大兴安岭北部次生林演替过程中物种多样性的变化及其影响因子

doi:10.17521/cjpe.2023.0254

摘要/Abstract

摘要：

物种多样性是生物多样性的重要组成部分。森林演替过程中物种多样性的变化会直接影响生态系统功能和稳定性。因此, 揭示森林演替过程中物种多样性变化及其与影响因子之间的关系, 对于准确预测森林生态过程和生物多样性格局具有重要意义。该研究选择大兴安岭北部白桦(Betula platyphylla)林(演替前期)、白桦-兴安落叶松(Larix gmelinii)混交林(演替中期)和兴安落叶松林(演替后期)为研究对象, 采用空间代替时间的研究方法, 分析我国寒温带地区森林演替过程中物种多样性的变化规律及其与影响因子之间的关系。结果表明: (1)大兴安岭北部森林演替过程中物种丰富度和多样性呈显著增加趋势, 演替初期Margalef丰富度指数和Shannon-Wiener多样性指数分别为2.42和2.69, 演替末期则分别为5.90和3.43; 而物种Pielou均匀度指数无显著差异。(2)随森林演替的进行, 植物群落的相似性逐渐降低, 差异性逐渐增强, Jaccard指数、Sorenson指数和Bray-Curtis指数均减小。(3)森林演替过程中土壤pH、有机质含量、全氮含量和全磷含量对物种多样性存在显著影响, 其中全氮含量和全磷含量是演替前期和中期物种多样性的主要影响因子, 土壤pH和有机质含量是影响演替后期物种多样性的主要因子。(4)随着演替的进行, 林分空间结构对物种多样性的影响逐渐增强, 角尺度和大小比数是影响物种多样性的主要空间结构因子。以上结果表明大兴安岭北部森林演替过程中土壤因子对物种多样性具有主导作用, 而林分空间结构对森林演替过程中物种多样性的驱动作用也不容忽视。

关键词: 大兴安岭, 次生演替, 物种多样性, 土壤性质, 林分结构

Abstract:

Aims Species diversity is a crucial component of biodiversity, directly affect ecosystem function and stability during forest succession. Therefore, it is of significant importance to investigate the changes in species diversity and their influencing factors during forest succession for accurately predicting forest ecological processes and biodiversity patterns.

Methods The study focused on three successional stages in the northern Da Hinggan Mountains: Betula platyphylla forest in the early stage, Betula platyphylla - Larix gmelinii mixed forest in the middle stage and Larix gmeliniiforest in the late stage. The study employed a space-for-time substitution to analyze the changes in species diversity during forest succession and explore the relationship between species diversity and environmental factors in cold temperate zone of China.

Important findings The results demonstrated that: (1) Species richness and diversity significantly increased during forest succession. The Margalef richness index and Shannon-Wiener index were 2.42 and 2.69 in the early stage of succession, and 5.90 and 3.43 in the late stage of succession. However, Pielou evenness index showed no significant difference. (2) As succession proceeded, the dissimilarity between plant communities gradually increased, indicated by the increased Jaccard, Sorenson and Bray-Curtis indices. (3) Soil pH, soil organic matter content, total nitrogen content, and total phosphorus content significantly influence species diversity during succession. Specifically, total nitrogen content and total phosphorus content were the primary factors affecting species diversity in the early and middle stages of succession, while soil pH and soil organic matter content were the dominate factors influencing species diversity in the late stage. (4) The influence of stand spatial structure on species diversity also increased over succession, where angular scale and size ratio emerge as the main spatial structure factors. These results underscore the dominant role of soil factors in shaping species diversity during forest succession in the northern Da Hinggan Mountains, while the influence of stand spatial structure on species diversity in the process of forest succession should not be overlooked.

Key words: Da Hinggan Mountains, secondary succession, species diversity, soil properties, stand structure

牛一迪, 蔡体久. 大兴安岭北部次生林演替过程中物种多样性的变化及其影响因子. 植物生态学报, 2024, 48(3): 349-363. DOI: 10.17521/cjpe.2023.0254

NIU Yi-Di, CAI Ti-Jiu. Changes in species diversity and influencing factors in secondary forest succession in northern Da Hinggan Mountains. Chinese Journal of Plant Ecology, 2024, 48(3): 349-363. DOI: 10.17521/cjpe.2023.0254

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2023.0254

https://www.plant-ecology.com/CN/Y2024/V48/I3/349

图/表 10

表1 大兴安岭3个演替阶段样地基本概况(平均值±标准差)

Table 1 Details of 3 successional stage plots in Da Hinggan Mountains (mean ± SD)

林分类型 Forest type	树种组成 Tree species	平均树高 Mean tree height (m)	平均胸径 Mean diameter at breast height (cm)	树木密度 Tree density (Ind·hm^-2)	叶面积指数 Leaf area index (m²·m^-2)	平均海拔 Mean altitude (m)
BF	白桦(100%) Betula platyphylla (100%)	12.91 ± 3.38	11.20 ± 3.17	2 214 ± 60	2.64 ± 0.17	384
BLF	白桦(60%)、兴安落叶松(40%) Betula platyphylla (60%), Larix gmelinii (40%)	12.81 ± 3.65	11.71 ± 4.52	2 878 ± 630	2.97 ± 0.19	387
LF	兴安落叶松(100%) Larix gmelinii (100%)	13.77 ± 3.96	12.08 ± 6.05	1 363 ± 97	2.50 ± 0.27	332

图1 大兴安岭北部次生林中草本层、灌木层、乔木层物种种类及重要值。BF, 白桦林; BLF, 白桦-兴安落叶松混交林; LF, 兴安落叶松林。

Fig. 1 Importance value and species of herb layer, shrub layer and tree layer in secondary forest in the northern Da Hinggan Mountains. BF, Betula platyphylla forest; BLF, Betula platyphylla - Larix gmelinii forest; LF, Larix gmelinii forest.

图2 大兴安岭北部次生林草本层、灌木层和乔木层物种多样性指数比较(平均值±标准差)。不同小写字母表示不同演替阶段差异显著(p < 0.05)。BF, 白桦林; BLF, 白桦-兴安落叶松混交林; LF, 兴安落叶松林。

Fig. 2 Comparison of species diversity index of herb layer, shrub layer and tree layer in secondary forest in the northern Da Hinggan Mountains (mean ± SD). Different lowercase letters indicate significant differences among different stages (p < 0.05). BF, Betula platyphylla forest; BLF, Betula platyphylla - Larix gmelinii forest; LF, Larix gmelinii forest.

表2 大兴安岭北部次生林不同演替阶段α多样性指数(平均值±标准差)

Table 2 α diversity index of secondary forest in the northern Da Hinggan Mountains at different successional stages (mean ± SD)

演替阶段 Stage of succession	Margalef丰富度指数 Margalef richness index	Simpson多样性指数 Simpson’s diversity index	Shannon-Wiener多样性指数 Shannon-Wiener’s diversity index	Pielou均匀度指数 Pielou evenness index
BF	2.42 ± 0.17^c	0.923 5 ± 0.040 6^c	2.689 ± 0.215 2^c	0.955 8 ± 0.089 2^a
BLF	3.62 ± 0.15^b	0.946 4 ± 0.012 8^b	3.040 ± 0.176 4^b	0.944 5 ± 0.028 3^a
LF	5.90 ± 0.45^a	0.960 1 ± 0.002 5^a	3.432 ± 0.279 9^a	0.922 7 ± 0.049 6^b

表3 大兴安岭北部次生林β多样性指数(平均值±标准差)

Table 3 β diversity index in secondary forest in the northern Da Hinggan Mountains (mean ± SD)

演替阶段 Stage of succession	Jaccard指数 Jaccard index		Sorenson指数 Sorenson index		Bray-Curtis指数 Bray-Curtis index
演替阶段 Stage of succession	BLF	LF	BLF	LF	BLF	LF
BF	0.65 ± 0.10^a	0.34 ± 0.06^c	0.74 ± 0.11^a	0.49 ± 0.07^b	0.89 ± 0.07^a	0.63 ± 0.08^b
BLF	-	0.52 ± 0.09^b	-	0.66 ± 0.08^a	-	0.66 ± 0.12^b

表4 大兴安岭北部次生林不同演替阶段土壤理化性质(平均值±标准差)

Table 4 Soil physical and chemical properties of secondary forest in the northern Da Hinggan Mountains at different successional stages (mean ± SD)

土壤因子 Soil factor	演替阶段 Succession stage
土壤因子 Soil factor	BF	BLF	LF
土壤pH Soil pH	5.54 ± 0.14^a	5.40 ± 0.17^a	5.06 ± 0.12^b
土壤含水量 Soil water content (%)	38.00 ± 11.00^b	41.00 ± 8.00^b	48.00 ± 11.00^a
土壤密度 Soil density (g·cm^-3)	1.01 ± 0.11^a	1.17 ± 0.14^a	0.83 ± 0.10^b
土壤有机质含量 Soil organic matter content (g·kg^-1)	40.14 ± 9.06^c	53.10 ± 9.00^b	64.15 ± 10.54^a
土壤全氮含量 Soil total nitrogen content (g·kg^-1)	1.39 ± 0.32^b	1.49 ± 0.40^b	2.72 ± 0.37^a
土壤全磷含量 Soil total phosphorus content (g·kg^-1)	0.66 ± 0.12^a	0.74 ± 0.12^a	0.67 ± 0.13^a
土壤全钾含量 Soil total potassium content (g·kg^-1)	1.07 ± 0.15^a	1.06 ± 0.16^a	0.97 ± 0.14^a

图3 大兴安岭北部次生林不同演替阶段林分空间结构参数(平均值±标准差)。不同小写字母表示不同取值间差异显著(p < 0.05)。BF, 白桦林; BLF, 白桦-兴安落叶松混交林; LF, 兴安落叶松林。

Fig. 3 Stand spatial structure parameters of secondary forest in the northern Da Hinggan Mountains at different successional stages (mean ± SD). Different lowercase letters indicate significant differences among different values (p < 0.05). BF, Betula platyphylla forest; BLF, Betula platyphylla - Larix gmelinii forest; LF, Larix gmelinii forest.

图4 大兴安岭北部次生林不同演替阶段土壤和林分结构对物种多样性的方差分解分析。a, 土壤因素; b, 林分空间结构因素; c, 土壤因素和林分空间结构因素二者共同解释作用。BF, 白桦林; BLF, 白桦-兴安落叶松混交林; LF, 兴安落叶松林。

Fig. 4 Variance decomposition analysis of soil and stand structure to species diversity of secondary forest in the northern Da Hinggan Mountains at different successional stages. a, soil factors; b, spatial structure factors of stand; c, soil factors and stand spatial structure factors explain the role together. BF, Betula platyphylla forest; BLF, Betula platyphylla - Larix gmelinii forest; LF, Larix gmelinii forest.

图5 大兴安岭北部次生林不同演替阶段影响因子与物种多样性指数的冗余分析(RDA)排序。A, 演替过程。B, 白桦林。C, 白桦-兴安落叶松混交林。D, 兴安落叶松林。D, Simpson多样性指数; E, Pielou均匀度指数; H', Shannon-Wiener多样性指数; M, 混交度; Mar, Margalef丰富度指数; pH, 土壤pH; SD, 土壤密度; SOM, 土壤有机质含量; SWC, 土壤含水量; TK, 土壤全钾含量; TN, 土壤全氮含量; TP, 土壤全磷含量; U, 大小比数; W, 角尺度。

Fig. 5 Redundancy analysis (RDA) ordinations between species diversity index and influencing factors of secondary forest in the northern Da Hinggan Mountains at different successional stages. A, Whole succession process. B, Betula platyphylla forest. C, Betula platyphylla - Larix gmelinii forest. D, Larix gmelinii forest. D, Simpson’s diversity index; E, Pielou evenness index; H', Shannon-Wiener diversity index; M, mingling; Mar, Margalef richness index; pH, soil pH; SD, soil density; SOM, soil organic matter content; SWC, soil water content; TK, soil total potassium content; TN, soil total nitrogen content; TP, soil total phosphorus content; U, dominance; W, uniform angle.

表5 大兴安岭北部次生林不同演替阶段物种多样性的因子贡献度及显著性

Table 5 Contribution and significance of factors to species diversity of secondary forest in the northern Da Hinggan Mountains at different successional stages

演替阶段 Stage of succession	参数 Index	贡献率 Contribution (%)	F	p
演替过程 Succession process	TN	33.2	16.2	0.002
	W	15.1	12.4	0.004
	U	12.6	13.9	0.003
	pH	10.4	4.8	0.049
	SOM	8.9	5.5	0.042
	TP	6.7	8.1	0.014
BF	TN	32.0	4.7	0.042
	TP	16.1	3.9	0.046
	U	14.2	2.3	0.052
BLF	TN	19.8	3.1	0.032
	TP	17.9	2.9	0.042
	M	17.8	3.8	0.056
LF	W	36.0	7.8	0.014
	SOM	16.5	27.5	0.002
	pH	14.8	20.0	0.008

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