Plant Diversity and Influencing Factors of Poplar-Birch Secondary Forests to Restoration Models

doi:10.16258/j.cnki.1674-5906.2026.04.001

Ecology and Environmental Sciences ›› 2026, Vol. 35 ›› Issue (4): 499-508.DOI: 10.16258/j.cnki.1674-5906.2026.04.001

• Research Article [Ecology] • Next Articles

Plant Diversity and Influencing Factors of Poplar-Birch Secondary Forests to Restoration Models

LI Yunhong¹^,²(), CHEN Yao¹^,², LIU Yulong¹^,³, ZHANG Hedong¹^,³, WANG Shuai¹^,², LIU Yankun¹^,³^,^*()

¹ Heilongjiang Institute of Ecology (Heilongjiang Wetland Research Center/Heilongjiang Grassland Research Center), Harbin 150081, P. R. China
² Mudanjiang Forest Ecosystem Research Station, Mudanjiang 157500, P. R. China
³ Key Laboratory of Forest Ecology and Forestry Ecological Engineering of Heilongjiang Province, Harbin 150081, P. R. China

Received:2025-08-25 Revised:2025-12-28 Accepted:2026-01-21 Online:2026-04-18 Published:2026-04-14

不同修复模式杨桦次生林植物多样性及其影响因子

李云红¹^,²(), 陈瑶¹^,², 刘玉龙¹^,³, 张鹤东¹^,³, 王帅¹^,², 刘延坤¹^,³^,^*()

¹ 黑龙江省生态研究所（黑龙江省湿地研究中心/黑龙江省草原研究中心），黑龙江哈尔滨 150081
² 黑龙江牡丹江森林生态系统国家定位观测研究站，黑龙江牡丹江 157500
³ 黑龙江省森林生态与林业生态工程重点实验室，黑龙江哈尔滨 150081

通讯作者: *E-mail: liuyankun1979@126.com
作者简介:李云红（1979年生），女，副研究员，硕士，研究方向为森林生态。E-mail: 704415@qq.com
基金资助:
黑龙江省自然科学基金联合引导项目(LH2023C107);黑龙江省省属科研院所科研业务费项目(CZKYF2025-1-B040);黑龙江省重点研发计划项目(GA21C030)

Abstract

Abstract:

Forest ecosystems play an irreplaceable ecological role in maintaining global biodiversity, regulating climate, and other aspects. Due to long-term disturbance, the Northeast Forest area is facing severe challenges from declining forest ecosystem function. Poplar-birch secondary forests are early or mid-stage forest types formed by natural succession on secondary bare land after natural or human disturbance and destruction of primary forests such as broad-leaved Pinus koraiensis natural forests and spruce forests. It is one of the most common secondary forest types in temperate and cold temperate forest areas. If subjected to high-intensity human or natural disturbance, it will form degraded forests, thereby affecting the positive succession process of forests. Therefore, it is particularly important to carry out active ecological restoration, such as replanting, nurturing, soil improvement, etc., to rebuild community structure and function and reactivate natural succession processes. This study aims to investigate the response of Poplar-birch secondary forest species diversity to different management and restoration modes by integrating two key dimensions: biodiversity and soil physicochemical characteristics to explore the following questions: 1) are there differences in species diversity among different restoration modes? 2) How do soil physicochemical characteristics respond to different restoration modes? 3) What is the relationship between biotic and abiotic factors in various restoration modes, specifically regarding species diversity and soil physicochemical characteristics? In response to the above three questions, we focused on three historical management and restoration models (with a management duration of about 15 years) of Poplar-birch secondary forests, namely tree species structure adjustment (I: Nurturing+Replanting of Pinus koraiensis, II: Nurturing+Replanting of Picea koraiensis), target-tree cultivation and management (III: nurturing management), and using the zonal top community broad-leaved Pinus koraiensis natural forest as a control (IV). The response of plant diversity and soil physicochemical characteristics to different management and restoration modes is analyzed. SPSS 21.0 software is used to perform one-way ANOVA on the species diversity index and soil physicochemical characteristics of different management and restoration modes. The R language Vegan package is used to perform redundancy analysis (RDA) on the species diversity index and soil physicochemical characteristics, and Monte Carlo permutation test is used to determine the main soil physicochemical factors affecting the species diversity index. The interaction mechanism between biotic and abiotic factors in the forest ecosystem restoration process is explored, to provide scientific basis for the restoration of degraded forest ecosystems in Northeast China. The results showed that: 1) the dominance index (D), Shannon-Wiener index (H), and evenness index (J) of the tree layer were significantly higher in the target tree management mode (III) compared to the tree structure adjustment mode I (p<0.05), while the number of shrub layer species (S) was significantly greater in the control treatment than in modes III and I. There was no significant difference in D and H among the four modes (p>0.05), and there was no significant difference in the S, D, H, and J index of the herb layer among different management and restoration modes (p>0.05); 2) The soil bulk density (SBD) of the three management and restoration modes was higher than the control, and the soil pH value and maximum water holding capacity (WHC) were lower than the control. However, there was no significant difference among the three modes (p>0.05). The total soil porosity (TPOR) tree species structure adjustment mode II was slightly higher than the control, but there was no significant difference (p>0.05). 3) There were significant differences in soil organic carbon (SOC), total nitrogen (TN), total phosphorus (TP), carbon nitrogen ratio (C/N), carbon phosphorus ratio (C/P), and nitrogen phosphorus ratio (N/P) among the three management and restoration models and between them and the control group. Among them, SOC, TP, and C/N ratio were significantly higher in model I and model II than in model III, and TN and N/P were significantly higher in model III than in model I; 4) The changes in soil TN, N/P, WHC, and TPOR in the tree layer are mainly reflected in the RDA1 axis, while S, H, D, and J all increase with the increase of soil TN, N/P, WHC, and TPOR. The changes in soil C/P are mainly reflected in the RDA2 axis, while D, J, S, and H all increase with the increase of soil C/P. The soil factors that significantly affect species diversity in the tree layer are mainly TN, C/N, and TPOR. The changes in soil SOC and WHC in the shrub layer are mainly reflected in the RDA1 axis. H, D, and S decrease with the decrease of soil SOC and WHC on the RDA1 axis, while J increases. The changes in N/P are mainly reflected in the RDA2 axis. On the RDA2 axis, S decreases as soil N/P decreases, while H, D, and J increase. The soil factors that significantly affect shrub layer species diversity are TN, SBD, and WHC, and there is no significant relationship between soil factors and herbaceous layer species diversity. The conclusion of this study is that: 1) the management and restoration mode of Poplar-birch secondary forests has differentiated effects, and the target tree management mode significantly improves the species diversity of the tree layer (with higher D, H, and J indices), and improves the nitrogen status of the soil. The tree species structure adjustment mode promotes uniform growth of the shrub layer (with higher J index of shrub layer uniformity), and increases the organic matter and phosphorus content of the soil, while also having a higher soil C/N ratio; 2) The core soil factors that affect species diversity at different levels are TN, C/N, and TPOR, which significantly affect species diversity in the tree layer under different management and restoration modes. TN, SBD, and WHC are soil factors that significantly affect species diversity in the shrub layer under different management and restoration modes. The scientific and technological problem that needs to be solved in the future: there is a clear “trade-off” between the target tree management model and the tree species structure adjustment model in terms of ecological benefits. How to quantify the comprehensive ecological benefits of these two models at the ecosystem scale? Is there an optimal collaborative solution that can simultaneously maximize the diversity of tree and shrub layers and the theoretical threshold of soil multifunctionality? How can we technically integrate the advantages of the two modes in practice? In addition, this study only focuses on middle-aged forests, and further research is needed to determine whether the restoration effect is consistent in other forest age stages.

Key words: polar-birch secondary forests, restoration models, species diversity, soil physical and chemical characteristics, redundancy analysis

摘要：

由于长期受到干扰，东北林区面临森林生态系统功能下降的严峻挑战，生态修复势在必行，而目前对于次生林不同修复模式下生物与非生物因子协同驱动生态恢复的机制仍缺乏深入认识。以3种修复模式的杨桦（Populus spp. & Betula spp.）次生林以及地带性顶级群落阔叶红松（Pinus koraiensis Sieb. et Zucc.）林为研究对象，分析植物多样性和土壤理化特征对不同经营修复模式的响应，采用冗余分析的方法，探讨森林生态系统恢复过程中生物-非生物因子相互作用机制，为东北林区的退化森林生态系统修复提供科学依据。结果表明：1）乔木层Simpson index（D）、Shannon-Wiener index（H）、Pielou index（J）均呈现目标树经营模式（Ⅲ）显著高于树种结构调整模式Ⅰ，灌木层Number of species（S）以对照处理显著高于模式Ⅲ和模式Ⅰ；2）3种修复模式土壤容重SBD均高于对照，土壤pH值、土壤最大持水率WHC均低于对照，土壤总孔隙度TPOR树种结构调整模式Ⅱ略高于对照，但无显著差异。3）3种经营修复模式之间及其与对照间土壤有机碳SOC、全氮TN、全磷TP、碳氮比C/N、碳磷比C/P以及氮磷比N/P均存在一定的显著差异；4）显著影响乔木层物种多样性的土壤因子主要为TN、C/N和TPOR，显著影响灌木层物种多样性的土壤因子为TN、SBD和WHC。在杨桦次生林经营修复模式选择中，需优先考虑能有效改良土壤理化性质，即通气透水性及养分状况的技术措施，以促进林分物种多样性的提升。

关键词: 杨桦次生林, 修复模式, 物种多样性, 土壤理化特征, 冗余分析

CLC Number:

LI Yunhong, CHEN Yao, LIU Yulong, ZHANG Hedong, WANG Shuai, LIU Yankun. Plant Diversity and Influencing Factors of Poplar-Birch Secondary Forests to Restoration Models[J]. Ecology and Environmental Sciences, 2026, 35(4): 499-508.

李云红, 陈瑶, 刘玉龙, 张鹤东, 王帅, 刘延坤. 不同修复模式杨桦次生林植物多样性及其影响因子[J]. 生态环境学报, 2026, 35(4): 499-508.

Figures/Tables 6

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修复模式	海拔/m	坡度/°	坡位	坡向	林龄组	郁闭度	平均胸径/cm	经营方式
Ⅰ	706.70±5.65	3.00±0.00	中	南	中龄林	0.8	13.29±0.24	抚育+林冠下补植红松
Ⅱ	664.70±9.62	3.00±0.00	中	南	中龄林	0.8	17.77±1.82	抚育+林冠下补植云杉
Ⅲ	712.17±5.12	3.00±0.00	中	南	中龄林	0.8	13.48±1.33	目标树抚育经营
Ⅳ	584.87±24.42	11.00±3.61	中	南	中龄林	0.8	17.21±1.18	对照

修复模式	海拔/m	坡度/°	坡位	坡向	林龄组	郁闭度	平均胸径/cm	经营方式
Ⅰ	706.70±5.65	3.00±0.00	中	南	中龄林	0.8	13.29±0.24	抚育+林冠下补植红松
Ⅱ	664.70±9.62	3.00±0.00	中	南	中龄林	0.8	17.77±1.82	抚育+林冠下补植云杉
Ⅲ	712.17±5.12	3.00±0.00	中	南	中龄林	0.8	13.48±1.33	目标树抚育经营
Ⅳ	584.87±24.42	11.00±3.61	中	南	中龄林	0.8	17.21±1.18	对照

植被层	多样性指数	Ⅰ	Ⅱ	Ⅲ	Ⅳ
乔木	S	8.00±0.00a	8.00±1.15a	9.33±1.20a	8.33±0.67a
	D	0.66±0.03b	0.78±0.04a	0.82±0.02a	0.78±0.02a
	H	1.46±0.06b	1.74±0.17ab	1.91±0.13a	1.74±0.17ab
	J	0.70±0.03b	0.84±0.02a	0.86±0.01a	0.83±0.01a
灌木	S	1.67±0.67b	2.00±0.58ab	1.67±0.33b	3.67±0.33a
	D	0.49±0.07a	0.45±0.06a	0.39±0.07a	0.54±0.07a
	H	0.78±0.17a	0.78±0.14a	0.65±0.12a	1.05±0.14a
	J	0.86±0.03a	0.74±0.04ab	0.67±0.04b	0.68±0.07b
草本	S	12.00±0.00a	12.00±0.58a	10.67±0.67a	13.00±2.65a
	D	0.74±0.01a	0.79±0.04a	0.79±0.04a	0.75±0.09a
	H	1.70±0.05a	1.91±0.16a	1.89±0.09a	1.82±0.34a
	J	0.66±0.02a	0.74±0.05a	0.77±0.05a	0.69±0.09a

植被层	多样性指数	Ⅰ	Ⅱ	Ⅲ	Ⅳ
乔木	S	8.00±0.00a	8.00±1.15a	9.33±1.20a	8.33±0.67a
	D	0.66±0.03b	0.78±0.04a	0.82±0.02a	0.78±0.02a
	H	1.46±0.06b	1.74±0.17ab	1.91±0.13a	1.74±0.17ab
	J	0.70±0.03b	0.84±0.02a	0.86±0.01a	0.83±0.01a
灌木	S	1.67±0.67b	2.00±0.58ab	1.67±0.33b	3.67±0.33a
	D	0.49±0.07a	0.45±0.06a	0.39±0.07a	0.54±0.07a
	H	0.78±0.17a	0.78±0.14a	0.65±0.12a	1.05±0.14a
	J	0.86±0.03a	0.74±0.04ab	0.67±0.04b	0.68±0.07b
草本	S	12.00±0.00a	12.00±0.58a	10.67±0.67a	13.00±2.65a
	D	0.74±0.01a	0.79±0.04a	0.79±0.04a	0.75±0.09a
	H	1.70±0.05a	1.91±0.16a	1.89±0.09a	1.82±0.34a
	J	0.66±0.02a	0.74±0.05a	0.77±0.05a	0.69±0.09a

环境因子	乔木			灌木			草本
环境因子	RDA1	RDA2	p	RDA1	RDA2	p	RDA1	RDA2	p
SOC	−0.999	0.039	0.604	−0.984	0.178	0.493	−0.374	0.927	0.129
TN	0.748	−0.664	0.008	−0.326	−0.946	0.027	0.503	0.864	0.793
C/N	−0.853	0.521	0.006	−0.058	0.998	0.086	−0.629	0.777	0.511
TP	−0.654	−0.757	0.235	−0.819	0.573	0.357	−0.867	0.498	0.105
C/P	0.311	0.951	0.166	0.250	−0.968	0.760	0.991	0.131	0.148
N/P	1.000	0.024	0.069	0.188	−0.982	0.127	0.897	−0.442	0.239
pH	−0.413	0.911	0.821	−0.816	0.579	0.155	−0.808	0.589	0.954
SBD	−0.949	−0.315	0.419	0.971	0.238	0.001	0.876	−0.483	0.935
WHC	0.994	−0.110	0.242	−0.860	−0.511	0.001	−0.108	0.994	0.919
TPOR	0.900	−0.435	0.050	−0.617	−0.787	0.087	0.956	0.294	0.540
特征值	3.268	0.691		2.496	1.164		2.461	0.474
累计解释变异	82.230	99.630		67.830	99.460		83.350	99.380

Plant Diversity and Influencing Factors of Poplar-Birch Secondary Forests to Restoration Models

不同修复模式杨桦次生林植物多样性及其影响因子

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