova, A. & Liska, J. 1993: De ontwikkeling van de vegetatie op stuifzand van de Veluwe. Gorteria 19: 73-79.
Pysek, P., Prach, K. & Mandak, B. 1998: Invasions of alien plants into habitats of Central European landscape: An historical pattern. In: Starfinger, U., Edwards, K., Kowarik I., Williarn-sorz, M. eds, Plant Invasions: Ecological Mechanisms and Human Responses, Backhuys Publ., Leiden, The Netherlands, pp. 23-32.
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Somsak, L. 1976: Fytocenozy borovicovych kultur a rubanisk viatych pieskov na Zahorskej nizine [Phytocoenoses of pine cultures and felled areas on drifting sands of the plain Zahorska nizina]. Biologia, Bratislava 31 (4): 241-251.
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Torok, P., Matus, G., Papp, M. & Tothmeresz, B. 2008: Secondary succession in overgrazed Pan-nonian sandy grasslands. Preslia 80: 73-85.
Tschöpe, O. & Tielbörger, K. 2010: The role of successional stage and small-scale disturbance for establishment of pioneer grass Corynephorus canescens. Applied Vegetation Science 13: 326-335.
Van der Putten, W.H., Mortimer, S.R., Hedlund, K., Van Dijk, C., Brown, V.K., Leps, J., Rod-ruguez-Barrueco, C., Roy, J., Diaz Len, T.A., Gormsen, D., Korthals, G.W., Lavorel, S., Santa Regina, I. & Smilauer, P. 2000: Plant species diversity as a driver of early succession in abandoned fields: a multi-site approach. Oecologia 124: 91-99.
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32	960811	10	190	75	50	50	Malacky, border of military range
33	010630	25	192	70	60	50	Malacky, border of military range
35	930731	12	215	60	40	60	Malacky, Orlovské vršky, open dunes
39	090526	16	206	80	30	75	Mikulašov, Bežnisko, milit. range
37	080516	7	196	80	75	10	Malacky, Križnica, air range
40	090607	16	194	65	17	55	Malacky, Križnica, air range
34	020630	25	190	75	60	40	Malacky, border of military range
47	100722	20	199	85	65	25	Malacky, Križnica, air range
46	100722	20	200	80	45	70	Malacky, Križnica, air range
38	090601	16	195	90	40	90	Malacky, border of military range
36	090526	16	169	90	20	80	Mikulašov, Bežnisko, milit. range
Column E, 10 rels., 5 rels. (1-10), made by Koubkova 2007 (orig. tab./rel.: 1/1, 2, 27, 62, 78), surroundings Bzenec and Hodonin in Moravia); 5 rels. (18-22) made by Hrsak 2004, orig. tab./ rel.: 1/6, 4, 9, 13, 17), all from Durdevac Sand Botanical Reservation in Croatia.
7. APPENDIX
Table heads:
Column A, 6 rels., all by Majekova 2005 (orig. tab./rel.: 12/23, 12/24, 13/3, 12/25, 12/28, 12/27, made during years 2003-2004, on abandoned fields N from Studienka village, Borska nizina Lowland. Column B, 13 rels., 3 rels. (27-29) Majekova 2005 (orig. tab./rel.:7/1, 8/1, 13/6, all on abandoned fields N from Studienka village + 10 rels. made by author during year 2010 - number in table, date (yymmdd), plot (m2), altitude (m a.s.l.), cover total (%), cover Ej and EO (%), specific locality:
52 101001 20 184 50 30 20 Sajdikove Humence, field near railway
49	100729 20 219 70 60 10 Laksarska Nova Ves, field W from village 51 100101 20 180 70 20 70 Sajdikove Humence, field near railway
44	100713 20 200 50 50 0 Laksarska Nova Ves, field towards Sisolaky
45	100713 20 199 50 50 0 Laksarska Nova Ves, field towards Sisolaky
41	100713 20 226 60 55 3 Studienka, field N from village
43 100713 20 203 60 60 3 Studienka, (Cechovec 198 m), aband. field
42	100713 20 199 50 50 1 Studienka, (Cechovec 198 m), aband. field 48 100729 20 165 50 50 2 Borsky Sv. Jur, towards Tomky settlement
50	100923 20 223 80 60 70 Studienka, field N from village
column c, 7 rels. (11-17) made by Lajer 2004 (orig. tab./rel.: 1/1-7, during 2000-2001 in Bolso-Somogy in Hungary). column D, 16 rels., 5 rels, Koubkova 2007 (orig. tab./rel.: 1/4, 8, 29, 58, 59) surroundings Bzenec in Moravia) + 11 rels. made by author - number in table, date (yymmdd), plot (m2), altitude (m a.s.l.), cover total (%), cover Ej and EO (%), specific locality:
Received 16. 5. 2011 Revision received 14. 10. 2011 Accepted 31. 10. 2011
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