Succession Model with Corynephorus Canescens in Abandoned Sandy Fields (W Slovakia)

HACQUETIA 11/1 • 2012, 5–15 DOI: 10.2478/v10028-012-0001-6

Succession model with Corynephorus canescens in abandoned sandy fields (W Slovakia)

Milan VALACHOVIČ1

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, oc- casionally 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, Borská nížina Lowland.

Izvleček: Proučevali smo sukcesijo vegetacije na opuščenih kislih peščenih poljih in primerjali rezultate s podobni- mi 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, Borská nížina.

1. Introduction omously regulate the soil pH by its root system (Rychnovská 1963). The Corynephorus-grassland represents widespread Beside dominant Corynephorus, another psam- pioneer vegetation on active mobile dunes along mophytes with constant frequency are present, the exposed Atlantic coasts from Portugal to such as Acetosella vulgaris, Agrostis tenuis, A. vinea- Denmark. It occur regularly along the southern lis, Hypochoeris radicata, Jasione montana, Logfia Baltic Sea (Hultén & Fries 1986: map 342) and minima, Pilosella officinarum, Spergula morisonii, decline as inland sand dune grasslands in the Teesdalia nudicaulis. This species assemblage well Central and Eastern European lowlands (Rych- defines the dry acidophilous and nutrient-poor novská 1963, Hršak 2004, Lájer 2004). The natu- plant communities of the alliance Corynephorion ral Corynephorus-grassland prefers acidic sands, canescentis Klika 1931 in larger European region but the particular species Corynephorus canescens (cf. Korneck 1978, p. 32, Dengler 2004). Some taxa occurs also on neutral or slightly basic dunes. For with atlantic and subatlantic distribution, e.g. Am- a growth of C. canescens is much more important mophila arenaria or Carex arenaria differ west Eu- open sandy substratum which is unstable, than ropean stands from those occurred more eastern, accurate pH value, because this grass can auton- but generally, the species composition of alliance

1 Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 9, SK-845 23 Bratislava, Slovak Republic, e-mail: [email protected]

5 Hacquetia 11/1 • 2012, 5–15 is conspicuously uniform. This fact is recently well 3. RESULTS known and achieved by analyses based on large data from whole Europe (cf. Dörsing 2008). The TWINSPAN classification separates in the Succession in abandoned fields and sandy first step the relevés from abandoned crop fields areas has been investigated in numerous older and group of those stands, which were under hu- studies (e.g. Prach 1985, Tilman 1987, Faliński et man impact but without agronomical and agro- al. 1993) but is still very attractive for recent in- chemical procedures. While the first group (col- vestigations (e.g. Putten et al. 2000, Csecserits & umn A-B in Table 1) represents the development Rédei 2001, Deng 2001, Ejrnæs et al. 2003, 2008, after abandonment of crop fields, the next three Koubková 2007, Török et al. 2008). The second- columns depict the succession on mechanically ary succession on sandy soils obviously heads disturbed sandy areas. The fertilization and cul- towards pine forest communities (Prach et al. tivation of fields in the past is manifested by pres- 1993), but in special situation gives a chance for ence of numerous weeds, e.g. Anthemis ruthenica, possible renaturalization of abandoned fields to- Conyza canadensis, Digitaria sanguinalis and many wards sandy semi-natural grasslands. A lower at- others. Especially in the earlier phases after aban- tention was given to the stands, where the sandy donment, the fields are covered by nitrogen de- vegetation was fully destroyed, and for longer manding plants, such as Agrostis gigantea, Chenopo- time managed as crop fields. dium album, Crepis tectorum, Elytrigia repens or Echi- The aim of the paper is (i) to describe process- um vulgare, later the nutrients are rather quickly es of succession on sandy soils, partly on natural and easily wash out and the abundance of weeds dunes, but predominately on secondary stands; changed in favour of more oligotrophic weeds, and (ii) to compare some succession models, such as Agrostis capillaris, Digitaria sanguinalis, Trifo- known from eastern part of Europe with situa- lium arvense etc. (see column B in Table 1). The dry tion in abandoned fields in Slovakia. sand on the upper layer of soil prefers psammophytes. The Corynephorus canescens, together with shallow rooted therophytes, such as Logfia minima, 2. Material and Methods Spergula morisonii, Teesdalia nudicaulis, Veronica dilenii, occupied the washed sand, but the nutrients A total of 21 phytosociological relevés were car- in the deeper layer of soils give a sufficient condi- ried out in the Borská nížina Lowland, equally tions for growth of some weeds, like Hypochoeris from abandoned fields (10 rels.) as well as near radicata. The percentage of therophytes in column natural dunes (11 rels.) in accordance of the A is conspicuously high (59%, see Figure 1, 2). Braun-Blanquet approach. The relevés were en- 70 tered to the Turboveg database (Hennekens & Schaminée 2001) and classified using modified 60 TWINSPAN classification (Hill 1979, Roleček et al. 2009) and carried out by using JUICE software 50 (Tichý 2002). All imput data are stored in Central 40 database at the Institute of Botany (http://ibot. sav.sk/cdf ). The final table depicts result of com- 30 parison of 52 relevés with similar stands found in East European regions (Hršak 2004, Lájer 2004, 20 Koubková 2007), and with earlier stadium from 10 the similar localities (Májeková 2005). The sort- ing of taxa to group of psammophytes and weeds 0 in large extends follows index prepared by Jurko A B C D E (1990). The spectra of life forms were calculated Therophytes Geophytes according this source as well. The abbreviations – T (therophytes = annuals and bienniales), H Hemicryptophytes Chamaephytes Lichens (hemicryptophytes), G (geophytes), C (chamae- Figure 1: Percentage spectra of selected life forms in columns phytes), P (phanerophytes), M (mosses), and L A–E. (lichens). The nomenclature of plant species fol- Slika 1: Spekter izbranih življenskih oblik v odstotkih v stolp- lows Marhold & Hindák (1998). cih A–E.

6 M. Valachovič: Succession model with Corynephorus canescens in abandoned sandy fields (W Slovakia)

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 percent - age 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 cyparissias, and Festuca vaginata, all plants typical for sandy soils with neutral or slightly basiphil- ous reaction.

4. DISCUSSION Figure 2: Weed Echium vulgare is replaced by oligotrophic Jasione montana and Logfia minima. Natural (near-natural) stands on sandy dunes Slika 2: Plevel Echium vulgare nadomestita oligotrofni vrsti The natural pioneer communities from the Jasione montana in Logfia minima. alliance Corynephorion canescentis are generally poor in vascular plants but rich in lichens, pri- The succession on open sandy dunes started marily of the genus Cladonia (Table 1, Figure 4). obviously with acidophilous mosses – Polytri- The floristic composition of herb layer is conspic- chum piliferum and lichens of the genus Cladonia. uously uniform in whole area of its distribution. This model on the abandoned fields was not ob- Among initial components of natural succession served. Only sporadic were found lichens, often belongs mostly Spergula morisonii, Teesdalia nudi- the Cladonia fimbriata. The result is partially mis- caulis, Veronica dilenii (cf. Juśkiewicz-Swaczyna represented, because in some published data the 2009); in Eastern Europe also Thymus serpyllum cryptogamic flora has been overlooked (cf. Láyer (Chytrý 2007). Stands of Corynephorus-grasslands 2004, column C, in Table 1). are obviously in this region named like separate In the column D are associated data from mili- association Thymo angustifolii-Corynephoretum tary grounds and ranges in Moravia and Slova- canescentis Krippel 1954 and more dense grass- kia, where traditionally absent agronomical in- lands than as the Festuco dominii-Corynephoretum terventions, but large areas (several hectares) are Borhidi (1958) 1996. These are differentiated by under extensive disturbance due military activi- Pannonian or continental species such as Carex ties. The number of psammophytes here is much ericetorum, Dianthus armeriastrum, D. serotinus,

Figure 3: Corynephorus-grasslands in military range near Figure 4: Corynephorus canescens growing in lichen carpet. Malacky, Krížnica. Figure 4: Vrsta Corynephorus canescens uspeva na preprogi Slika 3: Travišča z vrsto Corynephorus na vojaškem vadbišču lišajev. pri naselju Malacky, Krížnica.

7 Hacquetia 11/1 • 2012, 5–15

Festuca vaginata subsp. dominii, Thymus serpyllum subsp. angustifolius, Veronica dillenii, Viola kitaibe- liana and some others taxa (cf. Hršak 2004, Lájer 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 subcontinen- tal vicariate association. In recent studies (Chytrý 2007, Dörsing 2008, Juśkiewicz-Swaczyna 2009) the authors unified all stands, formerly described under the various names, into one association, the Cornicular- Figure 5: Abandoned field near Studienka with psammo- io aculeatae-Corynephoretum canescentis Steffen phyte vegetation. 1931 (Syn.: Spergulo morisonii-Corynephoretum Slika 5: Opuščena polja pri naselju Studienka s psamofitsko canescentis R. Tx. (1928) 1955; Spergulo vernalis- vegetacijo. Corynephoretum canescentis R. Tx. (1928) Libbert 1933) and certain differentiation lies only on level generally in this field very sparsely and after 12 of subassociations (Dörsing l.c.), variants or geo- years completely disappeared and whole field graphical races (Juśkiewicz-Swaczyna l.c.). returned to Calluna vulgaris heathland. Faliński et al. (1993) studied vegetation dynamic on the Secondary stands on sandy fields abandoned sandy farmlands during secondary The secondary grasslands with dominance of succession in Poland. The pioneer phase of veg- Corynephorus, sporadically developed on aban- etation after 2–3 years of the abandonment rep- doned crop-fields after several years, showed dif- resent weed plants such as Digitaria ischaemum, ferences against the above discussed stands of the Spergula arvensis accompanied by herbaceous Corniculario aculeatae-Corynephoretum canescentis perennials, among which psammophytes prevail association. – Jasione montana, Helichrysum arenarium, Hiera- There are noticeable differences between hab- cium pilosella, Hypericum perforatum, Hypochoeris itats, which were ploughed, and then after 2 or radicata, Acetosella vulgaris, Solidago virgaurea, 3 years afforested again by pine trees, or which etc. The occurrence of psammophilous grass were deforested, and for longer time exploited Corynephorus canescens as well as therophytes like as crop-fields. As an example of succession on Log fia minima is also characteristic (Figure 5). young pine plantings can be emphasized study The floristic correspondence with abandoned by Šomšák (1976). He observed fallows on sandy fields in Borská nížina lowland here is remark- soils in Borská nížina lowland and described able. The phase with Juniperus communis (cf. than typical plant community Setario viridis-Eri Faliński et al. 1993) is not common in our study geronetum Šomšák 1976. These stands are char - site, but final phase to stable pine forests after acterized by dominance of many weeds, like more than 100 years after the field abandonment Conyza canadensis, Agrostis tenuis, Setaria viridis, should by one alternative also in Slovakia, but Carex hirta, Apera spica-venti combined with lower required to stop human impact to the fields (ar- presence by psammophytes such as Corynephorus tificial forestation, repeated transformation into canescens, Festuca vaginata subsp. dominii, and crop-fields, urbanization, etc.) for so long time. Log fia arvensis. 22 years long succession on abandoned sandy Succession models fields in Denmark studied Deng (2001). Accord- Typical succession model on natural acidic ing his observation the annual weeds fade out as sandy dunes is generally described as (a) phase first (during 3–5 years) and Jasione montana and with open, shifting sand (dominate mosses and Acetosella vulgaris became dominant, slowly sup - lichens), practically without vascular plants ex- plement with persistent perennials such as Pilosel- cept the first tuft of grass Corynephorus canescens, la officinarum. The Corynephorus canescens grows (b) phase of dry grasslands with psammophytes

8 M. Valachovič: Succession model with Corynephorus canescens in abandoned sandy fields (W Slovakia) such as Pilosella officinarum Jasione montana, etc., agules in seed bank or from neighbouring habi- (c) phase of closed grasslands with dominate Fes- tats are present (cf. Ejrnæs et al. 2008, Šefferová tuca sp. div., and (d) development towards Callu- Stanová et al. 2011). Appropriate management on na vulgaris-heathland or direct to various oak pine similar stands give a chance to create conditions forests. The primary succession is relatively slow. for recover of rare plant populations and Natura Small-scale disturbances (anthills, molehills, and 2000 habitats typical for acidophilous open sandy holes) keep the process of succession towards dunes, such as 2340 – Pannonic inland dunes, sand grasslands in equilibrium for longer time and 6260 – Pannonic sand steppes (Stanová & (Jentsch et al. 2002, Tschöpe & Tielbörger 2010). Valachovič 2002). Different situation is on sandy fields. Accu- mulation of organic matter gives gradually the opportunity for more demanding plants and on- 5. ACKNOWLEDGEMENT set of next phases with various nitrophilous and mesophilous plants (Ejrnæs et al. 2003). Very fre- The author acknowledges to specialist for kindly quent situation is that fields are attack by rather determination of some hardly distinguishable specialized invasive aliens, in first stage annuals, taxa, P. Šmarda (Festuca), M. Peniašteková (Scle- herbaceous weeds and neophytes already present ranthus), I. Pišút and A. Dingová (Cladonia). The in the seed bank, latter also by persistent peren- research was supported from the Grant Agency nials (Van der Putten et al. 2000) and finally by of Slovak Academy of Sciences VEGA No. 0181 alien woody species, such as Acer negundo and (Dynamics and succesion of grassland vegeta- Robinia pseudoacacia (Pyšek et al. 1998). Hršak tion) and VEGA No. 0672 (Diversity changes in (2004) found direct relation between accumula- neighbouring zones with agrocoenoses). tion of organic matter and nutrients and changes in floristic pattern on sandy dunes in Croatia. Starting plants, like Corynephorus canescens and 6. REFERENCES Festuca vaginata are replaced by Calamagrostis epigeios, Conyza canadensis, Oenothera biennis, Soli- Csecserits, A. & Rédei, T. 2001: Secondary succes- dago gigantea, and shrubby Sambucus nigra. Very sion on sandy old-fields in Hungary. Applied similar succession model is common also in our Vegetation Science 4: 63–74. study area. Deng, H.J. 2001: Succession from farmland to In southern Moravia Koubková (2007) inves- heathland: a case for conservation of nature tigated floristic variability and secondary succes- and historic farming methods. Biological Con sion in sandy vegetation on disturbed open sands servation 97: 319–330. after three decades, when J. Vicherek started Dengler, J. 2004: 21. Klasse: Koelerio-Corynepho- study this vegetation. The results showed that retea Klika in Klika et V. Novák 1941 – soil pH did not change markedly, but the number Sandtrockenrasen und Felsgrusfluren von der of species with affinity to nutrients, soil moisture submeridionalen bis zur borealen Zone. In: and neutral reaction (in sense to Ellenberg indi- Berg, C., Dengler, J., Abdank, A. & Isermann, cation values) increased. M. (eds), Die Pflanzengesellschaften Meck- Secondary succession on abandoned fields lenburg-Vorpommerns und ihre Gefährdung with tendency to return toward more or less – Textband. Landesamt f. Umwelt, Natur- natural or near-natural vegetation is not typi- schutz und Geologie Mecklenburg-Vorpom- cal phenomenon. Nevertheless, on sandy fields, mern. Weissdorn-Verlag Jena, pp. 301–326 where the low amount of nutrients eliminate de- Dörsing, M. 2008: Classification of Corynephorion manding ruderal species during 10 years after vegetation. Abstracts and excursion guides. abandonment (Csecserits & Rédei 2001) the suc- 17th International Workshop European Vege- cession model head towards higher abundance of tation Survey, Brno, 1–5 May 2008. psammophytes. The nutrient poor soils support Ejrnæs, R., Hansen, D.N. & Aude, E. 2003: sand specialists and pioneer species of the open Changing course of secondary succession in dunes, such as Spergula morissonii and others. abandoned sandy fields. Biological Conserva- The abandonment of fields with nutrient poor tion 109: 343–350. soils may lead to the formation of semi-natural Ejrnæs, R., Liira, J., Poulsen, R.S. & Nygaard B. grasslands, especially in situation, when prop- 2008: When has an abandoned field become a

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Tschöpe, O. & Tielbörger, K. 2010: The role of 32 960811 10 190 75 50 50 Malacky, border of military range successional stage and small-scale disturbance 33 010630 25 192 70 60 50 Malacky, border of military range for establishment of pioneer grass Corynepho- 35 930731 12 215 60 40 60 Malacky, Orlovské vršky, open dunes rus canescens. Applied Vegetation Science 13: 39 090526 16 206 80 30 75 Mikulášov, Bežnisko, milit. range 326–335. 37 080516 7 196 80 75 10 Malacky, Krížnica, air range Van der Putten, W.H., Mortimer, S.R., Hedlund, 40 090607 16 194 65 17 55 Malacky, Krížnica, air range K., Van Dijk, C., Brown, V.K., Lepš, J., Rod- 34 020630 25 190 75 60 40 Malacky, border of military range ruguez-Barrueco, C., Roy, J., Diaz Len, T.A., 47 100722 20 199 85 65 25 Malacky, Krížnica, air range Gormsen, D., Korthals, G.W., Lavorel, S., 46 100722 20 200 80 45 70 Malacky, Krížnica, air range Santa Regina, I. & Šmilauer, P. 2000: Plant 38 090601 16 195 90 40 90 Malacky, border of military range species diversity as a driver of early succession 36 090526 16 169 90 20 80 Mikulášov, Bežnisko, milit. range in abandoned fields: a multi-site approach. Oecologia 124: 91–99. Column E, 10 rels., 5 rels. (1–10), made by Vicherek, J. 1972: Die Sandpflanzengesellschaf- Koubková 2007 (orig. tab./rel.: 1/1, 2, 27, 62, 78), ten des unteren und mittleren Dnjeprstromge- surroundings Bzenec and Hodonín in Moravia); bietes (die Ukraine). Folia Geobotanica et 5 rels. (18–22) made by Hršak 2004, orig. tab./ Phytotaxonomica 7: 9–46. rel.: 1/6, 4, 9, 13, 17), all from Ðurdevac Sand Bo- tanical Reservation in Croatia.

7. APPENDIX

Table heads:

Column A, 6 rels., all by Májeková 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, Borská nížina Lowland. Column B, 13 rels ., 3 rels. (27–29) Májeková 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 E1 and E O (%), specific locality:

52 101001 20 184 50 30 20 Šajdíkove Humence, field near railway 49 100729 20 219 70 60 10 Lakšárska Nová Ves, field W from village 51 100101 20 180 70 20 70 Šajdíkove Humence, field near railway 44 100713 20 200 50 50 0 Lakšárska Nová Ves, field towards Šišoláky 45 100713 20 199 50 50 0 Lakšárska Nová Ves, field towards Šišoláky 41 100713 20 226 60 55 3 Studienka, field N from village 43 100713 20 203 60 60 3 Studienka, (Čechovec 198 m), aband. field 42 100713 20 199 50 50 1 Studienka, (Čechovec 198 m), aband. field 48 100729 20 165 50 50 2 Borský 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 Lájer 2004 (orig. tab./rel.: 1/1–7, during 2000–2001 in Bolső- Somogy in Hungary). Column D, 16 rels., 5 rels, Koubková 2007 (orig. tab./rel.: 1/4, 8, 29, 58, 59) surroundings Bzenec in Moravia) + 11 rels. made by author – number in table, date (yymmdd), Received 16. 5. 2011 plot (m2), altitude (m a.s.l.), cover total (%), cover Revision received 14. 10. 2011

E1 and EO (%), specific locality: Accepted 31. 10. 2011

11 Hacquetia 11/1 • 2012, 5–15 ...... r r 2 2 2 + ...... r 2 1 2 5 2 ...... r 2 0 3 1 2 ...... r . r 1 9 2 2 2 ...... 1 1 8 2 3 ...... 1 4 0 2 2 + E + ...... 0 2 9 + + + + + ...... 0 3 5 2 + + + + ...... 0 2 2 + 2 + + ...... 0 5 1 + 2 + 2 2 + + ...... r r . r . r r r 3 2 6 2 2 1 + ...... r . . . r 1 3 2 8 2 2 + + ...... 2 4 2 3 6 2 1 + 2 + + 1 + + ...... 4 2 1 7 2 2 1 + + + + + + + + ...... r 2 3 3 4 2 1 2 2 1 + + + ...... 4 2 2 0 2 + + + + + ...... r r 1 3 1 2 7 3 3 2 + 1 + 1 + + ...... r r 3 1 2 2 9 2 1 + 2 + + ...... r 3 2 5 2 1 + + + D ...... 3 1 1 3 1 1 2 1 + 2 1 2 2 + ...... r 3 2 1 + + + ...... 2 0 8 1 + + + + + ...... r . . r . . 0 7 3 + + + ...... 0 3 6 2 + + + ...... 0 1 4 2 + 1 + + ...... 1 0 2 3 2 + r ...... 1 1 7 3 1 + + ...... 1 1 6 3 1 1 ...... 1 1 1 5 4 + + ...... 1 4 3 1 1 1 + 1 + + C ...... 1 2 3 1 3 1 + 1 1 1 + ...... 1 2 2 1 1 + 1 + ...... 1 1 2 1 1 + + + + ...... 2 5 1 0 1 2 1 2 2 2 + ...... r . . 1 4 8 2 3 2 + 1 + + + + ...... 4 2 3 2 2 + + + + + + + ...... r . r r 4 3 3 1 2 1 + + + + ...... r . 1 4 1 3 2 2 + + + + + ...... 4 5 2 2 2 2 + + + 1 + ...... r 1 4 4 2 2 2 2 + + + ...... 5 3 1 2 1 1 + + + B ...... 1 4 9 3 1 2 + + 1 + + + + ...... 2 5 1 2 1 2 + + ...... r 2 9 + + 2 + 3 + + ...... 1 2 2 5 8 1 2 + + + + . . . urejeni s programom TWINSPAN, od najbolj ruderaliziranih peščenih polj (stolpec A–B) do relativno naravnih in naravnih do relativno ruderaliziranih A–B) peščenih od(stolpec najbolj polj TWINSPAN, s programom urejeni ...... 1 2 2 7 + + + + + + + + + ...... 2 3 5 0 2 2 2 ...... 1 2 5 4 1 1 + + + -grasslands arranged in TWINSPAN order; from most ruderalised sandy fields (columns A–B) to relatively natural- or or natural- relatively to A–B) ruderalised most fields (columns from order; sandy in TWINSPAN arranged -grasslands ...... 2 3 2 1 1 2 4 2 2 + + + + ...... 1 2 6 3 + + + 1 + + + + A ...... 1 2 2 3 3 4 2 + ...... 2 2 5 1 + 2 + + + + + T T T T T T T T T C T T T C T T C H H G H H H H H H H H H M LF ) dominii ) guestfalica (incl.* (incl.* Weed species Weed Conyza canadensis Relevé number Column Peucedanum oreoselinum Thymus serpyllum Chondrilla juncea Anthemis ruthenica Polytrichum piliferum Dianthus carthusianorum Elytrigia repens Digitaria sanguinalis Armeria elongata Scleranthus polycarpos Psammophytes Corynephorus canescens Agrostis capillaris Agrostis Teesdalia nudicaulis Teesdalia Festuca ovina Jasione montana Crepis tectorum Crepis Erysimum diffusum agg. Festuca vaginata Logfia minima Helichrysum arenarium Trifolium arvense Trifolium Acetosella vulgaris Stipa borysthenica Koeleria glauca Logfia arvensis Veronica dillenii Veronica Spergula pentandra Spergula Apera spica-venti Spergula morisonii Spergula Relevés of the Corynephorus of 1: Relevés Table (1990). Jurko by (LF) in accordance forms C–E). Life (columns dunes the sandy on stands nature-related Corynephorus z vrsto travišč 1: Popisi Tabela (1990). (LF) so z Jurko v skladu oblike peščenih na sestojev C–E). Življenske dinah polnaravnih (stolpec

12 M. Valachovič: Succession model with Corynephorus canescens in abandoned sandy fields (W Slovakia) ...... r ...... r ...... 2 ...... 2 ...... r 1 ...... + + + ...... r + + + + + + ...... 2 2 2 + 2 3 + ...... 1 ...... r + ...... 3 + 2 2 + + ...... 3 + + 3 + ...... 2 1 1 1 1 2 + + ...... 1 + + 2 + + ...... 2 2 + 1 + ...... r ...... 3 1 + + 2 + + + ...... 1 1 2 + + 2 + ...... r 3 2 + + 1 + + ...... + 1 + ...... 3 + + + ...... 1 + + + + ...... r . . . . 2 1 ...... + + ...... + + 2 + ...... r 1 ...... + + ...... 1 + ...... 1 + + ...... 1 1 + ...... 1 + ...... 1 1 + ...... 1 + + ...... 1 1 + 4 + + + ...... + 1 ...... r ...... 1 ...... 1 + ...... r ...... + ...... 3 1 2 + + ...... r ...... 1 2 1 ...... 1 1 + + 1 + + + ...... + ...... + + + + + ...... 1 + + 3 + ...... 1 + 2 1 + + ...... 2 + + 1 + + + + + ...... + + + + + + 3 ...... + + ...... + 2 + + + ...... 5 + + L L T L T L T L L T T L L T L T L T T L T L L T T T H H G H C H C G H H H M Hypochaeris radicata Acetosa thyrsiflora Myosotis stricta Cladonia coniocraea Chenopodium album Cladonia -lichens Cladonia foliacea Equisetum arvense Vicia lathyroides Vicia Agrostis gigantea Agrostis Cladonia pleurota Cladonia subulata Ambrosia artemisiifolia Ambrosia Arabidopsis thaliana Solidago gigantea Cladonia phyllophora Cladonia fimbriata Cynodon dactylon Setaria viridis Tragopogon dubius Tragopogon Cladonia furcata Cladonia rei Bromus hordeaceus Bromus Echium vulgare Papaver dubium s.lat. Cladonia *verticillata Bromus tectorum Bromus Cladonia pyxidata Calamagrostis epigejos Calamagrostis Plantago lanceolata Artemisia campestris Cladonia chlorophaea Cladonia rangiformis Erodium cicutarium Erodium Artemisia absinthium Myosotis arvensis Others Ceratodon purpureus Viola arvensis Viola Cladonia *mitis

13 Hacquetia 11/1 • 2012, 5–15 ...... r 2 2 3 2 ...... r r r 2 1 + ...... 2 0 + + ...... r 1 9 + ...... 1 8 1 1 ...... 1 2 0 2 E + ...... r . 0 9 2 + 2 2 + 2 + + ...... r 0 5 + + ...... 0 1 2 + ...... 1 0 2 1 2 1 + 2 + + + ...... 3 6 + ...... 3 8 + + + + ...... r r r r 4 6 + + 2 + 1 + + ...... 4 1 7 2 1 2 1 + + + + + + + ...... 3 4 2 + + + + ...... r r 4 0 1 + + + + + ...... r 3 7 + 1 + + + ...... r 3 9 1 2 + ...... r r 3 5 + D ...... 2 3 3 1 + 2 + + + + + + ...... r 3 2 2 + 2 + 2 + + + + ...... 0 8 ...... 0 7 + ...... r . r 0 6 ...... 0 1 4 + ...... r 0 3 ...... 1 7 ...... 1 1 6 + ...... 1 1 5 + + ...... 1 4 + + + C ...... 1 1 3 + + ...... 1 2 1 ...... 1 1 1 1 1 ...... 5 0 2 ...... 4 8 + + ...... 4 2 + + ...... 4 3 1 ...... 4 1 ...... 4 5 + + ...... r r 4 4 + ...... 5 1 + B ...... r 4 9 ...... 5 1 2 2 + ...... 2 9 ...... 2 8 + + + ...... 2 7 4 + ...... 3 0 + ...... 2 4 + ...... 3 1 + ...... 2 6 + + A ...... 2 3 2 ...... 2 5 + P T C T T C C H H G G H G H C H G H H H H G H H H H H H H H H H H H H H H LF agg. agg. sp. Scabiosa ochroleuca Relevé number Column Scleranthus perennis Sedum sexangulare Carex hirta Carex Carex supina Carex Poa bulbosa Carex praecox Carex Acosta rhenana Pinus sylvestris Trifolium alpestre Trifolium Carex ericetorum Carex Pilosella officinarum Anthoxanthum odoratum Phleum phleoides Sarothamnus scoparius Sarothamnus Linaria genistifolia Tithymalus cyparissias Tithymalus Silene otites Teucrium chamaedrys Teucrium Luzula campestris Pilosella echioides Agrostis vinealis Agrostis Achillea millefolium Senecio jacobaea Festuca Hypericum perforatum Petrorhagia prolifera Petrorhagia Calluna vulgaris Poa pratensis agg. Psyllium arenarium Solidago virgaurea Pilosella cymosa Danthonia decumbens Potentilla argentea Holcus lanatus Cerastium semidecandrum Festuca rupicola

14 M. Valachovič: Succession model with Corynephorus canescens in abandoned sandy fields (W Slovakia)

...... r sp...... Silene Silene ...... [H] 1: +; ...... Petrorhagia Petrorhagia Elytrigia ...... r ...... + Cerastium [T] 39: r; Cerastium Saponaria [T] 24: +; Saponaria ...... + ...... Vulpia myuros [H] 50: 1; Vulpia

...... 2 ...... Seseli osseum osseum [T] 1: +; Seseli ...... + 1 + ...... [G] 32: r;32: [G] neglectum Muscari ...... Silene viscosa [H] 4: 1; Silene Vicia villosa [T] 24: +; Vicia ...... + + Brachythecium salebrosum+; 52: [M] [H] 18: r; ...... r 3 ...... + + ...... 3 1 + + ...... r + ...... podperae [T] 42: r;x podperae Festuca rubra agg. [C] 32: r;32: [C] arvensis Acinos ...... 1 Bromus erectus erectus [H] 4: 1; Bromus

[M] 1: 2; Cerastiumbrachypetalum

[T] 52: 1; 52: [T] foetida Crepis ...... sp. agg...... Veronica arvensis arvensis [T] 23: +; Veronica ...... + ...... r + ...... [H] 1: 2; Bryum ...... Tithymalus strictus[L] 40: +; Tithymalus macilenta [L] 40: +; Cladonia coccifera [L] 40: +; Cladonia sp...... [L] 32: +; 32: [L] didactyla Peltigera Poa nemoralis nemoralis [H] 35: r; Poa ...... Holosteum umbellatum2; 30: [T] [T] 15: +; Muscari15: [T] +; Bromus15: [G] comosum squarrosus arenarium Polygonum +; [T] 16: ...... Papaver *austromoravicum *austromoravicum [M] 23: 1; Papaver ...... Anthemis austriaca [T] 9: r; [T] 9: +; Anthemis [P] 10: [T] 9: +; MedicagoBetula pendula minimaincana [T] 9: +; Berteroa [H] 5: +; Onopordum acanthium [H] 9: 2; Thymus pannonicus [C] 9: 2; Cerastium glutinosum [T] 9: 2; Anchusa ......

...... sp...... 2 Linaria vulgaris vulgaris [M] 35: +; Linaria ...... Alyssum alyssoides1; 32: [T] ...... + ...... [H] 24: +; 24: [H] patula Campanula [T] 32: 2; 32: [T]

...... [T] 14: +; 14: [T] humifusum Hypericum ...... Bryum argenteum [T] 25: +; Bryumargenteum Verbascum phoeniceum phoeniceum [P] 8: r;[T] 8: +; Populussp. Verbascum ...... L T L T G C H H H M M M [C] 35: 1; Dicranumscoparium Thymus pulegioides [C] 33: +; Asperulapulegioides [H] 33: +; Thymus cynanchica cupressiforme[H] 33: +; Hypnum [M] lychnitis [H] 33: 1; Galium verum [H] 33: +; Verbascum . [H] 24: +; 24: [H] mollis Holcus Oenothera biennis biennis Oenothera Arenaria serpyllifolia agg. serpyllifolia Arenaria sp Achillea pannonica+; 14: [H] [H] 24: +; 24: [H] Lepidium ruderale ruderale [G] 9: +; Lepidium intermedia [T] 9: +; Elytrigia [H] 32: 2; 32: [H] Erophila verna [T] 37: +; Saccomorpha verna icmalea [L] 40: +; Diploschistes[C] 39: r; Erophila Scleranthus [H] 43: +; Acetosa pratensis [H] 43: r; [T] 45: r; Scleranthus arvensis [T] 52: r; glomerata Dactylis Spergula Avenula [G] 33: 1; Avenula leptopterum [T] 47: +; Corispermum [H] 1: r; collina Erigeron [T] 1: +; Potentilla segetum Cyanus Veronica officinalis officinalis 35: 2; Veronica officinalis saxifraga saxifraga arvense *alba *alba +; 13: [T] officinalis officinalis Secale cereale In one relevé only: one In [C] 18: 1; Alyssum *gmelinii sp. [T] 18: r; Hypericum montanum [P] 10: r; Teucrium robur [M] 10: +; Quercus polysetum 1; Dicranum Peltigera rufescens Asparagus officinalis Racomitrium canescens Senecio viscosus Lactuca serriola Cetraria sp. Potentilla arenaria Pleurozium schreberi Pleurozium Arrhenatherum elatius Arrhenatherum Genista pilosa Brachythecium albicans