阔叶红松林不同演替阶段凋落物产量及其稳定性的影响因素

doi:10.17521/cjpe.2020.0372

摘要/Abstract

摘要：

凋落物产量是生态系统净初级生产力的重要组分, 也是连接地下与地上生态过程的关键环节。但在森林演替进程中, 生物多样性、林分因子、功能性状如何共同影响凋落物产量及其时间稳定性, 其机理有待进一步研究。该文在阔叶红松(Pinus koraiensis)林分布的北界黑龙江胜山保护区, 选择从演替早期到红松原始林的4个演替(或发育)阶段, 连续3年测定凋落物产量, 以3年凋落物产量的变异系数的倒数(1/CV)反映其稳定性。采用层次划分和变异分离等方法研究林分因子(最大树高、胸径、胸高断面积、林冠空隙度)、群落水平功能性状(叶碳、氮含量及比叶面积)和乔木多样性(物种、功能、谱系多样性)对凋落物产量及其稳定性的相对作用大小。结果表明, 演替早期凋落物产量显著低于后3个演替阶段, 从演替中期至演替晚期凋落物产量无显著差异。凋落物产量的稳定性随演替进展显著提高。对变量重要性的评价表明, 凋落物产量主要受林分因子(最大树高、胸高断面积、林冠空隙度)和性状(叶碳含量)的影响, 物种丰富度也起一定作用; 而功能多样性对于凋落物稳定性的作用最大, 其次是林分因子(如最大胸径)。生物多样性对凋落物产量的独立解释力仅为0.41%, 而对其稳定性的独立解释力为33.12%, 说明多样性对凋落物稳定性有着独立于林分因子、性状之外的重要作用。同时, 林分因子、生物多样性、性状之间存在较强的协同作用(最高达53.8%), 说明这3种因素共同作用于凋落物产量及其稳定性。研究结果表明, 森林的恢复演替不仅能提高森林生产力, 还可有效提高生态系统稳定性, 因此, 保护原始林及促进森林恢复演替是提高生态系统功能的有效手段。

关键词: 凋落物产量, 稳定性, 生物多样性, 林分因子, 演替, 功能性状, 阔叶红松林

Abstract:

Aims Litterfall is an important component of ecosystem net primary productivity, and a key link between above- and below-ground carbon processes. However, it remains unclear how biodiversity, stand factors and functional traits work together in affecting litterfall production and its temporal stability during forest succession.

Methods We measured litterfall production for three years in plots from four successional stages of broad-leaf and Korean pine (Pinus koraiensis) mixed forests, at the northern limit of Korean pine forest in the Shengshan Reserve of Heilongjiang. Functional traits (leaf carbon and nitrogen contents, specific leaf area) were measured to quantify functional diversity and community weighted mean (CWM) of traits. Tree diameter and height, total basal area (TBA) and gap fraction were measured to quantify stand structure. We used hierarchy partitioning analysis and variance partitioning to evaluate the relative effects of stand factors, community level traits, and (species, functional and phylogenetic) diversity on annual litterfall production, and its temporal stability (= 1/coefficient of variation for annual litterfall production).

Important findings Litterfall production was significantly lower in the early successional stage, but did no vary from the middle to late successional stages. The litterfall stability increased continuously with forest succession. The variable importance of multivariate models also suggest that, the litterfall production was mainly affected by stand factors (e.g. height, TBA, gap fraction) and functional traits (leaf carbon content), with species richness also playing a role. For stability of litterfall production, however, functional diversity was the strongest predictor, followed by stand factors (such as maximum tree diameter). The independent effect of biodiversity on litterfall production was only 0.41%, but was as much higher (33.12%) for temporal stability of litterfall, suggesting that biodiversity have an important influence on litterfall stability that is independent of stand factors and traits. There was also a strong joint effect (up to 53.8%) among the stand factors, biodiversity and functional traits, indicating that these factors collectively affect litterfall production and its stability. Our results suggest that forest succession not only increases forest productivity but also improves ecosystem stability. Therefore, protecting primary forests and promoting forest restoration are effective ways to improve ecosystem functions.

Key words: litterfall production, stability, biodiversity, stand factors, succession, functional traits, broad-leaf and Korean pine mixed forest

孙浩哲, 王襄平, 张树斌, 吴鹏, 杨蕾. 阔叶红松林不同演替阶段凋落物产量及其稳定性的影响因素. 植物生态学报, 2021, 45(6): 594-605. DOI: 10.17521/cjpe.2020.0372

SUN Hao-Zhe, WANG Xiang-Ping, ZHANG Shu-Bin, WU Peng, YANG Lei. Abiotic and biotic modulators of litterfall production and its temporal stability during the succession of broad-leaf and Korean pine mixed forest. Chinese Journal of Plant Ecology, 2021, 45(6): 594-605. DOI: 10.17521/cjpe.2020.0372

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2020.0372

https://www.plant-ecology.com/CN/Y2021/V45/I6/594

图/表 5

表1 黑龙江胜山保护区阔叶红松林各演替阶段样地的林分特征

Table 1 Stand characteristics for plots of different succession stages of broadleaf Korean pine forest in the Shengshan Nature Reserve, Heilongjiang Province of China

演替阶段 Succession stage	平均树高 Mean tree height (m)	平均胸径 Mean DBH (cm)	最大树高 H_max (m)	最大胸径 DBH_max (cm)	林分密度 Stem density (tree·hm^-2)	林冠空隙度 Gap fraction (%)	胸高断面积 TBA (m²·hm^-2)
早期 Early stage	13.3	13.2	22.6	29.8	853	32.1	14.8
中期 Middle stage	8.9	11.1	24.3	43.8	1 577	13.1	30.9
中晚期 Mid-late stage	12.0	11.9	28.9	51.8	1 723	15.9	21.4
晚期 Late stage	13.1	16.6	28.6	67.7	1 180	6.6	42.4

图1 阔叶红松林不同演替阶段凋落物产量及其稳定性的差异。不同小写字母表示在p < 0.05水平上差异显著。

Fig. 1 Difference in annual litterfall production and its temporal stability among successional stages of Korean pine forest. Different lowercase letters indicate significant differences at p < 0.05 level.

表2 阔叶红松林不同演替阶段凋落物产量及其稳定性与各解释变量的决定系数(R2)

Table 2 Coefficients of determination (R2) for different factors in explaining annual litterfall production and its temporal stability across succesional stages of Korean pine forest

变量 Variable		凋落物产量 Annual litterfall production	凋落物产量稳定性 Stability of litterfall production
林分因子 Stand factor	最大树高 H_max	0.57^**	0.39^*
	最大胸径 DBH_max	0.45^*	0.55^**
	胸高断面积 TBA	0.43^*	0.57^**
	林冠空隙度 Gap fraction	-0.68^***	-0.53^**
功能性状 Functional trait	叶片碳含量 CWM_LC	-0.26	0.00
	叶片氮含量 CWM_LN	0.05	0.05
	比叶面积 CWM_SLA	-0.11	-0.54^**
生物多样性 Biodiversity	物种丰富度 Richness	0.65^**	0.42^*
	物种均匀度 Evenness	0.59^**	0.28
	功能多样性 Rao’s Q	0.45^**	0.16
	系统发育多样性 PD	0.32	0.24

图2 最终多元模型中保留的各变量对于凋落物产量(A)和凋落物产量稳定性(B)的相对重要性的层次划分分析。CWMLC, 叶片碳含量; CWMLN, 叶片氮含量; CWMSLA, 比叶面积; DBHmax, 最大胸径; Hmax, 最大树高; PD, 系统发育多样性; Rao’s Q, Rao二次熵; Richness, 物种丰富度; TBA, 胸高断面积。

Fig. 2 Relative importance of variables retained in the models explaining litterfall production (A) and temporal stability of litterfall (B), as obtained by the hierarchical partitioning analyses. CWMLC and CWMLN, community weighted mean of leaf carbon and nitrogen content, respectively; CWMSLA, community weighted mean of specific leaf area; DBHmax, maximum diameter at breast height; Hmax, maximum tree height; PD, phylogenetic diversity; Rao’s Q, Rao’s quadratic entropy; TBA, total basal area.

图3 最终多元模型中保留的林分因子、功能性状和生物多样性变量对凋落物产量(A)和凋落物产量稳定性(B)影响的变异分离图。a, b, c, 3种因素各自的独立作用; d, e, f, 两因素之间的协同作用; g, 3种因素之间的协同作用。

Fig. 3 Variance partitioning for the effects of stand factors, functional traits and biodiversity factors retained in the models on litterfall production (A) and temporal stability of litterfall (B). a, b, c, the independent effects by each of the three factors; d, e, f, the joint effects between two factors; g, the joint effect among three factors.

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