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Research Article

Subalpine tall-herb vegetation in Bulgaria: diversity and ecology

ORCID Icon, & ORCID Icon
Pages 490-510 | Received 20 Sep 2023, Accepted 26 Feb 2024, Published online: 09 Apr 2024

Figures & data

Figure 1. Map of main Bulgarian Mountain ranges with the occurrence of vegetation plots.

Figure 1. Map of main Bulgarian Mountain ranges with the occurrence of vegetation plots.

Figure 2. Alluvial diagram showing differences in classifications between created expert system (ESy) with 17 distinguished groups, modified twinspan (mTwinspan) for 23 clusters, Ward’s agglomerative algorithm (Ward.D2) for 17 clusters, and EUNIS expert system (EUNIS); ADE01 – Ranunculo platanifolii-Adenostyletum alliariae, ARU01 – Astrantio majoris-Calamagrostietum arundinaceae, ARU02 – Roso pendulinae-Calamagrostietum arundinaceae, CAL01 – Crepido conyzifoliae-Calamagrostietum villosae, CIC01 – Moehringio pendulae-Cicerbitetum alpinae, CIR01 – Angelico sylvestris-Heracleetum verticillati, DES01 – Carici lazarei-Deschampsietum cespitosae, DOR01 – Aquilegio aureae-Doronicetum columnae, FOR01 – Saxifrago rotundifoliae-Athyrietum distentifolii, FOR02 – Cardamino acri-Athyrietum filicis-feminae, MOL01 – Sphagno compacti-Molinietum cearuleae, PET01 – Petasitetum albi, PET02 – Petasitetum hybridi, RUM01 – Senecioni rupestris-Rumicetum alpini, RUM02 – Rumicetum alpini, TRO01 – Trollius europaeus communities; C21a – base-poor spring and spring brook, R – grasslands, R35 – moist to wet mesotrophic to eutrophic hay meadows, R37 – temperate and boreal moist or wet oligotrophic grassland, R43 – temperate acidophilous alpine grassland, R55 – lowland moist or wet tall-herb and fern fringe, R56 – montane to subalpine moist or wet tall-herb and fern fringe, R57 – herbaceous Forest clearing vegetation, V – man-made habitats.

Figure 2. Alluvial diagram showing differences in classifications between created expert system (ESy) with 17 distinguished groups, modified twinspan (mTwinspan) for 23 clusters, Ward’s agglomerative algorithm (Ward.D2) for 17 clusters, and EUNIS expert system (EUNIS); ADE01 – Ranunculo platanifolii-Adenostyletum alliariae, ARU01 – Astrantio majoris-Calamagrostietum arundinaceae, ARU02 – Roso pendulinae-Calamagrostietum arundinaceae, CAL01 – Crepido conyzifoliae-Calamagrostietum villosae, CIC01 – Moehringio pendulae-Cicerbitetum alpinae, CIR01 – Angelico sylvestris-Heracleetum verticillati, DES01 – Carici lazarei-Deschampsietum cespitosae, DOR01 – Aquilegio aureae-Doronicetum columnae, FOR01 – Saxifrago rotundifoliae-Athyrietum distentifolii, FOR02 – Cardamino acri-Athyrietum filicis-feminae, MOL01 – Sphagno compacti-Molinietum cearuleae, PET01 – Petasitetum albi, PET02 – Petasitetum hybridi, RUM01 – Senecioni rupestris-Rumicetum alpini, RUM02 – Rumicetum alpini, TRO01 – Trollius europaeus communities; C21a – base-poor spring and spring brook, R – grasslands, R35 – moist to wet mesotrophic to eutrophic hay meadows, R37 – temperate and boreal moist or wet oligotrophic grassland, R43 – temperate acidophilous alpine grassland, R55 – lowland moist or wet tall-herb and fern fringe, R56 – montane to subalpine moist or wet tall-herb and fern fringe, R57 – herbaceous Forest clearing vegetation, V – man-made habitats.

Figure 3. DCA ordination diagram with ESy groups. Projected are sample scores. RUM01 – Senecioni rupestris-Rumicetum alpini, RUM02 – Rumicetum alpini, PET01 – Petasitetum albi, PET02 – Petasitetum hybridi, CAL01 – Crepido conyzifoliae-Calamagrostietum villosae, MOL01 – Sphagno compacti-Molinietum cearuleae, DES01 – Carici lazarei-Deschampsietum cespitosae, ARU01 – Astrantio majoris-Calamagrostietum arundinaceae, ARU02 – Roso pendulinae-Calamagrostietum arundinaceae, ADE01 – Ranunculo platanifolii-Adenostyletum alliariae, TRO01 – Trollius europaeus communities, CIC01 – Moehringio pendulae-Cicerbitetum alpinae, FOR01 – Saxifrago rotundifoliae-Athyrietum distentifolii, FOR02 – Cardamino acri-Athyrietum filicis-feminae, CIR01 – Angelico sylvestris-Heracleetum verticillati, DOR01 – Aquilegio aureae-Doronicetum columnae.

Figure 3. DCA ordination diagram with ESy groups. Projected are sample scores. RUM01 – Senecioni rupestris-Rumicetum alpini, RUM02 – Rumicetum alpini, PET01 – Petasitetum albi, PET02 – Petasitetum hybridi, CAL01 – Crepido conyzifoliae-Calamagrostietum villosae, MOL01 – Sphagno compacti-Molinietum cearuleae, DES01 – Carici lazarei-Deschampsietum cespitosae, ARU01 – Astrantio majoris-Calamagrostietum arundinaceae, ARU02 – Roso pendulinae-Calamagrostietum arundinaceae, ADE01 – Ranunculo platanifolii-Adenostyletum alliariae, TRO01 – Trollius europaeus communities, CIC01 – Moehringio pendulae-Cicerbitetum alpinae, FOR01 – Saxifrago rotundifoliae-Athyrietum distentifolii, FOR02 – Cardamino acri-Athyrietum filicis-feminae, CIR01 – Angelico sylvestris-Heracleetum verticillati, DOR01 – Aquilegio aureae-Doronicetum columnae.

Figure 4. Distribution of Senecioni rupestris-Rumicetum alpini in Bulgaria.

Figure 4. Distribution of Senecioni rupestris-Rumicetum alpini in Bulgaria.

Figure 5. Distribution of Rumicetum alpini in Bulgaria.

Figure 5. Distribution of Rumicetum alpini in Bulgaria.

Figure 6. Distribution of Petasitetum albi in Bulgaria.

Figure 6. Distribution of Petasitetum albi in Bulgaria.

Figure 7. Distribution of Petasitetum hybridi in Bulgaria.

Figure 7. Distribution of Petasitetum hybridi in Bulgaria.

Figure 8. Distribution of Crepido conyzifoliae-Calamagrostietum villosae in Bulgaria.

Figure 8. Distribution of Crepido conyzifoliae-Calamagrostietum villosae in Bulgaria.

Figure 9. Distribution of Sphagno compacti-Molinietum caeruleae in Bulgaria.

Figure 9. Distribution of Sphagno compacti-Molinietum caeruleae in Bulgaria.

Figure 10. Distribution of Carici lazarei-Deschampsietum cespitosae in Bulgaria.

Figure 10. Distribution of Carici lazarei-Deschampsietum cespitosae in Bulgaria.

Figure 11. Distribution of Astrantio majori-Calamagrostietum arundinaceae in Bulgaria.

Figure 11. Distribution of Astrantio majori-Calamagrostietum arundinaceae in Bulgaria.

Figure 12. Distribution of Roso pendulinae-Calamagrostietum arundinaceae in Bulgaria.

Figure 12. Distribution of Roso pendulinae-Calamagrostietum arundinaceae in Bulgaria.

Figure 13. Distribution of Ranunculo platanifolii-Adenostyletum alliariae in Bulgaria.

Figure 13. Distribution of Ranunculo platanifolii-Adenostyletum alliariae in Bulgaria.

Figure 14. Distribution of Trollius europaeus communities in Bulgaria.

Figure 14. Distribution of Trollius europaeus communities in Bulgaria.

Figure 15. Distribution of Saxifrago rotundifoliae-Athyrietum distentifolii in Bulgaria.

Figure 15. Distribution of Saxifrago rotundifoliae-Athyrietum distentifolii in Bulgaria.

Figure 16. Distribution of Cardamino acri-Athyrietum filicis feminae in Bulgaria.

Figure 16. Distribution of Cardamino acri-Athyrietum filicis feminae in Bulgaria.

Figure 17. Distribution of Moehringio pendulae-Cicerbitetum alpinae in Bulgaria.

Figure 17. Distribution of Moehringio pendulae-Cicerbitetum alpinae in Bulgaria.

Figure 18. Distribution of Angelico sylvestris-Heracleetum verticillati in Bulgaria.

Figure 18. Distribution of Angelico sylvestris-Heracleetum verticillati in Bulgaria.

Figure 19. Distribution of Aquilegio aureae-Doronicetum columnae in Bulgaria.

Figure 19. Distribution of Aquilegio aureae-Doronicetum columnae in Bulgaria.

Figure 20. DCA ordination with vectors; cmi_max – maximum monthly climate moisture index, Dis_severity – disturbance severity, Dis_frequency – disturbance frequency, ngd0 – number of growing degree days > 0 °C, reaction – reaction; projected are sample scores (grey) and species scores (dark grey) with weights > 200; Alchemilla – Alchemilla sp., Calaru – Calamagrostis arundinacea, Cirapp – Cirsium appendiculatum, Desces – Deschampsia cespitosa, Doraus – Doronicum austriacum, Fesval – Festuca valida, Fesrub – Festuca rubra agg., Gersyl – Geranium sylvaticum, Geucoc – Geum coccineum, Herver – Heracleum sphondylium subsp. verticillatum, Hypmac – Hypericum maculatum, Luzalp – Luzula alpinopilosa, Rubida – Rubus idaeus, Rumalp – Rumex alpinus, Rumari – Rumex arifolius, Saxrot – Saxifraga rotundifolia, Sennem – Senecio nemorensis agg., Stelnem – Stellaria nemorum, Verlob – Veratrum lobelianum.

Figure 20. DCA ordination with vectors; cmi_max – maximum monthly climate moisture index, Dis_severity – disturbance severity, Dis_frequency – disturbance frequency, ngd0 – number of growing degree days > 0 °C, reaction – reaction; projected are sample scores (grey) and species scores (dark grey) with weights > 200; Alchemilla – Alchemilla sp., Calaru – Calamagrostis arundinacea, Cirapp – Cirsium appendiculatum, Desces – Deschampsia cespitosa, Doraus – Doronicum austriacum, Fesval – Festuca valida, Fesrub – Festuca rubra agg., Gersyl – Geranium sylvaticum, Geucoc – Geum coccineum, Herver – Heracleum sphondylium subsp. verticillatum, Hypmac – Hypericum maculatum, Luzalp – Luzula alpinopilosa, Rubida – Rubus idaeus, Rumalp – Rumex alpinus, Rumari – Rumex arifolius, Saxrot – Saxifraga rotundifolia, Sennem – Senecio nemorensis agg., Stelnem – Stellaria nemorum, Verlob – Veratrum lobelianum.

Figure 21. Differences in environmental factors between vegetation types. The ngd0 and cmi_max were tested with the ANOVA test, variables derived from EIV with the fourth corner permutation test (significance indicated at the upper right corner; *** < .001). The line in the boxplots represents the median, and the whiskers show maximum and minimum values at a maximum of 1.5 times the interquartile range. Boxes represent first and third quartiles, and dots represent outliers. The red dashed line represents the overall mean. ADE01 – Ranunculo platanifolii-Adenostyletum alliariae, ARU01 – Astrantio majoris-Calamagrostietum arundinaceae, ARU02 – Roso pendulinae-Calamagrostietum arundinaceae, CAL01 – crepido conyzifoliae-Calamagrostietum villosae, CIC01 – Moehringio pendulae-Cicerbitetum alpinae, CIR01 – Angelico sylvestris-Heracleetum verticillati, DES01 – Carici lazarei-Deschampsietum cespitosae, DOR01 – Aquilegio aureae-Doronicetum columnae, FOR01 – Saxifrago rotundifoliae-Athyrietum distentifolii, FOR02 – Cardamino acri-Athyrietum filicis-feminae, MOL01 – Sphagno compacti-Molinietum cearuleae, PET01 – Petasitetum albi, PET02 – Petasitetum hybridi, RUM01 – Senecioni rupestris-Rumicetum alpini, RUM02 – Rumicetum alpini, TRO01 – Trollius europaeus communities.

Figure 21. Differences in environmental factors between vegetation types. The ngd0 and cmi_max were tested with the ANOVA test, variables derived from EIV with the fourth corner permutation test (significance indicated at the upper right corner; *** < .001). The line in the boxplots represents the median, and the whiskers show maximum and minimum values at a maximum of 1.5 times the interquartile range. Boxes represent first and third quartiles, and dots represent outliers. The red dashed line represents the overall mean. ADE01 – Ranunculo platanifolii-Adenostyletum alliariae, ARU01 – Astrantio majoris-Calamagrostietum arundinaceae, ARU02 – Roso pendulinae-Calamagrostietum arundinaceae, CAL01 – crepido conyzifoliae-Calamagrostietum villosae, CIC01 – Moehringio pendulae-Cicerbitetum alpinae, CIR01 – Angelico sylvestris-Heracleetum verticillati, DES01 – Carici lazarei-Deschampsietum cespitosae, DOR01 – Aquilegio aureae-Doronicetum columnae, FOR01 – Saxifrago rotundifoliae-Athyrietum distentifolii, FOR02 – Cardamino acri-Athyrietum filicis-feminae, MOL01 – Sphagno compacti-Molinietum cearuleae, PET01 – Petasitetum albi, PET02 – Petasitetum hybridi, RUM01 – Senecioni rupestris-Rumicetum alpini, RUM02 – Rumicetum alpini, TRO01 – Trollius europaeus communities.

Table 1. Variables of CCA analysis tested by anova.cca permutation test on sample scores, significance * < .05, ** < .01.

Table 2. Variables of CCA analysis tested by anova.cca permutation test on species scores, significance *** < .001.

Combined synoptic table of fidelity and frequency.

Table B1. Aggregation of species.