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The Balkan wet grassland vegetation: a prerequisite to better understanding of European habitat diversity

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Abstract

The knowledge of broad-scale floristic variation in wet grasslands, which are endangered throughout Europe, is still limited and some regions have remained unexplored so far. In addition, hitherto published phytosociological studies were concentrated at the national level and therefore national vegetation classifications are not consistent with each other. In order to overcome these shortcomings of traditional phytosociology, we gathered original data from Bulgaria and analysed them together with the data from Central Europe. We further analysed major compositional gradients within Bulgarian wet grasslands and changes in species richness along them. We sampled 164 wet grassland vegetation plots throughout Bulgaria. We further prepared a restricted data set of wet grasslands from Central-European phytosociological databases. Both data sets were merged and classified by modified TWINSPAN. Four distinct vegetation types were differentiated. Even if they correspond with traditional alliances, which are primarily drawn as geographically defined units in Western and Central Europe (sub-Mediterranean Trifolion resupinati, sub-continental Deschampsion cespitosae and Molinion caeruleae and sub-oceanic Calthion palustris), they all occur in Bulgaria. When more precise classification was applied, two types of sub-Mediterranean wet grasslands and one high-altitude type of Calthion grasslands were detected solely in Bulgaria. DCA analysis showed that altitude is a dominant gradient controlling variation in Balkan wet grasslands. The second DCA axis was interpreted as the gradient of nutrient availability. Species richness shows skewed-unimodal trends along both major gradients, with the highest species richness in intermittently wet nutrient-limited grasslands. Tukey post-hoc test of altitudinal differences amongst vegetation types is significant for all pairs of clusters, suggesting that altitudinal differentiation is responsible for co-occurrence of nearly all European types of wet grasslands in Bulgaria. Our results suggest that (1) climate is an important factor for the diversity of wet grasslands; (2) Balkan vegetation of middle altitudes matches with that of Central Europe, whereas that of the lowest altitudes corresponds rather to the sub-Mediterranean region and high mountains are specific; (3) upward shift of Central-European vegetation types in southern Europe, so often described in forest vegetation is also evident for grassland vegetation and (4) the high diversity of Balkan vegetation is determined by a diverse relief enabling confluence of habitats possessing different climatic conditions.

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References

  • Aarssen LW, Schamp BS (2002) Predicting distributions of species richness and species size in regional floras: applying the species pool hypothesis to the habitat templet model. Perspect Plant Ecol Evol Syst 5:3–12

    Article  Google Scholar 

  • Al-Mufti MM, Sydes CL, Furness SB, Grime JP, Band SR (1977) A quantitative analysis of shoot phenology and dominance in herbaceous vegetation. J Ecol 65:759–791

    Article  Google Scholar 

  • Andreev N, Anchev M, Kozhuharov S, Markova M, Peev D, Petrova A (1992) Opredelitel na visshite rastenia v Bulgaria [Field guide to the vascular plants in Bulgaria]. Naouka and Izkoustvo, Sofia (in Bulgarian)

    Google Scholar 

  • Archibold OW (1995) Ecology of world vegetation. Chapman and Hall

  • Bachelet D, Neilson RP, Lenihan JM, Drapek RJ (2001) Climate change effects on vegetation distribution and carbon budget in the United States. Ecosystems 4:164–185

    Article  CAS  Google Scholar 

  • Bedford BL, Walbridge MR, Aldus A (1999) Patterns in nutrient availability and plant diversity of temperate North American wetlands. Ecology 80:2151–2169

    Article  Google Scholar 

  • Blackstock TH, Stevens DP, Stevens PA, Mockridge CP, Yeo MJM (1998) Edaphic relationships among Cirsio-Molinietum and related wet grassland communities in lowland Wales. J Veg Sci 9:431–444

    Article  Google Scholar 

  • Boeye D, Verhagen B, Van Haesebroeck V, Verheyen RF (1997) Nutrient limitation in species-rich lowland fens. J Veg Sci 8:415–424

    Article  Google Scholar 

  • Botta-Dukát Z, Chytrý M, Hájková P, Havlová M (2005) Vegetation of lowland wet meadows along a climatic continentality gradient in Central Europe. Preslia 77:89–111

    Google Scholar 

  • Boyer MLH, Wheeler BD (1989) Vegetation patterns in spring-fed calcareous fens: calcite precipitation and constrains on fertility. J Ecol 77:597–609

    Article  CAS  Google Scholar 

  • Bruelheide H, Chytrý M (2000) Towards unification of national vegetation classifications: a comparison of two methods for analysis of large data sets. J Veg Sci 11:295–306

    Article  Google Scholar 

  • Calef MP, McGuire AD, Epstein HE, Rupp TS, Shugart HH (2005) Analysis of vegetation distribution in Interior Alaska and sensitivity to climate change using a logistic regression approach. J Biogeogr 32:863–878

    Article  Google Scholar 

  • Chytrý M, Otýpková Z (2003) Plot sizes used for phytosociological sampling of European vegetation. J Veg Sci 14:563–570

    Article  Google Scholar 

  • Chytrý M, Rafajová M (2003) Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. Preslia 75:1–15

    Google Scholar 

  • Chytrý M, Danihelka J, Ermakov N, Hájek M, Hájková P, Kočí M, Kubešová S, Lustyk P, Otýpková Z, Popov D, Roleček J, Řezníčková M, Šmarda P, Valachovič M (2007) Plant species richness in continental southern Siberia: effects of pH and climate in the context of the species pool hypothesis. Glob Ecol Biogeogr Lett (in press)

  • Duarte MC, Rego F, Moreira I (2005) Distribution patterns of plant communities on Santiago Island, Cape Verde. J Veg Sci 16:283–292

    Article  Google Scholar 

  • Dwire KA, Kauffman JB, Brookshire ENJ, Baham J (2004) Plant biomass and species composition along an environmental gradient in montane riparian meadows. Oecologia 139:309–317

    Article  PubMed  Google Scholar 

  • Ejrnæs R, Bruun HH (2000) Gradient analysis of dry grassland in Denmark. J Veg Sci 11:573–584

    Article  Google Scholar 

  • Ewald J (2003a) A critique for phytosociology. J Veg Sci 14:291–296

    Article  Google Scholar 

  • Ewald J (2003b) The calcareous riddle: why are there so many calciphilous species in the Central European flora? Folia Geobot 38:357–366

    Article  Google Scholar 

  • Grootjans AP, Hunneman H, Verkiel H, Van Andel J (2005) Long-term effects of drainage on species richness of a fen meadow at different spatial scales. Basic Appl Ecol 6:185–193

    Article  Google Scholar 

  • Hájek M, Hájková P (2004) Environmental determinants of variation in Czech Calthion wet meadows: a synthesis of phytosociological data. Phytocoenologia 34:33–54

    Article  Google Scholar 

  • Hajek M, Hajkova P, Apostolova I (2005) Notes on the Bulgarian wetland flora, including new national and regional records. Phytol Balcan 11:173–184

    Google Scholar 

  • Hájek M, Horsák M, Hájková P, Dítě D (2006) Habitat diversity of central European fens in relation to environmental gradients and an effort to standardise fen terminology in ecological studies. Perspect Plant Ecol, Evol Syst. 8:97–114

    Google Scholar 

  • Hájková P, Hájek M (2007) Sphagnum distribution patterns along environmental gradients in Bulgaria. J Bryol 29:18–26

    Article  Google Scholar 

  • Hájková P, Hájek M, Apostolova I (2006) Diversity of wetland vegetation in the Bulgarian high mountains, main gradients and context-dependence of the pH role. Plant Ecol 184:111–130

    Article  Google Scholar 

  • Härdtle W, Redecker B, Assmann T, Meyer H (2006) Vegetation responses to environmental conditions in floodplain grasslands: prerequisites for preserving plant species diversity. Basic Appl Ecol 7:280–288

    Article  Google Scholar 

  • Havlová M (2006) Syntaxonomical revision of the Molinion meadows in the Czech Republic. Preslia 78:87–101

    Google Scholar 

  • Havlová M, Chytrý M, Tichý L (2004) Diversity of hay meadows in the Czech Republic: major types and environmental gradients. Phytocoenologia 34:551–567

    Article  Google Scholar 

  • Hill MO (1979) TWINSPAN. A FORTRAN program for arranging multivariate data in an ordered two-way table by classification of individuals and attributes. Ecology and Systematics, Cornell University, Ithaca, NY, US

    Google Scholar 

  • Hölzel N (2005) Seedling recruitment in flood-meadow species: the effects of gaps, litter and vegetation matrix. Appl Veg Sci 8:115–124

    Article  Google Scholar 

  • Horvat I, Glavač V, Ellenberg H (1974) Vegetation Südosteuropas. Gustav Fischer Verlag. Stuttgart

    Google Scholar 

  • Hubbell SP (2005) Neutral theory in community ecology and the hypothesis of functional equivalence. Funct Ecol 19:166–172

    Article  Google Scholar 

  • Jensen K, Meyer C (2001) Effects of light competition and litter on the performance of Viola palustris and on species composition and diversity of an abandoned fen meadow. Plant Ecol 155:169–181

    Article  Google Scholar 

  • Jensen K, Schrautzer J (1999) Consequences of abandonment for a regional fen flora and mechanisms of successional change. Appl Veg Sci 2:79–88

    Article  Google Scholar 

  • Knollová I, Chytrý M, Tichý L, Hájek O (2005) Stratified resampling of phytosociological databases: some strategies for obtaining more representative data sets for classification studies. J Veg Sci 16:479–486

    Article  Google Scholar 

  • Koch W (1926) Die Vegetationseinheiten der Linthebene unter Berücksichtigung der Verhältnisse in der Nordostschweiz. Systematisch-kritische Studie. Jahresber St Gallischen Naturwiss Ges 61/2:1–144

    Google Scholar 

  • Koleff P, Gaston KJ, Lennon JJ (2003) Measuring beta diversity for presence–absence data. J Anim Ecol 72:367–382

    Article  Google Scholar 

  • Kovacs-Lang E, Kroel-Dulay G, Kertesz M, Fekete G, Bartha S, Mika J, Dobi-Wantuch I, Redei T, Rajkai K, Hahn I (2000) Changes in the composition of sand grasslands along a climatic gradient in Hungary and implications for climate change. Phytocoenologia 30:385–407

    Google Scholar 

  • Mateeva Z (2002) Physical geographical factors. In: Kopralev I (ed) Geography of Bulgaria. Publ. House ForCom (in Bulgarian), pp 142–144

  • Melillo JM (1999) Warm, warm on the range. Science 283:183–184

    Article  CAS  Google Scholar 

  • Meshinev T, Apostolova I, Koleva E (2000) Influence of warming on timberline rising: a case study on Pinus peuce Griseb. in Bulgaria. Phytocoenologia 30:431–438

    Google Scholar 

  • Meshinev T, Apostolova I, Georgiev V, Dimitrov V, Petrova A, Veen P (2005) Grasslands of Bulgaria. Final report on the National Grasslands Inventory Project – Bulgaria, 2001–2004. Dragon 2003 Ltd. Publishers, Sofia

  • Micevski K (1964) Tipoloshki istrazhuvana na vegetacijata na nizinskite livadi vo Makedonija. [Typologische Untersuchungen der Vegetation der Niederungswiesen Mazedoniens]. Annuaire de la Faculté des Sciences de l’Université de Skopje 15/3:121–173 (in Macedonian)

    Google Scholar 

  • Mucina L, Rodwell JS, Schaminée JHJ, Dierschke H (1993) European Vegetation Survey: current state of some national programmes. J Veg Sci 4:429–439

    Article  Google Scholar 

  • Natcheva R, Ganeva A (2005) Check-list of the bryophytes of Bulgaria with data on their distribution. II. Musci. Cryptog Bryol 26:209–232

    Google Scholar 

  • Nekola JC (2004) Vascular plant compositional gradients within and between Iowa fens. J Veg Sci 15:771–780

    Article  Google Scholar 

  • Olde Venterink H, van der Vliet RE, Wassen MJ (2001) Nutrient limitation along a productivity gradient in wet meadows. Plant Soil 234:171–179

    Article  CAS  Google Scholar 

  • Pärtel M (2002) Local plant diversity patterns and evolutionary history at the regional scale. Ecology 83:2361–2366

    Google Scholar 

  • Peintinger M, Bergamini A (2006) Community structure and diversity of bryophytes and vascular plants in abandoned fen meadows. Plant Ecol 185:1–17

    Article  Google Scholar 

  • Petrů M (2005) Year-to-year oscillations in demography of the strictly biennial Pedicularis sylvatica and effects of experimental disturbances. Plant Ecol 181:289–298

    Article  Google Scholar 

  • Pither J, Aarssen LW (2005) The evolutionary species pool hypothesis and patterns of freshwater diatom diversity along a pH gradient. J Biogeogr 32:503–513

    Article  Google Scholar 

  • Prach K (1993) Vegetational changes in a wet meadow complex, South Bohemia, Czech Republic. Folia Geobot Phytotax 28:1–13

    Google Scholar 

  • Raus T (1983) Wechselnasse Wiesen in Griechenland. Tuexenia 3:259–270

    Google Scholar 

  • Rodwell J, Schaminée J, Mucina L, Pignatti S, Dring J, Moos D (2002) The diversity of European vegetation. An overview of phytosociological alliances and their relationships to EUNIS habitats. Wageningen, NL. Report EC-LNV nr. 2002/054

  • Schaffers AP (2002) Soil, biomass, and management of semi-natural vegetation Part I. Interrelationships. Plant Ecol 158:229–246

    Article  Google Scholar 

  • Schrautzer J, Asshoff M, Müller F (1996) Restoration strategies for wet grasslands in Northern Germany. Ecol Eng 7:255–278

    Article  Google Scholar 

  • Tallis JH (1991) Plant community history. Chapman and Hall, London

    Google Scholar 

  • Tichý L (2002) JUICE, software for vegetation classification. J Veg Sci 13:451–453

    Article  Google Scholar 

  • Tichý L (2005) New similarity indices for the assignment of relevés to the vegetation units of an existing phytosociological classification. Plant Ecol 179:67–72

    Article  Google Scholar 

  • Tichý L, Chytrý M (2006) Statistical determination of diagnostic species for site groups of unequal size. J Veg Sci 17:809–818

    Article  Google Scholar 

  • Tzonev R, Dimitrov M, Chytrý M, Roussakova V, Dimova D, Gussev Ch, Pavlov D, Vulchev V, Vitkova A, Gogoushev G, Nikolov I, Borisova D, Ganeva A (2006) Beech forest communities in Bulgaria. Phytocoenologia 36:247–279

    Article  Google Scholar 

  • Tüxen R (1937) Die Pflanzengesellschaften Nordwestdeutschlands. Mitt Florist-Soziol Arbeitsgem Niedersachsen 3:1–170

    Google Scholar 

  • van der Hoek D, Sýkora KV (2006) Fen-meadow succession in relation to spatial and temporal differences in hydrological and soil conditions. Appl Veg Sci 9:185–194

    Article  Google Scholar 

  • Virtanen R, Oksanen J, Oksanen L, Razzhivin VY (2006) Broad-scale vegetation-environment relationships in Eurasian high-latitude areas. J Veg Sci 17:519–528

    Article  Google Scholar 

  • Wheeler BD, Shaw SC (1991) Above-ground crop mass and species richness of the principal types of herbaceous rich-fen vegetation of lowland England and Wales. J Ecol 79:285–301

    Article  Google Scholar 

  • Wohlgemuth T, Schutz M, Keller W, Wildi O (1999) Computed ecograms of Weiss forests. Bot Helv 109:169–191

    Google Scholar 

Download references

Acknowledgements

We thank Hristo Pedashenko and Marcela Havlová for providing us unpublished relevés from Bulgaria. We thank Boryana Mihova for the English revision of the manuscript. Thanks are extended to Erwin Bergmeier and two anonymous reviewers, whose comments improved the manuscript. The research was performed within the long-term research plans of Masaryk University, Brno (No. MSM0021622416) and of Botanical Institute of Czech Academy of Sciences (No. AVZ0Z60050516). Field investigation was partly carried out within the exchange project of the Czech and Bulgarian Academies of Sciences (Vegetation diversity of the lowland wet meadows in Bulgaria, 2005–2007).

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Correspondence to Michal Hájek.

Appendices

Appendix 1

Shortened synoptic table of species percentage occurrence in seven major vegetation types of wet grasslands resulting from the Twinspan classification of the merged data set. Diagnostic species of individual vegetation types were determined using the phi-coefficient of association (* Φ > 0.35, ** Φ > 0.45). Species, the occurrence concentration probability of which, in the given vegetation type, does not differ from random at P < 0.05 are excluded from the list of diagnostic species. Only species with Φ > 0.35 or with percentage constancy >50% at least in one column are presented. Abbreviation “dif.” indicates the species having clear optimum outside wet grassland vegetation. The diagnostic (i.e. differentiate) value of such a species is valid only within our data set. Classified data sets contained 164 vegetation plots from Bulgaria and 164 vegetation plots from Central Europe.

No of cluster

1

2

3

4

5

6

7

No of relevés

21

22

70

81

46

29

59

Trifolion resupinati (cluster 1: salt-rich, lowest altitudes)

Hordeum secalinum

81**

14

.

.

.

.

.

Carex divisa

38**

.

3

.

.

.

.

Cichorium intybus

52**

27

.

1

.

.

.

Bromus secalinus

43**

23

.

.

.

.

.

Elymus repens

48**

.

23

1

2

7

.

Alopecurus myosuroides

33*

14

.

.

.

.

.

Lolium perenne

52*

45

6

1

.

.

.

Scorzonera laciniata

19*

.

.

.

.

.

.

Centaurea calcitrapa

19*

.

.

.

.

.

.

Trifolium resupinatum

19*

.

.

.

.

.

.

Geranium dissectum

19*

.

.

.

.

.

.

Trifolium michelianum

14*

.

.

.

.

.

.

Hordeum bulbosum

14*

.

.

.

.

.

.

Trifolion resupinati (cluster 2)

Ranunculus sardous

38

95**

1

.

.

.

.

Orchis laxiflora agg.

5

59**

10

5

.

3

3

Trifolium patens

5

59**

7

4

.

7

5

Galium debile

.

32**

.

.

.

.

.

Carex distans

29

59**

9

15

.

3

.

Gratiola officinalis

.

41**

14

5

.

3

.

Alopecurus rendlei

5

27*

.

.

.

.

.

Moenchia mantica

10

32*

.

1

.

.

.

Plantago lanceolata

43

86*

43

46

9

17

7

Rhinanthus angustifolius

.

18*

.

.

.

.

.

Euphrasia stricta

.

18*

.

.

.

.

.

Chrysopogon gryllus (dif.)

.

23*

.

6

.

.

.

Amblystegium riparium

.

18*

.

.

.

.

2

Rhinanthus rumelicus

.

36*

9

7

.

10

3

Ononis arvensis

14

41*

9

17

.

.

2

Cynosurus cristatus

5

59*

11

15

4

31

31

Mentha spicata

10

23*

.

.

.

.

.

Deschampsion (cluster 3)

Alopecurus pratensis

14

.

84**

12

54

24

5

Taraxacum sect Ruderalia

24

9

69**

17

2

24

8

Carex praecox

5

.

29*

1

.

.

.

Glechoma hederacea s. lat.

.

.

33*

4

.

14

.

Poa pratensis agg.

14

14

70*

42

39

31

3

Molinion (cluster 4)

Molinia caerulea s. lat.

.

.

.

80**

13

7

14

Serratula tinctoria

.

.

16

49**

2

.

3

Betonica officinalis

.

5

6

42**

2

3

3

Linum catharticum

.

.

.

31**

2

3

2

Centaurea jacea agg.

.

.

29

48**

4

.

3

Leontodon hispidus

.

5

10

36*

.

.

2

Succisa pratensis

.

.

6

49*

15

.

22

Carex flacca

.

.

1

28*

2

.

3

Filipendula vulgaris

.

18

9

41*

2

.

3

Sanguisorba officinalis

.

.

44

63*

30

10

7

Galium boreale

.

.

19

35*

7

.

.

Trifolium montanum

.

.

.

15*

.

.

.

Carex panicea

.

5

21

67*

41

38

24

Calthion (cluster 5: Central-European waterlogged Cirsium palustre grasslands)

Cirsium palustre

.

.

1

4

46**

3

.

Angelica sylvestris

.

.

6

5

48**

3

.

Myosotis palustris s. lat.

.

.

9

10

74**

21

51

Cardamine pratensis agg.

.

.

29

4

43*

3

.

Rhytidiadelphus squarrosus

.

.

.

2

24*

3

.

Cirsium rivulare

.

.

3

12

37*

10

2

Dactylorhiza majalis s. lat.

.

.

.

2

20*

.

.

Chaerophyllum hirsutum

.

.

.

.

26*

3

8

Calthion (cluster 6: sub-montane waterlogged grasslands on alkaline soils)

Mentha longifolia

.

.

3

.

11

55**

22

Hypericum tetrapterum

.

.

1

.

2

34**

7

Lythrum salicaria

.

.

19

25

11

55*

10

Juncus inflexus

.

9

6

15

4

41*

7

Ranunculus repens

5

9

61

20

43

79*

41

Plagiomnium undulatum

.

.

.

1

7

24*

3

Calthion (cluster 7: Balkan high-mountain waterlogged grasslands)

Geum coccineum

.

.

.

.

.

3

61**

Luzula sudetica

.

.

.

.

.

.

49**

Epilobium palustre

.

.

.

.

20

10

53**

Juncus thomasii

.

.

1

.

2

10

41**

Trifolium spadiceum

.

.

.

.

.

.

25**

Veratrum album s. lat.

.

.

.

5

11

.

39**

Parnassia palustris

.

.

.

4

4

.

32**

Dactylorhiza cordigera

.

.

.

.

2

10

36**

Carex echinata

.

.

.

2

11

24

49**

Hieracium caespitosum

.

.

.

2

.

21

41*

Galium palustre s. lat.

10

.

19

11

35

62

78*

Agrostis canina

.

9

.

9

22

24

53*

Potentilla erecta

.

5

.

47

41

41

73*

Oenanthe fistulosa

5

5

1

1

.

17

37*

Aulacomnium palustre

.

.

.

1

4

.

22*

Nardus stricta

.

.

.

15

17

.

37*

Alchemilla vulgaris agg.

.

.

10

4

46

28

58*

Carex nigra agg.

.

.

3

10

52

14

54*

Myosotis sicula

.

5

.

.

.

14

29*

Eriophorum latifolium

.

5

.

5

4

.

25*

Carex canescens

.

.

.

.

7

.

20*

Species diagnostic for two clusters

Poa sylvicola

95**

73*

11

.

.

28

14

Scirpus sylvaticus

.

.

4

5

80*

86*

63

Other frequent species

Ranunculus acris

5

.

66

72

72

62

69

Deschampsia caespitosa

5

9

46

65

46

34

83

Festuca pratensis agg.

33

82

63

36

41

62

19

Holcus lanatus

.

86

39

49

33

69

37

Festuca rubra agg.

.

5

29

52

57

38

71

Rumex acetosa

.

23

51

35

59

59

37

Anthoxanthum odoratum s. l.

.

73

34

36

37

45

39

Lathyrus pratensis

.

.

43

41

48

41

42

Lychnis flos-cuculi

.

.

61

23

57

62

17

Lysimachia nummularia

.

41

57

25

22

66

19

Prunella vulgaris

.

50

27

44

17

45

34

Achillea millefolium agg.

14

23

50

49

24

28

7

Poa trivialis

10

5

50

7

46

52

39

Cirsium canum

14

9

53

48

11

31

5

Trifolium pratense

38

73

37

22

13

38

19

Juncus effusus

.

23

11

6

43

55

68

Calliergonella cuspidata

.

32

11

16

22

59

63

Filipendula ulmaria

.

.

16

19

61

45

31

Caltha palustris

.

.

6

10

57

38

54

Plagiomnium affine agg.

.

.

1

11

39

52

54

Equisetum arvense

.

9

17

19

20

52

31

Appendix 2

Shortened synoptic table of species percentage occurrence in six major Bulgarian vegetation types of wet grasslands resulting from the Twinspan classification of the merged data set from Bulgaria and Central Europe. Diagnostic species of individual vegetation types were determined using the phi-coefficient of association (* Φ > 0.35, ** Φ > 0.45). Species, the occurrence concentration probability of which, in the given vegetation type, does not differ from random at P < 0.001 are excluded from the list of diagnostic species. Only species with Φ > 0.35 or with percentage constancy >50% at least in one column are presented. Cluster numbers are the same as in Appendix 1 and Fig. 1. Abbreviation “dif.” indicates the species having clear optimum outside wet grassland vegetation. The diagnostic (i.e. differentiate) value of such a species is valid only within our data set.

No. of cluster

1

2

3

4

6

7

No. of relevés

22

18

22

22

29

51

Trifolion resupinati (cluster 1: salt-rich, lowest altitudes)

Bromus secalinus

64**

.

.

.

.

.

Hordeum secalinum

77**

6

9

.

.

.

Cichorium intybus

68**

11

.

5

.

.

Lolium perenne

68**

28

18

5

.

.

Crepis setosa

50**

22

.

.

.

.

Carex divisa

36**

.

9

.

.

.

Plantago major

45**

6

9

.

7

2

Poa sylvicola

95*

61

55

5

31

12

Centaurea calcitrapa

18 *

.

.

.

.

.

Geranium dissectum

18 *

.

.

.

.

.

Scorzonera laciniata

18 *

.

.

.

.

.

Alopecurus myosuroides

32 *

17

.

.

.

.

Elymus repens

36 *

6

14

.

7

.

Juncus gerardii

36 *

28

.

.

.

.

Trifolion resupinati (cluster 2)

Ranunculus sardous

55

94**

.

.

.

.

Cirsium arvense (dif.)

5

39**

5

.

.

.

Orchis laxiflora s.l.

5

67**

32

14

14

.

Euphrasia stricta

.

22*

.

.

.

.

Sanguisorba minor

.

22*

.

.

.

.

Rhinanthus angustifolius

.

22*

.

.

.

.

Amblystegium riparium

.

22*

.

.

.

2

Galium debile

9

28*

.

.

.

.

Trifolium patens

18

56*

23

5

10

4

Gratiola officinalis

5

44*

14

9

10

.

Ononis arvensis

14

56*

14

32

3

.

Chrysopogon gryllus (dif.)

.

33*

.

18

.

.

Ophioglossum vulgatum

.

22*

.

5

.

.

Festuca nigrescens

.

22*

.

5

.

.

Deschampsion (cluster 3)

Taraxacum sect Ruderalia

.

6

73**

14

10

8

Alopecurus pratensis

.

.

68**

9

21

6

Carex otrubae

18

11

59**

.

.

.

Carex spicata

.

.

36**

.

3

.

Cirsium canum

.

6

86**

64

38

.

Bromus commutatus

.

.

36**

.

7

2

Juncus compressus

.

11

36**

.

.

.

Carex melanostachya

.

.

18*

.

.

.

Potentilla reptans

41

33

73*

5

31

2

Poa trivialis

.

.

68*

9

48

41

Molinion (cluster 4)

Serratula tinctoria

.

.

5

73**

.

.

Filipendula vulgaris

.

28

14

86**

3

.

Stachys officinalis

.

6

9

68**

3

.

Sanguisorba officinalis

.

.

18

73**

3

4

Molinia caerulea s.l.

.

.

.

68**

14

8

Carex caryophyllea

.

.

.

32**

.

.

Iris sibirica

.

.

.

27**

.

.

Luzula multiflora

.

.

.

27**

.

2

Luzula campestris

.

.

5

32*

.

4

Danthonia alpina

.

.

9

32*

.

.

Viola jordanii

.

.

.

18*

.

.

Gentiana pneumonanthe

.

.

.

18*

.

.

Rhinanthus wagneri

.

.

.

18*

.

.

Potentilla erecta

.

6

.

77*

45

71

Bistorta major

.

.

.

41*

7

22

Carex panicea

.

.

50

73*

41

22

Festuca rubra

.

6

32

82*

48

71

Centaurea jacea

.

.

9

27*

.

4

Juncus conglomeratus

.

.

23

45*

10

16

Calthion (cluster 6: sub-montane waterlogged grasslands on alkaline soils)

Hypericum tetrapterum

.

.

.

.

48**

2

Lythrum salicaria

.

.

23

9

69**

6

Mentha longifolia

.

.

5

.

59**

20

Ranunculus repens

.

6

55

5

90**

37

Lycopus europaeus

.

.

.

.

31**

4

Epilobium parviflorum

.

.

.

.

24**

.

Lysimachia vulgaris

.

.

9

14

45*

10

Plagiomnium affine agg.

.

.

.

14

59*

53

Calthion (cluster 7: Balkan high-mountain waterlogged grasslands)

Luzula sudetica

.

.

.

.

.

57**

Geum coccineum

.

.

.

.

14

65**

Carex nigra

.

.

.

.

14

59**

Epilobium palustre

.

.

.

.

17

57**

Myosotis nemorosa s.l.

.

.

.

.

21

57**

Caltha palustris

.

.

.

.

34

59**

Alchemilla vulgaris agg.

.

.

.

14

28

59**

Parnassia palustrs

.

.

.

.

7

33**

Crepis paludosa

.

.

.

.

7

33**

Carex rostrata

.

.

.

.

3

29**

Carex curta

.

.

.

.

.

24**

Trifolium spadiceum

.

.

.

.

3

27**

Juncus tomasii

.

.

5

9

14

43*

Carex echinata

.

.

.

5

38

51*

Veratrum lobelianum

.

.

.

18

7

41*

Eriophorum angustifolium

.

.

.

.

.

20*

Dactylorhiza cordigera

.

.

.

.

21

37*

Agrostis canina

5

6

.

18

28

55*

Climacium dendroides

.

.

.

9

38

47*

Cardamine rivularis

.

.

.

.

.

16*

Aulacomnium palustre

.

.

.

9

.

24*

Species diagnostic for two clusters (Calthion)

Scirpus sylvaticus

.

.

9

5

72**

69*

Galium palustre

.

.

27

14

76*

80*

Other frequent species

Deschampsia caespitosa

.

6

59

82

48

82

Holcus lanatus

18

78

55

82

72

31

Ranunculus acris

.

.

64

77

66

67

Anthoxanthum odoratum

14

78

59

68

45

35

Festuca pratensis

41

78

64

41

59

16

Calliergonella cuspidata

.

39

27

41

55

59

Juncus effusus

.

28

23

14

59

67

Trifolium pratense

50

67

45

41

38

16

Rumex acetosa

9

22

36

55

59

33

Plantago lanceolata

55

78

73

41

17

6

Cynosurus cristatus

14

56

45

55

34

27

Lysimachia nummularia

5

39

59

36

55

14

Carex hirta

14

44

59

32

34

18

Stellaria graminea

.

.

36

64

31

37

Equisetum arvense

.

11

27

23

59

25

Lychnis flos-cuculi

.

.

59

23

59

14

Carex pallescens

.

.

23

55

21

33

Agrostis stolonifera

.

.

55

45

38

6

Carex distans

27

61

36

23

7

.

Appendix 3

Species scores along the first DCA axis. Only species with the weight >5 in the analysis are presented.

Bromus secalinus

−0.8955

Hordeum secalinum

−0.5779

Crepis setosa

−0.3426

Carex divisa

−0.1794

Cichorium intybus

−0.1595

Alopecurus myosuroides

−0.1431

Vicia grandiflora

0.1025

Elymus repens

0.1780

Juncus gerardii

0.2227

Oenanthe silaifolia

0.2253

Lolium perenne

0.2733

Alopecurus rendlei

0.6686

Mentha spicata

0.7316

Convolvulus arvensis

0.8175

Poa sylvicola

0.8214

Moenchia mantica

0.8330

Ranunculus sardous

0.9300

Plantago major

0.9430

Daucus carota

0.9980

Cirsium arvense

1.2299

Rumex crispus

1.2796

Medicago lupulina

1.4163

Carex distans

1.5629

Lotus corniculatus

1.5642

Trifolium patens

1.5970

Trifolium repens

1.6102

Potentilla reptans

1.6704

Ononis arvensis

1.7301

Chrysopogon gryllus

1.7438

Plantago lanceolata

1.7890

Carex otrubae

1.7894

Trifolium pratense

1.9264

Festuca pratensis

1.9311

Galium verum

2.0918

Orchis laxiflora

2.0929

Dactylis glomerata

2.0938

Rhinanthus rumelicus

2.2552

Gratiola officinalis

2.2618

Juncus compressus

2.3433

Trifolium campestre

2.3920

Achillea millefolium

2.4135

Carex spicata

2.4149

Carex tomentosa

2.4543

Eleocharis palustris

2.4850

Festuca valesiaca

2.5343

Festuca arundinacea

2.5627

Leucanthemum vulgare

2.6597

Alopecurus pratensis

2.7119

Filipendula vulgaris

2.7377

Anthoxanthum odoratum

2.7386

Juncus articulatus

2.7387

Bromus commutatus

2.7697

Cynosurus cristatus

2.7786

Carex hirta

2.7921

Holcus lanatus

2.8180

Ranunculus polyanthemus

2.8408

Poa pratensis

2.8414

Eleocharis uniglumis

2.8532

Taraxacum sect. Ruderalia

2.8603

Cirsium canum

2.8776

Rhinanthus minor

2.8804

Leontodon hispidus

2.8877

Phleum pratense

2.9338

Sieglingia decumbens

3.0523

Agrostis stolonifera

3.0563

Stachys officinalis

3.0649

Lysimachia nummularia

3.0769

Drepanocladus aduncus

3.1030

Centaurea jacea

3.1626

Trifolium hybridum

3.1779

Leontodon autumnalis

3.2043

Juncus atratus

3.2093

Prunella vulgaris

3.2340

Serratula tinctoria

3.2415

Cerastium holosteoides

3.2438

Bromus racemosus

3.2758

Carex acuta

3.3125

Danthonia alpina

3.4155

Centaurea phrygia

3.4169

Rumex acetosa

3.4844

Sanguisorba officinalis

3.5560

Juncus inflexus

3.5592

Agrostis capillaris

3.6201

Lychnis flos-cuculi

3.6825

Carex panicea

3.7254

Briza media

3.8039

Luzula campestris

3.8283

Myosotis caespitosa

3.8519

Molinia caerulea

3.9568

Ranunculus repens

3.9577

Stellaria graminea

3.9830

Poa trivialis

3.9941

Calliergonella cuspidata

4.0122

Lythrum salicaria

4.0160

Carex ovalis

4.0278

Equisetum arvense

4.0354

Oenanthe banatica

4.0582

Cardamine matthioli

4.0720

Ranunculus acris

4.0936

Juncus conglomeratus

4.1269

Brachythecium rutabulum

4.1680

Deschampsia caespitosa

4.2354

Juncus effusus

4.2399

Vicia cracca

4.2577

Mentha arvensis

4.2923

Festuca rubra

4.2995

Equisetum palustre

4.3071

Carex pallescens

4.3205

Bryum pseudotriquetrum

4.3495

Lathyrus pratensis

4.3806

Blysmus compressus

4.3984

Oenanthe fistulosa

4.4194

Lysimachia vulgaris

4.4683

Hypericum tetrapterum

4.5527

Brachythecium rivulare

4.6075

Agrostis canina

4.6378

Lycopus europaeus

4.6380

Succisa pratensis

4.6526

Mentha longifolia

4.6639

Hieracium caespitosum

4.7176

Ajuga reptans

4.7360

Equisetum fluviatile

4.7896

Galium palustre

4.8112

Scirpus sylvaticus

4.8379

Ranunculus auricomus

4.8594

Potentilla erecta

4.9011

Climacium dendroides

4.9182

Nardus stricta

4.9254

Bistorta major

4.9378

Philonotis fontana

4.9383

Myosotis sicula

4.9498

Plagiomnium affine

4.9534

Ranunculus flammula

4.9727

Campylium stellatum

4.9995

Carex flava

5.0675

Veronica scutellata

5.0982

Trifolium spadiceum

5.1833

Cruciata glabra

5.2041

Carex echinata

5.2574

Juncus tomasii

5.3017

Eriophorum latifolium

5.3587

Hypericum maculatum

5.5243

Alchemilla vulgaris

5.6868

Filipendula ulmaria

5.6924

Aulacomnium palustre

5.7198

Caltha palustris

5.8021

Ranunculus nemorosus

5.8457

Carex rostrata

5.8802

Geum rivulare

5.9116

Crepis paludosa

5.9376

Dactylorhiza cordigera

5.9480

Veratrum lobelianum

5.9483

Carex curta

5.9882

Parnassia palustris

6.0211

Epilobium palustre

6.0849

Luzula sudetica

6.2168

Myosotis nemorosa

6.2762

Carex paniculata

6.3033

Geum coccineum

6.4584

Cardamine rivularis

6.5399

Carex nigra

6.5802

Silene asterias

6.7124

Chaerophyllum hirsutum

6.7303

Eriophorum angustifolium

6.8843

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Hájek, M., Hájková, P., Sopotlieva, D. et al. The Balkan wet grassland vegetation: a prerequisite to better understanding of European habitat diversity. Plant Ecol 195, 197–213 (2008). https://doi.org/10.1007/s11258-007-9315-8

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