Chin J Plant Ecol ›› 2019, Vol. 43 ›› Issue (2): 94-106.DOI: 10.17521/cjpe.2018.0312
Special Issue: 生物多样性
• Research Articles • Previous Articles Next Articles
WEN Chun1,JIN Guang-Ze1,2,*(
)
Received:2018-12-13
Accepted:2019-02-13
Online:2019-02-20
Published:2019-06-04
Contact:
JIN Guang-Ze
Supported by:WEN Chun, JIN Guang-Ze. Effects of functional diversity on productivity in a typical mixed broadleaved-Korean pine forest[J]. Chin J Plant Ecol, 2019, 43(2): 94-106.
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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2018.0312
| 功能性状 Functional trait | 功能意义 Functional significance |
|---|---|
| 木材密度 Wood density (WD) (g·mm-3) | 木材经济谱, 生长和生存之间的权衡, 水的运输和分配 Wood economic spectrum, trade-off between growth, transport and distribution of water |
| 最大高度 Maximum height (MH) (m) | 植物竞争活力与策略, 光生态位, 结构多样性 Plant competitive vigor and strategy, light niche, structural diversity |
| 比叶面积 Specific leaf area (SLA) (mm2·g-1) | 叶经济谱, 植物耐荫性 Leaf economic spectrum, plant shade tolerance |
| 叶面积 Leaf area (LA) (mm2) | 光竞争, 蒸腾速率 Light acquisition, transpiration rate |
| 叶片厚度 Leaf thickness (LT) (mm) | 储水能力, 蒸腾速率 Storage capacity, transpiration rate |
| 叶片干物质含量 Leaf dry matter content (LDMC) (mg·g-1) | 养分吸收, 结构物质 Nutrient absorption, structural substance |
| 叶片碳含量 Leaf carbon concentration (LCC) (mg·g-1) | 结构物质, 养分吸收 Structural substance, nutrient absorption |
| 叶片氮含量 Leaf nitrogen concentration (LNC) (mg·g-1) | 叶经济谱, 氮吸收 Leaf economic spectrum, nitrogen acquisition |
Table 1 Eight functional traits related to plant growth and community woody productivity and their significance
| 功能性状 Functional trait | 功能意义 Functional significance |
|---|---|
| 木材密度 Wood density (WD) (g·mm-3) | 木材经济谱, 生长和生存之间的权衡, 水的运输和分配 Wood economic spectrum, trade-off between growth, transport and distribution of water |
| 最大高度 Maximum height (MH) (m) | 植物竞争活力与策略, 光生态位, 结构多样性 Plant competitive vigor and strategy, light niche, structural diversity |
| 比叶面积 Specific leaf area (SLA) (mm2·g-1) | 叶经济谱, 植物耐荫性 Leaf economic spectrum, plant shade tolerance |
| 叶面积 Leaf area (LA) (mm2) | 光竞争, 蒸腾速率 Light acquisition, transpiration rate |
| 叶片厚度 Leaf thickness (LT) (mm) | 储水能力, 蒸腾速率 Storage capacity, transpiration rate |
| 叶片干物质含量 Leaf dry matter content (LDMC) (mg·g-1) | 养分吸收, 结构物质 Nutrient absorption, structural substance |
| 叶片碳含量 Leaf carbon concentration (LCC) (mg·g-1) | 结构物质, 养分吸收 Structural substance, nutrient absorption |
| 叶片氮含量 Leaf nitrogen concentration (LNC) (mg·g-1) | 叶经济谱, 氮吸收 Leaf economic spectrum, nitrogen acquisition |
| 变量 Variables | 范围 Range | 平均值 Mean | 标准偏差 SD |
|---|---|---|---|
| 凹凸度 Convexity | -1.91-2.36 | 0.01 | 0.40 |
| 海拔 Elevation (m) | 425.45-505.52 | 463.25 | 18.20 |
| 坡度 Slope (°) | 3.30-38.46 | 15.91 | 6.49 |
| 速效氮含量 Soil available nitrogen content (mg·kg-1) | 605.77-1 482.85 | 1 103.60 | 161.34 |
| 速效磷含量 Soil available phosphorus content (mg·kg-1) | 2.54-61.96 | 8.83 | 6.96 |
| 速效钾含量 Soil available potassium content (mg·kg-1) | 190.46-516.66 | 332.61 | 59.05 |
| 全氮含量 Soil total nitrogen content (g·kg-1) | 4.60-12.28 | 8.40 | 1.60 |
| 全磷含量 Soil total phosphorus content (g·kg-1) | 0.40-1.19 | 0.82 | 0.15 |
| pH值 Soil pH | 5.34-6.29 | 5.77 | 0.16 |
| 有机碳含量 Soil organic carbon content (%) | 31.29-216.77 | 75.75 | 31.25 |
| 土壤容重 Soil bulk density (mg·m-3) | 0.41-0.87 | 0.62 | 0.10 |
| 体积含水率 Soil bulk moisture content (%) | 13.72-48.40 | 30.35 | 5.91 |
| 质量含水率 Soil mass moisture content (%) | 0.54-1.51 | 0.91 | 0.23 |
Table 2 Summary of environment variables in a 9 hm2 mixed broadleaved-Korean pine forest plot
| 变量 Variables | 范围 Range | 平均值 Mean | 标准偏差 SD |
|---|---|---|---|
| 凹凸度 Convexity | -1.91-2.36 | 0.01 | 0.40 |
| 海拔 Elevation (m) | 425.45-505.52 | 463.25 | 18.20 |
| 坡度 Slope (°) | 3.30-38.46 | 15.91 | 6.49 |
| 速效氮含量 Soil available nitrogen content (mg·kg-1) | 605.77-1 482.85 | 1 103.60 | 161.34 |
| 速效磷含量 Soil available phosphorus content (mg·kg-1) | 2.54-61.96 | 8.83 | 6.96 |
| 速效钾含量 Soil available potassium content (mg·kg-1) | 190.46-516.66 | 332.61 | 59.05 |
| 全氮含量 Soil total nitrogen content (g·kg-1) | 4.60-12.28 | 8.40 | 1.60 |
| 全磷含量 Soil total phosphorus content (g·kg-1) | 0.40-1.19 | 0.82 | 0.15 |
| pH值 Soil pH | 5.34-6.29 | 5.77 | 0.16 |
| 有机碳含量 Soil organic carbon content (%) | 31.29-216.77 | 75.75 | 31.25 |
| 土壤容重 Soil bulk density (mg·m-3) | 0.41-0.87 | 0.62 | 0.10 |
| 体积含水率 Soil bulk moisture content (%) | 13.72-48.40 | 30.35 | 5.91 |
| 质量含水率 Soil mass moisture content (%) | 0.54-1.51 | 0.91 | 0.23 |
| 假说 Hypothesis | 指数 Index | 平均值 Mean | 范围 Range |
|---|---|---|---|
| 生物量比率假说 Mass ratio hypothesis | 木材密度的群落加权平均值 CWM of wood density (CWMWD) (g·mm-3) | 0.47 | 0.34-0.63 |
| 最大树高的群落加权平均值 CWM of maximum height (CWMMH) (m) | 17.24 | 4.59-43.01 | |
| 比叶面积的群落加权平均值 CWM of specific leaf area (CWMSLA) (mm2·g-1) | 254.30 | 158.29-392.23 | |
| 叶面积的群落加权平均值 CWM of leaf area (CWMLA) (mm2) | 33.87 | 8.20-74.54 | |
| 叶片厚度的群落加权平均值 CWM of leaf thickness (CWMLT) (mm) | 0.18 | 0.08-0.44 | |
| 叶干物质含量的群落加权平均值 CWM of leaf dry matter content (CWMLDMC) ( mg·g-1) | 0.29 | 0.19-0.38 | |
| 叶片碳含量的群落加权平均值 CWM of leaf carbon content (CWMLCC) ( mg·g-1) | 499.22 | 446.90-563.74 | |
| 叶片氮含量的群落加权平均值 CWM of leaf nitrogen content (CWMLNC) ( mg·g-1) | 23.56 | 17.49-32.38 | |
| 生态位互补假说 Niche complementarity hypothesis | 物种丰富度 Species richness (C) | 6.42 | 2-13 |
| 香农指数 Shannon index (H°) | 1.45 | 0.41-2.24 | |
| 辛普森指数 Simpson index (D) | 0.71 | 0.24-0.88 | |
| 均匀度指数 Evenness index (J) | 0.89 | 0.42-1.00 | |
| 多维功能均匀度指数 Multidimensional functional evenness index (FEve) | 0.72 | 0.21-0.99 | |
| Rao二次熵指数 Rao’s quadratic entropy index (FDQ) | 8.39 | 0.81-15.41 | |
| 8个性状组合的功能分散指数 FD is based on eight traits combined (FDcom) | 2.68 | 0.72-3.87 | |
| 木材密度的功能分散指数 FD is based on wood density (FDWD) | 0.48 | 0.01-1.13 | |
| 最大高度的功能分散指数 FD is based on maximum height (FDMH) | 0.78 | 0.07-1.71 | |
| 比叶面积的功能分散指数 FD is based on specific leaf area (FDSLA) | 0.63 | 0.02-1.61 | |
| 叶面积的功能分散指数 FD is based on leaf area (FDLA) | 0.66 | 0.02-1.61 | |
| 叶片厚度的功能分散指数 FD is based on leaf thickness (FDLT) | 0.58 | 0.03-1.51 | |
| 叶干物质含量的功能分散指数 FD is based on leaf dry matter content (FDLDMC) | 0.58 | 0.06-1.26 | |
| 叶片碳含量的功能分散指数 FD is based on leaf carbon content (FDLCC) | 0.93 | 0.03-2.37 | |
| 叶片氮含量的功能分散指数 FD is based on leaf nitrogen content (FDLNC) | 0.51 | 0.03-2.38 |
Table 3 Summary of functional diversity, species diversity indexes and community-weighted mean (CWM) functional traits in a 9 hm2 mixed broadleaved-?Korean pine forest plot
| 假说 Hypothesis | 指数 Index | 平均值 Mean | 范围 Range |
|---|---|---|---|
| 生物量比率假说 Mass ratio hypothesis | 木材密度的群落加权平均值 CWM of wood density (CWMWD) (g·mm-3) | 0.47 | 0.34-0.63 |
| 最大树高的群落加权平均值 CWM of maximum height (CWMMH) (m) | 17.24 | 4.59-43.01 | |
| 比叶面积的群落加权平均值 CWM of specific leaf area (CWMSLA) (mm2·g-1) | 254.30 | 158.29-392.23 | |
| 叶面积的群落加权平均值 CWM of leaf area (CWMLA) (mm2) | 33.87 | 8.20-74.54 | |
| 叶片厚度的群落加权平均值 CWM of leaf thickness (CWMLT) (mm) | 0.18 | 0.08-0.44 | |
| 叶干物质含量的群落加权平均值 CWM of leaf dry matter content (CWMLDMC) ( mg·g-1) | 0.29 | 0.19-0.38 | |
| 叶片碳含量的群落加权平均值 CWM of leaf carbon content (CWMLCC) ( mg·g-1) | 499.22 | 446.90-563.74 | |
| 叶片氮含量的群落加权平均值 CWM of leaf nitrogen content (CWMLNC) ( mg·g-1) | 23.56 | 17.49-32.38 | |
| 生态位互补假说 Niche complementarity hypothesis | 物种丰富度 Species richness (C) | 6.42 | 2-13 |
| 香农指数 Shannon index (H°) | 1.45 | 0.41-2.24 | |
| 辛普森指数 Simpson index (D) | 0.71 | 0.24-0.88 | |
| 均匀度指数 Evenness index (J) | 0.89 | 0.42-1.00 | |
| 多维功能均匀度指数 Multidimensional functional evenness index (FEve) | 0.72 | 0.21-0.99 | |
| Rao二次熵指数 Rao’s quadratic entropy index (FDQ) | 8.39 | 0.81-15.41 | |
| 8个性状组合的功能分散指数 FD is based on eight traits combined (FDcom) | 2.68 | 0.72-3.87 | |
| 木材密度的功能分散指数 FD is based on wood density (FDWD) | 0.48 | 0.01-1.13 | |
| 最大高度的功能分散指数 FD is based on maximum height (FDMH) | 0.78 | 0.07-1.71 | |
| 比叶面积的功能分散指数 FD is based on specific leaf area (FDSLA) | 0.63 | 0.02-1.61 | |
| 叶面积的功能分散指数 FD is based on leaf area (FDLA) | 0.66 | 0.02-1.61 | |
| 叶片厚度的功能分散指数 FD is based on leaf thickness (FDLT) | 0.58 | 0.03-1.51 | |
| 叶干物质含量的功能分散指数 FD is based on leaf dry matter content (FDLDMC) | 0.58 | 0.06-1.26 | |
| 叶片碳含量的功能分散指数 FD is based on leaf carbon content (FDLCC) | 0.93 | 0.03-2.37 | |
| 叶片氮含量的功能分散指数 FD is based on leaf nitrogen content (FDLNC) | 0.51 | 0.03-2.38 |
Fig. 1 Relationships between coarse woody productivity (ln transformed) and species diversity index of communities in a typical mixed broadleaved-Korean pine forest. CWP, C, H°, D represent coarse woody productivity, species richness, Shannon index and Simpson index, respectively.
Fig. 2 Relationships between coarse woody productivity (ln transformed) and functional diversity index of communities in a typical mixed broadleaved-Korean pine forest. CWP, coarse woody productivity; FEve, multidimensional functional evenness index; FDQ, Rao’s quadratic entropy index; FDcom, functional dispersion indices of eight trait combinations; FDWD, functional dispersion indices of wood density; FDMH, functional dispersion indices of maximum height; FDSLA, functional dispersion indices of specific leaf area; FDLA, functional dispersion indices of leaf area; FDLDMC, functional dispersion indices of leaf dry matter content; FDLCC, functional dispersion indices of leaf carbon content; FDLNC, functional dispersion indices of leaf nitrogen content.
Fig. 3 Relationships between coarse woody productivity (ln transformed) and community weighted mean of different traits of communities in a typical mixed broadleaved-Korean pine forest. CWP, coarse woody productivity; CWMWD, CWM of wood density; CWMMH, CWM of maximum height; CWMSLA, CWM of specific leaf area; CWMLA, CWM of leaf area; CWMLDMC, CWM of leaf dry matter content; CWMLCC, CWM of leaf carbon content; CWMLNC, CWM of leaf nitrogen content.
Fig. 4 Relationships between coarse woody productivity (ln transformed, CWP) and environmental factors in a typical mixed broadleaved-Korean pine forest.
| 模型参数 Model parameters | 物种多样性模型 Species diversity model | 功能多样性模型 Functional diversity models | ||
|---|---|---|---|---|
| A | B | C | ||
| R2 | 0.169 | 0.269 | 0.188 | 0.377 |
| AIC | 2 148.61 | 2 047.73 | 2 136.52 | 1 932.43 |
Table 4 Evaluation of linear regression models for forest productivity and diversity in a typical mixed broadleaved-Korean pine forest
| 模型参数 Model parameters | 物种多样性模型 Species diversity model | 功能多样性模型 Functional diversity models | ||
|---|---|---|---|---|
| A | B | C | ||
| R2 | 0.169 | 0.269 | 0.188 | 0.377 |
| AIC | 2 148.61 | 2 047.73 | 2 136.52 | 1 932.43 |
| 指标 Indicator | 解释变量 Explanatory variable | |||||||||||||||||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| CWMMH | CWMLCC | CWMSLA | CWMLNC | CWMLA | CWMWD | CWMLDMC | ||||||||||||||||||||
| 变量重要性 Variable importance | 1.00 | 1.00 | 0.90 | 0.67 | 0.48 | 0.39 | 0.14 | |||||||||||||||||||
| 含变量模型数 Containing models | 10 | 10 | 9 | 6 | 5 | 4 | 2 | |||||||||||||||||||
| 指标 Indicator | 解释变量 Explanatory variable | |||||||||||||||||||||||||
| FDMH | FDQ | FDcom | FDSLA | C | D | H° | FDLA | FDWD | FDLCC | FEve | FDLDMC | |||||||||||||||
| 变量重要性 Variable importance | 1.00 | 1.00 | 0.88 | 0.88 | 0.88 | 0.69 | 0.69 | 0.69 | 0.69 | 0.36 | 0.21 | 0.19 | ||||||||||||||
| 含变量模型数 Containing models | 8 | 8 | 7 | 7 | 7 | 5 | 5 | 5 | 5 | 3 | 2 | 2 | ||||||||||||||
| 指标 Indicator | 解释变量 Explanatory variable | |||||||||||||||||||||||||
| 海拔 Elevation | 全磷含量 Total P content | 凹凸度 Convexity | 速效磷含量 Available P content | 有机碳含量 Organic C content | 速效氮含量 Available N content | 体积含水率 Bulk moisture | 坡度 Slope | pH | ||||||||||||||||||
| 变量重要性 Variable importance | 1.00 | 0.95 | 0.85 | 0.68 | 0.53 | 0.26 | 0.21 | 0.09 | 0.04 | |||||||||||||||||
| 含变量模型数 Containing models | 17 | 16 | 14 | 11 | 9 | 4 | 4 | 2 | 1 | |||||||||||||||||
Table 5 Relative importance of variables in linear model constructed with productivity
| 指标 Indicator | 解释变量 Explanatory variable | |||||||||||||||||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| CWMMH | CWMLCC | CWMSLA | CWMLNC | CWMLA | CWMWD | CWMLDMC | ||||||||||||||||||||
| 变量重要性 Variable importance | 1.00 | 1.00 | 0.90 | 0.67 | 0.48 | 0.39 | 0.14 | |||||||||||||||||||
| 含变量模型数 Containing models | 10 | 10 | 9 | 6 | 5 | 4 | 2 | |||||||||||||||||||
| 指标 Indicator | 解释变量 Explanatory variable | |||||||||||||||||||||||||
| FDMH | FDQ | FDcom | FDSLA | C | D | H° | FDLA | FDWD | FDLCC | FEve | FDLDMC | |||||||||||||||
| 变量重要性 Variable importance | 1.00 | 1.00 | 0.88 | 0.88 | 0.88 | 0.69 | 0.69 | 0.69 | 0.69 | 0.36 | 0.21 | 0.19 | ||||||||||||||
| 含变量模型数 Containing models | 8 | 8 | 7 | 7 | 7 | 5 | 5 | 5 | 5 | 3 | 2 | 2 | ||||||||||||||
| 指标 Indicator | 解释变量 Explanatory variable | |||||||||||||||||||||||||
| 海拔 Elevation | 全磷含量 Total P content | 凹凸度 Convexity | 速效磷含量 Available P content | 有机碳含量 Organic C content | 速效氮含量 Available N content | 体积含水率 Bulk moisture | 坡度 Slope | pH | ||||||||||||||||||
| 变量重要性 Variable importance | 1.00 | 0.95 | 0.85 | 0.68 | 0.53 | 0.26 | 0.21 | 0.09 | 0.04 | |||||||||||||||||
| 含变量模型数 Containing models | 17 | 16 | 14 | 11 | 9 | 4 | 4 | 2 | 1 | |||||||||||||||||
Fig. 5 Structural equation model (SEM) analysis of the effects of environmental factors, functional diversity, community-weight mean of functional traits and forest initial biomass on community woody productivity in a typical mixed broadleaved-Korean pine forest. Solid lines indicate significant paths, while dashed lines indicate insignificant paths. R² indicates the total variation in a dependent variable that is explained by the combined independent variables. ***, p < 0.001; **, p < 0.01; *, p < 0.05.
| 解释变量 Explanatory variable | 对生产力的作用途径 Pathway to productivity | 结构方程模型 SEM | |||
|---|---|---|---|---|---|
| 影响效应 Effect | p | ||||
| 地形因子 Topography factors | 直接效应 Direct effect | -0.102 | 0.003 | ||
| 功能多样性的间接效应 Indirect effect via functional diversity | -0.027 | 0.009 | |||
| 群落加权平均值的间接效应 Indirect effect via CWMs | 0.008 | 0.043 | |||
| 初始生物量的间接效应 Indirect effect via initial biomass | 0.055 | <0.001 | |||
| 土壤因子的间接效应 Indirect effect via soil factors | -0.024 | 0.002 | |||
| 总效应 Total effect | -0.090 | <0.001 | |||
| 土壤因子 Edaphic factors | 直接效应 Direct effect | 0.105 | <0.001 | ||
| 功能多样性的间接效应 Indirect effect via functional diversity | -0.021 | 0.045 | |||
| 群落加权平均值的间接效应 Indirect effect via CWMs | -0.015 | 0.008 | |||
| 总效应 Total effect | 0.069 | 0.019 | |||
| 功能多样性 Functional diversity | 直接效应 Direct effect | 0.302 | <0.001 | ||
| 初始生物量的间接效应 Indirect effect via initial biomass | 0.081 | <0.001 | |||
| 总效应 Total effect | 0.383 | <0.001 | |||
| 群落加权平均值 Community-weighted mean functional traits | 直接效应 Direct effect | 0.090 | 0.002 | ||
| 初始生物量的间接效应 Indirect effect via initial biomass | 0.152 | <0.001 | |||
| 总效应 Total effect | 0.242 | <0.001 | |||
| 地上初始生物量 Initial standing biomass | 直接效应 Direct effect | 0.449 | <0.001 | ||
| 总效应 Total effect | 0.449 | <0.001 | |||
Table 6 Direct, indirect, and total standardized effects on the forest productivity, based on the structural equation models (SEM)
| 解释变量 Explanatory variable | 对生产力的作用途径 Pathway to productivity | 结构方程模型 SEM | |||
|---|---|---|---|---|---|
| 影响效应 Effect | p | ||||
| 地形因子 Topography factors | 直接效应 Direct effect | -0.102 | 0.003 | ||
| 功能多样性的间接效应 Indirect effect via functional diversity | -0.027 | 0.009 | |||
| 群落加权平均值的间接效应 Indirect effect via CWMs | 0.008 | 0.043 | |||
| 初始生物量的间接效应 Indirect effect via initial biomass | 0.055 | <0.001 | |||
| 土壤因子的间接效应 Indirect effect via soil factors | -0.024 | 0.002 | |||
| 总效应 Total effect | -0.090 | <0.001 | |||
| 土壤因子 Edaphic factors | 直接效应 Direct effect | 0.105 | <0.001 | ||
| 功能多样性的间接效应 Indirect effect via functional diversity | -0.021 | 0.045 | |||
| 群落加权平均值的间接效应 Indirect effect via CWMs | -0.015 | 0.008 | |||
| 总效应 Total effect | 0.069 | 0.019 | |||
| 功能多样性 Functional diversity | 直接效应 Direct effect | 0.302 | <0.001 | ||
| 初始生物量的间接效应 Indirect effect via initial biomass | 0.081 | <0.001 | |||
| 总效应 Total effect | 0.383 | <0.001 | |||
| 群落加权平均值 Community-weighted mean functional traits | 直接效应 Direct effect | 0.090 | 0.002 | ||
| 初始生物量的间接效应 Indirect effect via initial biomass | 0.152 | <0.001 | |||
| 总效应 Total effect | 0.242 | <0.001 | |||
| 地上初始生物量 Initial standing biomass | 直接效应 Direct effect | 0.449 | <0.001 | ||
| 总效应 Total effect | 0.449 | <0.001 | |||
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