长白山物种丰富度与物种组成对森林生产力的影响及其随演替的变化

doi:10.17521/cjpe.2024.0060

摘要/Abstract

摘要：

生物多样性是生态系统功能形成与维持的重要影响因素, 但其背后潜在的生态学机制仍存在较大争议。生态位互补效应和生物量比率效应是解释生物多样性与生态系统功能关系的两种主要假说, 但在温带森林中生态位互补效应和生物量比率效应对生态系统功能的相对贡献是否随森林演替而变化, 以及如何变化目前仍不清楚。该研究以长白山地区处于不同演替阶段的森林为研究对象, 分别在次生山杨(Populus davidiana) -白桦(Betula platyphylla)林(演替早期)、次生针阔混交林(演替中期)和原始紫椴(Tilia amurensis) -红松(Pinus koraiensis)林(演替后期)内设置3块面积为5.2 hm²的固定监测样地。基于两期样地调查数据, 分别采用物种丰富度和物种组成验证生态位互补效应和生物量比率效应, 采用地上生物量和森林生产力作为反映生态系统功能的指标, 通过结构方程模型检验生态位互补效应和生物量比率效应对生态系统功能的相对贡献及其随演替的变化。研究显示: 物种丰富度与生态系统功能关系随森林演替而变化, 演替初期物种丰富度的作用并不显著, 而在演替中后期物种丰富度的促进作用逐渐增强, 表明随着森林演替生态位互补效应逐渐增强。相比物种丰富度, 物种组成在森林演替的各个阶段都显著影响着生态系统功能, 表明生物量比率效应在森林演替过程中始终发挥着重要作用。此外, 地上生物量也是影响森林生产力的重要因素。该研究揭示了长白山森林生物多样性与生态系统功能关系及其随演替的变化规律, 研究结果能够为东北地区退化森林的生态恢复以及生物多样性保护提供一定的科学依据。

关键词: 物种丰富度, 物种组成, 地上生物量, 森林生产力, 森林演替

Abstract:

Aims Biodiversity is an important driver in the formation and maintenance of ecosystem functions, but the underlying ecological mechanisms behind it are still highly controversial. The niche complementarity and mass-ratio effects are two main hypotheses that explain the relationship between biodiversity and ecosystem function. However, it is still unclear whether the relative contributions of niche complementarity and mass-ratio effects to ecosystem function in temperate forests change with succession.

Methods Based on the observations from three forest plots in Changbai Mountains, including a secondary Populus davidiana- Betula platyphylla forest (early successional stage), a secondary needleleaf-broadleaf mixed forest (middle successional stage), and a primary Tilia amurensis- Pinus koraiensisforest (late successional stage), this study estimated how biodiversity-ecosystem function relationships change with forest succession. Aboveground biomass and productivity were used as two indicators reflecting ecosystem function. Species richness and species composition were used to represent the niche complementarity and mass-ratio effects, respectively. The relative contributions of niche complementarity and mass-ratio effects to ecosystem function were tested using structural equation modeling.

Important findings The results showed that the relationships between species richness and ecosystem functions change with forest succession. The complementary effect of species richness was not significant in the early stages of succession, but gradually increased in the middle and late stages. Compared with species richness, species composition significantly affected ecosystem functions at all stages of forest succession, indicating that the mass-ratio effect plays an important role during the whole forest succession process. In addition, the results showed that aboveground biomass is also an important factor affecting forest productivity. This study elucidates how the relationships between biodiversity and ecosystem functions change with succession in temperate forests, which provide a scientific support for the ecological restoration and biodiversity protection of secondary forests in northeast China.

Key words: species richness, species composition, aboveground biomass, forest productivity, forest succession

袁鹤洋, 郝珉辉, 何怀江, 张春雨, 赵秀海. 长白山物种丰富度与物种组成对森林生产力的影响及其随演替的变化. 植物生态学报, 2024, 48(12): 1602-1611. DOI: 10.17521/cjpe.2024.0060

YUAN He-Yang, HAO Min-Hui, HE Huai-Jiang, ZHANG Chun-Yu, ZHAO Xiu-Hai. Impact of species richness and composition on productivity and its changes with forest succession in Changbai Mountains, China. Chinese Journal of Plant Ecology, 2024, 48(12): 1602-1611. DOI: 10.17521/cjpe.2024.0060

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0060

https://www.plant-ecology.com/CN/Y2024/V48/I12/1602

图/表 6

图1 长白山不同演替阶段森林环境因子的主成分分析结果排序图。ENV1, 环境第一主成分; ENV2, 环境第一主成分。PTPF, 原始紫椴-红松林; SCBF, 次生针阔混交林; SPBF, 次生山杨-白桦林。ASP, 坡度; ELE, 海拔; N, 土壤全氮含量; OM, 土壤有机质含量; P, 土壤全磷含量; SD, 土壤厚度; SW, 土壤含水量。

Fig. 1 Principal component analysis of environmental factors of forests at different succession stages in Changbai Mountains. ENV1, the first principal component of the environment; ENV2, the second principal component of the environment. PTPF, primary Tilia amurensis - Pinus koraiensis forest; SCBF, secondary needleleaf-broadleaf mixed forest; SPBF, secondary Populus davidiana - Betula platyphylla forest. ASP, aspect; ELE, elevation; N, soil nitrogen content; OM, soil organic matter content; P, soil phosphorus content; SD, soil depth; SW, soil water content.

图2 长白山不同演替阶段森林物种组成的主坐标分析结果排序图。SComp1, 物种组成第一主坐标; SComp2, 物种组成第二主坐标。PTPF, 原始紫椴-红松林; SCBF, 次生针阔混交林; SPBF, 次生山杨-白桦林。AcPs, 紫花槭; AlSi, 辽东桤木; BeCo, 硕桦; BePl, 白桦; FrMa, 水曲柳; PiJe, 鱼鳞云杉; PiKo, 红松; PoDa, 山杨; QuMo, 蒙古栎; TiAm, 紫椴。

Fig. 2 Principal coordinate analysis of species composition of forests at different succession stages in Changbai Mountains. SComp1, the first principal coordinate of species composition; SComp2, the second principal coordinate of species composition. PTPF, primary Tilia amurensis - Pinus koraiensis forest; SCBF, secondary needleleaf-broadleaf mixed forest; SPBF, secondary Populus davidiana - Betula platyphylla forest. AcPs, Acer pseudosieboldianum; AlSi, Alnus sibirica; BeCo, Betula costata; BePl, Betula platyphylla; FrMa, Fraxinus mandshurica; PiJe, Picea jezoensis; PiKo, Pinus koraiensis; PoDa, Populus davidiana; QuMo, Quercus mongolica; TiAm, Tilia amurensis.

图3 长白山不同演替阶段物种丰富度与物种组成的变化。SComp1, 物种组成第一主坐标; SComp2, 物种组成第二主坐标; SR, 物种丰富度。PTPF, 原始紫椴-红松林; SCBF, 次生针阔混交林; SPBF, 次生山杨-白桦林。不同小写字母表示不同演替阶段间差异显著(p < 0.05)。

Fig. 3 Changes in species diversity and composition during forest succession in Changbai Mountains. SComp1, the first principal coordinate of species composition; SComp2, the second principal coordinate of species composition; SR, species richness. PTPF, primary Tilia amurensis - Pinus koraiensis forest; SCBF, secondary needleleaf-broadleaf mixed forest; SPBF, secondary Populus davidiana - Betula platyphylla forest. Different lowercase letters indicate significant differences between different succession stages (p < 0.05).

图4 长白山不同演替阶段物种丰富度和物种组成对生态系统功能的影响。A, 次生山杨-白桦林。B, 次生针阔混交林。C, 原始紫椴-红松林。AGB, 地上生物量; ENV1, 环境第一主成分; ENV2, 环境第二主成分; FP, 森林生产力; SComp1, 物种组成第一主坐标; SComp2, 物种组成第二主坐标; SR, 物种丰富度。红色实线代表显著正相关路径, 黑色实线代表显著负相关路径, 灰色虚线代表不显著路径, 线条上数值表示标准化路径系数, 线条粗细反映路径系数的大小。

Fig. 4 Impact of species richness and composition on ecosystem functions of forests at different succession stages in Changbai Mountains. A, Secondary Populus davidiana - Betula platyphylla forest. B, Secondary needleleaf -broadleaf mixed forest. C, Primary Tilia amurensis - Pinus koraiensis forest. AGB, aboveground biomass; ENV1, the first principal component of the environment; ENV2, the second principal component of the environment; FP, forest productivity; SComp1, the first principal coordinate of species composition; SComp2, the second principal coordinate of species composition; SR, species richness. Red solid lines represent significantly positively correlated paths, black solid lines represent significantly negatively correlated paths, and gray dashed lines represent insignificant paths, the values next to the lines represent the standardized path coefficients, and the line thickness is proportional to the standardized path coefficient.

附录I 长白山森林样地基本信息

Supplement I Basic information of the three forest plots in Changbai Mountains

样地 Plot	样地位置 Location	海拔 Altitude (m)	主要树种 Main species
次生山杨-白桦林 Secondary Populus davidiana - Betula platyphylla forest	42.32° N, 128.13° E	899	白桦、山杨、水曲柳、紫椴 Betula platyphylla, Populus davidiana, Fraxinus mandschurica, Tilia amurensis
次生针阔混交林 Secondary needleleaf-broadleaf mixed forest	42.35° N, 128.13° E	748	白桦、蒙古栎、紫椴、臭冷杉 Betula platyphylla, Quercus mongolica, Tilia amurensis, Abies nephrolepis
原始紫椴-红松林 Primary Tilia amurensis - Pinus koraiensis	42.23° N, 128.08° E	784	紫椴、红松、臭冷杉、大青杨 Tilia amurensis, Pinus koraiensis, Tilia amurensis, Populus ussuriensis

附录II 东北地区主要树种地上生物量异速生长方程

Supplement II Allometric equations of aboveground biomass of main tree species in northeast China

编号 No.	物种 Species	异速生长方程 Allometric equation
1	红松 Pinus koraiensis	Biomass = exp (-3.394 + 2.582 × ln (DBH))
2	臭冷杉 Abies nephrolepis	Biomass = exp (-2.989 + 2.613 × ln (DBH))
3	蒙古栎 Quercus mongolica	Biomass = exp (-2.797 + 2.571 × ln (DBH))
4	胡桃楸 Juglans mandshurica	Biomass = exp (-2.466 + 2.381 × ln (DBH))
5	五角槭 Acer pictum subsp. mono	Biomass = exp (-2.164 + 2.336 × ln (DBH))
6	东北槭 Acer mandshuricum	Biomass = exp (-2.111 + 2.310 × ln (DBH))
7	白桦 Betula platyphylla	Biomass = exp (-1.941 + 2.286 × ln (DBH))
8	水曲柳 Fraxinus mandschurica	Biomass = exp (-2.301 + 2.443 × ln (DBH))
9	山杨 Populus davidiana	Biomass = exp (-2.507 + 2.358 × ln (DBH))
10	紫椴 Tilia amurensis	Biomass = exp (-2.364 + 2.323 × ln (DBH))
11	水榆花楸 Sorbus alnifolia	Biomass = exp (-2.001 + 2.198 × ln (DBH))
12	春榆 Ulmus japonica	Biomass = exp (-2.058 + 2.271 × ln (DBH))

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