Chin J Plant Ecol ›› 2019, Vol. 43 ›› Issue (2): 94-106.doi: 10.17521/cjpe.2018.0312

• Research Articles • Previous Articles     Next Articles

Effects of functional diversity on productivity in a typical mixed broadleaved-Korean pine forest

WEN Chun1,JIN Guang-Ze1,2,*()   

  1. 1 Center for Ecological Research, Northeast Forestry University, Harbin 150040, China
    2 Key Laboratory of Sustainable Forest Ecosystem Management- Ministry of Education, Northeast Forestry University, Harbin 150040, China
  • Received:2018-12-13 Accepted:2019-02-13 Online:2019-06-04 Published:2019-02-20
  • Contact: JIN Guang-Ze
  • Supported by:
    Supported by the National Natural Science Foundation of China(31730015);the Fundamental Research Funds for the Central Universities(2572017EA02)

Abstract: <i>Aims</i>

Exploring the relationship between biodiversity and ecosystem productivity has become a hot topic in ecological research. The results support the mass ratio hypothesis and niche complementarity hypothesis, but their relative importance is still controversial. Our aim is to test the relative importance of these two hypotheses in explaining the variability of productivity, and to explore whether the relationship between biodiversity and productivity is influenced by both biotic and abiotic factors.


We used the data of woody plants in a 9 hm2 typical mixed broadleaved-Korean pine forest. By calculating the initial biomass, species diversity, functional diversity, community-weighted mean functional traits and measuring environmental factors, we analyzed the relationship between diversity and productivity by the linear regression model and structural equation model.

<i>Important findings</i>

The results showed that: (1) Both species diversity and functional diversity played a significant role in productivity, and functional diversity was more closely related to productivity than species diversity; (2) the functional diversity index could better explain the variation of productivity than community-weighted mean functional traits. It suggested that the niche complementarity hypothesis was more suitable for explaining the variation of productivity in the mixed broadleaved-Korean pine forest; (3) the relationship between biodiversity and productivity was affected by biotic and abiotic factors, and compared with diversity and functional character composition (vegetation quality), initial stand biomass (vegetation quantity) could explain the variation of productivity more effectively. Our study suggests that, it is important to maintain forest functional diversity and strengthen the protection of plants and soil environments to increase productivity and biodiversity effectively.

Key words: functional diversity, forest productivity, niche complementarity hypothesis, mass ratio hypothesis, structural equation model

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
Leaf dry matter content (LDMC) (mg·g-1)
养分吸收, 结构物质 Nutrient absorption, structural substance
Leaf carbon concentration (LCC) (mg·g-1)
结构物质, 养分吸收 Structural substance, nutrient absorption
Leaf nitrogen concentration (LNC) (mg·g-1)
叶经济谱, 氮吸收 Leaf economic spectrum, nitrogen acquisition

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

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
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
Niche complementarity
物种丰富度 Species richness (C) 6.42 2-13
香农指数 Shannon index () 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."

Table 4

Evaluation of linear regression models for forest productivity and diversity in a typical mixed broadleaved-Korean pine forest"

Model parameters
Species diversity model
功能多样性模型 Functional diversity models
R2 0.169 0.269 0.188 0.377
AIC 2 148.61 2 047.73 2 136.52 1 932.43

Table 5

Relative importance of variables in linear model constructed with productivity"

解释变量 Explanatory variable
Variable importance
1.00 1.00 0.90 0.67 0.48 0.39 0.14
Containing models
10 10 9 6 5 4 2
解释变量 Explanatory variable
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
Containing models
8 8 7 7 7 5 5 5 5 3 2 2
解释变量 Explanatory variable
Total P
Available P
Organic C
Available N
Bulk moisture
Variable importance
1.00 0.95 0.85 0.68 0.53 0.26 0.21 0.09 0.04
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."

Table 6

Direct, indirect, and total standardized effects on the forest productivity, based on the structural equation models (SEM)"

Explanatory variable
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
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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