Composition of live, dead and downed trees in Järvselja old-growth forest

Kangur, Ahto; Nigul, Kristi; Padari, Allar; Kiviste, Andres; Korjus, Henn; Laarmann, Diana; Põldveer, Eneli; Mitt, Risto; Frelich, Lee E.; Jõgiste, Kalev; Stanturf, John A.; Paluots, Teele; Kängsepp, Vivika; Jürgenson, Harli; Noe, Steffen M.; Sims, Allan; Metslaid, Marek

Composition of live, dead and downed trees in Järvselja old-growth forest

Volume 75 (2021): Issue 1 (December 2021)

By:

Ahto Kangur, Kristi Nigul, Allar Padari, Andres Kiviste, Henn Korjus, Diana Laarmann, Eneli Põldveer, Risto Mitt, Lee E. Frelich, Kalev Jõgiste, John A. Stanturf, Teele Paluots, Vivika Kängsepp, Harli Jürgenson, Steffen M. Noe, Allan Sims and Marek Metslaid

Open Access

|Jun 2022

Figures & Tables

H-D model calculation scheme based on the Näslund function.

The measured areas (light green colour) in Järvselja old-growth forest (compartment JS226) following the numbered sub-compartments. The locations of mapped downed trees and stems (A) and standing live and dead trees (B) are presented (green dots - live trees, blue dots - dead standing trees, red dots - standing snags, blue lines – downed trees and stems).

Model residual comparison according to main tree species. Tree species and tree species codes are as follows: CA – common aspen; EL – elm family; BI – silver birch and downy birch; NS – Norway spruce; BAR – black alder; CAR – common alder; SP – Scots pine; WI – willow family; ER – European rowan; LI – linden family; AH – European ash; CH – Common hazel; BC – bird cherry; NM – Norway maple.

The long-term dynamics of stand composition and average standing live stem-wood volume. Tree species and tree species codes are as follows: CA – common aspen; EL – elm family; BI – silver birch and downy birch; NS – Norway spruce; BAR – black alder; SP – Scots pine; LI – linden family; AH – European ash; NM – Norway maple; DF – Douglas fir; FF – fir family; OD – other deciduous.

Wood density and related model parameters for Equation 14_

Species	Fresh wood basic density d0, g/cm³	Equation constants

		a₀	a₁
Pedunculate oak	584.4	529.58	−88.546
European ash	574.7	529.58	−88.546
Elm family	550.4	529.58	−88.546
Larch family	547.7	433.39	−55.003
Fir family	317.0	431.07	−56.649
Willow family	387.0	435.23	−58.931

Earlier studies on Järvselja old-growth forest_

Author(s)	Publishing year	Measurement year/period	Inventory type^*	Citation
Kollom, M.	2011	1922–2010	3	(Kollom, 2011)
Kasesalu, H.	2001	2001	1	(Kasesalu, 2001)
Kasesalu, A.	1985, 1993	1982, 1983	1	(Kasesalu, 1985), (Kasesalu, 1993)
Irdt, E., Rebane, H.	1985	1954–1985	1; 2	(Irdt & Rebane, 1985)
Pettai, T.	1985	1922–1984	3	(Pettai, 1985)
Raukas, A.	1967	1958–1959	1; 3	(Raukas, 1967)
Krigul, T.	1961	1958, 1959	3	(Krigul, 1961)
Krigul, T.	1940	1939	3	(Krigul, 1940)
Visnapuu, M.	1927	1927	3	(Krigul, 1940)

Stand characteristics in sub-compartments_

Sub-compartment	Area, ha	Cohort	Stand composition¹	N	G	M	S	C

				per hectare²
1	0.60	Live	50NS25BAR19SP2BI2WI1AH1LI	1505	21.2	198.0	4696	38.263
		Snags	87NA7NS6BAR	127	6.4	7.5	119	0.623

		Downed	51NS2SP+BAR47NA	281		137.8	1916	21.968

2	0.74	Live	62SP38NS	905	31.4	339.5	6585	65.224
		Dead	57NS26BI17SP	151	4.4	44.7	1038	6.918
		Snags	74SP22NS4NA	82	1.5	9.9	246	1.579

		Downed	30SP27BI13NS30NA	164		23.5	499	3.795

3	1.13	Live	58SP29NS9BAR4BI	952	31.9	309.0	6509	60.798
		Dead	59NS41SP	167	1.3	8.8	340	1.464
		Snags	35SP34BI30NS1NA	134	2.8	10.3	264	1.595

		Downed	70NS10BI1SP17NA	184		16.8	441	2.545

4	3.37	Live	59SP26NS9BI6BAR	921	33.2	330.2	6656	66.027
		Dead	80SP20NS	63	2.6	24.6	500	4.319
		Snags	44SP38NA10BI8NS	183	7.8	37.7	655	5.999

		Downed	28NS17SP16BI2CA37NA	310		53.2	1197	7.167

5	1.11	Live	33WI22LI11AH11BI10NM5RE1CA1DF	1435	36.6	360.3	7063	77.277
		Dead	100NS	14	2.8	37.4	406	6.401
		Snags	67CA29NS3NA1RE	225	35.3	105.3	805	11.803

		Downed	40NS9BI3NM1BAR+WI+EL47NA	394		318.0	3650	37.188

6	0.25	Live	68LI28NS3NM1AH	501	38.1	497.6	7276	97.085
		Dead	100NS	4	0.1	0.6	22	0.123
		Snags	62NS22NA12LI4BAR	61	8.8	7.8	71	1.070

		Downed	59NS7LI4BAR3BI27NA	151		85.8	1044	11.600

7	1.37	Live	56NS41SP2BAR1LI	1032	29.9	310.8	6170	58.481
		Dead	55NS45SP	146	6.8	74.0	1480	11.615
		Snags	99NS1NA	167	7.4	19.4	303	3.035

		Downed	87NS7SP6NA	375		56.7	1325	7.928

8	1.71	Live	43NS33SP13BAR10BI1OD	828	37.1	369.5	6886	72.923
		Dead	62NS38SP	79	3.6	35.1	665	6.179
		Snags	62NS18SP10BAR4NA4FF2BI	152	6.0	19.8	367	3.120

		Downed	76NS15BI5SP4NA	308		48.2	1012	6.774

9	2.52	Live	56NS15LI14BAR7BI5CA2NM1SP	711	40.0	497.7	7249	96.452
		Dead	97NS2NM1LI	26	1.3	16.4	235	3.033
		Snags	44NS17BI10NA9BAR7CA6LI6CAR1RE	131	16.6	40.2	367	5.717

		Downed	56NS9BI6BAR3LI+CA+NM+SP+AH26NA	268		160.8	1883	18.707

10	0.64	Live	68NS15BAR8BI5SP1NM1CA1BC1OD	819	25.5	260.6	5449	49.877
		Dead	80NS15SP4BAR1NM	50	2.0	20.8	401	3.514
		Snags	76NS11NA7BI4BAR2SP	233	19.0	33.0	452	3.806

		Downed	75NS10BI6FF3BAR2SP4NA	439		138.5	2213	15.252

11	0.89	Live	71NS11SP10BAR5BI1NM1FF1OD	661	26.0	296.7	5288	56.064
		Dead	91NS6SP3AH	34	2.0	23.6	397	4.135
		Snags	81NS8SP6NA2BI1BAR1FF1AH	157	15.6	23.4	272	3.106

		Downed	73NS8BI6SP2AH1BAR+FF10NA	384		151.8	2334	17.569

12	2.31	Live	50BAR31NS7BI6SP5LI1OD	578	32.7	361.9	5530	72.801
		Dead	66NS15BAR9BI8SP1LI1AH	20	1.2	12.7	201	2.266
		Snags	36BAR28NA22NS10BI1CAR1AH1OD	83	7.1	20.7	195	2.393

		Downed	48NS14BI11BAR4SP2LI2AH1NM+FF18NA	394		98.4	1807	14.534

13	0.95	Live	61LI29NS5NM2AH2BAR1BI	661	38.0	467.8	7076	92.608
		Dead	99NS1BAR	15	0.6	7.1	120	1.161
		Snags	59NS24NA11LI4BAR2OD	77	10.7	13.1	132	1.686

		Downed	57NS8BAR7LI3SP1BI1AH1NM+EL+FF+ER+BC22NA	256		154.0	1768	20.462

14	0.15	Live	38BAR26LI17NS10BI4AH2BC2FF1NM	907	23.5	214.5	4877	44.431
		Snags	69NS21NM5BI5NA	172	16.1	16.6	219	1.735

		Downed	56NS20BI5BAR2NM+BC+EL+LI17NA	687		163.7	2395	15.524

15	1.64	Live	48NS23LI11BAR6CA4NM4BI3AH1OD	562	31.4	386.6	5784	76.092
		Dead	45NS20BI18AH13NM4LI	12	1.2	15.0	179	3.005
		Snags	56NS13BI9NA8LI8BAR2AH1CA1RE1OD	111	11.4	24.5	257	3.467

		Downed	50NS6LI5BI5NM4BAR3AH3CA+FF+BC+WI+ER+EL24NA	308		136.6	1898	19.182

Compartment	19.38	Live	46NS20LI18BAR6SP5BI2NM2CA 1AH+WI+ER+FF+EL+BC+DF+OD	837	33.3	363.5	6375	71.717
		Dead	75NS10SP6BI3BAR3NM2AH1LI	55	2.3	24.0	440	4.115
		Snags	49NS11BAR10BI5CA4LI3SP1AH1RE2OD14NA	142	10.9	29.7	372	4.084

		Downed	53NS9BI4BAR3SP2LI1NM1AH1CA+FF+WI+EL+BC+ER26NA	315		107.6	1624	13.889

j_fsmu-2021-0009_apptab_002

Site type	Tree species¹	N	a	b	x	R²	RMSE	p-value
Oxalis drained swamp	NS	707	2.01436	0.284572	0.72	0.8727	2.4486	<0.0001
	SP	20	2.16767	0.284568	0.66	0.2679	3.1509	0.0706
	AC	738	2.00648	0.285206	0.71	0.8771	2.4693	<0.0001

	BI	110	1.15131	0.310943	0.67	0.7346	2.5354	<0.0001
	BAR	147	1.24516	0.316697	0.75	0.7501	2.5076	<0.0001
	ER	15	0.57528	0.442944	0.65	0.2384	1.2038	0.1951
	LI	11	1.47998	0.342744	0.69	0.4544	2.51	0.0886
	NM	40	0.9162	0.347308	0.65	0.7481	1.5143	<0.0001
	AD	345	1.24048	0.315907	0.69	0.6945	2.7383	<0.0001

Alder fen	NS	475	2.25391	0.290099	0.65	0.8603	2.2956	<0.0001
	SP	34	1.21007	0.324187	0.62	0.2978	2.2989	0.0042
	AC	513	2.18471	0.293724	0.64	0.8618	2.3175	<0.0001

	BI	40	2.01465	0.301563	0.67	0.4511	2.6922	<0.0001
	DAR	346	1.48036	0.310482	0.68	0.8368	2.7971	<0.0001
	CAR	12	1.51185	0.316172	0.78	0.8124	2.3561	0.0005
	LI	57	1.84125	0.300121	0.72	0.7165	3.1607	<0.0001
	AH	10	1.36098	0.305205	0.77	0.885	1.9276	0.0005
	AD	476	1.52893	0.30999	0.71	0.8227	2.8396	<0.0001

Myrtillus drained swamp	NS	95	1.91242	0.296379	0.65	0.8897	2.2089	<0.0001
	SP	21	0.1853	0.346608	0.68	0.0349	1.5099	0.7264
	AC	117	1.85238	0.298818	0.6	0.9137	2.1902	<0.0001

	BI	10	1.44643	0.302228	1.12	0.8936	1.8118	0.0004
	DAR	22	1.18785	0.32696	0.5	0.9197	1.4993	<0.0001
	AD	48	1.34335	0.319018	0.61	0.872	1.8657	<0.0001

Aegopodium	NS	1088	2.08325	0.277529	0.68	0.8852	2.9114	<0.0001
	AC	1091	2.0843	0.277633	0.69	0.8834	2.9281	<0.0001

	CA	28	1.11678	0.287548	0.52	0.7055	3.4818	<0.0001
	SE	22	1.29345	0.347246	0.63	0.6247	2.2132	0.0001
	BI	49	1.84504	0.29027	0.82	0.565	4.3707	<0.0001
	DAR	209	1.6667	0.297854	1.03	0.8081	3.218	<0.0001
	WI	13	1.74562	0.30235	0.75	0.9448	1.7704	<0.0001
	ER	31	1.3131	0.331834	0.66	0.5718	2.8427	<0.0001
	LI	1155	1.68554	0.29393	0.67	0.7871	3.2555	<0.0001
	AH	58	1.30368	0.300885	0.7	0.7939	3.1191	<0.0001
	CH	38	1.04191	0.366278	0.67	0.1382	2.7214	0.0741
	BC	13	0.52782	0.389661	0.45	0.4197	2.1269	0.0658
	NM	257	1.36259	0.30576	0.9	0.7314	2.7437	<0.0001
	AD	1870	1.64682	0.294705	0.73	0.7861	3.3405	<0.0001

Transitional bog	NS	122	1.95335	0.30006	0.66	0.7957	2.4476	<0.0001
	SP	29	1.24269	0.324479	0.73	0.3303	1.9695	0.0055
	AC	159	1.84773	0.306233	0.64	0.8308	2.3759	<0.0001

	BI	18	1.28842	0.325971	0.73	0.6217	2.5596	0.0007
	DAR	35	1.48154	0.310051	0.66	0.8894	1.9883	<0.0001
	AD	59	1.40218	0.316723	0.71	0.8469	2.2027	<0.0001

H-D model parameters from Nigul et al_ (2021) study_

Tree species	Chapman-Richards function (Eq. 5)			Näslund function (Eq. 1)

	a	b	c	a	b
SP	30.67342	0.032552	0.80312	1.59472	0.30835
NS	39.02975	0.033294	1.18444	2.16428	0.27970
BI	33.57059	0.023891	0.62549	1.19810	0.31378
CA	37.72567	0.043817	0.89476	1.28707	0.28380
BAR	30.46570	0.036539	0.85649	1.43745	0.30966
LI	33.70126	0.040448	1.07656	1.69100	0.29469
NM	33.17741	0.030304	0.76981	1.26131	0.31403
Other	34.06000	0.035000	0.91100	1.51900	0.30000

Summary statistics of the measured data_

Variable	Live trees	Standing dead wood	Snags	Downed dead wood
Number of measured trees, (N)	6205	270	1119	2983
Number of estimated trees in the compartment JS226	16227	1073	2749	6112
Dominating species^*	NS, SP, WI, LI, BAR	NS, SP	NS, SP, CA, BAR	NS, SP
Mean diameter, D (cm)	22.5	22.8	31.3	26.9
Mean height/length, H/L (m)	21.3	21.8	3.9	8.6
Mean decay class	-	1.91	2.60	3.03
Mean stem volume, V (m³)	0.43	0.43	0.21	0.34
Mean stem lateral surface area, SLSA (m²)	7.61	7.95	2.62	5.15

j_fsmu-2021-0009_apptab_001

Point	x	y	Point	x	y
1	6 464 594,87	694 981,72	37	6 464 890,85	694 924,91
2	6 464 586,41	694 931,92	38	6 464 877,95	694 876,23
3	6 464 577,35	694 882,21	39	6 464 869,38	694 828,03
4	6 464 567,75	694 833,29	40	6 464 861,24	694 778,04
5	6 464 558,40	694 784,62	41	6 464 854,12	694 727,95
6	6 464 548,39	694 737,17	42	6 464 843,08	694 680,60
7	6 464 643,70	694 972,09	43	6 464 939,49	694 916,71
8	6 464 635,09	694 923,09	44	6 464 929,00	694 868,38
9	6 464 624,73	694 873,60	45	6 464 913,88	694 814,16
10	6 464 616,65	694 826,37	46	6 464 891,17	694 763,67
11	6 464 604,62	694 776,79	47	6 464 902,60	694 723,56
12	6 464 597,74	694 725,18	48	6 464 890,52	694 673,42
13	6 464 690,72	694 962,87	49	6 464 988,72	694 907,52
14	6 464 683,63	694 914,06	50	6 464 980,91	694 868,03
15	6 464 673,81	694 863,77	51	6 464 967,85	694 810,65
16	6 464 666,04	694 817,53	52	6 464 960,56	694 757,21
17	6 464 657,10	694 765,99	53	6 464 947,26	694 710,16
18	6 464 648,22	694 716,03	54	6 464 948,30	694 659,10
19	6 464 743,26	694 954,20	55	6 465 037,07	694 895,42
20	6 464 735,45	694 907,09	56	6 465 035,31	694 856,40
21	6 464 722,88	694 855,44	57	6 465 009,53	694 810,44
22	6 464 715,35	694 807,84	58	6 465 014,67	694 752,37
23	6 464 706,62	694 755,87	59	6 464 998,40	694 719,68
24	6 464 695,19	694 709,06	60	6 464 984,42	694 657,38
25	6 464 793,81	694 943,93	61	6 465 093,76	694 887,58
26	6 464 779,58	694 894,90	62	6 465 075,21	694 841,74
27	6 464 772,83	694 845,93	63	6 465 062,54	694 782,18
28	6 464 766,20	694 797,11	64	6 465 061,73	694 739,59
29	6 464 755,54	694 748,05	65	6 465 047,56	694 695,83
30	6 464 745,35	694 699,82	66	6 465 039,32	694 648,56
31	6 464 839,77	694 936,05	67	6 465 136,17	694 877,23
32	6 464 832,38	694 886,11	68	6 465 128,02	694 829,38
33	6 464 822,55	694 838,31	69	6 465 122,07	694 782,05
34	6 464 809,65	694 791,79	70	6 465 122,45	694 743,17
35	6 464 806,20	694 734,42	71	6 465 103,39	694 671,84
36	6 464 794,40	694 690,46	72	6 465 086,71	694 631,43

Studies carried out in Estonia containing information about the volume of coarse woody debris (CWD) in unmanaged forests (located often in nature protection areas), or locating in both, unmanaged and commercial forests_

	Location	Citation	Site description	Dominant tree species¹	Protection/management regime²	Average amount of dead wood, m³/ha
A	Karula National Park	(Köster et al., 2005)	water swamp forest, dry boreal forest	NS, BI, SP, BAR	NR and RMZ	27.6
B	Alam-Pedja Nature Reserve	(Lõhmuset al., 2005)	mostly forests on wet soils	BI, SP, NS, CA, CAR, BAR, other	MAN	9.0
B	Alam-Pedja Nature Reserve	(Lõhmuset al., 2005)	mostly forests on wet soils	BI, SP, NS, CA, CAR, BAR, other	NR	6.2
C	Lahemaa National Park	(Köster et al., 2005)	dry boreal, heath and ombro-trophic bog forests	NS, BI, SP	NR	85.9
C	Lahemaa National Park	(Köster et al., 2005)	dry boreal, heath and ombro-trophic bog forests	NS, BI, SP	RMZ	63.2
D	Estonia	(Lõhmus & Kraut, 2010)	dry boreal forest	SP	mature MAN	10
			meso-eutrophic forest	conifer/deciduous mixtures		62
			eutrophic boreo-nemoral forest	conifer/deciduous mixtures		43
			water swamp forest	BAR, BI		52
			dry boreal forest	SP	NAT	36
			meso-eutrophic forest	conifer/deciduous mixtures		144
			eutrophic boreo-nemoral forest	conifer/deciduous mixtures		198
			water swamp forest	BAR, BI		140
		(Põldveer et al., 2020)	mixed oligo-mesotrophic and mesotrophic forests on mineral soils	SP	MAN	28.9
					REC	33.9
					NAT	49.3
				NS	MAN	15.3
					REC	46.6
					NAT	85.1

Wood density and C relative content according to tree species and decay class (Köster et al_, 2015; Saarman & Veibri, 2006; Uri et al_, 2019, 2017, 2014)_

Variable	Tree species	Decay class

		fresh	1	2	3	4	5
Basic density, g/cm³	SP	422.7¹	381.1²	337.2²	258.8²	233.7²	141.8²
	NS, DF	375.0¹	410.7²	354.2²	280.7²	191.3²	124.8²
	BI, ER	538.2¹	466.6²	326.5²	230.0²	175.9²	112.1²
	CA, LI, NA	419.2¹	391.3²	330.6²	230.6²	161.1²	60.7²
	BAR	440.5¹	422.4²	289.4²	212.9²	158.9²	95.6²
	CAR, CH, BC, OD	396.0³	426.9²	345.3²	220.7²	184.9²	153.6²
	PO, NM	584.4¹	486.7	389.0	291.3	193.5	95.8
	AH	574.7¹	478.6	382.5	286.4	190.3	94.2
	EL	550.4¹	458.4	366.3	274.3	182.3	90.2
	WI	387.0¹	334.6	282.2	229.8	177.4	125.0
	LF	547.7¹	478.2	408.7	339.2	269.7	200.1
	FF	317.0¹	275.3	233.7	192.0	150.4	108.7

Carbon relative content, %	SP, LF	47.46⁴	49.03²	49.26²	49.56²	49.58²	50.21²
	NS, DF, FF	47.46	48.35²	48.31²	47.93²	49.60²	51.33²
	BI, ER, WI, PO, AH, NM, EL	47.00⁵	47.16²	47.69²	47.45²	48.80²	50.12²
	CA, LI, NA	47.00	47.19²	47.37²	47.38²	46.56²	46.31²
	BAR	47.30	47.89²	48.24²	48.07²	48.35²	48.11²
	CAR, CH, BC, OD	47.30³	48.02²	48.07²	48.71²	47.95²	48.04²

Volume and C content of live and dead trees in sub-compartments_

Sub-compartment	Volume, m³/ha	Live, %	Dead standing, %	Dead laying, %	C content MgC/ha	Live, %	Dead standing, %	Dead laying, %
1	343.4	57.7	2.2	40.1	60.85	62.9	1.0	36.1
2	417.5	81.3	13.1	5.6	77.52	84.1	11.0	4.9
3	345.0	89.6	5.5	4.9	66.40	91.6	4.6	3.8
4	445.7	74.1	14.0	11.9	83.51	79.1	12.3	8.6
5	820.9	43.9	17.4	38.7	132.67	58.3	13.7	28.0
6	591.7	84.1	1.4	14.5	109.88	88.4	1.1	10.5
7	460.9	67.4	20.3	12.3	81.06	72.1	18.1	9.8
8	472.6	78.2	11.6	10.2	89.00	81.9	10.5	7.6
9	715.2	69.6	7.9	22.5	123.91	77.8	7.1	15.1
10	452.8	57.5	11.9	30.6	72.45	68.8	10.1	21.1
11	495.4	59.9	9.5	30.6	80.87	69.3	9.0	21.7
12	493.7	73.3	6.8	19.9	91.99	79.1	5.1	15.8
13	641.9	72.9	3.1	24.0	115.92	79.9	2.5	17.6
14	394.7	54.3	4.2	41.5	61.69	72.0	2.8	25.2
15	562.7	68.7	7.0	24.3	101.75	74.8	6.4	18.8

Compartment	524.9	69.3	10.2	20.5	93.81	76.5	8.7	14.8

DOI: https://doi.org/10.2478/fsmu-2021-0009 | Journal eISSN: 1736-8723 | Journal ISSN: 1406-9954

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Language: English

Page range: 15 - 40

Submitted on: Dec 2, 2021

Accepted on: Dec 28, 2021

Published on: Jun 4, 2022

Published by: Estonian University of Life Sciences

In partnership with: Paradigm Publishing Services

Publication frequency: 2 times per year

Keywords:

CWD,

Related subjects:

Ecology,

© 2022 Ahto Kangur, Kristi Nigul, Allar Padari, Andres Kiviste, Henn Korjus, Diana Laarmann, Eneli Põldveer, Risto Mitt, Lee E. Frelich, Kalev Jõgiste, John A. Stanturf, Teele Paluots, Vivika Kängsepp, Harli Jürgenson, Steffen M. Noe, Allan Sims, Marek Metslaid, published by Estonian University of Life Sciences
This work is licensed under the Creative Commons Attribution 4.0 License.

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Composition of live, dead and downed trees in Järvselja old-growth forest

Figures & Tables

Figure 1

Figure 2

Figure 3

Figure 4

Wood density and related model parameters for Equation 14_

Earlier studies on Järvselja old-growth forest_

Stand characteristics in sub-compartments_

j_fsmu-2021-0009_apptab_002

H-D model parameters from Nigul et al_ (2021) study_

Summary statistics of the measured data_

j_fsmu-2021-0009_apptab_001

Studies carried out in Estonia containing information about the volume of coarse woody debris (CWD) in unmanaged forests (located often in nature protection areas), or locating in both, unmanaged and commercial forests_

Wood density and C relative content according to tree species and decay class (Köster et al_, 2015; Saarman & Veibri, 2006; Uri et al_, 2019, 2017, 2014)_

Volume and C content of live and dead trees in sub-compartments_

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