Corresponding author: Kamila Reczyńska (
Academic editor: Jürgen Dengler
Montane to subalpine tall-herb communities of Europe, Siberia and Greenland are classified in the class
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
Besides the above-mentioned associations, from the Polish side of the Sudetes Mts. and their foothills the following association-level syntaxa were reported:
The aims of this study are thus: i) to conduct a comprehensive analysis of diversity of tall-herb communities of the
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
Area of the investigation. Red dots show the distribution of the analyzed relevés of tall-herb communities (
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
Additionally, we used all the available relevés from the literature (
The relevés are available
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,
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 (
To identify the main ecological drivers affecting the diversity of distinct groups, a distance-based redundancy analysis (
In the analyzed data we could distinguish nine groups of tall-herb communities (Table
Cl.
O.
All.
Group 1:
Group 2:
Group 3:
Group 4:
O.
All.
Group 5:
O.
All.
Group 6:
Group 7:
Group 8:
O.
All.
Group 9:
The nine associations clearly differ in terms of diagnostic species (Table
Distribution maps of the distinguished associations of tall-herb communities of the Sudetes Mts. and their foothills. The numbers refer to the group IDs used throughout this paper. Background from © MapTiler © OpenStreetMap contributors.
Tall-forb vegetation belonging to the
Tall-forb vegetation belonging to the
Communities of
Summarized synoptic table with percentage frequency and fidelity values derived from 399 relevés of tall-herb associations of the
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 |
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Alliance |
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14 | 100 ** | 7 | 1 | . | . | . | . | . | . |
4.5 | 55 ** | 12 | 5 | . | . | . | . | . | . | |
2.5 | 52 ** | 20 | 5 | 4 | . | . | . | . | . | |
3.1 | 42 ** | 14 | . | . | . | . | . | . | . | |
5.9 | 39 * | 2 | 4 | 6 | . | . | . | 2 | 7 | |
3.3 | 36 * | 7 | 11 | 4 | . | . | . | 2 | . | |
17 | 30 ** | 2 | . | . | . | . | . | . | . | |
17 | 30 ** | 2 | . | . | . | . | . | . | . | |
5.0 | 27 * | 2 | 5 | . | . | . | . | . | . | |
100 | 21* | |||||||||
|
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24 | . | 100 ** | 4 | . | . | . | . | . | . |
1.5 | 9 | 56 * | 36 | 14 | . | 8 | . | 2 | ||
|
100 | . | 41 ** | . | . | . | . | . | . | . |
1.8 | 9 | 31 * | 7 | . | . | . | 2 | . | 17 | |
1.9 | 12 | 24 * | 1 | . | . | . | . | . | . | |
|
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5.7 | 3 | 12 | 99 ** | 69 | . | 8 | . | 8 | 7 |
1.3 | 12 | 17 | 46 * | 35 | . | 17 | . | 3 | . | |
1.6 | 27 | 25 | 45* | 24 | . | . | . | . | . | |
3.2 | 9 | 12 | 45 * | 14 | . | . | . | . | . | |
1.6 | . | 2 | 28 * | 18 | . | 17 | . | 2 | . | |
2.6 | 6 | 10 | 27 * | 8 | . | . | . | 2 | . | |
|
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1.8 | . | . | 16 | 53 * | . | 25 | . | 27 | 7 | |
1.8 | . | 5 | 24 | 49 * | 5 | 42 | . | . | . | |
|
1.3 | . | 2 | 18 | 47* | . | 17 | 11 | 34 | 17 |
2.5 | . | . | 8 | 47 * | . | 17 | 7 | 19 | . | |
1.6 | . | . | 4 | 41 * | . | 8 | 7 | 25 | 10 | |
4.0 | . | . | 4 | 33 * | . | 8 | . | 3 | 7 | |
1.5 | . | 1 | 15 | 24 * | . | . | . | . | 2 | |
2.1 | . | . | 7 | 22 * | . | . | . | 10 | . | |
|
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51 | . | . | . | 2 | 100** | . | . | . | . |
|
100 | . | . | . | . | 35** | . | . | . | . |
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2.9 | . | 3 | 8 | 6 | 20 | 58 ** | . | 3 | . | |
100 | . | . | . | . | . | 42 ** | . | . | . | |
100 | . | . | . | . | . | 33 ** | . | . | . | |
|
1.3 | 4 | 10 | 33 * | 15 | 27 | ||||
17 | . | 2 | 1 | 2 | . | 33 * | . | . | . | |
6.5 | . | 2 | 3 | . | . | 33 * | . | 5 | . | |
|
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100 | . | . | . | . | . | . | 64 ** | . | . | |
|
9.7 | . | . | . | . | . | . | 61 ** | 2 | 7 |
|
100 | . | . | . | . | . | . | 54 ** | . | . |
31 | . | . | . | . | . | . | 54 ** | 2 | . | |
22 | . | . | . | 2 | . | . | 44 ** | 2 | . | |
1.3 | . | . | 22 | 12 | . | 17 | 41 * | 12 | 30 | |
25 | . | . | 1 | . | . | . | 34 ** | . | . | |
2.5 | . | . | 3 | 14 | . | . | 34 * | 14 | . | |
1.7 | . | . | 3 | . | 20 | . | 34 * | 5 | 20 | |
|
100 | . | . | . | . | . | . | 31 ** | . | . |
|
100 | . | . | . | . | . | . | 21 * | . | . |
100 | . | . | . | . | . | . | 21 * | . | . | |
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4.7 | . | 2 | 11 | 2 | . | . | 15 | 3 | 70 ** |
2.3 | 9 | 24 | 8 | 2 | 20 | . | 7 | 2 | 57 * | |
|
5.3 | . | . | 1 | . | 10 | . | . | . | 53 ** |
3.5 | 3 | 2 | 4 | 6 | 15 | 8 | . | 12 | 53 * | |
3.6 | . | . | . | . | 5 | . | 15 | 5 | 53 * | |
|
5.2 | . | 5 | 9 | 2 | . | 8 | . | . | 50 * |
2.1 | 12 | . | 20 | 18 | . | 8 | 11 | 19 | 43 * | |
4.7 | . | 2 | 4 | . | . | 8 | 7 | 3 | 40 * | |
1.4 | . | 20 | 1 | 6 | . | . | . | 3 | 30 * | |
|
8.1 | . | . | . | . | . | . | 3 | 2 | 27 * |
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1.9 | 85** | 68 * | 35 | 14 | . | . | . | . | . | |
23 | 39* | 32 * | 1 | . | . | . | . | . | . | |
1.3 | 9 | 34 * | 7 | 2 | 25 | 8 | . | 2 | 3 | |
1.4 | 27* | 3 | 19 | 6 | . | . | . | . | . | |
1.3 | 18 | 25 * | 4 | . | . | . | . | . | . | |
3.6 | 6 | 2 | 22* | 6 | . | . | . | . | . | |
2.8 | 6 | 3 | 17* | 4 | . | . | . | . | . | |
1 | 24 | 7 | 36 * | 37 | 5 | . | . | 8 | . | |
1 | 48 | 8 | 53 * | 61 * | . | 17 | . | 2 | . | |
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3.9 | . | . | . | . | 65 ** | 17 | 5 | 7 | 3 | |
4.1 | . | . | . | . | 35 * | 8 | 5 | . | . | |
1.5 | 15 | 24 | 1 | . | 35 * | . | . | . | . | |
2 | 6 | 10 | 22 | 22 | 45 * | 8 | . | 8 | 3 | |
3 | 3 | 3 | . | 2 | 30 * | . | 2 | . | 9 | |
|
7.5 | . | . | . | . | 25** | . | . | . | 3 |
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1.7 | . | . | . | 4 | 5 | 17 | 95 ** | 58 * | 33 | |
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1.3 | . | . | 15 | 69 | 20 | 50 | 92 * | 92 * | 50 |
1.6 | . | . | 4 | 20 | . | 33 | 90 ** | 56 * | 6 | |
|
1.4 | . | . | . | 4 | 5 | . | 57 * | 49 * | 33 |
1.9 | . | . | . | . | . | 8 | 67 * | 37 * | . | |
2.5 | . | 3 | 1 | 6 | 15 | 25 | 92 ** | 27 | 37 | |
3.4 | . | . | 4 | 10 | . | 8 | 82 ** | 24 | 7 | |
2.7 | . | . | 1 | 18 | 5 | 8 | 79 ** | 29 | . | |
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. | . | . | 4 | . | 8 | 2 | 8 | . | |
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. | . | . | . | . | . | 11 | 3 | 3 | |
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4.7 | . | . | 11 | 84 * | . | 8 | 15 | 69 * | 87 * |
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1.9 | . | 2 | 7 | 10 | . | 8 | 74 * | 37 | 73 * |
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1.7 | . | . | 1 | . | . | 33 | 21 | 14 | 87** |
|
1.9 | . | . | 1 | . | 20 | . | 26 | 44 | 87 ** |
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3.8 | . | . | 4 | 2 | . | . | 13 | 7 | 50 ** |
1.4 | 3 | 2 | 3 | 2 | . | . | 30 | 5 | 43 * | |
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2.7 | . | 15 | 7 | . | . | 17 | 28 | 6 | 77 ** |
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19 | . | 2 | 4 | 2 | 85* | . | 79* | 78* | 100 * |
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6.3 | . | . | 4 | 8 | 10 | . | 64* | 97 ** | 67 * |
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2.2 | . | 3 | 18 | 31 | . | 8 | 82* | 71 * | 77 * |
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2.9 | 12 | 17 | 7 | 2 | 10 | 8 | 15 | 51 * | 63 * |
1.6 | . | . | . | . | 5 | 8 | 57 * | 5 | 37 * | |
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1.9 | 15 | 5 | 14 | 8 | . | 8 | . | . | 30 * |
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1.7 | . | . | . | . | 30 | 25 | . | 3 | 53 * |
4.6 | 3 | . | 11 | 4 | . | . | 2 | 5 | 50 * | |
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2.0 | 9 | 3 | 11 | 20 | 5 | . | 25 | 2 | 50 * |
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3.0 | . | 2 | 4 | . | . | 8 | 11 | 12 | 37* |
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1.4 | 21 | 34 | 69 | 69 | 70 | 100 * | 2 | 36 | 60 |
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1.5 | . | . | 19 | 27 | . | . | 46* | 27 | 30 |
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1.1 | . | 2 | 9 | . | 65 | 75 | 75 | 86 * | 93 * |
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21 | 49 | 77 | 84 | 50 | 83 | . | 14 | 77 | |
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. | . | 15 | 33 | . | 8 | 20 | 24 | 40 * | |
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21 | 20 | 27 | 18 | . | 8 | 2 | . | 23 | |
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. | 2 | 9 | 4 | . | 17 | . | 2 | 3 | |
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. | 3 | 3 | 2 | . | . | . | . | . | |
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. | 2 | 5 | 2 | . | . | . | . | 3 | |
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1.3 | 21 | 7 | 59 | 90 * | 10 | 92 * | 11 | 83 * | 1 |
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4.0 | . | 14 | . | . | 15 | 8 | 92* | 61* | 63* |
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1.6 | 30 | 19 | 69 * | 71 * | 25 | 42 | 2 | 24 | 7 |
1.3 | 6 | 7 | 72 * | 69 * | 5 | 50 | . | 10 | . | |
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1.4 | 9 | 15 | 9 | 14 | 85 * | 92 * | 48 | 39 | 60 |
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3 | 27 | 41 | 49 | 15 | 17 | 3 | 8 | 33 | |
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9 | 8 | 42 | 41 | 25 | 8 | 30 | 61 | 60 | |
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. | 10 | 4 | 4 | 35 | 17 | 41 | 15 | 30 | |
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1 | 21 | 36 | 30 | 29 | 55 | 58 * | . | 8 | 3 |
1.2 | . | 17 | 26 | 27 | 15 | 67 * | 5 | 8 | 53 | |
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1.1 | 91 * | 78 * | 65 | 35 | 70 | 25 | . | 5 | 20 |
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1.1 | . | 3 | 14 | 35 | . | 42 * | 3 | 15 | 7 |
9 | 14 | 3 | 4 | . | . | 2 | 14 | 12 | ||
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1.1 | 27 | 39 | 15 | 8 | 45 * | 25 | . | 3 | . |
|
. | 14 | 41 | 29 | 15 | 33 | . | 7 | 30 |
Significance of Spearman’s rank correlation of mean EIVs with three main
Axis 1 | Axis 2 | Axis 3 | ||||
---|---|---|---|---|---|---|
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|
|
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|
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- |
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0.32 | 0.220 | -0.08 | 0.696 | |
- |
|
0.34 | 0.164 | 0.32 | 0.108 | |
- |
|
0.29 | 0.300 | 0.39 | 0.052 | |
ElV light | 0.46 | 0.192 |
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0.33 | 0.056 | |
-0.48 | 0.164 | -0.12 | 0.624 |
|
1 Spearman’s
Summary box-and-whisker plots of mean Ellenberg indicator values (EIVs) for clusters recognized within
In the following, among the diagnostic species, the character species (in bold) are highlighted (see Table
Number of relevés: 33
Number of relevés: 59
Number of relevés: 74
Number of relevés: 51
Number of relevés: 20
Number of relevés: 12
Number of relevés 61
Number of relevés: 59
Number of relevés 30
As shown in the
The
The
The simple term and conditional effects of analyzed environmental variables on species composition of the distinguished montane tall-herb communities, identified using
Environmental variable | Simple Term Effects | Conditional effects | ||||
---|---|---|---|---|---|---|
Lambda | pseudo- |
Lambda | pseudo- |
|||
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
Hitherto, in synthetic studies concerning the
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
1.
2.
3.
4.
Here we adopted the concept of four orders of
The main discrepancies occur for the communities classified here in the
An additional point of debate is the placement of the
The next two associations that we distinguished (
We propose to include both these associations in the order
Group 5 embraces montane, nitrophilous phytocoenoses with a dominance of
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
Group 9 includes relevés from the highest parts of the Karkonosze Mts. We classified the communities from this group to the
It should be noted that we had no data of the
In this comprehensive regional typology of the
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.
Krzysztof Świerkosz (krzysztof.swierkosz@uwr.edu.pl), ORCID: https://orcid.org/0000-0002-5145-178X
Kamila Reczyńska (Corresponding author, kamila.reczynska@uwr.edu.pl), ORCID: https://orcid.org/0000-0002-0938-8430
List of species distinguishing communities of the class
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)