群落物种及结构多样性对森林地上生物量的影响及其尺度效应: 以巴拿马BCI样地为例

doi:10.17520/biods.2017155

摘要/Abstract

摘要：

随着全球物种多样性丧失速率的日益加剧, 森林群落(特别是占陆地碳储量一半的热带森林)内物种多样性与地上生物量之间的关系愈加受到关注。一般认为森林群落的物种多样性能够促进地上生物量的累积。但随着研究的深入, 研究人员发现物种多样性对地上生物量的促进作用可能会受到结构多样性和非生物环境因子的干扰或抵消, 而且对于影响地上生物量的各类因素的作用强度是否会随空间尺度的不同而发生变化也还存在争议。本文基于巴拿马Barro Colorado岛(BCI) 50 ha大型森林动态监测样地的调查数据, 采用结构方程模型对影响其地上生物量的直接及间接因素进行分析。结果表明: 随着空间尺度的增加, 物种多样性对地上生物量的作用由不显著转为显著负作用, 且强度逐渐增加; 而结构多样性对地上生物量均具有显著正作用, 且强度逐渐减弱。非生物因子与两种多样性之间均具有一定的相关性, 并进而对地上生物量产生间接影响。这一结果暗示未考虑结构多样性而得到的物种多样性与地上生物量的正相关关系可能是由结构多样性介导或受非生物因素的间接作用所致, 而在森林群落特殊的林窗及强烈的非对称性竞争中, 物种多样性与地上生物量更可能呈现负相关关系。

关键词: 地上生物量, 物种多样性, 结构多样性, 空间尺度, 非生物因素, 结构方程模型, 大型森林动态样地

Abstract

With global decreases in species diversity, the effects of species diversity on aboveground biomass of forest communities, especially tropical forests which store nearly half of terrestrial carbon, have received much attention. It is commonly assumed that species diversity can promote the accumulation of aboveground biomass in forest communities. However, increasing evidence suggests that this positive effect of species diversity can be influenced or neutralized by structural diversity and various abiotic environmental variables. It is still far from clear whether scale dependent effects of various drivers of aboveground biomass exist. Using structural equation modeling, we examined direct and indirect drivers of aboveground biomass in a 50 ha tropical forest dynamics plot on Barro Colorado Island (BCI), Panama. Our results showed that the effect of species diversity on aboveground biomass was non-significant at the smallest scale and became significantly negative at larger scales. The strength of this negative effect increased with spatial scales. Conversely, structural diversity had a significant positive effect on aboveground biomass and the strength of this effect decreased with scale. Abiotic variables had some relationships with both types of diversity and can indirectly affect aboveground biomass. These results suggest that positive relationships between species diversity and aboveground biomass in conventional studies may be caused by structural diversity. The relationship between species diversity and aboveground biomass is more likely to be negative, due to under the gap dynamics and strong asymmetric competition in forest communities.

Key words: aboveground biomass, species diversity, structural diversity, spatial scale, abiotic factors, structural equation modeling, large-scale forest dynamics plot

谭珊珊, 王忍忍, 龚筱羚, 蔡佳瑶, 沈国春 (2017) 群落物种及结构多样性对森林地上生物量的影响及其尺度效应: 以巴拿马BCI样地为例. 生物多样性, 25, 1054-1064. DOI: 10.17520/biods.2017155.

Shanshan Tan, Renren Wang, Xiaoling Gong, Jiayao Cai, Guochun Shen (2017) Scale dependent effects of species diversity and structural diversity on aboveground biomass in a tropical forest on Barro Colorado Island, Panama. Biodiversity Science, 25, 1054-1064. DOI: 10.17520/biods.2017155.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2017155

https://www.biodiversity-science.net/CN/Y2017/V25/I10/1054

图/表 4

图1 热带雨林BCI样地内不同空间尺度下物种多样性与地上生物量的关系。实线及灰色区间为线性回归模型得到的最优拟合直线及其95%的置信区间, R2表示调整后的拟合系数, P值表示整个模型的显著度。

Fig. 1 Relationships between species diversity and total aboveground biomass at different spatial scales in a 50 ha tropical forest dynamics plot (BCI plot). Solid lines and their corresponding grey area are the regression lines and 95% confidence intervals from the best fitted model. R2 and P are the adjusted R2 and significance of the whole regression model, respectively.

图2 热带雨林BCI样地内不同空间尺度下结构多样性与地上生物量的关系。实线及灰色区间为线性回归模型下得到的最优拟合直线及其95%的置信区间。R2表示调整后的拟合系数, P值表示整个模型的显著度。

Fig. 2 Relationships between structural diversity and total aboveground biomass at different spatial scales in a 50 ha tropical forest dynamics plot (BCI plot). Solid lines and their corresponding grey area are the regression lines and 95% confidence intervals from the best fitted model. R2 and P are the adjusted R2 and significance of the whole regression model, respectively.

表1 最优结构方程模型中立木密度、物种丰富度和结构多样性对地上生物量的标准化影响效应

Table 1 The standardized effects of stem density, species richness and structural diversity on the aboveground biomass in our best structural equation model

影响因子 Predictor	对地上生物量的效应 Pathway to aboveground biomass	空间尺度 Spatial scale
影响因子 Predictor	对地上生物量的效应 Pathway to aboveground biomass	10 m × 10 m	20 m × 20 m	50 m × 50 m
立木密度 Stem density	直接效应 Direct effect	-	-	-0.210
	间接效应 Indirect effect	0.273	-0.005	-0.137
	总效应 Total effect	0.273	-0.005	-0.347
物种丰富度 Species richness	直接效应 Direct effect	-	-0.113	-0.284
	间接效应 Indirect effect	0.046	0.029	-
	总效应 Total effect	0.046	-0.084	-0.284
结构多样性 Structural diversity	直接效应 Direct effect	0.650	0.541	0.181
	间接效应 Indirect effect	-	-0.012	-
	总效应 Total effect	0.650	0.529	0.181

图3 不同空间尺度物种多样性和结构多样性与地上生物量的最佳结构方程模型, 以及土壤养分和地形因子对地上生物量的间接影响。实线和虚线箭头分别表示显著的正负作用; 线段粗细表示作用强弱; 线上的数值表示标准通径系数。*P < 0.05; ** P < 0.01; *** P < 0.001。模型评价采用比较拟合指数(CFI)、Tucker-Lewis指数(TLI)、渐进残差均方和平方根(RMSEA)和标准化残差均方和平方根(SRMR)。不同空间尺度最佳结构方程模型评价指标的具体数值详见附录2。

Fig. 3 The best fitted structural equation models (SEM) relating aboveground biomass to species diversity, structural diversity, soil nutrient and topographic factors at different spatial scales. Solid and dashed arrow lines represent the positive and negative effects, respectively. Width of arrow line indicates the strength of the path. Number and its associated asterisks are standardized path coefficient and significant level (*P < 0.05; ** P < 0.01; *** P < 0.001) for each path. Comparative fit index (CFI), Tucker-Lewis index (TLI), Root mean square error of approximation (RMSEA) and standardized root mean square residual (SRMR) were used to compare different alternative models. The corresponding values of these indices for the best fitted model were given in Appendix 2.

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