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Diversity of Pinus pumila-dominated communities on the Kamchatka Peninsula

Jaanus Paal
Jaanus Paal
, 
Anton Pesterov
Anton Pesterov
 et 
Valentina Neshataeva
Valentina Neshataeva
   | 01 déc. 2021
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Article Category: Research paper
Publié en ligne: 01 déc. 2021
Pages: 26 - 57
Reçu: 09 juil. 2021
Accepté: 31 août 2021
DOI: https://doi.org/10.2478/fsmu-2021-0003
© 2021 by the authors., published by Sciendo
This work is licensed under the Creative Commons Attribution 4.0 International License.

Figure 1.

Location of sample areas on the Kamchatka Peninsula.
Location of sample areas on the Kamchatka Peninsula.

Figure 2.

Species and environmental variables ordination biplot by the DCA axes 1 and 2 (A), and axes 2 and 3 (B). Notations: Fert, Moist – habitat fertility and moisture level, Incl – habitat inclination, Alt – altitude, Exp – exposition; Shrub, Herb, DwShrub, Moss and Lichen – total cover of shrubs, herbs, dwarf-shrubs, mosses and lichens, respectively; Ppum – cover of Pinus pumila. The full names of the species are presented in Appendix 1.
Species and environmental variables ordination biplot by the DCA axes 1 and 2 (A), and axes 2 and 3 (B). Notations: Fert, Moist – habitat fertility and moisture level, Incl – habitat inclination, Alt – altitude, Exp – exposition; Shrub, Herb, DwShrub, Moss and Lichen – total cover of shrubs, herbs, dwarf-shrubs, mosses and lichens, respectively; Ppum – cover of Pinus pumila. The full names of the species are presented in Appendix 1.

Figure 3.

Community type centroids and environmental variables ordination biplot. Notations: group 1 – pure dwarf-pine communities, group 2 – shrub-rich communities, group 3 – dwarf-shrub-rich communities, group 4 – herb-grass-rich communities, group 5 – moss-rich communities, group 6 – lichen-rich communities; other notations as in Figure 2.
Community type centroids and environmental variables ordination biplot. Notations: group 1 – pure dwarf-pine communities, group 2 – shrub-rich communities, group 3 – dwarf-shrub-rich communities, group 4 – herb-grass-rich communities, group 5 – moss-rich communities, group 6 – lichen-rich communities; other notations as in Figure 2.

Environmental variables affecting the total number of species and projective cover of different vegetation layers according to the multiple regression analyses by stepwise entry of variables. Notations: β – standardized regression coefficient, SEβ – its standard error, B – unstandardized regression coefficient (slope), SEβ – its standard error, t – value of t-criterion, p – significance level, Mu R2 – multiple R2, Ad R2 – adjusted R2, F – value of F-criterion. Other notations as in Figure 2.

Variable β SEβ B SEβ t p Summary statistics
Mu R2 Ad R2 F p
Total number of species
Intercept 14.68 2.073 7.08 <0.001 0.156 0.134 7.23 <0.001
Ppum -0.22 0.057 -0.06 0.014 -3.84 <0.001
Moist2 1.29 0.370 1.42 0.407 3.50 0.001
Shrub -0.17 0.060 -0.04 0.013 -2.73 0.006
Alt 0.14 0.062 0.01 0.001 2.21 0.025
Moist -1.22 0.382 -6.816 2.125 -3.22 0.002
Fert 1.26 0.355 6.23 1.750 3.56 <0.001
Fert2 -1.10 0.332 -1.11 0.337 -3.30 0.001
Total cover of shrubs, square root transformed
Intercept -1.32 0.912 -1.44 0.150 0.136 0.126 14.55 <0.001
Alt 0.29 0.058 0.01 0.000 4.92 <0.001
Fert 0.93 0.282 2.61 0.795 3.28 0.001
Fert2 -0.74 0.284 -0.43 0.165 -2.59 0.010
Total cover of dwarf shrubs, square root transformed
Intercept 8.39 0.745 11.26 <0.001 0.344 0.335 36.38 <0.001
Fert -0.35 0.051 -1.20 0.173 -6.93 0.001
Ppum -0.27 0.049 -0.05 0.009 -5.49 <0.001
Shrub -0.23 0.052 -0.04 0.017 -4.42 <0.001
Alt 0.23 0.053 0.01 0.001 4.32 <0.001
Total cover of herbs, square root transformed
Intercept 3.32 0.663 5.00 <0.001 0.502 0.493 55.59 <0.001
Fert 0.44 0.047 1.28 0.137 9.32 <0.001
Alt -0.34 0.045 -0.01 0.000 -7.60 <0.001
Ppum -0.18 0.045 -0.03 0.007 -3.97 <0.001
DwShrub -0.16 0.049 -0.02 0.006 -3.22 0.001
Incl -0.10 0.044 -0.02 0.008 -2.38 0.018
Total cover of mosses, square root transformed
Intercept 2.55 0.625 4.08 <0.001 0.238 0.230 28.96 <0.001
Moist 0.41 0.053 1.70 0.222 7.67 <0.001
Herb -0.22 0.053 -0.04 0.008 -4.14 <0.001
Exp -0.15 0.053 -0.01 0.003 -2.87 0.004
Total cover of lichens, square root transformed
Intercept 10.34 0.806 12.83 <0.001 0.504 0.492 39.81 <0.001
Fert -0.79 0.272 -1.97 0.682 -2.89 0.004
Moist -1.27 0.294 -3.58 0.831 -4.31 <0.001
Moist2 1.01 0.284 0.57 0.159 3.57 <0.001
Ppum -0.20 0.043 -0.03 0.006 -4.65 <0.001
Alt 0.19 0.047 <0.01 0.000 4.08 <0.001
Shrub -0.16 0.047 -0.14 0.041 -3.44  0.001
Fert2 0.56 0.255 0.29 0.131 2.18  0.030

UT2

Species Max p Community type
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
Pinus pumila 1 0.001 7 6 6 5 5 5 5 6 5 6 6 6 5 6 7 6 5 4
Spiraea beauverdiana 2 <0.001 2 32 3 4 3 5 2 0 1 3 1 6 6 7 5 4 1 0
Plagiomnium cuspidatum 2 0.029 0 12 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Maianthemum dilatatum 3 <0.001 2 3 37 0 18 0 0 0 0 0 1 0 0 0 0 0 0 0
Oxalis acetosella 3 0.004 2 0 22 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0
Dryopteris expansa 3 0.009 0 0 20 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0
Brachythecium reflexum 3 0.017 0 0 13 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0
Phegopteris connectilis 3 0.028 0 0 12 0 0 0 0 0 0 0 0 4 0 0 0 0 0 0
Polytrichum commune 4 <0.001 0 1 0 38 6 1 0 0 1 2 1 0 2 6 18 3 1 1
Sorbus sambucifolia 5 <0.001 1 1 27 1 47 1 0 0 0 1 0 1 0 0 0 1 0 0
Gymnocarpium dryopteris 5 <0.001 0 0 10 0 35 0 0 0 0 0 0 0 0 0 0 0 0 0
Linnaea borealis 5 0.001 2 1 2 1 27 1 2 0 2 1 2 0 0 3 2 4 0 1
Cladonia squamosa 5 0.002 0 0 0 0 23 0 0 0 0 0 0 0 0 0 0 0 0 0
Lonicera chamissoi 5 0.003 0 0 0 0 23 0 0 0 0 0 0 0 0 7 0 0 0 0
Dicranum majus 5 0.010 0 0 1 3 17 0 0 0 0 0 2 0 0 0 10 2 0 0
Sanionia uncinata 5 0.022 1 0 7 0 14 3 2 0 0 0 0 0 0 0 11 1 0 2
Rhododendron aureum 6 <0.001 0 1 0 11 0 47 2 1 1 0 0 0 0 2 1 0 0 1
Dicranum scoparium 6 0.026 1 4 0 3 6 12 0 0 1 2 0 0 6 0 7 0 2 1
Vaccinium uliginosum 7 <0.001 0 1 0 1 0 1 26 4 11 2 0 0 1 1 2 5 17 5
Ceratodon purpureus 7 <0.001 0 0 0 0 0 0 31 0 1 0 0 0 0 0 0 0 0 0
Salix pulchra 7 <0.001 0 0 0 0 0 0 28 0 5 0 0 0 0 0 0 0 0 0
Saussurea pseudo-tilesii 7 0.001 0 0 0 0 0 0 30 0 0 0 4 0 0 0 1 0 0 0
Atragene ochotensis 7 0.001 0 0 0 0 0 0 27 0 1 0 0 0 0 0 0 0 0 0
Pyrola incarnata 7 0.003 0 0 0 0 0 0 21 0 3 0 0 0 0 0 0 0 0 0
Ribes triste 7 0.018 0 0 0 0 0 0 14 0 0 0 0 0 0 0 0 0 0 0
Carex koraginensis 7 0.024 0 0 2 0 0 0 14 1 0 0 0 0 0 0 0 0 0 0
Polytrichum juniperinum 7 0.026 0 0 0 0 0 0 14 1 0 4 0 0 0 0 0 0 0 0
Salix hastata 7 0.030 0 0 0 0 0 0 13 0 1 0 0 0 0 0 0 0 0 0
Racomitrium lanuginosum 7 0.036 0 0 0 0 0 0 14 0 0 0 0 0 0 0 0 0 0 0
Aulacomnium turgidum 7 0.037 0 0 0 0 0 0 13 0 0 1 0 0 0 0 0 0 4 0
Dicranum fulvum 7 0.045 0 0 0 0 0 0 14 0 0 0 0 0 0 0 0 0 0 0
Salix arctica 7 0.046 0 0 0 0 0 0 11 4 10 0 0 0 0 0 0 0 0 0
Vaccinium vitis-idaea 8 <0.001 0 0 0 5 1 6 5 29 5 4 0 0 0 5 1 3 11 5
Empetrum nigrum 9 <0.001 0 0 0 0 0 0 6 6 52 0 0 0 5 3 0 2 10 0
Cladonia gracilis 9 0.002 0 0 0 0 0 0 0 0 26 0 0 0 0 0 0 0 0 0
Sphagnum lenense 10 0.046 0 0 0 0 0 0 0 0 0 9 0 0 0 0 0 0 0 0
Lerchenfeldia flexuosa 11 0.001 0 0 0 0 1 0 0 0 0 0 28 0 0 2 0 0 0 0
Chamerion angustifolium 11 0.032 0 0 1 0 1 0 2 0 0 0 14 4 3 1 0 0 2 0
Calamagrostis purpurea 12 <0.001 2 11 13 0 1 0 0 0 0 1 7 39 7 0 2 1 0 0
Chamaepericlymenum suecicum 13 <0.001 1 0 0 0 12 0 0 0 1 0 1 0 44 0 0 0 0 0
Equisetum sylvaticum 13 <0.001 1 0 0 0 1 0 1 0 2 2 1 2 36 0 4 0 0 0
Rubus chamaemorus 13 <0.001 3 0 0 0 0 2 0 0 0 1 0 14 37 0 0 0 2 0
Carex globularis 13 0.002 0 0 0 0 0 0 0 0 0 3 0 0 30 0 0 0 4 0
Betula exilis 13 0.008 2 0 0 0 0 0 2 0 1 1 0 0 17 0 0 0 3 0
Dicranum sp. 14 <0.001 0 0 0 0 0 0 0 0 1 1 1 0 0 75 0 0 0 0
Mnium sp. 14 0.001 0 0 0 0 0 0 0 0 1 0 0 0 0 27 0 0 0 0
Cladonia uncialis 14 0.003 0 0 0 0 0 0 0 2 0 1 0 0 0 28 0 0 1 1
Dicranum flexicaule 15 0.011 1 0 0 8 0 0 0 0 0 0 0 0 0 0 15 0 1 0
Larix cajanderi 16 <0.001 0 0 0 0 0 0 6 0 4 1 0 0 0 0 0 35 0 0
Oxycoccus microcarpus 16 <0.001 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 33 0 0
Sphagnum fuscum 16 <0.001 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 33 0 0
Sphagnum girgensohnii 16 <0.001 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 88 0 0
Sphagnum capillifolium 16 <0.001 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 33 0 0
Poa pratensis 16 0.001 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 24 0 0
Plagiothecium laetum 16 0.002 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 25 0 0
Ledum palustre 16 0.002 0 0 0 2 0 2 9 1 5 7 0 0 0 4 1 19 16 17
Cladonia gonecha 16 0.009 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 17 0 0
Poa platyantha 16 0.016 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 13 4 0
Alnus fruticosa 16 0.020 0 0 0 0 0 3 0 3 0 0 1 0 0 0 0 11 0 0
Pleurozium schreberi 17 <0.001 0 2 0 7 10 5 1 0 3 16 3 0 1 6 4 4 26 1
Cladonia rangiferina 18 <0.001 0 0 0 0 0 1 2 3 3 2 1 0 0 1 0 0 1 41
Cladonia arbuscula 18 <0.001 0 1 0 0 0 1 0 1 3 1 1 0 0 0 0 0 1 54
Thamnolia vermicularis 18 <0.001 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 46
Polytrichum piliferum 18 0.001 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 0 32
Cladonia stellaris 18 0.001 0 0 0 0 0 0 0 0 0 0 5 0 0 0 0 0 0 25
Cetraria islandica 18 0.004 0 0 0 0 0 1 0 0 1 0 0 0 0 0 0 0 0 22
Nephroma arcticum 18 0.004 0 0 0 0 0 0 0 0 2 0 0 0 0 0 0 0 0 21
Flavocetraria nivalis 18 0.007 0 0 0 0 0 2 2 1 0 0 0 0 0 0 0 0 0 16
Aconogonon tripterocarpum 18 0.017 0 0 0 0 0 0 1 0 8 1 0 0 0 3 3 0 1 16
Hierochloë alpina 18 0.025 0 0 0 0 0 0 4 0 0 0 0 0 0 0 0 0 0 17

UT1

Species Groups of community types
Ppum pure Shrub-rich Dwarf-shrub-rich Herb-grass-rich Moss-rich Lichen-rich
Community types
1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18
Statistics
N 42 14 23 7 15 15 14 10 14 9 14 30 12 12 13 5 15 8
S 10 10 11 13 11 10 16 10 15 16 12 8 12 13 11 15 14 13
SDS 3 4 4 3 4 4 4 4 4 4 5 2 3 3 2 3 4 4
E 0.38 0.67 0.67 0.73 0.73 0.72 0.71 0.63 0.70 0.77 0.62 0.63 0.68 0.72 0.68 0.66 0.64 0.77
SDF 0.14 0.10 0.07 0.12 0.11 0.11 0.05 0.10 0.08 0.05 0.12 0.09 0.06 0.07 0.07 0.10 0.05 0.07
Trees
Pinus pumila 81.3 68.0 64.7 62.5 55.7 57.0 61.4 65.0 52.9 70.9 63.8 69.0 58.7 69.0 80.7 68.3 54.4 47.9
Betula ermanii – – 0.3 – – – – – – 0.3 0.8 – – – – – 0.6 –
Larix cajanderi 0.1 – – – – – 2.1 – 1.4 0.8 0.4 – – 0.5 0.4 5.0 – –
Shrubs
Alnus fruticosa 0.2 – – – – 0.9 0.4 0.8 0.4 0.1 0.8 – – – – 1.7 – –
Betula divaricata 0.1 – – – 0.7 – – 0.7 – 0.4 – 0.1 – – – – 0.6 –
Betula exilis 0.7 – – – – – 0.9 0.4 0.3 0.4 – – 2.4 <0.1 – – 2.2 –
Lonicera caerulea 0.3 0.8 – 0.6 0.1 – 0.6 – – 0.3 2.8 1.0 0.2 – – – 0.7 –
Lonicera chamissoi <0.1 – – 1.9 – – – – – – <0.1 – – 1.1 – – – –
Rhododendron aureum 1.0 2.7 0.7 0.1 22.9 41.7 6.1 2.4 2.0 1.4 – – – 4.3 2.5 – 0.2 1.8
Ribes triste – – – – – – 1.1 – – – – – – – – – – –
Salix hastata – – – – – – 0.9 – 0.1 – – – – – – – – –
Salix arctica <0.1 – – – – 0.4 1.6 1.3 1.7 <0.1 – – – – – – – –
Salix pulchra – – – – – – 2.6 – 0.8 – – – – – – – – –
Salix caprea – – – – – 0.4 – – – – – – – – – – – –
Salix crassijulis – – – – – – – – – – – – – 1.0 – – – –
Sorbus sambucifolia 1.1 1.7 15.7 25.1 2.1 1.7 – – – 1.3 – 2.1 <0.1 <0.1 0.7 1.7 0.6 0.1
Juniperus sibirica <0.1 – – – – – – – <0.1 <0.1 0.8 – – – – – 1.1 –
Spiraea beauverdiana 2.8 37.3 5.7 6.3 5.5 5.8 3.1 0.4 1.4 5.3 1.3 9.5 7.2 10.8 6.3 5.2 1.9 0.7
Dwarf shrubs
Aconogon tripterocarpum <0.1 <0.1 – – – – <0.1 <0.1 0.5 <0.1 – – – 0.3 0.2 – 0.1 0.7
Arctous alpina – <0.1 – – – 0.9 1.4 1.9 1.0 0.1 – – – – – – 0.2 2.0
Empetrum nigrum 0.6 0.8 – <0.1 – 0.2 5.1 4.9 32.1 0.8 – – 3.8 2.7 1.4 2.4 6.9 –
Ledum palustre 0.3 0.7 – – 2.3 2.3 7.9 1.8 4.4 7.6 <0.1 – 0.8 4.0 1.3 15.7 13.6 14.4
Loiseleuria procumbens – – – – – – 0.9 0.4 <0.1 – – – – – – – – –
Phyllodoce caerulea – 0.7 – – – 0.9 1.1 – – – – – – – – – – –
Vaccinium vitis-idaea 0.6 0.8 – 1.5 5.1 5.8 4.5 22.1 4.5 4.6 <0.1 – 0.8 6.6 2.7 2.6 9.7 4.4
Vaccinium uliginosum 0.4 2.5 – – 1.4 1.7 20.0 5.5 9.4 3.1 <0.1 – 1.1 1.4 3.7 5.6 14.6 5.4
Herbs and grasses
Aconogonon tripterocarpum <0.1 <0.1 – – – – <0.1 <0.1 0.5 <0.1 – – – 0.3 0.2 – 0.1 0.7
Artemisia arctica – – – – – – 0.4 <0.1 <0.1 – <0.1 – – – – – – –
Artemisia opulenta – – 0.3 – – – – – – – – – – – – – – –
Aruncus kamtschaticus – – 0.3 – – – – – – – – – – – – – – –
Atragene ochotensis – – – – – – <0.1 – <0.1 – – – – – – – – –
Bromopsis pumpelliana <0.1 – – – – – 0.7 <0.1 – – – – – – – – – –
Calamagrostis purpurea 1.9 9.7 11.3 1.5 0.9 0.5 0.4 0.5 <0.1 1.1 8.5 31.5 8.3 0.6 2.4 2.3 – <0.1
Carex globularis <0.1 – – – – – – – <0.1 0.9 – – 2.6 – – – 0.4 –
Carex koraginensis – – – – – – 0.1 – – – – – – – – – – –
Carex pallida – – – – – – 1.5 <0.1 0.2 0.2 1.5 – – – <0.1 – – –
Carex sp. – – – – – – – – – – – – 0.3 0.1 – – – –
Chamaepericlymenum suecicum 1.0 – – 10.0 – – <0.1 – 1.1 0.1 1.2 0.2 18.0 0.7 – – – –
Chamerion angustifolium 0.2 <0.1 0.3 0.2 – – 0.5 <0.1 <0.1 <0.1 1.9 1.3 0.5 0.7 – – 0.6 –
Cassiope lycopodioides – – – – – – – – – – – – – – – – – 0.7
Cirsium kamtschaticum – – 0.3 – – – – – – – – – – – – – – –
Cacalia kamtschatica – – 0.3 – – – – – – – – – – – – – – –
Dryopteris expansa 0.1 – 6.0 – – – – – – – – <0.1 <0.1 – – – – –
Deschampsia borealis – – – – – – – – – – 1.9 – – – – – – –
Equisetum arvense <0.1 – <0.1 <0.1 <0.1 – <0.1 – – – 0.2 0.1 1.1 0.1 0.7 <0.1 1.3 –
Equisetum sylvaticum 0.9 0.7 – 1.3 0.6 – 1.1 0.4 1.5 1.5 0.6 1.1 10.2 – 2.9 – 0.1 –
Glyceria alnasteretum – – <0.1 – – – – – – – – 0.5 – – – – – –
Gymnocarpium dryopteris <0.1 <0.1 5.1 13.8 0.4 – – – – 0.2 – – – <0.1 – – – –
Hierochloë alpina – – – – – – 0.5 – – – – – – – – – – 0.5
Lerchenfeldia flexuosa 0.1 – 0.3 0.3 – – – – <0.1 0.2 5.0 – 0.1 0.7 – – – –
Linnaea borealis 1.3 0.8 1.5 12.5 0.7 0.8 1.4 – 1.2 1.1 1.3 <0.1 0.2 1.7 1.2 1.5 0.2 1.0
Lycopodium annotinum 0.5 3.1 2.0 1.6 1.3 <0.1 – – 0.2 0.5 0.8 – – 1.5 <0.1 1.2 – –
Lycopodium juniperoideum – 0.7 – – – 0.1 – – – <0.1 – – – 0.5 1.4 – – –
Maianthemum dilatatum 1.8 1.0 9.6 4.6 <0.1 0.1 – – <0.1 0.1 0.5 0.3 – 0.2 0.2 0.3 0.2 –
Oxalis acetosella 0.1 – 0.5 <0.1 – – – – – – – 0.1 – – – – – –
Oxycoccus microcarpus – – – – – – – – – – – – – – – 0.1 – –
Phegopteris connectilis – <0.1 5.0 – – – – – – – – 3.0 – – – – – –
Poa malacantha – – – – – – 0.8 <0.1 – – – – 0.7 – – – – –
Poa platyantha – – – – – – – <0.1 <0.1 0.1 – – – 0.1 – 0.7 0.2 –
Poa pratensis – – – – – – – – – – – – – – – 0.1 – –
Pyrola incarnata 0.1 – – – – – 0.4 – 0.4 – – – – – – – – –
Rubus chamaemorus 1.8 0.7 – – 0.4 1.4 – 0.8 0.5 1.1 0.1 4.8 8.1 – – – 0.7 –
Rubus arcticus 0.2 <0.1 – – – – – – – <0.1 0.9 0.2 <0.1 1.5 1.2 – – <0.1
Saussurea pseudo-tilesii <0.1 <0.1 <0.1 – – – 1.2 <0.1 <0.1 – 0.5 – – – 0.1 – – –
Thalictrum minus – – <0.1 – – – – – – – 1.5 – – – – – – –
Trisetum sibiricum – – 0.3 – – – – – – – – – – – – – – –
Trientalis europaea 0.5 1.1 1.2 1.0 0.1 <0.1 – – <0.1 0.6 0.5 0.3 <0.1 <0.1 <0.1 – <0.1 –
Bryophytes
Aulacomnium turgidum <0.1 – – – – – 0.7 <0.1 – 0.3 – – – – – – 0.6 –
Brachythecium oedipodium <0.1 – – – – – – <0.1 – 0.1 <0.1 1.0 2.1 – – – – –
Brachythecium reflexum <0.1 – 1.1 – – – – – <0.1 – 0.4 0.3 – <0.1 0.4 – – –
Ceratodon purpureus 0.2 – – – – – 7.9 – 1.1 – – – – – – – – –
Dicranum bergeri – – – – – – 1.1 – <0.1 0.6 – – – – – – – –
Dicranum elongatum – – – – – – – <0.1 0.4 1.1 0.4 – – – – – – –
Dicranum flexicaule 0.3 – – – 1.3 – – – – 0.1 – – – – 2.5 – 0.6 –
Dicranum fragilifolium – – 0.1 – – – – – <0.1 0.3 0.8 0.2 – – 0.4 0.7 – –
Dicranum fulvum – – – – – – 0.7 – <0.1 – – – – – – – – –
Dicranum fuscescens – – 0.7 – – 0.2 – 1.7 – 0.1 1.2 0.6 – – – – <0.1 1.6
Dicranum majus – – 0.6 8.1 1.9 0.6 – – 0.4 0.4 1.4 0.2 – – 3.6 0.9 – 0.4
Dicranum scoparium 1.2 4.7 0.2 6.9 3.6 11.5 0.4 0.4 1.4 2.9 0.5 0.7 6.2 – 7.9 – 3.1 1.8
Dicranum bonjeanii – – – – – – – – – – – 0.3 – – – – – –
Dicranum sp. 0.7 0.7 0.4 – 0.4 0.7 – 0.8 1.6 1.2 1.3 – 0.3 28.5 0.4 – 1.1 –
Hylocomium splendens – – – – 0.1 0.9 <0.1 – 1.4 – <0.1 – – – – <0.1 – –
Hypnum sibiricum – – – – – – – – – – – – – 0.5 – – – –
Hypnum plicatulum – – – – 0.1 1.3 – – – – <0.1 – – – – 0.9 – –
Hypnum cupressiforme – – – – – – – – – – – – – 0.5 – – – –
Mnium thomsonii – – – – – – – – 0.4 – – – – 3.5 – – – –
Plagiomnium cuspidatum – 0.5 – – – – – – – <0.1 – – – – – – – –
Plagiothecium laetum – – – – – – – – – – – – – – – 0.4 – –
Pleurozium schreberi 1.4 8.3 1.1 25.6 16.4 14.6 5.4 0.4 14.6 39.9 6.9 1.0 3.2 17.1 12.1 10.5 65.6 4.3
Polytrichastrum alpinum – – – – – – – – 0.4 – <0.1 – – – – – – –
Polytrichum jensenii – – <0.1 – – – 0.9 – 2.5 – – – – – – – – –
Polytrichum juniperinum <0.1 – – – – – 1.1 0.4 <0.1 1.1 – – – 0.1 – – – –
Polytrichum piliferum – – – – – – – 0.4 – 0.2 <0.1 – – – – – – 1.2
Polytrichum commune 1.3 4.3 1.0 10.0 50.0 3.4 0.4 2.5 3.1 4.5 2.4 1.3 4.3 9.5 23.6 5.4 2.0 1.6
Ptilidium ciliare 0.2 – – – – – – – <0.1 <0.1 0.1 – – 0.1 0.7 0.3 – –
Racomitrium lanuginosum – – – – – – 0.8 <0.1 <0.1 – – – – – – – – –
Rhytidium rugosum – – – – – – – <0.1 – – – – – – – – 1.1 –
Sanionia uncinata 0.5 – 3.3 4.6 – 3.5 1.2 0.4 <0.1 0.3 0.1 – – – 4.4 0.7 – 1.1
Sphagnum capillifolium – – – – – – – – – – <0.1 – – – – 4.6 – –
Sphagnum fuscum – – – – – – – – – – – – – – – 3.3 – –
Sphagnum girgensohnii – – – – 2.9 1.0 – – 0.4 0.5 1.5 – 0.3 0.5 0.7 60.8 0.6 –
Sphagnum lenense – – – – – – – – – 1.8 – – – – – – – –
Sphagnum fimbriatum – – – – – – – – – – – – – – – – 4.4 –
Lichens
Cetraria islandica – – – – – 0.3 – – 0.2 <0.1 – – – – – – – 2.2
Cladonia gonecha – – – – – – – – – – – – – – – 0.1 – –
Cladonia gracilis – <0.1 – – – – <0.1 – 1.2 – – – – – – – <0.1 –
Cladonia grisea – – – – – – – – – 0.2 – – – – – – – –
Cladonia rangiferina <0.1 0.7 – – 0.4 1.3 1.7 2.4 2.8 1.8 1.2 – – 1.5 0.4 0.5 0.4 13.9
Cladonia stellaris – – – – – – – <0.1 <0.1 0.3 3.5 – – 0.5 – – – 6.1
Cladonia arbuscula <0.1 1.7 – – 0.4 1.5 0.6 1.1 2.3 1.4 1.8 – – 1.0 0.1 0.4 0.7 18.2
Cladonia ecmocyna – – – – – – 0.4 – <0.1 0.3 <0.1 – – – 0.2 <0.1 0.3 0.1
Cladonia uncialis – – <0.1 – – – <0.1 0.1 – <0.1 – – – 1.1 – – 0.1 0.2
Cynodontium strumiferum – – – – – – <0.1 – – – – – – 1.0 – – – –
Flavocetraria nivalis – – – – – 0.7 0.6 0.7 – <0.1 <0.1 – – – – – – 2.5
Nephroma arcticum – – – – – – – – 0.4 – – – – – – – – 1.1
Stereocaulon alpinum – – – – – <0.1 – 0.4 2.9 – – – – – – – – 1.9
Stereocaulon paschale – <0.1 – – <0.1 – – 3.3 <0.1 0.4 <0.1 – – 1.1 – – – 0.7
Thamnolia vermicularis – 0.3 – – – <0.1 – <0.1 0.3 <0.1 <0.1 – – <0.1 – – <0.1 3.4

Median values of exposition, moisture and fertility, and average values of other environmental variables of community types. Notations: Ppum – Pinus pumila pure, Shrub – shrub-rich, DwShrub – dwarf-shrub-rich, HGrass – herb- and grass-rich, Moss – moss-rich, Lichen – li-chen-rich; S – average number of species in communities, E – evenness coefficient; Alt, Expos, Slope, Moist, Trophy – habitats’ exposition, slopeness, humidity and fertility, respectively; CovShr, CovHerb, CovDwShr, CovMoss, CovLich – average cover of shrubs, herbs+grasses, dwarf-shrubs, mosses and lichens in communities. With uppercase letters are marked similar average values according to the Fisher LSD post-hoc tests or similar medians according to the Duncan’s post-hoc tests (for Expos, Moist and Trophy); pANOVA – significance level by the univariate ANOVA, pKW – significance level by the Kruskal-Wallis post-hoc median test.

Community type Type group Variables
S E Alt Expos Slope Moist Trophy CovShr CovHerb CovDwShr CovMoss CovLich
1 Ppum 10ab 0.382a 359bce E 8ab 2b 3efgh 6ab 11bc 2a 6a 0a
2 Shrub 10abce 0.666bcde 536dfg E 6ab 2bc 3efghi 41g 17cd 6ab 19bcd 3ab
3 Shrub 11abcde 0.668bcde 244ab E 8ab 2bc 3jk 22ef 43e 0a 8abc 0a
4 Shrub 13bcdefgh 0.727def 253abc NE 9abc 2b 3hij 34g 44e 2a 55f 0a
5 Shrub 11abcdef 0.730def 564defg E 11abc 2bc 2cdef 31fg 4ab 9abc 77gh 1ab
6 Shrub 10abce 0.724df 868i SW 17c 2b 2cdef 48h 3a 12bc 38e 4abc
7 DShrub 16i 0.707cdef 754hi SW 8ab 2b 2.5efgh 13de 10abc 41e 20cd 3abc
8 DShrub 10abc 0.633bc 497cdef E 14bc 1a 1ab 5abc 2a 37e 7abc 8bc
9 DShrub 15hi 0.697cdef 558dfgh SE 6ab 2b 2bcde 6abc 6ab 51f 28de 10c
10 DShrub 12cdef 0.653bcd 393bcde NE 10ab 2bc 2def 9abcd 8ab 16c 55f 4abc
11 HGrass 12bcdefg 0.616b 396bcde SW 14bc 2b 4j 5abc 22d 0a 17abcd 5abc
12 HGrass 8a 0.633bc 115a NE 6ab 2bc 4k 13bcde 42e 0a 6abc 0a
13 HGrass 12bcdefg 0.678bcde 81a plain 4a 2bc 3ghij 9abcd 50e 6ab 16abd 0a
14 Moss 13defghi 0.715cdef 510cdef NE 10ac 2b 2bcd 16de 9abc 15bc 60fg 5abc
15 Moss 11bcdef 0.678bcde 671fgh NE 6ab 2bc 3fghi 9abcd 11abc 9abc 57f 1a
16 Moss 15ghi 0.660bcde 450bcde SW 27d 4d 2bc 6abcd 6ab 26d 86h 1ab
17 Moss 14fghi 0.643bce 251ab W 12abc 2c 2efg 5abc 4ab 45ef 73gh 2ab
18 Lichen 13dfgh 0.769f 737ghi SE/E 13bc 1a 1a 3a 2a 26d 11abc 51d
pANOVA <0.001 <0.001 <0.001 <0.001 <0.001 <0.002 <0.003 <0.004 <0.005
pKW 0.741 <0.001 <0.001

Correlation of environmental variables with DCA ordination axes. Notations: r – Pearson correlation coefficient, τ – Kendall rank order correlation coefficient; other notations as in Figure 2.

Variable Axis 1 Axis 2 Axis 3
Statistics
r r2 τ r r2 τ r r2 τ
Moist  0.107  0.012  0.124 -0.223  0.050 -0.136 -0.455  0.207 -0.385
Fert  0.605  0.366  0.490 -0.067  0.004 -0.051 -0.294  0.087 -0.196
Exp  0.203  0.041  0.160 -0.013  0.000  0.005 -0.128  0.016  0.067
Incl -0.188  0.035 -0.159  0.014  0.000  0.019  0.046  0.002  0.036
Alt -0.346  0.119 -0.245  0.324  0.105  0.204  0.200  0.040  0.108
Shrub  0.218  0.047  0.187  0.649  0.421  0.405 -0.143  0.020 -0.122
DwShrub  0.593  0.352  0.423 -0.184  0.034 -0.146 -0.135  0.018 -0.131
Herb -0.778  0.605 -0.663 -0.234  0055 -0.116  0.214  0.046  0.149
Moss -0.306  0.094 -0.211  0.151  0.023  0.166 -0.708  0.501 -0.524
Lichen -0.463  0.215 -0.461  0.084  0.007  0.004  0.470  0.221  0.207

Exposition of habitats of different type communities. Type group notations as in Figure 3.

Community type Type group Exposition Plain Total
N NW W SW S SE E NE
1 P. pumila 2 3 4 8 3 2 1 21 44
2 Shrub 2 1 3 1 1 1 6 15
3 Shrub 5 3 2 15 25
4 Shrub 1 4 2 1 2 2 5 17
5 Shrub 1 1 1 1 1 2 7
6 Shrub 2 1 1 4 8
7 DwShrub 5 2 1 3 1 9 21
8 DwShrub 2 1 2 1 1 1 1 6 15
9 DwShrub 2 1 3 1 8 15
10 DwShrub 1 4 5 4 1 1 3 3 9 31
11 HGrass 7 1 3 1 2 14
12 HGrass 1 3 1 1 1 5 12
13 HGrass 1 2 1 1 5
14 Moss 4 2 1 1 1 3 1 13
15 Moss 1 1 1 1 1 9 14
16 Moss 4 1 1 1 1 8
17 Moss 1 2 2 2 1 1 1 10
18 Lichen 4 1 2 1 8
Total 26 18 39 22 17 9 23 22 106 282
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