植物生态学报 ›› 2008, Vol. 32 ›› Issue (5): 1031-1040.DOI: 10.3773/j.issn.1005-264x.2008.05.008 cstr: 32100.14.j.issn.1005-264x.2008.05.008
张新平1, 王襄平1, 朱彪1, 宗占江2, 彭长辉3, 方精云1,*(
)
收稿日期:2008-01-25
接受日期:2008-04-15
出版日期:2008-01-25
发布日期:2008-09-30
作者简介:*(jyfang@urban.pku.edu.cn)基金资助:
ZHANG Xin-Ping1, WANG Xiang-Ping1, ZHU Biao1, ZONG Zhan-Jiang2, PENG Chang-Hui3, FANG Jing-Yun1,*(
)
Received:2008-01-25
Accepted:2008-04-15
Online:2008-01-25
Published:2008-09-30
摘要:
在中国东北长白山、帽儿山、凉水、根河的主要森林类型中设置27个样地, 连续3年(2004~2006年)观测森林凋落物的生产量, 以研究我国东北地区森林凋落物产量及其与环境因子的关系。结果表明, 不同森林类型凋落物年产量存在显著差异, 针阔叶混交林显著高于落叶针叶林和常绿针叶林, 落叶针叶林、常绿针叶林、落叶阔叶林和针阔叶混交林的年平均产量分别为2 337、2 472、3 130和4 146 kg·hm-2; 树叶、枝条、繁殖器官和其它组分占总凋落量的平均比例为71%、22%、6%和1%, 不同森林类型凋落物组分的比例差异较大。森林凋落物产量主要受温度限制, 降水、森林类型和群落结构无显著影响。不同组分凋落物量的影响因素不同: 树叶凋落量主要受温度和森林类型的影响; 枝条凋落量主要受降水和蓄积量的影响; 而繁殖器官凋落量则与树种的繁殖特性以及年降水有关。各组分占总凋落量的比例主要受降水影响, 树叶占凋落物比例随降水增加而下降, 枝条所占比例很小, 表现出与叶相反的变化趋势。
张新平, 王襄平, 朱彪, 宗占江, 彭长辉, 方精云. 我国东北主要森林类型的凋落物产量及其影响因素. 植物生态学报, 2008, 32(5): 1031-1040. DOI: 10.3773/j.issn.1005-264x.2008.05.008
ZHANG Xin-Ping, WANG Xiang-Ping, ZHU Biao, ZONG Zhan-Jiang, PENG Chang-Hui, FANG Jing-Yun. LITTER FALL PRODUCTION IN RELATION TO ENVIRONMENTAL FACTORS IN NORTHEAST CHINA’S FORESTS. Chinese Journal of Plant Ecology, 2008, 32(5): 1031-1040. DOI: 10.3773/j.issn.1005-264x.2008.05.008
| 地点 Site | 纬度 Latitude (N) | 经度 Longitude (E) | 海拔 Altitude (m) | 年均温 MAT (℃) | 年降水 MAP (mm) | 主要群落类型 Main community type | 样地数 No. plots |
|---|---|---|---|---|---|---|---|
| 长白山 Mt. Changbai | 42°03′~42°24′ | 128°04′~128°15′ | 650~1 940 | -3.7~2.9 | 600~1 340 | 岳桦林、常绿针叶林、针阔叶混交林、落叶针叶林 BE, EN, DNM, DN | 14 |
| 帽儿山 Mt. Mao’er | 45°16′~45°17′ | 127°34′~127°35′ | 300~420 | 1.9~2.5 | 724 | 落叶阔叶林、兴安落叶松林DB, LG | 7 |
| 凉水 Liangshui | 47°11′~47°12′ | 128°53′~128°54′ | 340~430 | 0.7~1.1 | 680 | 红松林、杨桦林 PK, BP | 4 |
| 根河 Genhe | 50°56′ | 121°30′~121°31′ | 810~840 | -4.8~-4.6 | 450~550 | 兴安落叶松林 LG | 2 |
表1 研究地点的地理位置、气候和植被概况
Table 1 General information on locations, climate, and vegetation types for the study sites
| 地点 Site | 纬度 Latitude (N) | 经度 Longitude (E) | 海拔 Altitude (m) | 年均温 MAT (℃) | 年降水 MAP (mm) | 主要群落类型 Main community type | 样地数 No. plots |
|---|---|---|---|---|---|---|---|
| 长白山 Mt. Changbai | 42°03′~42°24′ | 128°04′~128°15′ | 650~1 940 | -3.7~2.9 | 600~1 340 | 岳桦林、常绿针叶林、针阔叶混交林、落叶针叶林 BE, EN, DNM, DN | 14 |
| 帽儿山 Mt. Mao’er | 45°16′~45°17′ | 127°34′~127°35′ | 300~420 | 1.9~2.5 | 724 | 落叶阔叶林、兴安落叶松林DB, LG | 7 |
| 凉水 Liangshui | 47°11′~47°12′ | 128°53′~128°54′ | 340~430 | 0.7~1.1 | 680 | 红松林、杨桦林 PK, BP | 4 |
| 根河 Genhe | 50°56′ | 121°30′~121°31′ | 810~840 | -4.8~-4.6 | 450~550 | 兴安落叶松林 LG | 2 |
| 森林类型 Forest type | 群落类型 Community type | 主要树种 Major species | 样地数量 No. plots | 平均胸径 Mean DBH (cm) | 平均树高 Mean tree height (m) | 蓄积量 Volume (m3) | 立木密度 Density (/hm2) |
|---|---|---|---|---|---|---|---|
| 落叶阔叶林 DB | 平均值 Mean | 9 | 14.8 | 12.1 | 200.4 | 1 154 | |
| 岳桦林 BE | Betula ermanii(88%) | 3 | 14.2 | 9.2 | 110.6 | 1 393 | |
| 杨桦林 BP | B. platyphylla(27%) Populus davidiana(20%) | 3 | 14.6 | 14.5 | 264.3 | 1 207 | |
| 硬阔叶杂木林 DBM | Quercus mongolica(25%) Tilia amurensis(15%) | 3 | 15.7 | 12.6 | 226.2 | 863 | |
| 落叶针叶林 DN | 平均值 Mean | 7 | 21.6 | 17.5 | 305.7 | 694 | |
| 兴安落叶松林 LG | Larix gmelini(94%) | 5 | 16.6 | 14.6 | 244.5 | 820 | |
| 长白落叶松林 LO | L. olgensis(74%) | 2 | 34.1 | 24.7 | 458.7 | 380 | |
| 针阔叶混交林 DNM | 平均值 Mean | 6 | 25.5 | 16.6 | 573.1 | 617 | |
| 红松林 PK | Pinus koraiensis(40%) | 3 | 28.4 | 16.7 | 584.5 | 487 | |
| 红松阔叶林 PKDM | P. koraiensis(29%) T. amurensis(20%) | 3 | 22.5 | 16.5 | 561.6 | 747 | |
| 常绿针叶林 EN | 云冷杉林 PA | Picea jezoensis(38%) Abies nephrolepis (35%) | 5 | 19.3 | 14.2 | 518.8 | 1 352 |
表2 各森林类型的群落特征
Table 2 Community characteristics of all forest types
| 森林类型 Forest type | 群落类型 Community type | 主要树种 Major species | 样地数量 No. plots | 平均胸径 Mean DBH (cm) | 平均树高 Mean tree height (m) | 蓄积量 Volume (m3) | 立木密度 Density (/hm2) |
|---|---|---|---|---|---|---|---|
| 落叶阔叶林 DB | 平均值 Mean | 9 | 14.8 | 12.1 | 200.4 | 1 154 | |
| 岳桦林 BE | Betula ermanii(88%) | 3 | 14.2 | 9.2 | 110.6 | 1 393 | |
| 杨桦林 BP | B. platyphylla(27%) Populus davidiana(20%) | 3 | 14.6 | 14.5 | 264.3 | 1 207 | |
| 硬阔叶杂木林 DBM | Quercus mongolica(25%) Tilia amurensis(15%) | 3 | 15.7 | 12.6 | 226.2 | 863 | |
| 落叶针叶林 DN | 平均值 Mean | 7 | 21.6 | 17.5 | 305.7 | 694 | |
| 兴安落叶松林 LG | Larix gmelini(94%) | 5 | 16.6 | 14.6 | 244.5 | 820 | |
| 长白落叶松林 LO | L. olgensis(74%) | 2 | 34.1 | 24.7 | 458.7 | 380 | |
| 针阔叶混交林 DNM | 平均值 Mean | 6 | 25.5 | 16.6 | 573.1 | 617 | |
| 红松林 PK | Pinus koraiensis(40%) | 3 | 28.4 | 16.7 | 584.5 | 487 | |
| 红松阔叶林 PKDM | P. koraiensis(29%) T. amurensis(20%) | 3 | 22.5 | 16.5 | 561.6 | 747 | |
| 常绿针叶林 EN | 云冷杉林 PA | Picea jezoensis(38%) Abies nephrolepis (35%) | 5 | 19.3 | 14.2 | 518.8 | 1 352 |
| 森林类型 Forest type | 群落类型 Community type | 凋落物产量 Litter production | 平均 Average | 标准差 SD | 变异系数 CV (%) | ||
|---|---|---|---|---|---|---|---|
| 2004 | 2005 | 2006 | |||||
| 落叶阔叶林 DB | 平均值 Mean | 3 255 | 2 868 | 3 267 | 3 130ab | 227 | 7.2 |
| 岳桦林 BE | 1 332 | 2 302 | 2 293 | 1 976A | 557 | 28.2 | |
| 杨桦林 BP | 3 803 | 3 377 | 4 019 | 3 733AB | 327 | 8.8 | |
| 硬阔叶杂木林 DBM | 3 669 | 2 926 | 3 488 | 3 361AB | 387 | 11.5 | |
| 落叶针叶林 DN | 平均值 Mean | 2 803 | 2 095 | 2 113 | 2 337a | 404 | 17.3 |
| 兴安落叶松 LG | 3 018 | 2 363 | 2 130 | 2 504A | 460 | 18.4 | |
| 长白落叶松 LO | 2 266 | 1 560 | 2 086 | 1 971A | 367 | 18.6 | |
| 针阔叶混交林 DNM | 平均值 Mean | 4 312 | 4 066 | 4 060 | 4 146b | 144 | 3.5 |
| 红松林 PK | *** | 3 207 | 3 800 | 3 504AB | 420 | 12.0 | |
| 红松阔叶林 PKDM | 4 312 | 4 925 | 4 319 | 4 519B | 352 | 7.8 | |
| 常绿针叶林 EN | 云冷杉林 PA | 3 018 | 2 110 | 2 288 | 2 472aA | 481 | 19.5 |
表3 东北地区各森林类型凋落物年产量(kg·hm-2)
Table 3 Litter production for different forest types in Northeast China (kg·hm-2)
| 森林类型 Forest type | 群落类型 Community type | 凋落物产量 Litter production | 平均 Average | 标准差 SD | 变异系数 CV (%) | ||
|---|---|---|---|---|---|---|---|
| 2004 | 2005 | 2006 | |||||
| 落叶阔叶林 DB | 平均值 Mean | 3 255 | 2 868 | 3 267 | 3 130ab | 227 | 7.2 |
| 岳桦林 BE | 1 332 | 2 302 | 2 293 | 1 976A | 557 | 28.2 | |
| 杨桦林 BP | 3 803 | 3 377 | 4 019 | 3 733AB | 327 | 8.8 | |
| 硬阔叶杂木林 DBM | 3 669 | 2 926 | 3 488 | 3 361AB | 387 | 11.5 | |
| 落叶针叶林 DN | 平均值 Mean | 2 803 | 2 095 | 2 113 | 2 337a | 404 | 17.3 |
| 兴安落叶松 LG | 3 018 | 2 363 | 2 130 | 2 504A | 460 | 18.4 | |
| 长白落叶松 LO | 2 266 | 1 560 | 2 086 | 1 971A | 367 | 18.6 | |
| 针阔叶混交林 DNM | 平均值 Mean | 4 312 | 4 066 | 4 060 | 4 146b | 144 | 3.5 |
| 红松林 PK | *** | 3 207 | 3 800 | 3 504AB | 420 | 12.0 | |
| 红松阔叶林 PKDM | 4 312 | 4 925 | 4 319 | 4 519B | 352 | 7.8 | |
| 常绿针叶林 EN | 云冷杉林 PA | 3 018 | 2 110 | 2 288 | 2 472aA | 481 | 19.5 |
| 森林类型 Forest type | 群落类型 Community type | 组分 Component production (%) | |||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 树叶 Leaf | 枝条 Branch | 繁殖器官 Fruit | 其它 Others | ||||||||||
| 落叶阔叶林 DB | 平均值 Mean | 2 549b(76) | 531a(18) | 122a(4) | 59(2) | ||||||||
| 岳桦林 BE | 1 545(66) | 551(26) | 177(7) | 19(1) | |||||||||
| 杨桦林 BP | 3 320(83) | 476(12) | 152(4) | 54(1) | |||||||||
| 硬阔叶杂木林 DBM | 2 781(80) | 565(15) | 37(2) | 103(3) | |||||||||
| 落叶针叶林 DN | 平均值 Mean | 1 428a(73) | 441a(22) | 74a(4) | 33(1) | ||||||||
| 兴安落叶松 LG | 1 589 (82) | 272(15) | 38(2) | 34(2) | |||||||||
| 长白落叶松 LO | 1 023(50) | 865(41) | 165(8) | 32(1) | |||||||||
| 针阔叶混交林 DNM | 平均值 Mean | 3 033b(74) | 679a(16) | 325b(8) | 83 (2) | ||||||||
| 红松林 PK | 2 897(76) | 444(12) | 350(9) | 110(3) | |||||||||
| 红松阔叶林 PKDM | 3 169(72) | 913(20) | 301(7) | 56(1) | |||||||||
| 常绿针叶林 EN | 云冷杉林 PA | 1 280a(54) | 764a(35) | 211ab(9) | 33(1) | ||||||||
表4 2006年各森林类型凋落组分产量(kg·hm-2)及其占总凋落量的比例(%)
Table 4 Productions of different litter components in 2006 (kg·hm-2) (Data in parenthesis are percentages)
| 森林类型 Forest type | 群落类型 Community type | 组分 Component production (%) | |||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 树叶 Leaf | 枝条 Branch | 繁殖器官 Fruit | 其它 Others | ||||||||||
| 落叶阔叶林 DB | 平均值 Mean | 2 549b(76) | 531a(18) | 122a(4) | 59(2) | ||||||||
| 岳桦林 BE | 1 545(66) | 551(26) | 177(7) | 19(1) | |||||||||
| 杨桦林 BP | 3 320(83) | 476(12) | 152(4) | 54(1) | |||||||||
| 硬阔叶杂木林 DBM | 2 781(80) | 565(15) | 37(2) | 103(3) | |||||||||
| 落叶针叶林 DN | 平均值 Mean | 1 428a(73) | 441a(22) | 74a(4) | 33(1) | ||||||||
| 兴安落叶松 LG | 1 589 (82) | 272(15) | 38(2) | 34(2) | |||||||||
| 长白落叶松 LO | 1 023(50) | 865(41) | 165(8) | 32(1) | |||||||||
| 针阔叶混交林 DNM | 平均值 Mean | 3 033b(74) | 679a(16) | 325b(8) | 83 (2) | ||||||||
| 红松林 PK | 2 897(76) | 444(12) | 350(9) | 110(3) | |||||||||
| 红松阔叶林 PKDM | 3 169(72) | 913(20) | 301(7) | 56(1) | |||||||||
| 常绿针叶林 EN | 云冷杉林 PA | 1 280a(54) | 764a(35) | 211ab(9) | 33(1) | ||||||||
图2 凋落物产量随海拔、纬度的变化 a: 长白山海拔梯度上凋落物产量的变化 Litter production along the altitudinal gradient of Mt. Changbai b: 凋落物产量随纬度的变化, 图中为各研究地点海拔小于1 000 m的样地数据 Changes of litter production along the latitudinal gradients in NE China, data shown were from plots with a similar altitude (307-970 m)
Fig. 2 Changes of litter production with altitude and latitude
图3 凋落物产量和年均温、年降水的关系 a、b: 凋落物总量 Litter production c、d: 叶、枝凋落量 Production of leaf and branch litter 图中回归线在p<0.05水平显著 The regression lines were shown for relationships significant at p<0.05 MAT、MAP: 见表5 See Table 5
Fig. 3 Relationship between litter production and climatic indices
| 凋落物总量 Litter production | 组分 Component production | 百分比 Percentage | |||||
|---|---|---|---|---|---|---|---|
| 树叶 Leaf | 枝条 Branch | 繁殖器官 Fruit | 树叶 Leaf | 枝条 Branch | 繁殖器官 Fruit | ||
| 气候 Climate | |||||||
| 年均温 MAT | 0.75*** | 0.57** | 0.06 | 0.11 | 0.30 | -0.32 | -0.15 |
| 年降水 MAP | -0.30 | -0.44 * | 0.48* | 0.21 | -0.75*** | 0.68*** | 0.51** |
| 群落结构特征 Community structure | |||||||
| 平均胸径 Da | -0.08 | -0.03 | 0.26 | 0.49** | -0.31 | 0.18 | 0.50** |
| 平均树高 Ha | -0.03 | -0.02 | 0.27 | 0.22 | -0.24 | 0.19 | 0.22 |
| 立木密度 Density | -0.17 | -0.22 | 0.12 | -0.20 | -0.25 | 0.31 | -0.05 |
| 蓄积量 Volume | 0.26 | 0.14 | 0.55** | 0.65*** | -0.39* | 0.26 | 0.55** |
表5 凋落物产量和气候、群落结构指标的相关系数
Table 5 Correlation between litter production and climate, community structure
| 凋落物总量 Litter production | 组分 Component production | 百分比 Percentage | |||||
|---|---|---|---|---|---|---|---|
| 树叶 Leaf | 枝条 Branch | 繁殖器官 Fruit | 树叶 Leaf | 枝条 Branch | 繁殖器官 Fruit | ||
| 气候 Climate | |||||||
| 年均温 MAT | 0.75*** | 0.57** | 0.06 | 0.11 | 0.30 | -0.32 | -0.15 |
| 年降水 MAP | -0.30 | -0.44 * | 0.48* | 0.21 | -0.75*** | 0.68*** | 0.51** |
| 群落结构特征 Community structure | |||||||
| 平均胸径 Da | -0.08 | -0.03 | 0.26 | 0.49** | -0.31 | 0.18 | 0.50** |
| 平均树高 Ha | -0.03 | -0.02 | 0.27 | 0.22 | -0.24 | 0.19 | 0.22 |
| 立木密度 Density | -0.17 | -0.22 | 0.12 | -0.20 | -0.25 | 0.31 | -0.05 |
| 蓄积量 Volume | 0.26 | 0.14 | 0.55** | 0.65*** | -0.39* | 0.26 | 0.55** |
图4 树叶枝条比例和年均温、年降水的关系 图中回归线在p<0.05水平显著 The regression lines were shown for relationships significant at p<0.05 MAP、MAT: 见表5 See Table 5
Fig. 4 Relationship between climate indices and percentage of leaf and branch litter
| 变量 Variable | Df | Mean Sq | p | %SS | Df | Mean Sq | p | %SS |
|---|---|---|---|---|---|---|---|---|
| 凋落物总量 Litter production | 树叶 Leaf | |||||||
| 年均温 MAT | 1 | 18 994 353 | 0.000 | 56.17 | 1 | 8 420 356 | 0.000 | 32.55 |
| 年降水 MAP | 1 | 12 101 | 0.883 | 0.04 | 1 | 1 540 973 | 0.045 | 5.96 |
| 森林类型 Forest type | 3 | 568 994 | 0.395 | 5.05 | 3 | 2 908 711 | 0.001 | 33.73 |
| 群落类型 Community type | 4 | 613 267 | 0.374 | 7.25 | 4 | 303 022 | 0.471 | 4.68 |
| 蓄积量 Volume | 1 | 2 028 856 | 0.070 | 6.00 | 1 | 762 445 | 0.145 | 2.95 |
| 残差 Residuals | 16 | 538 843 | 25.49 | 16 | 325 606 | 20.14 | ||
| 枝条 Branch | 繁殖器官 Fruit | |||||||
| 年均温 MAT | 1 | 10 670 | 0.631 | 0.42 | 1 | 5 241 | 0.390 | 1.20 |
| 年降水 MAP | 1 | 758 214 | 0.001 | 29.66 | 1 | 32 402 | 0.043 | 7.41 |
| 森林类型 Forest type | 3 | 53 574 | 0.340 | 6.29 | 3 | 73 215 | 0.000 | 50.20 |
| 群落类型 Community type | 4 | 103 302 | 0.101 | 16.16 | 4 | 13 158 | 0.150 | 12.03 |
| 蓄积量 Volume | 1 | 501 654 | 0.004 | 19.62 | 1 | 20 203 | 0.102 | 4.62 |
| 残差 Residuals | 16 | 44 502 | 27.85 | 16 | 6 715 | 24.55 | ||
表6 气候、森林类型、群落结构指标对凋落物产量的影响
Table 6 Effects of climate, forest type and community structure on litter production
| 变量 Variable | Df | Mean Sq | p | %SS | Df | Mean Sq | p | %SS |
|---|---|---|---|---|---|---|---|---|
| 凋落物总量 Litter production | 树叶 Leaf | |||||||
| 年均温 MAT | 1 | 18 994 353 | 0.000 | 56.17 | 1 | 8 420 356 | 0.000 | 32.55 |
| 年降水 MAP | 1 | 12 101 | 0.883 | 0.04 | 1 | 1 540 973 | 0.045 | 5.96 |
| 森林类型 Forest type | 3 | 568 994 | 0.395 | 5.05 | 3 | 2 908 711 | 0.001 | 33.73 |
| 群落类型 Community type | 4 | 613 267 | 0.374 | 7.25 | 4 | 303 022 | 0.471 | 4.68 |
| 蓄积量 Volume | 1 | 2 028 856 | 0.070 | 6.00 | 1 | 762 445 | 0.145 | 2.95 |
| 残差 Residuals | 16 | 538 843 | 25.49 | 16 | 325 606 | 20.14 | ||
| 枝条 Branch | 繁殖器官 Fruit | |||||||
| 年均温 MAT | 1 | 10 670 | 0.631 | 0.42 | 1 | 5 241 | 0.390 | 1.20 |
| 年降水 MAP | 1 | 758 214 | 0.001 | 29.66 | 1 | 32 402 | 0.043 | 7.41 |
| 森林类型 Forest type | 3 | 53 574 | 0.340 | 6.29 | 3 | 73 215 | 0.000 | 50.20 |
| 群落类型 Community type | 4 | 103 302 | 0.101 | 16.16 | 4 | 13 158 | 0.150 | 12.03 |
| 蓄积量 Volume | 1 | 501 654 | 0.004 | 19.62 | 1 | 20 203 | 0.102 | 4.62 |
| 残差 Residuals | 16 | 44 502 | 27.85 | 16 | 6 715 | 24.55 | ||
图5 不同森林类型凋落物产量的差异 a: 叶凋落量 Production of leaf litter b: 繁殖器官凋落量 Production of fruit litter 在分析之前已消除了气候指标(年均温和年降水)对凋落物产量的影响(详见研究方法), 上标字母相同者在p<0.05水平上无显著差异 Effects of climate indices on litter production have been removed before analysis. Forest types that shared the same superscript letter were not significantly different at 0.05 DB、DN、DNM、EN: 见表1 See Table 1
Fig. 5 Different litter production between forest types
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