Chin J Plant Ecol ›› 2022, Vol. 46 ›› Issue (6): 642-655.DOI: 10.17521/cjpe.2021.0454
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WEI Long-Xin, GENG Yan(), CUI Ke-Da, QIAO Xue-Tao, YUE Qing-Min, FAN Chun-Yu, ZHANG Chun-Yu, ZHAO Xiu-Hai
Received:
2021-12-06
Accepted:
2022-02-24
Online:
2022-06-20
Published:
2022-03-15
Contact:
GENG Yan
Supported by:
WEI Long-Xin, GENG Yan, CUI Ke-Da, QIAO Xue-Tao, YUE Qing-Min, FAN Chun-Yu, ZHANG Chun-Yu, ZHAO Xiu-Hai. Responses of tree growth to harvesting intensity among forest strata and growth stages in a broadleaved Korean pine forest[J]. Chin J Plant Ecol, 2022, 46(6): 642-655.
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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2021.0454
Fig. 1 Schematic diagram of vertical stratification in mixed-species and uneven-aged stands. I, large/mature canopy trees; II, small/immature canopy trees; III, understory tree species and shrubs of any size.
样地 Sample plot | 海拔 Altitude (m) | 坡度 Slope (°) | 坡向 Aspect | 郁闭度 Canopy density | 平均胸径 Mean diameter at breast height (cm) | 平均树高 Mean tree height (m) | 主要乔木树种组成 Composition of main tree species |
---|---|---|---|---|---|---|---|
CK | 453 | 1 | 东北 Northeast | 0.9 | 14.6 | 9.7 | 3春榆; 2水曲柳; 1紫椴; 1胡桃楸; 1五角枫; +红松; +蒙古栎; +黄檗; -山杨; -朝鲜槐; -三花槭; -硕桦; -白桦; -东北槭; -稠李; -裂叶榆 3 Ulmus davidiana var. japonica; 2 Fraxinus mandshurica; 1 Tilia amurensis; 1 Juglans mandshurica; 1 Acer pictum subsp. mono; + Pinus koraiensis; + Quercus mongolica; + Phellodendron amurense; - Populus davidiana; - Maackia amurensis; - Acer triflorum; - Betula costata; - Betula platyphylla; - Acer mandshuricum; - Padus racemosa; - Ulmus laciniata |
Low | 443 | 4 | 东北 Northeast | 0.9 | 13.9 | 9.6 | 2水曲柳; 1紫椴; 1红松; 1五角枫; 1蒙古栎; 1白桦; 1春榆; +东北槭; +胡桃楸; -香杨; -黄檗; -千金榆; -朝鲜槐; -稠李; -茶条槭; -裂叶榆 2 Fraxinus mandshurica; 1 Tilia amurensis; 1 Pinus koraiensis; 1 Acer pictum subsp. mono; 1 Quercus mongolica; 1 Betula platyphylla; 1 Ulmus davidiana var. japonica; + Acer mandshuricum; + Juglans mandshurica; - Populus koreana; - Phellodendron amurense; - Carpinus cordata; - Maackia amurensis; - Padus racemosa; - Acer ginnala; - Ulmus laciniata |
Medium | 430 | 5 | 东北 Northeast | 0.9 | 14.8 | 9.7 | 2红松; 2水曲柳; 1紫椴; 1蒙古栎; 1五角枫; 1三花槭; 1春榆; 1白桦; +山杨; +东北槭; +胡桃楸; -朝鲜槐; -硕桦; -稠李; -茶条槭; -青楷槭; -香杨; -黄檗 2 Pinus koraiensis; 2 Fraxinus mandshurica; 1 Tilia amurensis; 1 Quercus mongolica; 1 Acer pictum subsp. mono; 1 Acer triflorum; 1 Ulmus davidiana var. japonica; 1 Betula platyphylla; + Populus davidiana; + Acer mandshuricum; + Juglans mandshurica; - Maackia amurensis; - Betula costata; - Padus racemosa; - Acer ginnala; - Acer tegmentosum; - Populus koreana; - Phellodendron amurense |
High | 447 | 3 | 东北 Northeast | 0.9 | 12.4 | 8.8 | 2水曲柳; 2白桦; 1红松; 1紫椴; 1蒙古栎; 1五角枫; 1三花槭; 1春榆; +香杨; -山杨; -朝鲜槐; -硕桦; -东北槭; -稠李; -辽椴; -胡桃楸; -茶条槭; -裂叶榆; -黄檗 2 Fraxinus mandshurica; 2 Betula platyphylla; 1 Pinus koraiensis; 1 Tilia amurensis; 1 Quercus mongolica; 1 Acer pictum subsp. mono; 1 Acer triflorum; 1 Ulmus davidiana var. japonica; + Populus koreana; - Populus davidiana; - Maackia amurensis; - Betula costata; - Acer mandshuricum; - Padus racemosa; - Tilia mandshurica; - Juglans mandshurica; - Acer ginnala; - Ulmus laciniate; - Phellodendron amurense |
Table 1 Plot information of the broadleaved Korean pine forest in Jiaohe, Jilin Province
样地 Sample plot | 海拔 Altitude (m) | 坡度 Slope (°) | 坡向 Aspect | 郁闭度 Canopy density | 平均胸径 Mean diameter at breast height (cm) | 平均树高 Mean tree height (m) | 主要乔木树种组成 Composition of main tree species |
---|---|---|---|---|---|---|---|
CK | 453 | 1 | 东北 Northeast | 0.9 | 14.6 | 9.7 | 3春榆; 2水曲柳; 1紫椴; 1胡桃楸; 1五角枫; +红松; +蒙古栎; +黄檗; -山杨; -朝鲜槐; -三花槭; -硕桦; -白桦; -东北槭; -稠李; -裂叶榆 3 Ulmus davidiana var. japonica; 2 Fraxinus mandshurica; 1 Tilia amurensis; 1 Juglans mandshurica; 1 Acer pictum subsp. mono; + Pinus koraiensis; + Quercus mongolica; + Phellodendron amurense; - Populus davidiana; - Maackia amurensis; - Acer triflorum; - Betula costata; - Betula platyphylla; - Acer mandshuricum; - Padus racemosa; - Ulmus laciniata |
Low | 443 | 4 | 东北 Northeast | 0.9 | 13.9 | 9.6 | 2水曲柳; 1紫椴; 1红松; 1五角枫; 1蒙古栎; 1白桦; 1春榆; +东北槭; +胡桃楸; -香杨; -黄檗; -千金榆; -朝鲜槐; -稠李; -茶条槭; -裂叶榆 2 Fraxinus mandshurica; 1 Tilia amurensis; 1 Pinus koraiensis; 1 Acer pictum subsp. mono; 1 Quercus mongolica; 1 Betula platyphylla; 1 Ulmus davidiana var. japonica; + Acer mandshuricum; + Juglans mandshurica; - Populus koreana; - Phellodendron amurense; - Carpinus cordata; - Maackia amurensis; - Padus racemosa; - Acer ginnala; - Ulmus laciniata |
Medium | 430 | 5 | 东北 Northeast | 0.9 | 14.8 | 9.7 | 2红松; 2水曲柳; 1紫椴; 1蒙古栎; 1五角枫; 1三花槭; 1春榆; 1白桦; +山杨; +东北槭; +胡桃楸; -朝鲜槐; -硕桦; -稠李; -茶条槭; -青楷槭; -香杨; -黄檗 2 Pinus koraiensis; 2 Fraxinus mandshurica; 1 Tilia amurensis; 1 Quercus mongolica; 1 Acer pictum subsp. mono; 1 Acer triflorum; 1 Ulmus davidiana var. japonica; 1 Betula platyphylla; + Populus davidiana; + Acer mandshuricum; + Juglans mandshurica; - Maackia amurensis; - Betula costata; - Padus racemosa; - Acer ginnala; - Acer tegmentosum; - Populus koreana; - Phellodendron amurense |
High | 447 | 3 | 东北 Northeast | 0.9 | 12.4 | 8.8 | 2水曲柳; 2白桦; 1红松; 1紫椴; 1蒙古栎; 1五角枫; 1三花槭; 1春榆; +香杨; -山杨; -朝鲜槐; -硕桦; -东北槭; -稠李; -辽椴; -胡桃楸; -茶条槭; -裂叶榆; -黄檗 2 Fraxinus mandshurica; 2 Betula platyphylla; 1 Pinus koraiensis; 1 Tilia amurensis; 1 Quercus mongolica; 1 Acer pictum subsp. mono; 1 Acer triflorum; 1 Ulmus davidiana var. japonica; + Populus koreana; - Populus davidiana; - Maackia amurensis; - Betula costata; - Acer mandshuricum; - Padus racemosa; - Tilia mandshurica; - Juglans mandshurica; - Acer ginnala; - Ulmus laciniate; - Phellodendron amurense |
Fig. 2 Diameter at breast height-tree height bivariate density graphs used to distinguish mature and immature individuals of dominant tree species in the canopy in the sampling plots of broadleaved Korean pine forest. The red dots are mature individuals, and the blue dots are immature individuals. The grey lines from the inside out represent the 0.05, 0.25, 0.5, 0.75 and 0.95 quantiles, respectively.
物种 Species | 林层 Forest strata | 划分I和II组的方法 Method used to differentiate between mature (I) and immature (II) trees | 组别 Cohort | 监测株数 Number of stems | 耐阴性 Shade- tolerance | |
---|---|---|---|---|---|---|
白桦 Betula platyphylla | C | 树高胸径分布法 DBH-H distribution | I (53.5): DBH ≥ 22 cm | II (43.5): DBH < 22 cm | 178 | 喜光 LD |
东北槭 Acer mandshuricum | C | 树高胸径分布法 DBH-H distribution | I (87.4): DBH ≥ 8 cm | II (55.6): DBH < 8 cm | 270 | 喜光 LD |
春榆 Ulmus davidiana var. japonica | C | 树高胸径分布法 DBH-H distribution | I (67.8): DBH ≥ 12 cm | II (39.2): DBH < 12 cm | >627 | 喜光且耐阴 LD and ST |
硕桦 Betula costata | C | 最大树高法 Hmax | I: H ≥ 10.2 m | II: H < 10.2 m | 7 | 喜光 LD |
红松 Pinus koraiensis | C | 树高胸径分布法 DBH-H distribution | I (62.5): DBH ≥ 20 cm | II (42.4): DBH < 20 cm | 387 | 喜光 LD |
胡桃楸 Juglans mandshurica | C | 最大树高法 Hmax | I (50.0): H ≥ 11.9 m | II (35.9): H < 11.9 m | 90 | 喜光 LD |
朝鲜槐 Maackia amurensis | C | 树高胸径分布法 DBH-H distribution | I (60.3): DBH ≥ 12 cm | II (47.3): DBH < 12 cm | 94 | 喜光 LD |
黄檗 Phellodendron amurense | C | 最大树高法 Hmax | I (66.1): H ≥ 9.3 m | II (52.9): H < 9.3 m | 61 | 喜光 LD |
辽椴 Tilia mandshurica | C | 最大树高法 Hmax | I: H ≥ 11 m | II: H < 11 m | 16 | 喜光 LD |
裂叶榆 Ulmus laciniata | C | 最大树高法 Hmax | I (68.4): H ≥ 10.2 m | II (52.1): H < 10.2 m | >15 | 耐阴 ST |
蒙古栎 Quercus mongolica | C | 树高胸径分布法 DBH-H distribution | I (59.2): DBH ≥ 20 cm | II (44.2): DBH < 20 cm | 177 | 喜光 LD |
三花槭 Acer triflorum | C | 树高胸径分布法 DBH-H distribution | I (86.6): DBH ≥ 13 cm | II (50.0): DBH < 13 cm | >379 | 耐阴 ST |
五角枫 Acer pictum subsp. mono | C | 树高胸径分布法 DBH-H distribution | I (81.3): DBH ≥ 13 cm | II (60.8): DBH < 13 cm | 851 | 喜光 LD |
山杨 Populus davidiana | C | 最大树高法 Hmax | I (28.0): H ≥ 11.7 m | II (20.6): H < 11.7 m | 20 | 喜光 LD |
水曲柳 Fraxinus mandschurica | C | 树高胸径分布法 DBH-H distribution | I (60.1): DBH ≥ 21 cm | II (49.6): DBH < 21 cm | 323 | 喜光 LD |
香杨 Populus koreana | C | 最大树高法 Hmax | I: H ≥ 13.7 m | II: H < 13.7 m | 10 | 喜光 LD |
紫椴 Tilia amurensis | C | 树高胸径分布法 DBH-H distribution | I (68.3): DBH ≥ 18 cm | II (43.6): DBH < 18 cm | 493 | 喜光 LD |
暴马丁香 Syringa reticulata subsp. amurensis | U | - | III | >171 | 耐阴 ST | |
茶条槭 Acer ginnala | U | - | III | 20 | 喜光 LD | |
稠李 Padus racemosa | U | - | III | 32 | 耐阴 ST | |
簇毛槭 Acer barbinerve | U | - | III | 34 | 耐阴 ST | |
花楸树 Sorbus pohuashanensis | U | - | III | 1 | 耐阴 ST | |
楤木 Aralia elata | U | - | III | 1 | 喜光 LD | |
毛榛 Corylus mandshurica | U | - | III | 16 | 耐阴 ST | |
千金榆 Carpinus cordata | U | - | III | 1 | 耐阴 ST | |
青楷槭 Acer tegmentosum | U | - | III | 44 | 喜光 LD | |
山荆子 Malus baccata | U | - | III | 68 | 耐阴 ST | |
鼠李 Rhamnus davurica | U | - | III | 9 | 耐阴 ST | |
卫矛 Euonymus alatus | U | - | III | >12 | 喜光且耐阴 LD and ST |
Table 2 Cohort specification and shade-tolerance of tree species in the plots of broadleaved Korean pine forest
物种 Species | 林层 Forest strata | 划分I和II组的方法 Method used to differentiate between mature (I) and immature (II) trees | 组别 Cohort | 监测株数 Number of stems | 耐阴性 Shade- tolerance | |
---|---|---|---|---|---|---|
白桦 Betula platyphylla | C | 树高胸径分布法 DBH-H distribution | I (53.5): DBH ≥ 22 cm | II (43.5): DBH < 22 cm | 178 | 喜光 LD |
东北槭 Acer mandshuricum | C | 树高胸径分布法 DBH-H distribution | I (87.4): DBH ≥ 8 cm | II (55.6): DBH < 8 cm | 270 | 喜光 LD |
春榆 Ulmus davidiana var. japonica | C | 树高胸径分布法 DBH-H distribution | I (67.8): DBH ≥ 12 cm | II (39.2): DBH < 12 cm | >627 | 喜光且耐阴 LD and ST |
硕桦 Betula costata | C | 最大树高法 Hmax | I: H ≥ 10.2 m | II: H < 10.2 m | 7 | 喜光 LD |
红松 Pinus koraiensis | C | 树高胸径分布法 DBH-H distribution | I (62.5): DBH ≥ 20 cm | II (42.4): DBH < 20 cm | 387 | 喜光 LD |
胡桃楸 Juglans mandshurica | C | 最大树高法 Hmax | I (50.0): H ≥ 11.9 m | II (35.9): H < 11.9 m | 90 | 喜光 LD |
朝鲜槐 Maackia amurensis | C | 树高胸径分布法 DBH-H distribution | I (60.3): DBH ≥ 12 cm | II (47.3): DBH < 12 cm | 94 | 喜光 LD |
黄檗 Phellodendron amurense | C | 最大树高法 Hmax | I (66.1): H ≥ 9.3 m | II (52.9): H < 9.3 m | 61 | 喜光 LD |
辽椴 Tilia mandshurica | C | 最大树高法 Hmax | I: H ≥ 11 m | II: H < 11 m | 16 | 喜光 LD |
裂叶榆 Ulmus laciniata | C | 最大树高法 Hmax | I (68.4): H ≥ 10.2 m | II (52.1): H < 10.2 m | >15 | 耐阴 ST |
蒙古栎 Quercus mongolica | C | 树高胸径分布法 DBH-H distribution | I (59.2): DBH ≥ 20 cm | II (44.2): DBH < 20 cm | 177 | 喜光 LD |
三花槭 Acer triflorum | C | 树高胸径分布法 DBH-H distribution | I (86.6): DBH ≥ 13 cm | II (50.0): DBH < 13 cm | >379 | 耐阴 ST |
五角枫 Acer pictum subsp. mono | C | 树高胸径分布法 DBH-H distribution | I (81.3): DBH ≥ 13 cm | II (60.8): DBH < 13 cm | 851 | 喜光 LD |
山杨 Populus davidiana | C | 最大树高法 Hmax | I (28.0): H ≥ 11.7 m | II (20.6): H < 11.7 m | 20 | 喜光 LD |
水曲柳 Fraxinus mandschurica | C | 树高胸径分布法 DBH-H distribution | I (60.1): DBH ≥ 21 cm | II (49.6): DBH < 21 cm | 323 | 喜光 LD |
香杨 Populus koreana | C | 最大树高法 Hmax | I: H ≥ 13.7 m | II: H < 13.7 m | 10 | 喜光 LD |
紫椴 Tilia amurensis | C | 树高胸径分布法 DBH-H distribution | I (68.3): DBH ≥ 18 cm | II (43.6): DBH < 18 cm | 493 | 喜光 LD |
暴马丁香 Syringa reticulata subsp. amurensis | U | - | III | >171 | 耐阴 ST | |
茶条槭 Acer ginnala | U | - | III | 20 | 喜光 LD | |
稠李 Padus racemosa | U | - | III | 32 | 耐阴 ST | |
簇毛槭 Acer barbinerve | U | - | III | 34 | 耐阴 ST | |
花楸树 Sorbus pohuashanensis | U | - | III | 1 | 耐阴 ST | |
楤木 Aralia elata | U | - | III | 1 | 喜光 LD | |
毛榛 Corylus mandshurica | U | - | III | 16 | 耐阴 ST | |
千金榆 Carpinus cordata | U | - | III | 1 | 耐阴 ST | |
青楷槭 Acer tegmentosum | U | - | III | 44 | 喜光 LD | |
山荆子 Malus baccata | U | - | III | 68 | 耐阴 ST | |
鼠李 Rhamnus davurica | U | - | III | 9 | 耐阴 ST | |
卫矛 Euonymus alatus | U | - | III | >12 | 喜光且耐阴 LD and ST |
林木株数 Number of stems | 胸高断面积 Basal area (m2·hm-2) | |||||
---|---|---|---|---|---|---|
I | II | III | I | II | III | |
伐前 Pre-harvesting | ||||||
CK | 354 | 516 | 76 | 23.01 | 3.63 | 0.36 |
Low | 345 | 621 | 60 | 25.77 | 3.36 | 0.26 |
Medium | 339 | 589 | 71 | 25.86 | 3.67 | 0.39 |
High | 296 | 595 | 134 | 23.63 | 3.41 | 0.35 |
伐后 Post-harvesting | ||||||
CK | 354 | 516 | 76 | 23.01 | 3.63 | 0.36 |
Low | 303 (12%) | 580 (7%) | 48 (20%) | 21.24 (18%) | 3.11 (7%) | 0.25 (4%) |
Medium | 251 (26%) | 504 (14%) | 58 (18%) | 16.94 (34%) | 3.25 (11%) | 0.27 (31%) |
High | 149 (50%) | 379 (36%) | 68 (49%) | 8.47 (64%) | 2.61 (23%) | 0.21 (40%) |
Table 3 Number of stems, basal area of diameter at breast height (DBH) pre- and post-harvesting, and harvesting intensity of each cohort in each plot of broadleaved Korean pine forest
林木株数 Number of stems | 胸高断面积 Basal area (m2·hm-2) | |||||
---|---|---|---|---|---|---|
I | II | III | I | II | III | |
伐前 Pre-harvesting | ||||||
CK | 354 | 516 | 76 | 23.01 | 3.63 | 0.36 |
Low | 345 | 621 | 60 | 25.77 | 3.36 | 0.26 |
Medium | 339 | 589 | 71 | 25.86 | 3.67 | 0.39 |
High | 296 | 595 | 134 | 23.63 | 3.41 | 0.35 |
伐后 Post-harvesting | ||||||
CK | 354 | 516 | 76 | 23.01 | 3.63 | 0.36 |
Low | 303 (12%) | 580 (7%) | 48 (20%) | 21.24 (18%) | 3.11 (7%) | 0.25 (4%) |
Medium | 251 (26%) | 504 (14%) | 58 (18%) | 16.94 (34%) | 3.25 (11%) | 0.27 (31%) |
High | 149 (50%) | 379 (36%) | 68 (49%) | 8.47 (64%) | 2.61 (23%) | 0.21 (40%) |
Fig. 3 Diameter at breast height (DBH) distribution in control plot (A) and plots subjected to low intensity harvesting (B), medium intensity harvesting (C), and high intensity harvesting plot (D) before and after harvesting of broadleaved Korean pine forest.
变异来源 Source of variation | 生长速率 Growth rate | 相对生长速率 Relative growth rate | ||||
---|---|---|---|---|---|---|
df | F | p | df | F | p | |
采伐强度 Harvesting intensity (HI) | 3 | 24.772 | <0.001 | 3 | 23.690 | <0.001 |
组别 Cohort (C) | 2 | 215.467 | <0.001 | 2 | 100.555 | <0.001 |
恢复时期 Period (P) | 2 | 91.112 | <0.001 | 2 | 19.289 | <0.001 |
采伐强度×组别 HI × C | 6 | 2.456 | 0.023 | 6 | 3.858 | <0.001 |
采伐强度×恢复时期 HI × P | 6 | 7.500 | <0.001 | 6 | 5.639 | <0.001 |
组别×恢复时期 C × P | 4 | 6.691 | <0.001 | 4 | 4.822 | <0.001 |
采伐强度×组别×恢复时期 HI × C × P | 12 | 1.435 | 0.145 | 12 | 0.934 | 0.512 |
Table 4 Multi-factor analysis of variance of affecting the growth of residual trees of broadleaved Korean pine forest
变异来源 Source of variation | 生长速率 Growth rate | 相对生长速率 Relative growth rate | ||||
---|---|---|---|---|---|---|
df | F | p | df | F | p | |
采伐强度 Harvesting intensity (HI) | 3 | 24.772 | <0.001 | 3 | 23.690 | <0.001 |
组别 Cohort (C) | 2 | 215.467 | <0.001 | 2 | 100.555 | <0.001 |
恢复时期 Period (P) | 2 | 91.112 | <0.001 | 2 | 19.289 | <0.001 |
采伐强度×组别 HI × C | 6 | 2.456 | 0.023 | 6 | 3.858 | <0.001 |
采伐强度×恢复时期 HI × P | 6 | 7.500 | <0.001 | 6 | 5.639 | <0.001 |
组别×恢复时期 C × P | 4 | 6.691 | <0.001 | 4 | 4.822 | <0.001 |
采伐强度×组别×恢复时期 HI × C × P | 12 | 1.435 | 0.145 | 12 | 0.934 | 0.512 |
Fig. 4 Relative growth rate (mean ± SE) of diameter at breast height (DBH) of each group of trees under different harvesting intensities in different recovery periods of broadleaved Korean pine forest. A, First two years after harvested. B, 2-4 years after harvested. C, 4-7 years after harvested. D, Over 7 years after harvested. Different lowercase letters indicate significant differences among different harvesting intensities within the same cohort; different uppercase letters indicate significant differences among tree cohorts within the same harvesting intensity (p < 0.05). I, large/mature canopy trees; II, small/immature canopy trees; III, understory tree species and shrubs of any size. CK, control plot; High, heavily harvesting plot; Low, light harvesting plot; Medium, moderate harvesting plot.
Fig. 5 Relationship between the relative growth rate of diameter at breast height (DBH) of each group of trees and their initial size under different harvesting intensities of broadleaved Korean pine forest. I, large/mature canopy trees; II, small/immature canopy trees; III, understory tree species and shrubs of any size. CK, control plot; High, heavily harvesting plot; Low, light harvesting plot; Medium, moderate harvesting plot. Shadow volume represents 95% confidence interval of the fitted line.
Fig. 6 Relative growth rate of diameter at breast height (DBH)(mean ± SE) of mature (I) and immature (II) individuals of dominant, subdominant and rare species in the forest canopy under different harvesting intensities of broadleaved Korean pine forest. Different uppercase letters indicate significant differences in relative growth rate of DBH of trees in cohort I among different harvesting intensities (p < 0.05); different lowercase letters indicate significant differences in relative growth rates of DBH of trees in cohort II among different harvesting intensities (p < 0.05). CK, control plot; High, heavily harvesting plot; Low, light harvesting plot; Medium, moderate harvesting plot.
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