Abiotic and biotic modulators of litterfall production and its temporal stability during the succession of broad-leaf and Korean pine mixed forest

doi:10.17521/cjpe.2020.0372

Abstract

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

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[J]. Chin J Plant Ecol, 2021, 45(6): 594-605.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2020.0372

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

Figures/Tables 5

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

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.

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

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.

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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