Diversity of Pinus pumila-dominated communities on the Kamchatka Peninsula

Jaanus Paal; Anton Pesterov; Valentina Neshataeva

doi:10.2478/fsmu-2021-0003

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Diversity of Pinus pumila-dominated communities on the Kamchatka Peninsula

Forestry Studies

Volume 74 (2021): Issue 1 (December 2021)

By: Jaanus Paal, Anton Pesterov and Valentina Neshataeva

Open Access

|Dec 2021

Figures & Tables

Location of sample areas on the Kamchatka Peninsula.

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.

Table 1.

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

Table 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, SE_β – its standard error, B – unstandardized regression coefficient (slope), SE_β – 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	SE_β	B	SE_β	t	^p		Summary statistics
Variable	β	SE_β	B	SE_β	t	^p	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

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.

Table 3.

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); p_ANOVA – significance level by the univariate ANOVA, p_KW – significance level by the Kruskal-Wallis post-hoc median test.

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

Table 4.

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

Community type	Type group				Exposition					Plain	Total
Community type	Type group	N	NW	W	SW	S	SE	E	NE		Total
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

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
SD_S	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
SD_F	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

Species	Max	p	Community type
Species	Max	p	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

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DOI: https://doi.org/10.2478/fsmu-2021-0003 | Journal eISSN: 1736-8723 | Journal ISSN: 1406-9954

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Language: English

Page range: 26 - 57

Submitted on: Jul 9, 2021

Accepted on: Aug 31, 2021

Published on: Dec 1, 2021

Published by: Estonian University of Life Sciences

In partnership with: Paradigm Publishing Services

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© 2021 Jaanus Paal, Anton Pesterov, Valentina Neshataeva, published by Estonian University of Life Sciences
This work is licensed under the Creative Commons Attribution 4.0 License.

Volume 74 (2021): Issue 1 (December 2021)

Diversity of Pinus pumila-dominated communities on the Kamchatka Peninsula

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Figure 1.

Figure 2.

Table 1.

Table 2.

Figure 3.

Table 3.

Table 4.

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