H-7000 Sárbogárd, Ady E. út 162, Hungary; (Received 13 January, 2012; Accepted 15 June, 2012)

Transcription

1 Acta Botanica Hungarica 55(1 2), pp , 2013 DOI: /ABot STEPPE WOODLANDS WITH TATARIAN MAPLE (ACERI TATARICI-QUERCETUM PUBESCENTIS-ROBORIS) ON THE GREAT HUNGARIAN PLAIN AND ITS NEIGHBOURHOOD. AN UNFINISHED SYNTHESIS WITH SUPPLEMENTARY NOTES B. Zólyomi, A. Horváth 1, B. Kevey 2 and G. Lendvai 3 1 Institute of Ecology and Botany, Centre for Ecological Research, Hungarian Academy of Sciences H-2163 Vácrátót, Alkotmány u. 2 4, Hungary; ahorvath@botanika.hu 2 Department of Botany and Phytogeography, and Department of Agrobotany and Viticulture University of Pécs, H-7624 Pécs, Ifjúság útja 6, Hungary; keveyb@ttk.pte.hu 3 H-7000 Sárbogárd, Ady E. út 162, Hungary; gaborlendvai@hotmail.com (Received 13 January, 2012; Accepted 15 June, 2012) This paper presents the so far only partially published research material of the late Bálint Zólyomi on one of his major fields of interest, the forest steppe vegetation. The phytosociological tables presented here were found in his bequest indicating that he was going to publish a grand synthesis on this topic, which, however, has not come true. Since no written text accompanied the original tables, a historical overview and a short explanation to the tables are also provided. Key words: forest steppe, phytosociology, Hungary INTRODUCTION A major part of the lifelong scientific contribution of Bálint Zólyomi is his work on the forest steppe vegetation found on the Great Hungarian Plain. The comparative analysis of steppe woodlands in the forest steppe zone growing on chernozem soils is one of his most frequently cited and widely known works (Zólyomi 1957). In this paper, he was the first to identify several forest stands as representatives of steppe woodlands on the loess plateaux in Hungary he named Acereto tatarici-quercetum pubescentis-roboris, and to point out the uniform nature of the steppe woodlands on loess across the European forest steppe zone. The forests described in the study were mostly restricted to the southern foothills of the mountain ranges. Among them was the forest Kerecsendierdő, a new discovery of two ambitious and then young botanists, Pál Jakucs /$ Akadémiai Kiadó, Budapest

2 168 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. and Tamás Pócs (see Pócs 2001), which later became the standard of the Pedunculate Oak-Tatarian Maple steppe woodlands in Hungary. Several stands of the Lovasberény Forest on the eastern side of the hills Velencei-hegység, which were studied by Gábor Fekete in 1955 as part of his master s thesis (Fekete 1955), were also included. Zólyomi also collected samples from the hill Somlyó at Fót, and included one relevé from each, the Mezőcsát and Ohat forests, both on the Great Hungarian Plain. The number of the studied stands thus was relatively small. In a work published a year later, Zólyomi (1958) reported the analytical results of a different set of stands, but unfortunately, again in the form of a synoptic table. From the text it is apparent that he included samples from new locations, such as Vácrátót and Budaörs. In the only review on steppe woodlands on loess occurring in Hungary (Zólyomi 1967), he once again used only a synoptic table, which now contained a much larger sample of forests. According to the list of locations, the samples were collected exclusively east of the Danube, and included all relevés published in his first paper under the subassociation hungaricum. However, he removed the two forests on the plain from the samples, and placed them in steppe woodlands on alkali soils (Galatello-Quercetum roboris Zólyomi et Tallós 1967). As indicated by these publications, Zólyomi kept working on steppe woodlands over several years. The true scope of this work, however, could not be appreciated from these publications. His work on oak steppe woodlands had been even more extensive than what was perceived from the published data and his comments on the subject. Between 1957 and 1961, he intensively searched for existing remnants, and recorded in detail their phytosociological characteristics. He, however, did not work alone. His most faithful companions were his former students and young colleagues, namely Pál Jakucs, Gábor Fekete and Tamás Pócs from the Herbarium of the Natural History Museum, and Gábor Vida, who not only participated in the surveys of several forests, but also sampled some stands completely alone. During this period, Zólyomi and his team surveyed and sampled the forests in the central (around the city of Gödöllő) and southern (the neighbourhood of Albertirsa and Pánd) parts of the hills Gödöllői-dombság, on the Great Hungarian Plain, and on the southern foothills of the North Hungarian Mountain Range. We have found unpublished phytosociological tables from Gödöllő and Dormánd in his heritage, which perhaps would be gladly published, as new finds, by current botanists. Zólyomi also did research west of the Danube. In addition to the hills Velencei-hegység, where material was collected by Gábor Fekete, he extended the search for steppe woodland remnants to the Mezőföld. The unpublished vegetation samples collected at Érd and Dunaszentgyörgy are evidence of this.

3 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 169 Apparently, Zólyomi was preparing to publish a major comprehensive synthesis of steppe woodlands in Hungary. This is evident from the two largesized annotated tables and the accompanying field notes and data sheets found in his bequest, which summarised the results of the more than forty vegetation samples collected in an area that stretches from the Mezőföld in eastern Transdanubia to the river Sajó in northeastern Hungary. Unfortunately, the majority of his data collected over several years has never been published, although the study of the forest steppe vegetation was one of the main research projects of the Botanical Institute at Vácrátót, the workplace of Zólyomi at the time (Zólyomi 1959). The reason of this is not yet clear. During the forty years that have passed since then, no significant progress has been made in the study of the forest steppe woodlands growing on chernozem soils in Hungary. In fact, no-one studied this community in detail, and not even at the descriptive level. Regrettably, Gábor Fekete did not discuss steppe woodlands in his book on the forests of the hills Gödöllői-dombság (Fekete 1965), which apparently occurred there. As he indicated, he omitted these forests from his book, because a large analytical work by Bálint Zólyomi had been in progress at that time. Clearly, he referred to the material that has never been published in full. The only paper known to us appeared in 1982 on the thermophilous forests of the Somlyó Hill at Fót (Fekete and Kovács 1982), in which the authors reported phytosociological relevés also on steppe woodlands, among which two were the original samples collected by Zólyomi in Thus, our recent knowledge on these forests is based exclusively on the works of Zólyomi discussed above. Recently, scientific attention has gradually shifted to the forest steppe vegetation again owing to its importance in biogeography and vegetation history. However, a considerable portion of the once known woodland stands has suffered drastic changes since the time passed. Gábor Fekete reported in the late eighties that the stands he surveyed in the hills Velencei-hegység had mostly been spoiled by planting Turkey Oak in them. The once remnant woodlands of the cold continental forest steppe in the hills Gödöllői-dombság (Fekete 1965) were also destroyed by forest mismanagement (Fekete, ex verb.), although there are reports that a few fragments have recently been located (Molnár 1999, Molnár and Kun 2000). The unfavourable changes observed in the Kerecsend Forest are also significant. In the absence of recent surveys, the fate and current condition of several other, once studied woodlands (the forests at Mezőcsát, Demjén, Sajólád, and the Somlyó Hill at Fót) are unknown. By the end of the nineties, it became apparent that steppe oak woodlands with Tatarian Maple represent one of the most endangered and rarest vegetation types of the forest steppe in Hungary (Molnár 1999, Molnár and Kun

4 170 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. 2000, Bölöni et al. 2008). The field surveys launched at the same time were aimed at providing the foundation for future research by locating all the still existing steppe woodland remnants besides the once known stands. The first catalogue of this data base has been put together (Molnár 1999), from which, however, several, once studied stands, and even entire regions were omitted. Our knowledge on steppe woodlands on chernozem soils cannot be complete without publishing the survey material of the once studied stands. Following the suggestions and advice of Prof. Gábor Fekete, and also as a tribute to the author and first researcher of the association, we decided therefore to make available for the public the whole research material of Bálint Zólyomi as known to us, of which a considerable portion has never been published, and a part of it has been available only in synoptic tables. We believe that by this we can pay one of the largest and very long lasting debts in the field of vegetation science in Hungary, and thereby draw the attention to the importance of the works and accomplishments of our forerunners. Our primary goal is to make public even the data and information on this subject that Bálint Zólyomi accumulated during his scientific career, but were never scientifically analysed and made available for scientific research. It follows that it is by no means our goal to evaluate the data in any way in this research material. Instead, we made great effort to accurately reproduce the strictly original material. We preserved the scientific names of the species used in the manuscripts and species grouping, the subassociation names applied by the author, as well as the abbreviations used in his tables. Deviations from the original manuscript occur only in three details. First, we placed the accidental species back into the tables for better reading as opposed to Zólyomi, who summarised them in a list at the end of each section of his tables. Second, we listed the A D and K values of each subassociation not at the right end of the table in separate columns, but directly next to the columns of the corresponding syntaxonomic unit. Third, we corrected the apparent typographical errors in the tables written by a conventional typewriter. We hope that the amount and details of the research material published here will themselves convince everyone about the lifelong achievements of Bálint Zólyomi on this subject. The floristical data that may be derived from these relevés and the published locations of the studied stands may further stimulate the search for even further remnant stands of steppe woodlands and the study of their flora and vegetation. ABOUT THE RESEARCH MATERIAL IN THE MANUSCRIPT Zólyomi summarised the results of his research on steppe woodlands on loess bedrock in two typed and annotated large-sized tables. The first table

5 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 171 contains the data of 21 vegetation samples collected east of the Danube. They were named by Zólyomi as the regional association Aceri tatarici-quercetum pubescentis-roboris hungaricum (Appendices 1 & 2). The second table includes 22 relevés that were collected in part west of the Danube in the Velence Hills, the Tétény Plateau, and the Mezőföld, and in part at the southern foothills of the Cserehát Hills (Fót, Kosd, Vácrátót). They were named the Aceri tatarici- Quercetum pubescentis-roboris submatricum association (Appendices 3 & 4). Zólyomi included all of the first table in the synoptic table in his 1967 paper. The better part of the second table, however, is essentially unpublished, as only a small fraction was used in the 1958 study, also in the form of a synoptic table. The only relevés published individually are the ones from Fót (Fekete and Kovács 1982). The two tables are further divided into parts, which Zólyomi distinguished as subassociations. Both tables contain the subassociations festucetosum and lithospermetosum, which mainly differ in the density of the forest canopy and, as a consequence, the composition of the understory. In addition, the first table also contains the subassociation galietosum, in which the proportion of mesophilous forest herbs indicating more mesic conditions is higher. These categories reflect the original view of Zólyomi on the regional and habitat variation of the association. With the inclusion of TWR values and categories of geographical distribution and life-forms, the tables provide ample information on the then current view on the ecological features, distribution patterns, and phytosociological affinities of the included species. For the latter, Zólyomi used various symbols next to species names, which, however, are not always defined in the heading of the appropriate section. While editing the tables, we noticed that Zólyomi reported the occurrence of Pannonian Ash (Fraxinus angustifolia subsp. pannonica) for the Kerecsend forest. However, one of the authors (B. K.) searched for this species in vain, while working recently at the same location, and found only the related Common Ash (F. excelsior). As the former species is restricted mainly to mesicmoist habitats and thus occurs on river floodplains in Hungary, it is likely that Zólyomi has mistaken common ash for Pannonian Ash, an error easily made because of the strong resemblance of the two species. The supplementary data on the relevés are often incomplete. Unfortunately, Zólyomi often made his notes on a piece of paper happened to be at hand, some of which may still be found in his bequest. However, even these do not show the date or other circumstantial information. His lack of care is indicated by his later notes, in which he made an attempt to collect all the data for every relevé, and where huge question marks indicate the missing data. Occasionally, even the sample location was uncertain for Zólyomi. Thus, even

6 172 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. with the help of his often deficient diary and field notes we managed to collect only part of the supplementary data. We have been unable to recover any additional information on the rest so far. * Acknowledgements We wish to express our thanks to the Bálint Zólyomi Foundation and the Institute of Ecology and Botany at Vácrátót for allowing and helping our necessary research in the bequest in every way. We also wish to express our thanks to Prof. Gábor Fekete, who not only drew our attention to this material and provided us with its copies, but also personally asked us to make the complete material public before we publish our own results. Here we gladly fulfil his request. Without his encouragement, support, and very useful advice, this work could not have been made possible. REFERENCES Bölöni, J., Molnár, Zs., Biró, M. and Horváth, F. (2008): Distribution of the (semi-)natural habitats in Hungary II. Woodlands and shrublands. Acta Bot. Hung. 50(Suppl.): Fekete, G. (1955): Die Vegetation des Velenceer Gebirges. Annls hist.-nat. Mus. natn hung. 7: Fekete, G. (1965): Die Waldvegetation im Gödöllőer Hügelland. Akadémiai Kiadó, Budapest, 223 pp. Fekete, G. and Kovács, M. (1982): A fóti Somlyó vegetációja. Bot. Közlem. 69: Molnár, Zs. (ed.) (1999): Erdőssztyeppmozaikok az Alföldön. WWF MTA ÖBKI, Vácrátót Budapest (manuscript). Molnár, Zs. and Kun, A. (eds) (2000): Alföldi erdőssztyepp-maradványok Magyarországon. WWF füzetek 15: Pócs, T. (2001): Hej, azok a növénytári évek Kitaibelia 6(2): Zólyomi, B. (1957): Der Tatarenahorn-Eichen-Lösswald der zonalen Waldsteppe. Acta Bot. Acad. Sci. Hung. 3: Zólyomi, B. (1958): Budapest és környékének természetes növénytakarója. In: Pécsi, M. (ed.): Budapest természeti képe. Akadémiai Kiadó, Budapest, pp Zólyomi, B. (1959): Beszámoló az MTA Botanikus Kertje és Geobotanikai Laboratóriuma munkájáról. (Bericht über die Arbeit des Botanischen Gartens und Geobotanischen Laboratoriums der Ungarischen Akademie der Wissenschaften). MTA Biol. Csop. Közlem. 3: Zólyomi, B. (1967): Aceri tatarico-quercetum pubescenti-roboris (hungaricum). In: Zólyomi, B. (ed.): Guide der Exkursionen des Internationalen Geobotanischen Symposiums. Ungarn, Eger Vácrátót, 5 10, Juni 1967, pp

7 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 173 Appendix 1. Description of the relevés of the Aceri tatarici-quercetum pubescentis-roboris hungaricum samples Location Date of relevés Elevation a.s.l. (m) 1: Pánd: Hársas Area Exposition Made by first second (m 2 ) first second Declination ( ) NNW Zólyomi Ulbrich Stieber 2: Alberti, 2nd valley NOO Zólyomi Máthé Kovács Fekete 3: Kerecsend: Fácános, SW corner E Zólyomi-(V) Zólyomi Pócs Vajda 4: Demjén Zólyomi Pócs Vajda 5: Demjén SWW Zólyomi Fekete 6: Felsővadász (Cserehát): Nagy-erdő 1: Alberti: Hársas, 3rd valley SE Zólyomi Jakucs Futák NE Zólyomi 2: Alberti, 2nd valley Zólyomi Máthé Kovács 3: Alberti, 2nd valley NE Zólyomi Máthé Kovács 4: Alberti: Hársas, 1st valley 5: Kerecsend a, NW corner 6: Kerecsend b, NE corner (útbejáró) Zólyomi Kárpáti Újvárosi Zólyomi Pócs Vajda 7: Kerecsend c, S corner S Zólyomi Pócs Vajda

8 174 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Location Date of relevés Elevation a.s.l. (m) Appendix 1 (continued) Area Exposition Made by first second (m 2 ) first second Declination ( ) 8: Kerecsend g Zólyomi Pócs Medvecka Kornas Zólyomi Futák Mihalkó 9: Vécs-Feldebrő Zólyomi Fekete Zólyomi 10: Alberti? SSW Zólyomi Máthé Kovács Fekete 1: Aszaló: Pap-erdő S Zólyomi Fekete 2: Aszaló: Pap-erdő Zólyomi Fekete 3: Sajóvámos: Frank-h S Zólyomi Fekete 4: Sajóvámos: Frank-h S Zólyomi Fekete 5: Kisnémedi Szórivölgy NNW Zólyomi

9 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 175 Appendix 2. Aceri tatarici-quercetum pubescentis-roboris hungaricum festucetosum lithospermetosum galietosum T W R K A D K A D K A D Aceri-Quercion (Prunion spinosae-fruticosae) species 6k 4k 4 P-Pann MM A2 Acer tataricum 1 ( +) (1) 1( 2) k 4 4 P-Pann MM A2 Acer tataricum 1 ( +) (1) 1( 2) M B x (+) V + 2 +(2) V V + 3 M B III II III + 1 6k 2 4 Kt M B1 0 (Cerasus fruticosa sst.) + 1(2) I + 2 (+) (+) + + IV + 6k 2 4 Eua-Kt M B1 0 (Amygdalus nana sst.) 1 I 1 M B2 + I + Quercetalia petraeae-pubescentis Quercetea pubescenti-petraeae (= common with Carpino-Fagetea) species 6a 2 5 Subm MM A1-(A2) Quercus pubescens V (x) + 2 (+) V (+) III + 1 M B II I + M B III + 5a 4 3 Eua M B1 Crataegus monogyna x 1 + V V + 1 +(2) + + III + 5a 3 3 Eua M B1 Prunus spinosa + + x + IV + + (+) x V III + 2 5a 4 4 Balk-Eu M B1 Euonymus verrucosa + 1 x III IV IV + 2 5a 3 3 Subm(or) MM A1-(A2) Quercus cerris IV (+ 1) III I 1 M B III II I 2 M B V II I + 5a 3 4 Eua M B1 Rhamnus catharticus + I (+) + + x + III IV Eu MM A2 Pyrus achras (+) I + (+) + I II + M B1 + + II + + x I II + 6a 2 4 P-Subm M B1 Rosa gallica III I II + 6a 4 4 Subm-Em M B1 Viburnum lantana (+) I + + (+) (+) II III + MM A1-(A2) Quercus hybr. 1 I (+) II + 1 6a 3 4 Subm-Em M B1 Cornus mas + I Eua M B1 0 Rosa rubiginosa + I + + I + Querceo-Fagetea s. l. (0 = Carpino-Fagetea) species 5a 6 0 Eu MM A1-(A2) Quercus robur III + 2 (+ 2) x 1( 2) IV V + 3 M B1 + I II III + 2 M B2 + I II + 5a 4 4 Eu MM A2 Acer campestre 1 1 II (x) 1 IV + 2 1( 2) IV + 2 M B1 Acer campestre III + 1 3( 4) V V 1 2 M B2 + I II I + 5a 5 3 Eu M B1 Euonymus europaea x + + V (+) V IV + 1 5a 4 3 Eu M B1 Ligustrum vulgare + 1 x 1 1 V + 1 1( 2) + + +( 2) V V 1 2 5a 3 3 Eua M B1 Rosa canina s. l. + + x 1 + IV III II + 5a 4 4 Eu-Subm M B1 Cornus sanguinea III + (+) III x 1( 2) 3 IV + 3 5a 6 4 Eu MM A2 Ulmus carpinifolia + + I + + I + (+) I + M B1 (+) + + III I + + I + M B1 Crataegus oxyacantha + + I III + 5a 4 0 Eu MM A1-(A2) Quercus petraea (+) I (2 3) III III + 1 M B1 + I I 1 M B2 + I +

10 176 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Appendix 2 (continued) festucetosum lithospermetosum galietosum T W R K A D K A D K A D P-Pann MM A1 0 Fraxinus angustifolia subsp. pannonica (+) I I 2 M B1 + I + + I Eua H B1 Rubus caesius (+) I + (+) (+) II Subm(or) E A1 Loranthus europaeus 1 I 1 + I + + I + 6a 4 4 Subm(or)-Em M B1 Staphylea pinnata + I + (+) I + 5a 5 3 Em MM B1 Carpinus betulus + I + 5a 6 4 Eu M A2 Malus sylvestris + I + 5a 5 3 Eua M A2 0 Cerasus avium + I + 5a 4 3 Balk-Kauk M B1 0 Tilia cf. rubra + I + 5a 5 3 Em-Subm N-E B1 Clematis vitalba + I + Aceri-Quercion and local character species (! = common with Festuco-Brometea) 6k 2 5 P-Pann G C Iris variegata sst. 1( 3) I 1 (+) + (+) (+) + 1 (0 2) 1 + V II Kt H C 0 Phlomis tuberosa sst. +( 3) (1 2) 1 V (+) II III Kt H C Pulmonaria mollissima sst II + 1 (+) (+) + 1(2) IV (1) IV + 1 5a 3 4 Subm(o) H C Carex michelii sst IV (+) I + (+) (+) + III Kt H C Melica altissima sst. + I x 0 1 III (+) 0 2 IV Pann-Balk H C Euphorbia polychroma + I III + 1 (+) (+) 1 III + 1 6k 3 4 Balk-Pann G C 0 Doronicum hungaricum sst. + + II (+ 1) 1 II + 1 (+) I Eua H C 0 Nepeta pannonica sst. 1 + II II I 1 6k 3 3 Pann-Balk H C 0 Cytisus albus sst. (+) I 6k 2 5 Kt G C Lathyrus pannonicus subsp. collinus sst. (x) I + (+) I P-Subm H C 0! Peucedanum officinale sst. + I + (+) (+) I + 6k 4 4 Eua-Kt H C 0! Trinia ramosissima + I P H C 0 Viola suavis 2 I 2 Quercetalia petraeae-pubescentis Quercetea pubescenti-petraeae species (0 = Quercion petraeae-group species,! = common with Festuco-Brometea, = common with Carpino-Fagetea) 6k 2 5 Eua H C Dictamnus albus sst III V III + 6a 2 4 Subm-Em Ch(H) C! Teucrium chamaedrys sst V I 1 (+) + II + 5a 2 4 Eua H C Brachypodium pinnatum sst IV II IV + 1 5k 4 3 Eua H C Inula salicina 0 1 I II (+) +( 2) + IV + 5a 3 4 Eua H C Sedum maximum sst. + + II III (+) II Eua H C! Silene cucubalus + + II (+ 1) IV + 1 (+) I + 5k 3 4 Kt H C 0 Thalictrum minus sst. + +(2) + + IV + + (+) II II + 6k 3 4 Eua G C 0! Asparagus officinalis + + II + + (+) III + (+) I Eua H C! Betonica officinalis sst III II II Cp H C Calamintha clinopodium III + + (+) II (+) + III + 5a 3 4 Em-Subm H C Chrysanthemum corymbosum III III (+) + III + 6k 3 4 Eua H C Hirundinaria vincetoxicum (+) 1 + II + 1 (+) III + 1 (+) (+) + III + 6k 4 4 P-Pann TH C Lactuca quercina + + II III III + 5a 4 3 Em H C 0 Lathyrus niger + + II II III Balk-Pann Th-H C Verbascum austriacum III + 1 5k 4 3 Eu H C 0 Vicia cassubica 2( 3) + + III II Eu H C Astragalus glycyphyllos + + II I + (+) + II + 6k 3 4 Kt H C 0 Campanula bononiensis + I II + + I +

11 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 177 Appendix 2 (continued) festucetosum lithospermetosum galietosum T W R K A D K A D K A D 5k 4 4 Kt H C 0 Carex tomentosa + II + 1 5a 3 4 Eua H C! Galium mollugo 1 + II I I 1 6a 2 5 P-Subm G C Iris graminea + (+) 1 + II + 1 (+) I + 5k 3 5 Eu-Kt Th C 0! Melampyrum cristatum sst. + I I (+) II + 6a 4 4 Subm G C 0 Muscari botryoides II Eua H C Origanum vulgare + + II II + 1 (+) I + 5a 2 5 Kt H C! Peucedanum cervaria sst II II + 1 (+) + II + 5a 4 3 Em H C 0 Potentilla alba + (+) II x II + 1 5k 4 3 Eua H C 0 Ranunculus polyanthemos sst. + + II Eua H C Serratula tinctoria + I II + + I + 5a 3 4 Eu H C Trifolium alpestre (+) I + + I + (+) + 1 II + 6a 3 5 P-Subm H C Lithospermum III ( 3) + 3 (+) 1 2 V I + purpureo-coeruleum (4) 5k 3 4 Em H C Vicia pisiformis + I + (+) + (+) + 1 II II + 5a 3 4 Eua H C! Viola hirta + 1 II II I + 5a 3 2 Eua H C Viscaria vulgaris + + II I + 5a 4 3 Eua H C 0 Carex montana sst. I (+) + I 5a 3 4 Em H C! Coronilla varia I I + 5k 3 4 Kt H C! Fragaria viridis sst. + I + + I + + I + 5a 2 5 Em-Subm H C Geranium sanguineum sst. + I I H C 0 Hieracium umbellatumracemosum + (+) I + (+) I + 5k 3 3 Eua H C 0 Hypochoeris maculata sst. + I + (+) I + 5a 3 4 Subm-Em H C Inula conyza + I + (+) I + 5k 2 4 Kt H C! Inula hirta sst. 2 I 2 + I + 6a 4 3 Subm-Em H C 0 Lychnis coronaria I Kt G C! Polygonatum odoratum sst. + 1 I + 1 I 1 5a 3 3 Eua H C Silene nutans + I + + I + 4k 4 4 Eua H C 0 Thalictrum aquilegiifolium sst. + (+) I + + I Cp Th C Turritis glabra (+) I + + I + 7a 3 3 Atl-Subm TH C! Campanula rapunculus 6k 3 5 Kt H C Clematis recta sst. (+) I 6k 4 2 Em N C Cytisus nigricans + I + 5a 4 3 Em H C Festuca heterophylla + I + 5a 3 2 Em-Subm H C Hieracium sabaudum + I Subm-Em H C Lathyrus latifolius 5k 2 3 Eua H C! Lithospermum officinale (+) I + 5k 3 5 Em H C 0 Primula veris + I + 5a 3 3 Eua H C Trifolium medium + I + 5a 3 3 Em-Subm H C Trifolium ochroleucon (+) I + 5k 3 4 Kt H C! Veronica teucrium + I + Querco-Fagetea s. l. (0 = Carpino-Fagetea) species Eua H C Fragaria vesca V (0 1) I IV + 6k 5 4 P-Pann G C Polygonatum latifolium III + 3 1( 4) V (+) 1 + 2(3) IV + 3 6k 4 4 Em H C Viola cyanea III (+) + IV V Cp H C 0 Agropyron caninum + 1 II V (+ 1) + III + 1

12 178 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Appendix 2 (continued) festucetosum lithospermetosum galietosum T W R K A D K A D K A D 5a 5 4 Eua H C Brachypodium sylvaticum + I IV III + 1 5a 6 4 Eua H C Dactylis glomerata III (+) IV III Cp H C Geum urbanum II III IV Cp H C Poa nemoralis III IV II + 5a 5 3 Eua H-G C Ranunculus ficaria + 1( 2) (+) (+) IV (+) (+) + (+ 1) IV + 2 5a 4 4 Eua H(-Ch) C Veronica chamaedrys II IV I + 5a 4 3 Em H C Galium schultesii I 1 (+) III Kozm Th C 0 Geranium robertianum (x) II (+) II + 5a 6 3 Eua H C Heracleum sphondylium + + I (+) III Cp Th C Polygonum dumetorum + + II III (+) III Eu-Kt H C Sisymbrium strictissimum III + 1 5a 6 4 Em G C Arum maculatum (+) II (+) II + 1 5a 6 3 Em H C 0 Bromus ramosus subsp. benekenii + I (+) + II II + 5a 4 0 Eu H C Carex spicata (+) I II I + 5a 6 4 Eua G C 0 Corydalis cava (+ 1) I + (2 3) (+ 1) (+ 2) II + 3 5a 5 4 Eua G C Corydalis solida 1 I 1 (+ 1) II 1 5a 3 0 Eua H C Hypericum perforatum + + II III + 1 5a 4 3 Eua Th C Lapsana communis + I + + (+) + 1 I (+) II + 5k 6 4 Eua Th C 0 Myosotis sparsiflora (1 2) II + 2 5a 4 3 Subm-Em Ch C 0 Vinca minor +( 2) (+) II Eua-Kt H C Calamagrostis epigeios +(1) I + 1 I 1 5a 4 3 Cp G C Convallaria majalis + 2 II + 2 5k 4 3 Eua H C Campanula persicifolia I + + I + 6a 5 3 Em G C 0 Isopyrum thalictroides (+ 1) + I + 1 H C 0 Pulmonaria officinalis + I + + 5a 6 3 Eua H C 0 Scrophularia nodosa + I Cp H C Solidago virga-aurea + I + + I Eu H C 0 Campanula rapunculoides + I Eua H C 0 Campanula trachelium + I + 5a 3 4 Eu H C Carex pairae + I + 5a 5 3 Eua H C 0 Epilobium montanum + I + 5a 5 3 Em Th C 0 Galeopsis pubescens + I + 5a 8 4 Eua Ch C Lysimachia nummularia + I + 5a 4 3 Eua Th C 0 Moehringia trinervia + I + 5a 6 3 Eua G C Platanthera bifolia + I + 5a 5 3 Cp G C 0 Polygonatum multiflorum + I + 5a 6 3 Eua-Kt H C 0 Ranunculus auricomus + I + 5a 5 3 Eua H-Ch C 0 Stellaria holostea 1( 2) I 1 5a 5 4 Eua H C Viola elatior + I + 6a 4 4 Subm-Eu H C Viola odorata (+) I + 5a 5 3 Em-Subm H C 0 Viola sylvestris (+) I + Festucion sulcatae Festuco-Brometea species (! = occurring also in Quercetea pubescenti-petraeae associations, = Salvio-Festucetum sulcatae species) 6k 1 4 Eua H C Festuca hirsuta-valesiaca (2) V + 3 (+ 1) I 1 (+) I + 5k 3 3 Eua H C! Filipendula vulgaris V III + 1 (+) 1 II + 1

13 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 179 Appendix 2 (continued) festucetosum lithospermetosum galietosum T W R K A D K A D K A D Cp H C Poa pratensis subsp. angustifolia V III III + 6k 2 4 P-Pann H C 0! Vinca herbacea sst ( 2) 1 2 V + 2 (+) I I Eua-Kt H C 0 Adonis vernalis IV + 1 (+) + I + + I + 5k 2 4 Eua-Kt H C 0 Arenaria micradenia (+) (+) (+) III + 5k 3 4 Eua-Kt H C Galium verum III II + 6k 2 4 Kt-Eu H C Verbascum phoeniceum + (+) + III + 6k 2 4 Kt-Eu H C Achillea pannonica + + II + (+) I Eua H C Agrimonia eupatoria + + II + I 2 (+) + II Eua G C Agropyron intermedium II 1 2 (3) 2 + I + 3 5k 3 3 Eua-Kt G C 0 Carex praecox II I + 5a 6 1 Em H C Centaurea pannonica II + I 2 5k 1 4 Eua H C Phleum phleoides 1 + II + 1 Cp H C Potentilla argentea + + I + + I + 6k 2 4 P-Subm H C 0 Ranunculus illyricus + 1 II + 1 5k 2 4 Eua-Kt H-G C 0 Ranunculus pedatus (+) (+) II + (+) I + 6a 3 4 P-Subm H C 0! Aster linosyris + I + (+) I + 5k 3 4 Eua H C 0 Astragalus cicer (+) (+) I + 5k 3 4 Eua-Kt H C Bromus inermis + I + + I + 5k 4 4 Eua-Kt H C 0 Campanula glomerata + I + + I + 6k 6 5 Eua-Kt H C 0 Clematis integrifolia + I + + I + (+) I + 5a 3 3 Em-Subm Th-TH C 0 Dianthus armeria + I + + I + 6k 3 3 P H C Euphorbia salicifolia 1 I 1 + I Eua TH C 0 Falcaria vulgaris + I + (+) I + 6k 2 5 P-Subm H C Galium glaucum + I + + I + 6a 2 0 Subm-Em H C Muscari comosum 1 I I I + 6k 3 4 Kt H C 0! Peucedanum alsaticum sst. (x) + I + 1 I 1 5a 3 3 Eua H C Pimpinella saxifraga + I + + I P-Subm H C! Stachys recta 1 I 1 + I + + I Pann H C 0 Thlaspi jankae (+) I + (+) I + 5k 1 4 Eua H C Veronica spicata 1 I 1 C Veronica spicata + I + 6k 2 4 P H C Ajuga laxmannii (+) I Eua G C Allium paniculatum + I Eua H C Artemisia pontica 5k 3 4 P-Pann H C! Cirsium pannonicum + 1 I + 6a 3 3 Subm H C Danthonia calycina 1 2 I Pann-Karp H C Dianthus collinus-glabriusculus + I Balk-Pann H C Dianthus pontederae + I + 6a 2 4 Subm-Em H C Dorycnium herbaceum + I Cp H C Galium boreale (+) I + 5a 1 3 Eu H C Hieracium pilosella + I + 6k 2 4 P-Pann H C Inula germanica 2 I Cp H C Koeleria cristata + I Eua-Kt H C Lavatera thuringiaca + I + 5a 4 0 Eua H C Leontodon hispidus + I +

14 180 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Appendix 2 (continued) festucetosum lithospermetosum galietosum T W R K A D K A D K A D 6a 2 5 Subm G C Muscari racemosum + I P-Subm G C Ornithogalum tenuifolium + I + 6a 2 4 P-Subm Th C Orthantha lutea + I Eua H C Plantago media + I Eua H C! Potentilla recta + I P-Subm H C! Salvia pratensis (+) I + 6k 2 4 P-Subm H C Scorzonera hispanica (+) I + 5a 1 3 Em-Subm Ch C Sedum sexangulare + I + Accompanying species 5k 3 0 Eua H-G C Euphorbia cyparissias V II + (+) + II + 5a 4 4 Eua Th-TH C Alliaria petiolata + + II II + 1 x IV + 1 5a 4 4 Eua Th C Veronica hederifolia 1 (0 1) II ( 2) (+) IV II Eua Th C Galium aparine + + I (+) III II Eua H-Ch C 0 Glechoma hederacea (et hirsuta) + I I II + 5a 3 2 Cp H C 0 Agrostis tenuis + + III I Eua G C Agropyron repens + I + + I Eua H C Alopecurus pratensis + I + 5a 4 3 Eu-Kt H C Campanula cervicaria + I + 5a 3 3 Eu H C Cruciata chersonensis + I + 5a 7 4 Eua H C Cucubalus baccifer + I G C Gagea sp. (+) I Eua H C Inula britannica + I + Subm-Em H C Stachys germanica + I Eua H C Stellaria graminea + I + 5k 5 5 Kt H C Thalictrum lucidum x I 6k 4 4 Eua H C Ajuga genevensis + I I + + I + Culta et ruderalia Cult MM A2 Robinia pseudo-acacia 1 2 I II + 3 M B1 Robinia pseudo-acacia 1 + II III I + 5a 5 3 Eu M B1 Sambucus nigra 2 1 II (+) III I 1 Cult M B1 Celtis occidentalis + I + + (+) + II + + I Eua Th-TH C Melandrium album + I III IV Eua TH C Arctium lappa III + + I 1 + I + 7k 3 4 Subm-Em H C Ballota nigra III II III + 5a 5 3 Eua TH C Conium maculatum + I + + I II + 1 6k 5 0 Eua-Kt H C Euphorbia villosa + + II Eua Th C Lamium purpureum + I II + + I Kozm Th C Stellaria media + + II + + I II Eua H C Taraxacum officinale + II + + I Eua Th C Thlaspi perfoliatum + + II + + I Eua Th-TH C Torilis japonica + + II (+) + II + 5a 6 3 Em MM A2 Acer pseudo-platanus + I + Cult MM A2 Ailanthus altissima + I + 5a 5 3 Eu M B1 Acer platanoides + I +

15 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 181 Appendix 2 (continued) festucetosum lithospermetosum galietosum T W R K A D K A D K A D Cult M A2 Morus alba (+) I + Eua Th C Aethusa cynapium + I Eua Th C Bromus sterilis + I + + (+) I Eua H C Chelidonium majus + I Kozm Th C Chenopodium album + I + (+) I Eua H C Euphorbia esula + I + + I Eua Th-TH C Lactuca serriola + I + 5a 3 3 Eua H C Linaria vulgaris + I + + I Kozm H-G C Urtica dioica 1 2 I 2 1 I 1 5a 6 4 Eu TH C Arctium minus + I + Appendix 3. Description of the relevés of the Aceri tatarici-quercetum submatricum samples 1: Valkó 1960 first second (a.s.l.) (m) Area (m 2 ) Location Date of relevés Elevation Declination ( ) Exposition Made by first second Zólyomi Fekete Zólyomi Jakucs Hoffman 2: Pusztaszentjakab (Haraszti-erdő) Fekete 3: Gödöllő-Iharos 1959 Zólyomi 4: Fót SW Zólyomi Futák Kárpáti 5: Kosd: Urasági-erdő SSW Vida 6: Dunaszentgyörgy Zólyomi 7: Érd NNE Zólyomi 8: Meleg-hegy (1) Fekete (1955) sub Querceto-Lithospermetum (1) 9: Meleg-hegy (2) Fekete (1955) sub Querceto-Lithospermetum (2)

16 182 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Appendix 3 (continued) Location Date of relevés El- Area Declination Expo- Made by first second eva- (m 2 ) ( ) sition first second tion (a.s.l.) (m) 10: Meleg-hegy (3) Fekete (1955) sub Querceto-Lithospermetum (3) 11: Meleg-hegy (4) Fekete (1955) sub Querceto-Lithospermetum (4) 1: Gödöllő-Őrház 1960 Fekete-(Zólyomi) 2: Gödöllő 1960 Fekete 3: Gödöllő-Erzsébet 1960 Zólyomi Jakucs 4: Fót NW Zólyomi Kárpáti 5: Vácrátót W Zólyomi Kárpáti 6: Kamara-erdő NNW Pócs Zólyomi 7: Kamara-erdő N Pócs Zólyomi 8: Meleg-hegy (sensu Zólyomi, non Fekete 1955) 9: Alcsút Zólyomi 10: Templom-hegy (5) Fekete (1955) sub Querceto-Lithospermetum (5) 11: Lovasberényi-erdő (6) Fekete (1955) sub Querceto-Lithospermetum (6)

17 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 183 Appendix 4. Aceri tatarici-quercetum submatricum. Abbreviations: V = Valkó, P = Pusztaszentjakab, GI = Gödöllő-Iharos, F = Fót, K = Kosd, D = Dunaszentgyörgy; E = Érd; M = Meleg-hegy, Gő = Gödöllő Őrház, G = Gödöllő,? = unknown, Fh = Fóti H, Vt = Vácrátót, Bp = Budapest, Kamara-erdő, A = Alcsút festucetosum lithospermetosum V P GI F K D E M M M M Gő G? Fh Vt Bp Bp M A M M T W R A-D K A-D K Association, local association and Aceri-Quercion species (Prunion fruticosae species) 6k 4 4 P-pann MM A2 Acer tataricum II 1(2) (+) + 1 III M B III III 6k 2 4 Kt M B Cerasus fruticosa sst II 6k 2 5 Kt M B Rosa pimpinellifolia I Quercetalia petraeae-pubescentis and Quercetea pubescentis-petraeae species ( = occurring subordinately also in Carpino-Fagetea associations, + = species transgreding from Orno-Cotinetalia) 6a 2 5 Subm MM A1 Quercus pubescens V (2) IV M B IV II 5a 3 3 Subm(or) MM A1 Quercus cerris 1 (+) III IV M B II III 5a 3 4 Eua M B Crataegus monogyna V V 6a 2 5 Subm MM A2 + Fraxinus ornus 1(2) IV IV M B IV III 5a 4 4 Balk-Em M B Euonymus verrucosa IV II 5a 3 3 Eu M B Prunus spinosa II (2) IV Eu MM A2 Pyrus achras I I M B IV III 5a 3 4 Eua M B Rhamnus cathartica III IV 6a 4 4 Subm-Em M B Viburnum lantana III IV 6a 3 5 Subm M B + Colutea arborescens III 6a 3 4 Subm-Em M B Cornus mas + + I 2 + x III 5a 4 4 Subm-Em MM A2 Sorbus torminalis II + + I M B II II 5a 3 4 Eu M B Berberis vulgaris + + (+) + II I P-pann MM A1-(2) Quercus hybr. virg. 1 (+) + 1 I II 6a 2 4 P-subm M B Rosa gallica I + (+) (+) II 6a 2 4 Subm M B + Sorbus domestica I + + I 6a 2 5 P-subm M B + Cotinus coggygria I 6a 5 3 Subm M B + Lonicera caprifolium 1(4) 1 I??? Eua M B Rosa rubiginosa + + I Querco-Fagetea (Θ = Fagetalia, Carpinio-Fagetea species) 5a 4 4 Eu MM A2 Acer campestre IV III M B IV V 5a 5 3 Eu M B Euonymus europaea IV x IV 5a 4 3 Eu M B Ligustrum vulgare V V 5a 6 0 Eu MM A1 Quercus robur III III M B II 5a 6 4 Eu MM A2 Ulmus carpinifolia III (+) 2(3) III M B III III 5a 4 4 Eu-Subm M B Cornus sanguinea II III 5a 4 0 Eu MM A1-(2) Quercus petraea 2 3 (1 2) III 1(2) (1 2) III M B I 5a 3 3 Eua M B Rosa canina s. l III + + 1(+) III

18 184 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Appendix 4 (continued) festucetosum lithospermetosum V P GI F K D E M M M M Gő G? Fh Vt Bp Bp M A M M T W R A-D K A-D K 5a 5 3 Eu MM A2 Θ Tilia cordata 1(2) I M B II 5a 5 3 Em-Subm MM A2 Θ Carpinus betulus I (+) + I 5a 5 3 Eu-Subm? M B Corylus avellana I Subm(or) MM A1 Loranthus europaeus I + + I 5a 6 4 Eu M B Malus silvestris + + I 6a 4 4 Subm(or) M B Θ Staphylea pinnata I 5a 4 4 Em-Subm M B Θ Tilia platyphyllos + + I 5a 5 3 Eua M B Θ Cerasus avium + + I 5a 4 4 M sapling Acer campestre + + I + + I 6k 4 4 M sapling Acer tataricum + + I + + I 6a 2 5 M sapling Fraxinus ornus I 1 1 I M sapling Pyrus achras + + I 5a 4 0 M sapling Quercus petraea + + I 6a 2 5 M sapling Quercus pubescens I 5a 6 0 M sapling Quercus robur + + I + + I Association, local association and Aceri-Quercion species (Prunion fruticosae species)! = occurring also in Festucetalia associations 5k 4 4 Kt H C Pulmonaria mollissima sst IV IV 6k 2 5 P-pann G Iris variegata sst + (+) III I 5a 3 4 Subm(or) H Carex michelii sst II II Pann-balk H Euphorbia polychroma II Kt H Phlomis tuberosa sst II (+) II 6k 3 4 Balk-pann G Doronicum hungaricum sst I + + I 6k 2 4 P-pann H! Inula germanica sst x + I I 6k 2 5 Kt G Lathyrus pannonicus subsp. collinus sst + + I 5k 3 4 Eua H Nepeta pannonica sst (+) (x) + I (+) + I 5k 4 5 Kt H Melica altissima sst (+) + I Eua H Veronica paniculata subsp. foliosa sst (+) + I Quercetalia petraeae-pubescentis and Quercetea pubescenti-petraeae species (+ = species transgreding from Orno-Cotinetalia, = subordinately occurring also in Carpino-Fagetea associations,! = occurring also in Festuco-Brometea associations) 5a 2 4 Eua H Brachypodium pinnatum V II Eua H Calamintha clinopodium V x IV 6a 2 4 Subm-Em Ch(H)! Teucrium chamaedrys sst V III 5a 3 4 Em H! Coronilla varia (+) IV (+) IV 6k 2 5 Eua H Dictamnus albus IV III 5a 3 4 Eua H! Galium mollugo x III IV 6k 4 4 Eua H Cynanchum vincetoxicum sst + (+) IV II 5a 3 3 Em H Lathyrus niger II x IV 6a 3 5 P-Subm H Lithospermum purpureo-coeruleum IV IV Kt G! Polygonatum odoratum sst III III 5a 3 4 Eua H Sedum maximum sst II IV 5k 3 4 Kt H Thalictrum minus sst IV III

19 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 185 Appendix 4 (continued) festucetosum lithospermetosum V P GI F K D E M M M M Gő G? Fh Vt Bp Bp M A M M T W R A-D K A-D K 5a 3 4 Eu H Trifolium alpestre II (+) + + III 6k 3 4 Eua G! Asparagus officinalis III II Eua H! Betonica officinalis sst II (+) + 1 III 5a 2 4 Em-Subm H Chrysanthemum corymbosum III II 5a 3 5 Em-Subm H Geranium sanguineum sst III II 5k 2 5 Eu-Kt Th! Melampyrum cristatum sst III I 5a 3 5 Kt H! Peucedanum cervaria sst III II 5a 3 3 Eua H Silene nutans III II 5a 3 4 Eua H! Viola hirta II II 5k 3 4 Kt-(Eua) H Anemone sylvestris sst II Eu H Astragalus glycyphyllos I II P-SUbm H Erysimum odoratum sst II I 5a 4 0 Em-Kt H Fragaria moschata sst II I 5k 4 3 Eua H Inula salicina II ( 2) + 1 II 6a 2 5 P-Subm G Iris graminea I 6a 4 3 Subm-Em H Lychnis coronaria + + I II 5a 4 5 Subm-Em H Melittis grandiflora II II 6a 3 5 Subm(or) H Oryzopsis virescens + + I II Eua H! Silene cucubalus II I Eua H Valeriana officinalis II Em H Vicia pisiformis II + (+) + I Karp-Balk H Achillea distans I 6a 3 4 Subm-Em Th-H Arabis turrita + + I + + I 6k 3 4 Kt H Campanula bononiensis + + I + + I Em N Cytisus nigricans I + + I 5a 4 3 Eu H Digitalis grandiflora + + I + + I 5a 4 3 Em H Festuca heterophylla I 5k 3 4 Kt H! Fragaria viridis sst (+) I 5a 5 4 Em H Hypericum montanum I 6k 4 4 P-pann TH Lactuca quercina + + I 2 2 I 5k 2 3 Eua H! Lithospermum officinale + + I + + I 6a 3 4 Em-Subm G Orchis purpurea + + I I Eua H Origanum vulgare + + I (+) I 5k 3 5 Em H Primula veris subsp. canescens + + I + + I 5k 4 3 Eua H Ranunculus polyanthemos sst I I Eua H Serratula tinctoria I I Subm H + Smyrnium perfoliatum + + I I 4k 4 4 Eua H Thalictrum aquilegiifolium sst + + I (+) + I Cp Th Turritis glabra + + I + + I Balk-pann Th-H Verbascum austriacum I + + I 5k 3 4 Kt H! Veronica teucrium I Eua H! Vicia cracca subsp. tenuifolia I I 5a 3 2 Eua H Viscaria vulgaris I 6a 3 4 Kt H C! Bupleurum falcatum + + I 5a 4 3 Eua H Carex montana + + I

20 186 ZÓLYOMI, B., HORVÁTH, A., KEVEY, B. and LENDVAI, G. Appendix 4 (continued) festucetosum lithospermetosum V P GI F K D E M M M M Gő G? Fh Vt Bp Bp M A M M T W R A-D K A-D K 6k 3 5 Kt H Clematis recta + + I 5k 5 4 Kt H Crepis praemorsa + + I 5a 3 3 Eu H-N Genista tinctoria subsp. elatior + + I Subm-Em H Hieracium racemosum + + I 5a 3 2 Subm-Em H Hieracium sabaudum + + I Cp H Hieracium umbellatum + + I 5a 3 4 Subm-Em H Inula conyza? 5a 4 3 Em H Potentilla alba + + I 6a 3 3 Subm-Em H Viola alba + + I Festuco-Brometea species (! = occurring also in Quercetea pubescenti-petreae associations) Kt H C Adonis vernalis V 6k 1 4 Eua H Festuca hirsuta et valesiaca (+) (1 2) V + + (+) II 6k 2 4 P-pann H! Vinca herbacea V II P-Subm H! Stachys recta IV II 5k 2 5 Eua-Kt G Carex humilis III 5k 3 3 Eua H! Filipendula vulgaris III III 6k 3 4 Eua H! Medicago falcata III + + I Cp H Poa pratensis subsp. angustifolia III II Eua H Agrimonia eupatoria II + (+) + I 5a 3 4 Em-Subm G! Anthericum ramosum II I 6k 2 5 P H Astragalus asper II 6k 3 4 P H! Cytisus austriacus II 6a 2 5 P-Subm H Melica ciliata II 6a 2 5 Subm G Muscari racemosum II I 6k 3 4 Kt H! Peucedanum alsaticum II 5k 1 4 Eua H Veronica spicata I (+) + + I 6k 2 4 Kt H Achillea pannonica I I 6k 2 4 P H Ajuga laxmanni I 6k 0 5 Subm G Allium flavum + + I + + I 5k 3 4 EuaKt H Bromus inermis I 5k 4 4 Eua-Kt H Campanula glomerata (+) + I + + I Cp H Campanula rotundifolia I 5k 3 3 Eua-Kt G Carex praecox + + I + + I Subm G Cleistogenes serotina I Pann-Balk H Dianthus pontederae I 6k 3 4 Pann-Balk H Euphorbia pannonica (+) + I (+) + I Eua TH Falcaria vulgaris + + I 1 1 I 6k 2 5 P-Subm H Galium glaucum I 5k 2 3 Kt(-Eua) H! Hieracium bauhinii I 5a 3 3 Eua H Pimpinella saxifraga + + I + + I Eu H Poa compressa I 5k 2 4 Eu-Kt H Potentilla heptaphylla + + I + + I 6k 2 4 P-Subm H Ranunculus illyricus + + I + + I P-Subm H! Salvia pratensis I + + I

21 STEPPE WOODLANDS WITH TATARIAN MAPLE ON THE HUNGARIAN PLAIN 187 Appendix 4 (continued) festucetosum lithospermetosum V P GI F K D E M M M M Gő G? Fh Vt Bp Bp M A M M T W R A-D K A-D K Eua-Subm Th-H Sideritis montana + + I (+) + I 6k 2 4 Kt-Eu H Verbascum phoeniceum + + I + + I Eua G Allium paniculatum + + I Eua H Botriochloa ischaemum + + I Cp Th-H! Arabis hirsuta sst + + I Eua H Artemisia pontica 1 1 I 6a 3 4 P-Subm H! Aster linosyris + + I 6k 2 4 P-pann H Carduus hamulosus I 6k 3 4 Eua-Kt G Carex liparicarpos I 5a 2 4 Eua H Carlina vulgaris subsp. intermedia + + I 5a 6 0 Em H Centaurea pannonica + + I 6k 2 4 Pann H Centaurea sadleriana 1 1 I 6a 2 4 Cucubalus baccifer II I Alp-Balk N Dorycnium germanicum + + I 6k 1 4 P-Subm H Eryngium campestre + + I 6k 1 4 Kt H Euphorbia seguieriana + + I 5k 3 4 Eua-Kt H Euphorbia virgata + + I Eua H Galium mollugo verum (pomeranicum) + + I Eua-Kt H Galium verum + + I P-pann H! Inula ensifolia 1 1 I 6k 1 4 Kt H! Linaria genistifolia + + I 6k 1 5 P-Subm H Melica transsilvanica + + I 6k 2 4 Eua-Kt H Onobrychis arenaria + + I 6k 2 5 P H Onosma arenarium + + I 6k 2 4 Eua H Plantago media + + I Cp H Potentilla argentea + + I 6a 1 5 Eua H Potentilla recta + + I P H Salvia austriaca + + I 6a 2 4 Eua-Kt H Salvia nemorosa + + I 6k 2 4 Kt H Scabiosa ochroleuca + + I 6k 2 4 P-Subm Scorzonera hispanica + + I 6k 2 4 Kt? H Serratula radiata I 6k 2 5 P-pann Taraxacum serotinum + + I Eua TH Thlaspi perfoliatum + + I Kt Ch Thymus glabrescens + + I 6k 1 4 Em H Veronica austriaca subsp. dentata + + I 6k 2 4 Kt Th Veronica prostrata + + I Querco-Fagetea (0 = Fagetalia-Carpino-Fagetea species) 5a 5 4 Eua H C Brachypodium sylvaticum II x 1(3) V Cp H Geum urbanum II V 5a 4 4 Eua H(-Ch) Veronica chamaedrys II x V 5a 6 4 Eua H Dactylis glomerata IV 1(2) (2) IV