Research Paper |
Corresponding author: Kamila Reczyńska ( kamila.reczynska@uwr.edu.pl ) Academic editor: Jürgen Dengler
© 2022 Krzysztof Świerkosz, Kamila Reczyńska.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Świerkosz K, Reczyńska K (2022) Diversity of Mulgedio-Aconitetea communities in the Sudetes Mts. (SW Poland) in the Central European context. Vegetation Classification and Survey 3: 67-86. https://doi.org/10.3897/VCS.70200
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Aims: To describe the compositional and ecological diversity of Mulgedio-Aconitetea communities in the Sudetes Mts. and their foothills. Study area: The Sudetes Mts. (Southwestern Poland). Methods: A total of 399 vegetation relevés from own field studies and the literature were sorted into groups that match the higher syntaxa of the EuroVegChecklist and associations described in the literature. Diagnostic species of the so delimited associations were determined with the phi-coefficient of association, and maps of the associations produced. Direct ordination methods were applied to identify the main environmental gradients shaping the plant communities. Results: We distinguished nine associations, belonging to four alliances: submontane and colline communities (Petasition officinalis: Geranio phaei-Urticetum dioicae, Petasitetum hybridi, Chaerophyllo hirsuti-Petasitetum albi, Prenanthetum purpureae), upper montane nitrophilous communities (Rumicion alpini: Rumicetum alpini); subalpine communities with a dominance of graminoids and ferns (Calamagrostion villosae: Poo chaixii-Deschampsietum cespitosae, Crepido conyzifoliae-Calamagrostietum villosae, Athyrietum filicis-feminae) and subalpine tall-herb communities (Adenostylion alliariae: Cicerbitetum alpinae). Altitude, light availability, and bedrock type, which determines nutrient availability and soil reaction, played an important role in differentiating the studied communities. Conclusions: For convenience, we placed the four alliances in four separate orders as in the EuroVegChecklist. The fact that our ordination diagram separated only two main groups suggests the need of further research in this matter.
Taxonomic reference:
Syntaxonomic reference: Higher syntaxa follow
Abbreviations: db-RDA = distance-based redundancy analysis; EIV = Ellenberg indicator value; pANOVA = permutational analysis of variance; PCoA = principal coordinates analysis.
Adenostylion alliariae, Calamagrostion villosae, Central Europe, hygrophilous species, montane vegetation, Mulgedio-Aconitetea, Petasition officinalis, Poland, Rumicion alpini, synecology, syntaxonomy, tall-herb community
Montane to subalpine tall-herb communities of Europe, Siberia and Greenland are classified in the class Mulgedio-Aconitetea Hadač et Klika in Klika et Hadač 1944. They include communities of tall forbs, ferns and graminoids found close to high altitude watercourses or growing within the vast areas of subalpine grasslands. According to the most comprehensive syntaxonomic overview to date, the class encompasses tall-herb communities of eutrophic habitats on raw alluvia of streams in the upper colline to supramontane belts (
The discrepancy between the systems covering Europe or larger parts thereof and local classifications of tall-herb communities in individual countries prompted us to analyze the variability of this type of vegetation in the Sudetes and their foothills. In Poland, montane tall-herb communities are still classified within the class Betulo-Adenostyletea Br.-Bl. 1948 and the order Calamagrostietalia villosae Pawłowski et al. 1928, which includes two alliances (
Besides the above-mentioned associations, from the Polish side of the Sudetes Mts. and their foothills the following association-level syntaxa were reported: Geranio phaei-Urticetum
The aims of this study are thus: i) to conduct a comprehensive analysis of diversity of tall-herb communities of the Mulgedio-Aconitetea in the Sudetes Mts., assign the terminal units to described associations and place them into the higher syntaxa of the EuroVegChecklist (https://www.synbiosys.alterra.nl/evc/;
Our research was conducted on the whole area of the Sudetes, a mountain range in southwestern Poland extending over a length of 300 km and covering together with its foothills approximately 5,550 km2 (Figure
Between 1991 and 2020, we sampled 212 vegetation plots of the class in the Sudetes Mts. and their foothills (coordinates 15.32°E–17.23°E and 50.20°N–51.25°N), at elevations from 240 to 1400 m a.s.l. The plots were located in the terraces or banks of the stream valleys as well as within mires with tall-herbs. We chose stands that contained diagnostic species for the class Mulgedio-Aconitetea and its subordinate syntaxa as known from the literature (Suppl. material
Additionally, we used all the available relevés from the literature (
The relevés are available via the Polish Vegetation Database (Global Index of Vegetation-Plot Databases, ID: EU-PL-001;
In order to identify the ecological conditions of the tall-herb communities within the study area, different environmental variables were analyzed. Elevation (measured in m a.s.l. and divided by 1000 for presentation), slope, heat load and bedrock type were used as explanatory variables. The bedrock type at each site was obtained from the Detailed Geological Map of the Sudetes Mts. (Polish Geological Institute, National Research Institute, http://sudety.pgi.gov.pl). Based on the criterion of mineral composition and major geological processes (
Prior to the analyses, occurrences of the same species in different vertical layers were merged using the procedure implemented in JUICE, under the assumption that the overlap of layers is random (
We conducted an unsupervised classification with the modified TWINSPAN algorithm (
A principal coordinates analysis (PCoA) was performed in CANOCO (
To identify the main ecological drivers affecting the diversity of distinct groups, a distance-based redundancy analysis (db-RDA), embedded in CANOCO 5.0 (
In the analyzed data we could distinguish nine groups of tall-herb communities (Table
Cl. Mulgedio-Aconitetea Hadač et Klika in Klika et Hadač 1944
O. Petasito-Chaerophylletalia Morariu 1967 ex Kopecký
All. Petasition officinalis
Group 1: Geranio phaei-Urticetum dioicae
Group 2: Petasitetum hybridi Imchenetzky 1926
Group 3: Chaerophyllo hirsuti-Petasitetum albi Sýkora et Hadač 1984
Group 4: Prenanthetum purpureae
O. Senecioni rupestris-Rumicetalia alpini Mucina et Karner 2016
All. Rumicion alpini Scharfetter 1938
Group 5: Rumicetum alpini Beger 1922
O. Calamagrostietalia villosae Pawłowski et al. 1928
All. Calamagrostion villosae Pawłowski et al. 1928
Group 6: Poo chaixii-Deschampsietum cespitosae Pawłowski et Walas 1949
Group 7: Crepido conyzifoliae-Calamagrostietum villosae (Zlatník 1925) Jeník 1961
Group 8: Athyrietum filicis-feminae Wendelberger in Höfler et Wendelberger 1960
O. Adenostyletalia alliariae Br.-Bl. 1930
All. Adenostylion alliariae Br.-Bl. 1926
Group 9: Cicerbitetum alpinae
The nine associations clearly differ in terms of diagnostic species (Table
Tall-forb vegetation belonging to the Petasito-Chaerophylletalia. a–d. Geranio phaei-Urticetum dioicae components in colline location (330‒380 m a.s.l. in the Pełcznica river’s ravine (Wałbrzyskie Foothills, Central Sudetes) (a. Chaerophyllum hirsutum; b. Geranium robertianum; c. Stellaria nemorum; d. Geranium phaeum); e. Petasitetum hybridi in the valley of the Bystrzyca Dusznicka river, 725 m a.s.l. (Bystrzyckie Mts., Central Sudetes). (Photographs a–d. K. Reczyńska; e. K. Świerkosz).
Tall-forb vegetation belonging to the Petasito-Chaerophylletalia. a–b. Delphinium elatum in the Kleśnica valley (Śnieżnik Massif, Eastern Sudetes) where it occurs in the plots of the Petasitetum hybridi and Chaerophyllo hirsuti-Petasitetum albi; c‒d. Prenanthetum purpureae in the Stołowe Mts. (Eastern Sudetes) with the presence of Veratrum lobelianum. (Photographs a-d K. Świerkosz).
Communities of Mulgedio-Aconitetea in the Karkonosze Mts. and Orlickie Mts. a. Aconitum variegatum in the Athyrietum filicis-feminae in the Orlickie Mts. near Zieleniec; b. Rumicetum alpini in the vicinity of Hala Szrenicka montane hostel (1100 m a.s.l.) c‒d. Subendemic Aconitum plicatum in the Crepido-Calamagrostietum (c) and the Cicerbitetum alpinae (d) – one of the most spectacular species occurring in subalpine tall-herb communities (1170‒1260 m a.s.l.). (Photographs b, d. K. Reczyńska; a, c. K. Świerkosz).
Summarized synoptic table with percentage frequency and fidelity values derived from 399 relevés of tall-herb associations of the Mulgedio-Aconitetea in the Sudetes Mts. and their foothills (SW Poland). The positive Φ coefficient values (multiplied by 100) are presented as superscripts (*: 25 to 49, **: ≥ 50). Species diagnostic for both one or several clusters (Φ ≥ 25 and constancy ratio ≥1.3) are shaded in grey. Character and differentiating species were sorted according to constancy, while the other species were sorted according to the number of occurrences in clusters or resemblance to the diagnostic species in neighboring cluster. Among accompanying species, only the most common ones, which occurred in at least seven clusters, were included in the table. Abbreviations: Ch: character species; D*: regionally differentiating species (marked in the table with asterisk). Shortened names of syntaxa: Se.Ru.: order Senecioni rupestris-Rumicetalia alpini; Cal.vill.: order Calamagrostietalia villosae; Ad.al.: order Adenostyletalia alliariae; R.al.: alliance Rumicion alpini; Ca.vi.: alliance Calamagrostion villosae; Ad.: alliance Adenostylion alliariae.
Group No. | Constancy | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 |
---|---|---|---|---|---|---|---|---|---|---|
No. of relevés | ratio | 33 | 59 | 74 | 51 | 20 | 12 | 61 | 59 | 30 |
Order | Petasito-Chaerophylletalia | Se.Ru | Cal.vill. | Ad. al | ||||||
Alliance | Petasition officinalis | R.al. | Ca. vi. | Ad. | ||||||
Ch. and D.* Geranio phaei-Urticetum | ||||||||||
Geranium phaeum | 14 | 100 ** | 7 | 1 | . | . | . | . | . | . |
Geum urbanum* | 4.5 | 55 ** | 12 | 5 | . | . | . | . | . | . |
Chaerophyllum aromaticum* | 2.5 | 52 ** | 20 | 5 | 4 | . | . | . | . | . |
Glechoma hederacea* | 3.1 | 42 ** | 14 | . | . | . | . | . | . | . |
Poa nemoralis* | 5.9 | 39 * | 2 | 4 | 6 | . | . | . | 2 | 7 |
Schedonorus giganteus* | 3.3 | 36 * | 7 | 11 | 4 | . | . | . | 2 | . |
Alliaria petiolata* | 17 | 30 ** | 2 | . | . | . | . | . | . | . |
Stellaria holostea* | 17 | 30 ** | 2 | . | . | . | . | . | . | . |
Ulmus glabra* | 5.0 | 27 * | 2 | 5 | . | . | . | . | . | . |
Chaerophyllum temulum* | 100 | 21* | ||||||||
Ch. and D.* Petasitetum hybrydi | ||||||||||
Petasites hybridus | 24 | . | 100 ** | 4 | . | . | . | . | . | . |
Cirsium oleraceum* | 1.5 | 9 | 56 * | 36 | 14 | . | 8 | . | 2 | |
Phalaroides arundinacea | 100 | . | 41 ** | . | . | . | . | . | . | . |
Filipendula ulmaria* | 1.8 | 9 | 31 * | 7 | . | . | . | 2 | . | 17 |
Anthriscus sylvestris* | 1.9 | 12 | 24 * | 1 | . | . | . | . | . | . |
Ch. and D.* Chaerophylo hirsuti-Petasitetum albi | ||||||||||
Petasites albus | 5.7 | 3 | 12 | 99 ** | 69 | . | 8 | . | 8 | 7 |
Impatiens noli-tangere* | 1.3 | 12 | 17 | 46 * | 35 | . | 17 | . | 3 | . |
Stachys sylvatica* | 1.6 | 27 | 25 | 45* | 24 | . | . | . | . | . |
Carex sylvatica* | 3.2 | 9 | 12 | 45 * | 14 | . | . | . | . | . |
Lysimachia nemorum* | 1.6 | . | 2 | 28 * | 18 | . | 17 | . | 2 | . |
Fraxinus excelsior* | 2.6 | 6 | 10 | 27 * | 8 | . | . | . | 2 | . |
Ch and D.* Prenanthetum purpureae | ||||||||||
Dryopteris carthusiana* | 1.8 | . | . | 16 | 53 * | . | 25 | . | 27 | 7 |
Equisetum sylvaticum* | 1.8 | . | 5 | 24 | 49 * | 5 | 42 | . | . | . |
Prenanthes purpurea | 1.3 | . | 2 | 18 | 47* | . | 17 | 11 | 34 | 17 |
Sorbus aucuparia subsp. aucuparia* | 2.5 | . | . | 8 | 47 * | . | 17 | 7 | 19 | . |
Dryopteris dilatata* | 1.6 | . | . | 4 | 41 * | . | 8 | 7 | 25 | 10 |
Gymnocarpium dryopteris* | 4.0 | . | . | 4 | 33 * | . | 8 | . | 3 | 7 |
Fagus sylvatica* | 1.5 | . | 1 | 15 | 24 * | . | . | . | . | 2 |
Phegopteris connectilis* | 2.1 | . | . | 7 | 22 * | . | . | . | 10 | . |
Ch. Rumicetum alpini | ||||||||||
Rumex alpinus | 51 | . | . | . | 2 | 100** | . | . | . | . |
Peucedanum ostruthium | 100 | . | . | . | . | 35** | . | . | . | . |
Ch. and D.* Poo-Deschampsietum cespitosae | ||||||||||
Juncus effusus * | 2.9 | . | 3 | 8 | 6 | 20 | 58 ** | . | 3 | . |
Viola lutea subsp. sudetica | 100 | . | . | . | . | . | 42 ** | . | . | . |
Cardamine opizii * | 100 | . | . | . | . | . | 33 ** | . | . | . |
Poa chaixi | 1.3 | 4 | 10 | 33 * | 15 | 27 | ||||
Viola palustris* | 17 | . | 2 | 1 | 2 | . | 33 * | . | . | . |
Stellaria alsine * | 6.5 | . | 2 | 3 | . | . | 33 * | . | 5 | . |
Ch. and D.* Crepido conyzifoliae-Calamagrostietum villosae | ||||||||||
Anthoxanthum odoratum* | 100 | . | . | . | . | . | . | 64 ** | . | . |
Potentilla aurea | 9.7 | . | . | . | . | . | . | 61 ** | 2 | 7 |
Pulsatilla alpina | 100 | . | . | . | . | . | . | 54 ** | . | . |
Calluna vulgaris* | 31 | . | . | . | . | . | . | 54 ** | 2 | . |
Melampyrum sylvaticum* | 22 | . | . | . | 2 | . | . | 44 ** | 2 | . |
Calamagrostis arundinacea* | 1.3 | . | . | 22 | 12 | . | 17 | 41 * | 12 | 30 |
Vaccinium vitis-idaea* | 25 | . | . | 1 | . | . | . | 34 ** | . | . |
Maianthemum bifolium* | 2.5 | . | . | 3 | 14 | . | . | 34 * | 14 | . |
Achillea millefolium* | 1.7 | . | . | 3 | . | 20 | . | 34 * | 5 | 20 |
Anemonastrum narcissiflorum | 100 | . | . | . | . | . | . | 31 ** | . | . |
Thesium alpinum | 100 | . | . | . | . | . | . | 21 * | . | . |
Nardus stricta* | 100 | . | . | . | . | . | . | 21 * | . | . |
Ch. and D.* Cicerbitetum alpinae | ||||||||||
Thalictrum aquilegiifolium | 4.7 | . | 2 | 11 | 2 | . | . | 15 | 3 | 70 ** |
Heracleum sphondylium* | 2.3 | 9 | 24 | 8 | 2 | 20 | . | 7 | 2 | 57 * |
Epilobium alpestre | 5.3 | . | . | 1 | . | 10 | . | . | . | 53 ** |
Milium effusum* | 3.5 | 3 | 2 | 4 | 6 | 15 | 8 | . | 12 | 53 * |
Epilobium angustifolium* | 3.6 | . | . | . | . | 5 | . | 15 | 5 | 53 * |
Carduus personata | 5.2 | . | 5 | 9 | 2 | . | 8 | . | . | 50 * |
Dryopteris filix-mas* | 2.1 | 12 | . | 20 | 18 | . | 8 | 11 | 19 | 43 * |
Daphne mezereum* | 4.7 | . | 2 | 4 | . | . | 8 | 7 | 3 | 40 * |
Angelica sylvestris* | 1.4 | . | 20 | 1 | 6 | . | . | . | 3 | 30 * |
Hieracium prenanthoides | 8.1 | . | . | . | . | . | . | 3 | 2 | 27 * |
D*. All. Petasition officinalis | ||||||||||
Aegopodium podagraria* | 1.9 | 85** | 68 * | 35 | 14 | . | . | . | . | . |
Galium aparine* | 23 | 39* | 32 * | 1 | . | . | . | . | . | . |
Poa trivialis* | 1.3 | 9 | 34 * | 7 | 2 | 25 | 8 | . | 2 | 3 |
Mercurialis perennis* | 1.4 | 27* | 3 | 19 | 6 | . | . | . | . | . |
Lamium maculatum* | 1.3 | 18 | 25 * | 4 | . | . | . | . | . | . |
Ajuga reptans* | 3.6 | 6 | 2 | 22* | 6 | . | . | . | . | . |
Pulmonaria obscura* | 2.8 | 6 | 3 | 17* | 4 | . | . | . | . | . |
Acer pseudoplatanus* | 1 | 24 | 7 | 36 * | 37 | 5 | . | . | 8 | . |
Lamium galeobdolon agg.* | 1 | 48 | 8 | 53 * | 61 * | . | 17 | . | 2 | . |
D.* all. Rumicion alpini | ||||||||||
Agrostis capillaris* | 3.9 | . | . | . | . | 65 ** | 17 | 5 | 7 | 3 |
Festuca rubra* | 4.1 | . | . | . | . | 35 * | 8 | 5 | . | . |
Veronica chamaedrys* | 1.5 | 15 | 24 | 1 | . | 35 * | . | . | . | . |
Epilobium montanum* | 2 | 6 | 10 | 22 | 22 | 45 * | 8 | . | 8 | 3 |
Ranunculus acris* | 3 | 3 | 3 | . | 2 | 30 * | . | 2 | . | 9 |
Ochlopoa supina | 7.5 | . | . | . | . | 25** | . | . | . | 3 |
Ch and D* all. Calamagrostion villosae | ||||||||||
Avenella flexuosa * | 1.7 | . | . | . | 4 | 5 | 17 | 95 ** | 58 * | 33 |
Calamagrostis villosa | 1.3 | . | . | 15 | 69 | 20 | 50 | 92 * | 92 * | 50 |
Vaccinium myrtillus * | 1.6 | . | . | 4 | 20 | . | 33 | 90 ** | 56 * | 6 |
Gentiana asclepiadea | 1.4 | . | . | . | 4 | 5 | . | 57 * | 49 * | 33 |
Trientalis europaea * | 1.9 | . | . | . | . | . | 8 | 67 * | 37 * | . |
Luzula luzuloides * | 2.5 | . | 3 | 1 | 6 | 15 | 25 | 92 ** | 27 | 37 |
Solidago virgaurea * | 3.4 | . | . | 4 | 10 | . | 8 | 82 ** | 24 | 7 |
Homogyne alpina * | 2.7 | . | . | 1 | 18 | 5 | 8 | 79 ** | 29 | . |
Luzula sylvatica | . | . | . | 4 | . | 8 | 2 | 8 | . | |
Galium saxatile | . | . | . | . | . | . | 11 | 3 | 3 | |
Ch. and D* All Adenostylion alliariae | ||||||||||
Lactuca alpina | 4.7 | . | . | 11 | 84 * | . | 8 | 15 | 69 * | 87 * |
Ranunculus platanifolius | 1.9 | . | 2 | 7 | 10 | . | 8 | 74 * | 37 | 73 * |
Aconitum plicatum | 1.7 | . | . | 1 | . | . | 33 | 21 | 14 | 87** |
Adenostyles alliariae | 1.9 | . | . | 1 | . | 20 | . | 26 | 44 | 87 ** |
Viola biflora | 3.8 | . | . | 4 | 2 | . | . | 13 | 7 | 50 ** |
Lilium martagon* | 1.4 | 3 | 2 | 3 | 2 | . | . | 30 | 5 | 43 * |
Geranium sylvaticum | 2.7 | . | 15 | 7 | . | . | 17 | 28 | 6 | 77 ** |
Class Mulgedio-Aconitetea | ||||||||||
Senecio nemorensis | 19 | . | 2 | 4 | 2 | 85* | . | 79* | 78* | 100 * |
Athyrium distentifolium | 6.3 | . | . | 4 | 8 | 10 | . | 64* | 97 ** | 67 * |
Veratrum album | 2.2 | . | 3 | 18 | 31 | . | 8 | 82* | 71 * | 77 * |
Silene dioica | 2.9 | 12 | 17 | 7 | 2 | 10 | 8 | 15 | 51 * | 63 * |
Silene vulgaris * | 1.6 | . | . | . | . | 5 | 8 | 57 * | 5 | 37 * |
Anthriscus nitida | 1.9 | 15 | 5 | 14 | 8 | . | 8 | . | . | 30 * |
Alchemilla glabra | 1.7 | . | . | . | . | 30 | 25 | . | 3 | 53 * |
Valeriana excelsa subsp. sambucifolia | 4.6 | 3 | . | 11 | 4 | . | . | 2 | 5 | 50 * |
Phyteuma spicatum | 2.0 | 9 | 3 | 11 | 20 | 5 | . | 25 | 2 | 50 * |
Salix silesiaca | 3.0 | . | 2 | 4 | . | . | 8 | 11 | 12 | 37* |
Stellaria nemorum | 1.4 | 21 | 34 | 69 | 69 | 70 | 100 * | 2 | 36 | 60 |
Polygonatum verticillatum | 1.5 | . | . | 19 | 27 | . | . | 46* | 27 | 30 |
Rumex arifolius | 1.1 | . | 2 | 9 | . | 65 | 75 | 75 | 86 * | 93 * |
Chaerophyllum hirsutum | 21 | 49 | 77 | 84 | 50 | 83 | . | 14 | 77 | |
Streptopus amplexifolius | . | . | 15 | 33 | . | 8 | 20 | 24 | 40 * | |
Primula elatior | 21 | 20 | 27 | 18 | . | 8 | 2 | . | 23 | |
Aconitum variegatum | . | 2 | 9 | 4 | . | 17 | . | 2 | 3 | |
Delphinium elatum | . | 3 | 3 | 2 | . | . | . | . | . | |
Campanula latifolia | . | 2 | 5 | 2 | . | . | . | . | 3 | |
Diagnostic for more than one association | ||||||||||
Oxalis acetosella | 1.3 | 21 | 7 | 59 | 90 * | 10 | 92 * | 11 | 83 * | 1 |
Bistorta officinalis | 4.0 | . | 14 | . | . | 15 | 8 | 92* | 61* | 63* |
Senecio ovatus | 1.6 | 30 | 19 | 69 * | 71 * | 25 | 42 | 2 | 24 | 7 |
Athyrium filix-femina | 1.3 | 6 | 7 | 72 * | 69 * | 5 | 50 | . | 10 | . |
Deschampsia cespitosa | 1.4 | 9 | 15 | 9 | 14 | 85 * | 92 * | 48 | 39 | 60 |
Common accompanying species (at least in 7 groups) | ||||||||||
Crepis paludosa | 3 | 27 | 41 | 49 | 15 | 17 | 3 | 8 | 33 | |
Rubus idaeus | 9 | 8 | 42 | 41 | 25 | 8 | 30 | 61 | 60 | |
Hypericum maculatum | . | 10 | 4 | 4 | 35 | 17 | 41 | 15 | 30 | |
Ranunculus repens | 1 | 21 | 36 | 30 | 29 | 55 | 58 * | . | 8 | 3 |
Myosotis scorpioides agg. | 1.2 | . | 17 | 26 | 27 | 15 | 67 * | 5 | 8 | 53 |
Urtica dioica | 1.1 | 91 * | 78 * | 65 | 35 | 70 | 25 | . | 5 | 20 |
Picea abies | 1.1 | . | 3 | 14 | 35 | . | 42 * | 3 | 15 | 7 |
Alchemilla monticola agg. | 9 | 14 | 3 | 4 | . | . | 2 | 14 | 12 | |
Dactylis glomerata | 1.1 | 27 | 39 | 15 | 8 | 45 * | 25 | . | 3 | . |
Chrysosplenium alternifolium | . | 14 | 41 | 29 | 15 | 33 | . | 7 | 30 |
PCoA diagram for the 399 vegetation plots of tall-herb communities in the Sudetes Mts. and their foothills (SW Poland). The numbers of clusters are identical with Figure
Significance of Spearman’s rank correlation of mean EIVs with three main PCoA axes within the tall-herb communities in the Sudetes Mts. and their foothills using modified permutation test.
Axis 1 | Axis 2 | Axis 3 | ||||
---|---|---|---|---|---|---|
rho 1 | p 2 | rho | p | rho | p | |
EIV temperature | -0.77** | 0.004 | 0.32 | 0.220 | -0.08 | 0.696 |
EIV soil reaction | -0.73* | 0.012 | 0.34 | 0.164 | 0.32 | 0.108 |
EIV nutrients | -0.71* | 0.020 | 0.29 | 0.300 | 0.39 | 0.052 |
ElV light | 0.46 | 0.192 | 0.64** | 0.008 | 0.33 | 0.056 |
EIV moisture | -0.48 | 0.164 | -0.12 | 0.624 | 0.40* | 0.032 |
Summary box-and-whisker plots of mean Ellenberg indicator values (EIVs) for clusters recognized within Mulgedio-Aconitetea communities of the Sudetes and their foreland (SW Poland) produced by one-way permutational ANOVA. The central line of each box indicates the median value, box boundaries the lower (25%) and upper (75%) quartiles, and whiskers the range of values. pmodif was calculated using a modified permutation test of significance for analysis of mean Ellenberg indictor values (EIVs), F-test statistic. Numbers of groups are the same as in Figure
In the following, among the diagnostic species, the character species (in bold) are highlighted (see Table
Number of relevés: 33
Diagnostic species: Geranium phaeum, Alliaria petiolata, Chaerophyllum aromaticum, Ch. temulum, Geum urbanum, Glechoma hederacea, Poa nemoralis, Schedonorus giganteus, Stellaria holostea, Ulmus glabra, Aegopodium podagraria, Galium aparine, Mercurialis perennis
Constant species: Aegopodium podagraria, Chaerophyllum aromaticum, Geranium phaeum, Geum urbanum, Urtica dioica
Dominant species: Geranium phaeum
Ecology: The association was found at elevations between 200 and 750 m a.s.l. (mean 406 m a.s.l.), thus is submontane/montane in character. It occurred mainly on terraces of streams developing on Quaternary Pleistocene deposits, only exceptionally on Holocene sands and gravels. It usually accompanies nitrophilous alluvial forest communities of the Carpino-Fagetea Jakucs ex Passarge 1968 class, for example, in gaps of the stand or in places devoid of trees and shrubs.
Number of relevés: 59
Diagnostic species: Petasites hybridus, Phalaroides arundinacea, Anthriscus sylvestris, Filipendula ulmaria, Cirsium oleraceum, Aegopodium podagraria, Galium aparine, Poa trivialis, Rumex acetosa
Constant species: Aegopodium podagraria, Cirsium oleraceum, Petasites hybridus, Urtica dioica
Dominant species: Petasites hybridus
Ecology: The association was found at elevations between 300 and 964 m a.s.l. (mean 532 m a.s.l.), indicating its submountain and lower-mountain character. It was recorded mainly in stream valleys over Quaternary Holocene deposits, and only exceptionally over postglacial formations. Most of the documented stands occurred in the vicinity of submontane riparian forests, but in large, open and sunny clearings. The succession of trees and shrubs in such tall-forb patches is low, which may indicate regular floods of watercourses, increasing the mortality rate of seedlings of woody forms.
Number of relevés: 74
Diagnostic species: Petasites albus, Carex sylvatica, Fraxinus excelsior, Impatiens noli-tangere, Lysimachia nemorum, Ajuga reptans. Athyrium filix-femina, Senecio ovatus, Stachys sylvatica
Constant species: Athyrium filix-femina, Chaerophyllum hirsutum, Lamium galeobdolon agg., Oxalis acetosella, Petasites albus, Senecio ovatus, Stellaria nemorum, Urtica dioica
Dominant species: Petasites albus
Ecology: The association was found at elevations of 375‒995 m a.s.l. (mean 695 m a.s.l.), indicating its submontane/lower-montane character. It usually accompanies nitrophilous, deciduous forest communities of the Carpino-Fagetea class ‒ mainly alluvial forests, but also nitrophilous beech or ravine woods. This association was recorded from all types of analyzed bedrock, most often on Holocene gravels in stream valleys, but also on sedimentary and metamorphic rocks. Therefore, it does not appear to show any preferences regarding bedrock type and is one of the most widely distributed communities within the alliance. Stands over calcareous rocks are richer in species, with the inclusion of Delphinium elatum, Ranunculus platanifolius or Aconitum variegatum, and resemble the Delphinietum elatae Beger ex Sutter 1978. However, so far only one locality of that type is known in the Kleśnica Valley (Eastern Sudetes, Śnieżnik Massif, about 800 m a.s.l.).
Number of relevés: 51
Diagnostic species: Prenanthes purpurea, Equisetum sylvaticum, Dryopteris carthusiana, Dryopteris dilatata, Gymnocarpium dryopteris, Fagus sylvatica, Phegopteris connectilis, Sorbus aucuparia subsp. aucuparia, Athyrium filix-femina, Lactuca alpina, Oxalis acetosella, Senecio ovatus
Constant species: Athyrium filix-femina, Calamagrostis villosa, Chaerophyllum hirsutum, Lactuca alpina, Dryopteris carthusiana, Lamium galeobdolon agg., Oxalis acetosella, Petasites albus, Senecio ovatus, Stellaria nemorum
Dominant species: Lactuca alpina
Ecology: The association was found at elevations of (405‒) 650‒925 m a.s.l. (mean 758 m a.s.l.), therefore is montane in character. However, in contrast to the previous association, it accompanies acidophilous beech forests of the Luzulo-Fagetalia sylvaticeae Scamoni et Passarge 1959 order and spruce forest communities of natural or anthropogenic origin. Stands were mainly reported from sedimentary and metamorphic rocks, rarely from Holocene gravels or granites. The association is less frequent than the previous one and is restricted to a few mountain ranges (Śnieżnik Massif, Stołowe Mts., Orlickie Mts., Izerskie Mts. and Karkonosze Mts.).
Number of relevés: 20
Diagnostic species: Rumex alpinus, Peucedanum ostruthium, Ochlopoa supina, Agrostis capillaris, Deschampsia cespitosa, Epilobium montanum, Festuca rubra, Ranunculus acris, Veronica chamaedrys
Constant species: Agrostis capillaris, Deschampsia cespitosa, Ranunculus repens, Rumex alpinus, R. arifolius, Senecio nemorensis, Stellaria nemorum, Urtica dioica
Dominant species: Rumex alpinus
Ecology: The association was recorded at elevations of (840‒) 1055‒1190 m a.s.l. (mean 1063 m a.s.l.) within the lower alpine zone. It is connected to intensive human presence, for example near montane hostels and abandoned pastures. It was found mainly on granite (Karkonosze Mts.).
Number of relevés: 12
Diagnostic species: Viola lutea subsp. sudetica, Poa chaixii, Cardamine opizii, Juncus effusus, Stellaria alsine, Viola palustris, Agrostis canina, Deschampsia cespitosa, Oxalis acetosella, Stellaria nemorum
Constant species: Chaerophyllum hirsutum, Deschampsia cespitosa, Juncus effusus, Myosotis scorpioides agg., Oxalis acetosella, Ranunculus repens, Rumex arifolius, Stellaria nemorum
Dominant species: Deschampsia cespitosa, Doronicum austriacum
Ecology: The association was found at elevations of (650‒) 885‒1231 m a.s.l. (mean 1024 m a.s.l.), i.e., in the montane and subalpine zones. In the Sudetes Mts. it was reported from the Śnieżnik Massif, Izerskie Mts., Bystrzyckie and Orlickie Mts. mainly on metamorphic rocks (schists) and, rarely, Holocene deposits.
Number of relevés 61
Diagnostic species: Anemonastrum narcissiflorum, Potentilla aurea, Pulsatilla alpina, Thesium alpinum, Achillea millefolium, Anthoxanthum odoratum, Calluna vulgaris, Maianthemum bifolium, Melampyrum sylvaticum, Vaccinium vitis-idaea, Athyrium distentifolium, Calamagrostis arundinacea, Avenella flexuosa, Bistorta officinalis, Gentiana asclepiadea, Homogyne alpina, Luzula luzuloides, Nardus stricta, Polygonatum verticillatum, Ranunculus platanifolius, Silene vulgaris, Solidago virgaurea, Trientalis europaea, Veratrum album
Constant species: Anthoxanthum odoratum, Athyrium distentifolium, Bistorta officinalis, Calamagrostis villosa, Calluna vulgaris, Avenella flexuosa, Gentiana asclepiadea, Homogyne alpina, Luzula luzuloides, Potentilla aurea, Pulsatilla alpina, Ranunculus platanifolius, Rumex arifolius, Senecio nemorensis, Silene vulgaris, Solidago virgaurea, Trientalis europaea, Vaccinium myrtillus, Veratrum album
Dominant species: Calamagrostis villosa
Ecology: The association was found in the Karkonosze Mts. at elevations from 1150 to 1470 m a.s.l. (mean 1287 m a.s.l.) and is subalpine in character.
Number of relevés: 59
Diagnostic species: Athyrium distentifolium, Lactuca alpina, Avenella flexuosa, Gentiana asclepiadea, Oxalis acetosella, Silene dioica, Veratrum album
Constant species: Athyrium distentifolium, Calamagrostis villosa, Lactuca alpina, Avenella flexuosa, Gentiana asclepiadea, Oxalis acetosella, Bistorta officinalis, Rubus idaeus, Rumex arifolius, Senecio nemorensis, Vaccinium myrtillus, Veratrum album
Dominant species: Athyrium distentifolium
Ecology: The association was found at elevations from 770 to 1380 m a.s.l. (mean 1168 m a.s.l.), in the upper montane and subalpine zones. In the Sudetes Mts. it is reported from the Karkonosze Mts. on granites, mainly on steep slopes of the postglacial circles, and from the Orlickie and Izerskie Mts. on metamorphic bedrocks.
Number of relevés 30
Diagnostic species: Carduus personata, Epilobium alpestre, Hieracium prenanthoides, Thalictrum aquilegiifolium, Angelica sylvestris, Daphne mezereum, Dryopteris filix-mas, Epilobium angustifolium, Heracleum sphondylium subsp. sphondylium, Milium effusum, Aconitum plicatum, Adenostyles alliariae, Alchemilla glabra, Anthriscus nitida, Athyrium distentifolium, Bistorta officinalis, Lactuca alpina, Geranium sylvaticum, Lilium martagon, Phyteuma spicatum, Ranunculus platanifolius, Silene dioica, Salix silesiaca, Veratrum album, Viola biflora
Constant species: Aconitum plicatum, Adenostyles alliariae, Alchemilla glabra, Athyrium distentifolium, Chaerophyllum hirsutum, Lactuca alpina, Deschampsia cespitosa, Epilobium alpestre, Geranium sylvaticum, Heracleum sphondylium subsp. sphondylium, Myosotis scorpioides agg., Bistorta officinalis, Ranunculus platanifolius, Rubus idaeus, Rumex arifolius, Senecio nemorensis, Silene dioica, Stellaria nemorum, Thalictrum aquilegiifolium, Valeriana excelsa subsp. sambucifolia, Veratrum album
Dominant species: Adenostyles alliariae, Lactuca alpina
Ecology: The association was found at elevations from 1100 to 1400 m a.s.l. (mean 1277 m a.s.l.), in the upper montane and subalpine zones. In the Sudetes Mts. it is reported only from the Karkonosze Mts. on granite.
As shown in the PCoA ordination diagram, the nine associations fall within two clusters that were clearly separated along the first PCoA axis (Figure
The pANOVA revealed that temperature played a significant role in the floristic differentiation of the nine associations (p < 0.05). In contrast, the other analyzed EIVs for nutrients, soil reaction, moisture and light were not significant (Figure
The db-RDA (Figure
db-RDA plot of samples of the tall-herb communities of the Sudetes Mts. and their foothills (SW Poland) overlaid with environmental variables. Symbols indicate alliances: circles: Petasition officinalis (1: Geranio phaei-Urticetum, 2: Petasitetum hybrydi, 3: Chaerophylo hirsuti-Petasitetum albi; 4: Prenanthetum purpureae); triangles: Rumicion alpini (5: Rumicetum alpini); diamonds: Calamagrostion villosae (6: Poo chaixii-Deschampsietum, 7: Athyrietum filicis-feminae, 8: Crepido conyzifoliae-Calamagrostietum); squares: Adenostylion alliariae (9: Cicerbitetum alpinae).
The simple term and conditional effects of analyzed environmental variables on species composition of the distinguished montane tall-herb communities, identified using db-RDA with SQRT Jaccard binary distance and Monte Carlo permutation test. Lambda - variance explained by the environmental variable [in %], p adj.: p with Bonferroni correction.
Environmental variable | Simple Term Effects | Conditional effects | ||||
---|---|---|---|---|---|---|
Lambda | pseudo-F | p adj. | Lambda | pseudo-F | p adj. | |
Altitude | 7.7 | 21.3 | 0.018 | 7.73 | 21.3 | 0.018 |
Plutonic rocks | 6.7 | 18.3 | 0.018 | 1.06 | 3.0 | 0.018 |
Slope | 4.7 | 12.4 | 0.018 | 0.77 | 2.2 | 0.018 |
Calcicolous rocks | 2.8 | 7.3 | 0.018 | 0.87 | 2.4 | 0.018 |
Heat Load | 1.8 | 4.6 | 0.018 | 1.00 | 2.8 | 0.018 |
Metamorphic rocks | 1.2 | 3.0 | 0.018 | 0.67 | 1.9 | 0.018 |
Holocene deposits | 2.6 | 6.9 | 0.018 | 0.38 | 1.1 | > 0.05 |
Sandstones | 0.7 | 1.8 | 0.018 | 0.55 | 1.6 | > 0.05 |
Separation of communities of the class Mulgedio-Aconitetea from other tall-herb communities of the classes Epilobietea angustifolii Tx. et Preising ex von Rochow 1951, Trifolio-Geranietea sanguinei T. Müller 1962 as well as order Filipendulo ulmariae-Lotetalia uliginosi Passarge 1975 requires a detailed analysis of phytosociological relevés from all over Europe (or at least its central part) covering all the above-mentioned syntaxonomic units. Therefore, it significantly exceeds the scope of the presented study.
Hitherto, in synthetic studies concerning the Mulgedio-Aconitetea class, character species were distinguished based on a priori prepared lists (
Despite recognition at both regional and supra-regional scales, there is still no general agreement on the syntaxonomy of tall-herb communities of the class Mulgedio-Aconitetea. The synthesis carried out by
1. Adenostyletalia alliariae: Tall-herb vegetation with three Central European alliances Adenostylion alliariae (on siliceous substrates at high altitudes in the nemoral zone of Europe), Dryopterido filicis-maris-Athyrion distentifolii (Holub ex Sýkora et Štursa 1973) Jeník et al. 1980 (on fertile soils at high altitudes of the Alps, Carpathians, Hercynicum and Scandinavia), Delphinion elati Hadač in
2. Calamagrostietalia villosae: Tall-grass and herb-rich vegetation on nutrient-poor soils of the Alps, Carpathians and Hercynicum with three Central European alliances: Calamagrostion villosae (tall-herb and herb-rich vegetation on acidic soils in the subalpine and alpine belts of the Alps, Carpathians and Hercynicum), Trisetion fusci Krajina 1933 (on alluvial acidic soils along alpine streams of the Carpathians) and Calamagrostion arundinaceae (of tall-grass and herb-rich vegetation on dry acidic soils in the upper montane and subalpine belts of the mountain ranges of suboceanic Europe).
3. Petasito-Chaerophylletalia: Tall-herb vegetation on nutrient-rich soils along mountain streams of Central Europe, the Balkans and the Apennines of order with two Central European alliances: Petasition officinalis (vegetation on raw alluvia of streams in the upper colline to supramontane belts of the Carpathians and the Hercynicum) and Arunco dioici-Petasition albi (in the montane and supramontane belts of the Alps).
4. Senecioni rupestris-Rumicetalia alpini: Tall-herb anthropogenic vegetation on nutrient-rich soils in the upper montane to alpine belts with the single alliance Rumicion alpini.
Here we adopted the concept of four orders of
The main discrepancies occur for the communities classified here in the Petasito-Chaerophylletalia order (Groups 1‒4). The first problems concern the assignment of the Geranio phaei-Urticetum to the order Petasito-Chaerophylletalia. Phytocoenoses of this type were described for the first time by
An additional point of debate is the placement of the Petasitetum hybridi. Traditionally, this association is not placed in the class Mulgedio-Aconitetea, but considered as a lowland community, and thus placed in tall-herb classes of the lowlands. A typical stand of the Petasition officinalis according to
The next two associations that we distinguished (Chaerophyllo hirsuti-Petasitetum albi and Prenanthetum purpureae) overlap with the range of the broadly defined Chaerophyllo hirsuti-Cicerbitetum alpinae. However, in our opinion, differences in species composition and ecological characteristics fully justify their separation. The association Chaerophyllo hirsuti-Petasitetum albi is the central unit of the alliance, because of the large amplitude of the occupied habitat types, wide altitudinal range, and the species composition, determined by the dominance of Petasites albus, which is common in the Sudetes Mts., as well as the constant presence of species of the Carpino-Fagetea class. Since the Petasitetum albi is recognized as nomen ambiguum (
We propose to include both these associations in the order Petasito-Chaerophylletalia and in the alliance Petasition officinalis, instead of in the Calamagrostietalia villosae and the alliance Arunco dioici-Petasition albi, as proposed by
Group 5 embraces montane, nitrophilous phytocoenoses with a dominance of Rumex alpinus and corresponds to the Rumicetum alpini, an association regularly mentioned both in regional studies (
Group 6 includes relevés of mountain tall-grass communities that most often accompany local wetlands, spring zones and stream valleys in open areas. Kočí (
Group 7 represents alpine tall-grass stands in the Karkonosze Mts. The analyzed data was mainly obtained from the literature (
Group 8 embraces stands with Athyrium distentifolium and specific combination of acidophytes of the alliance Calamagrostion villosae and species of the Adenostylion alliariae. Following
Group 9 includes relevés from the highest parts of the Karkonosze Mts. We classified the communities from this group to the Cicerbitetum alpinae (within the Adenostylion alliariae alliance), which was also reported by
It should be noted that we had no data of the Arunco-Doronicetum austriaci from our study region (neither own records, nor literature data), which has been reported from other parts of the Sudetes Mts. (
In this comprehensive regional typology of the Mulgedio-Aconitetea in the Polish Sudetes Mts. we distinguished nine associations with relatively clear floristic and ecological separation. For convenience, we largely adopted the higher syntaxa of the current EuroVegChecklist (
All analyzed relevés (including environmental variables) are available on request through the VESTA Database (Global Index of Vegetation-Plot Databases, ID: EU–PL–004) and the Polish Vegetation Database (Global Index of Vegetation-Plot Databases, ID: EU-PL-001).
K.Ś. and K.R planned the research, K.Ś. and K.R conducted the field sampling, K.Ś. performed the statistical analyses and led the writing, while both authors critically revised the manuscript.
We are grateful to Jürgen Dengler, Kiril Vassilev and two anonymous Reviewers for their valuable comments on the manuscript.
List of species distinguishing communities of the class Mulgedio-Aconitetea from other non-forest communities based on literature sources
A detailed description of the TWINSPAN analysis used to distinguish the associations described in the present paper
Full, sorted relevé table of the studied tall-herb communities in the Sudetes Mts. (SW Poland)