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

Ahto Kangur
Ahto Kangur
, 
Kristi Nigul
Kristi Nigul
, 
Allar Padari
Allar Padari
, 
Andres Kiviste
Andres Kiviste
, 
Henn Korjus
Henn Korjus
, 
Diana Laarmann
Diana Laarmann
, 
Eneli Põldveer
Eneli Põldveer
, 
Risto Mitt
Risto Mitt
, 
Lee E. Frelich
Lee E. Frelich
, 
Kalev Jõgiste
Kalev Jõgiste
, 
John A. Stanturf
John A. Stanturf
, 
Teele Paluots
Teele Paluots
, 
Vivika Kängsepp
Vivika Kängsepp
, 
Harli Jürgenson
Harli Jürgenson
, 
Steffen M. Noe
Steffen M. Noe
, 
Allan Sims
Allan Sims
 e 
Marek Metslaid
Marek Metslaid
   | 04 giu 2022
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Article Category: Research paper
Pubblicato online: 04 giu 2022
Pagine: 15 - 40
Ricevuto: 02 dic 2021
Accettato: 28 dic 2021
DOI: https://doi.org/10.2478/fsmu-2021-0009
Parole chiave
© 2021 Ahto Kangur et al., published by Sciendo
This work is licensed under the Creative Commons Attribution 4.0 International License.

Figure 1

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

Figure 2

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).
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).

Figure 3

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.
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.

Figure 4

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.
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
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
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
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