东北阔叶红松林不同演替阶段树种分布与空间关联格局

doi:10.17521/cjpe.2025.0365

植物生态学报 ›› 2026, Vol. 50 ›› Issue (4): 833-845.DOI: 10.17521/cjpe.2025.0365 cstr: 32100.14.cjpe.2025.0365

东北阔叶红松林不同演替阶段树种分布与空间关联格局

王子平¹^,²^,³, 毛子昆¹^,², 贺涵¹^,²^,³, 姜鹏程¹^,²^,³, 闫瑞环¹^,²^,³, 王绪高¹^,²^,^*()

¹ 中国科学院沈阳应用生态研究所森林生态与保育重点实验室, 沈阳 110016
² 辽宁省陆地生态系统碳中和重点实验室, 沈阳 110016
³ 中国科学院大学, 北京 100049

收稿日期:2025-10-11 接受日期:2026-01-21 出版日期:2026-04-20 发布日期:2026-07-15
通讯作者: ^*王绪高(wangxg@iae.ac.cn)
基金资助:
国家重点研发计划(2022YFF1300501);国家自然科学基金(32525041);国家自然科学基金(32301344);辽宁省“兴辽英才计划”项目(XLYC2402003)

Spatial distribution and association patterns of tree species across successional stages in broadleaf Korean pine forests of Northeast China

WANG Zi-Ping¹^,²^,³, MAO Zi-Kun¹^,², HE Han¹^,²^,³, JIANG Peng-Cheng¹^,²^,³, YAN Rui-Huan¹^,²^,³, WANG Xu-Gao¹^,²^,^*()

¹ Key Laboratory of Forest Ecology and Silviculture, Institute of Applied Ecology, Chinese Academy of Sciences, Shenyang 110016, China
² Key Laboratory of Terrestrial Ecosystem Carbon Neutrality, Liaoning Province, Shenyang 110016, China
³ University of Chinese Academy of Sciences, Beijing 100049, China

Received:2025-10-11 Accepted:2026-01-21 Online:2026-04-20 Published:2026-07-15
Contact: ^*WANG Xu-Gao(wangxg@iae.ac.cn)
Supported by:
National Key R&D Program of China(2022YFF1300501);National Natural Science Foundation of China(32525041);National Natural Science Foundation of China(32301344);Liaoning Revitalization Talents Program(XLYC2402003)

摘要/Abstract

摘要： 阔叶红松(Pinus koraiensis)林是东北东部地区地带性顶极植物类型, 长期以来受到人类活动的强烈干扰, 大面积原生阔叶红松林退化为次生杨桦林, 但目前相关的系统对比研究仍然不足, 限制了对阔叶红松林自然恢复过程的深入理解。该研究基于长白山24 hm²固定森林监测样地, 利用成对相关函数g(r)和异质性泊松分布零模型, 对比分析了样地北部次生杨桦林(演替前期, 15.6 hm²)与南部阔叶红松林(演替后期, 8.4 hm²)的树种分布和空间关联格局。结果发现: (1)两种林型中大多数树种的空间分布具有尺度依赖性, 即在小尺度(0-10 m)上呈聚集分布, 在较大尺度(>20 m)趋于随机分布, 胸径(DBH) < 10 cm的小径级个体和中径级个体(10 cm ≤ DBH < 30 cm)的聚集程度均高于大径级个体(DBH ≥ 30 cm); 次生杨桦林的大径级个体普遍呈随机分布, 而阔叶红松林的大径级个体在0-10 m仍以聚集分布为主。(2)两种林型中树种空间关联以不相关为主; 大径级个体的空间负关联比例均高于小径级个体, 大径级个体对中径级个体存在明显抑制作用, 而其对小径级个体的作用具有较大变异性。(3)作为核心建群种的红松, 在两种林型中均呈现出聚集分布, 红松大树在阔叶红松林的聚集程度较低, 且与白桦(Betula platyphylla)、山杨(Populus davidiana)等先锋树种的种间关系表现为负关联, 而红松小树与白桦、山杨等显著正关联, 表明这些先锋树种对红松小树具有促进作用, 但是随着森林演替先锋树种逐渐受到抑制并退出群落。综上, 阔叶红松林和次生杨桦林的树种分布与空间关联格局大体一致, 但关键建群种存在明显差异。

关键词: 空间分布, 空间关联, 点格局分析, 阔叶红松林, 次生杨桦林

Abstract:

Aims The broadleaf Korean pine (Pinus koraiensis) mixed forest, represents the zonal climax vegetation in eastern Northeast China. Long-term human disturbances have led to widespread degradation of primary forests into secondary poplar-birch forests. However, systematic comparative studies between these two forest types remain insufficient, limiting our understanding of the natural recovery process of broadleaf Korean pine mixed forests.

Methods Based on a 24 hm² permanent forest dynamics plot in Changbai Mountains, we applied the paired correlation function g(r) combined with the heterogeneous Poisson null model to compare the spatial distribution and interspecific association patterns of tree species between the northern secondary poplar-birch forest (early successional stage, 15.6 hm²) and the southern broadleaf Korean pine mixed forest (late successional stage, 8.4 hm²).

Important findings (1) Most tree species in both forest types exhibited scale-dependent spatial patterns, characterized by aggregation at small scales (0-10 m) and random distribution at larger scales (>20 m). Small individuals with a diameter at breast height (DBH) below 10 cm and medium-sized individuals (10 cm ≤ DBH <  30 cm) generally showed stronger aggregation than large individuals (DBH ≥  30 cm). Large individuals in the secondary poplar-birch forest were mostly randomly distributed, whereas those in the broadleaf Korean pine forest remained aggregated at small scales (0-10 m). (2) Interspecific spatial associations were predominantly random in both forest types. Large individuals exhibited a higher proportion of negative interspecific associations than small ones. Large individuals exerted strong suppressive effects on medium-sized individuals, while their effects on small individuals were more variable. (3) As a key foundation species, Pinus koraiensis exhibited aggregated distribution patterns in both forest types, although its aggregation intensity was lower in the broadleaf Korean pine mixed forest. Large Pinus koraiensis individuals showed negative spatial associations with pioneer species such as Betula platyphylla and Populus davidiana in the late successional forest, whereas small Pinus koraiensis individuals displayed significant positive associations with these pioneers species in the secondary poplar-birch forest. These findings indicate that pioneer species facilitate the establishment of Pinus koraiensis at early successional stages but become progressively suppressed and are eventually excluded as forest succession proceeds. Overall, the two forest types exhibited broadly similar species distribution and interspecific association patterns, whereas pronounced differences were observed for key foundation species, such as Pinus koraiensis. These results enhance our understanding of the natural recovery process of the broadleaf Korean pine mixed forest and provide valuable insights for forest management and restoration of temperate forests of Northeast China.

Key words: spatial distribution, spatial association, spatial point pattern analysis, broadleaf Korean pine forest, secondary poplar-birch forest

王子平, 毛子昆, 贺涵, 姜鹏程, 闫瑞环, 王绪高. 东北阔叶红松林不同演替阶段树种分布与空间关联格局. 植物生态学报, 2026, 50(4): 833-845. DOI: 10.17521/cjpe.2025.0365

WANG Zi-Ping, MAO Zi-Kun, HE Han, JIANG Peng-Cheng, YAN Rui-Huan, WANG Xu-Gao. Spatial distribution and association patterns of tree species across successional stages in broadleaf Korean pine forests of Northeast China. Chinese Journal of Plant Ecology, 2026, 50(4): 833-845. DOI: 10.17521/cjpe.2025.0365

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2025.0365

https://www.plant-ecology.com/CN/Y2026/V50/I4/833

图/表 10

图1 东北阔叶红松林样地内主要树种的空间分布。A, 红松。B, 白桦。C, 山杨。图中虚线为两种林型的分界线。点的大小表示树木径级: 分别为胸径(DBH) < 10 cm, 10 cm ≤ DBH < 30 cm, DBH ≥ 30 cm。

Fig. 1 Spatial distribution of major tree species within the plot in broadleaf Korean pine forests of Northeastern China. A, Pinus koraiensis. B, Betula platyphylla. C, Populus davidiana. The dashed lines in the figure represent the boundary between the two forest types. The size of the dots indicates tree diameter classes: diameter at breast height (DBH) < 10 cm, 10 cm ≤ DBH < 30 cm, and DBH ≥ 30 cm.

表1 东北阔叶红松林不同演替阶段样地小树的空间格局分析

Table 1 Spatial pattern analysis of small trees across different succession stages in broadleaf Korean pine forests of Northeastern China

物种 Species	次生杨桦林 Secondary poplar-birch forests	阔叶红松林 Broadleaf Korean pine forests
暴马丁香 Syringa reticulata	1-26 (+)	1-18 (+)
稠李 Prunus padus	1-10 (+), 17-30 (-)	1-17 (+), 21-30 (-)
簇毛槭 Acer barbinerve	1-17 (+)	1-20 (+), 22 (+)
春榆 Ulmus japonica	1-16 (+), 21 (-), 28-29 (-)	1-22 (+)
东北山梅花 Philadelphus schrenkii	1-17 (+), 21-24 (+), 28-29 (+)	1-17 (+)
怀槐 Maackia amurensis	1-13 (+), 15-16 (+)	1-8 (+), 19-23 (+)
红松 Pinus koraiensis	1-20 (+)	1-11 (+), 18-21 (+)
假色槭 Acer pseudo-sieboldianum	1-15 (+), 20-21 (-), 25-28 (-)	1-20 (+), 27-30 (-)
瘤枝卫矛 Euonymus verrucosus	1-6 (+)	1-4 (+)
蒙古栎 Quercus mongolica	1-11 (+)	1-19 (+), 22-24 (+)
毛榛 Corylus mandshurica	1-19 (+)	1-22 (+)
拧筋槭 Acer triflorum	1-6 (+)	1-7 (+)
青楷槭 Acer tegmentosum	1-5 (+)	1-5 (+)
山丁子 Malus baccata	1-11 (+), 20-27 (+)	1-13 (+), 20-21 (-)
色木槭 Populus davidiana	1-14 (+), 16-17 (+), 19-28 (+), 30 (+)	1-12 (+), 18-19 (+)
水曲柳 Fraxinus mandshurica	1-14 (+), 24 (-), 26-30 (-)	1-5 (+), 8-10 (+), 12-13 (+)
紫椴 Tilia amurensis	1-14 (+), 28 (-)	1-15 (+), 22-23 (-)
黄檗 Phellodendron amurense	随机分布 Random distribution
糠椴 Tilia mandshurica	1-21 (+)

表2 东北阔叶红松林不同演替阶段样地中树的空间格局分析

Table 2 Spatial pattern analysis of medium-sized trees across different succession stages in broadleaf Korean pine forests of Northeastern China

物种 Species	次生杨桦林 Secondary poplar-birch forests	阔叶红松林 Broadleaf Korean pine forests
白桦 Betula platyphylla	1-12 (+), 18 (+), 21 (+)	1-26 (+)
春榆 Ulmus japonica	1-2 (+), 4-8 (+), 18-19 (+)	1-11 (+), 14-15 (+), 28-30 (-)
怀槐 Maackia amurensis	3-24 (+)	7 (+), 11 (+)
蒙古栎 Quercus mongolica	1-15 (+)	1-12 (+)
山丁子 Malus baccata	1-10 (+)	1-4 (+)
色木槭 Populus davidiana	1 (+), 5-8 (+), 10-13 (+)	随机分布 Random distribution
水曲柳 Fraxinus mandshurica	1-13 (+), 24-30 (-)	1-10 (+)
山杨 Populus davidiana	1-11 (+)	1-7 (+), 17-19 (+), 26-28 (+)
紫椴 Tilia amurensis	1-13 (+)	1-13 (+), 24 (-), 29-30 (-)
黄檗 Phellodendron amurense	1-6 (+)
红松 Pinus koraiensis	1-5 (+), 20-21 (-)
糠椴 Tilia mandshurica	1-11 (+)
拧筋槭 Acer triflorum		11-14 (+)

表3 东北阔叶红松林不同演替阶段样地大树的空间格局分析

Table 3 Spatial pattern analysis of big trees across different succession stages in broadleaf Korean pine forests of Northeastern China

物种 Species	次生杨桦林 Secondary poplar-birch forests	阔叶红松林 Broadleaf Korean pine forests
白桦 Betula platyphylla	随机分布 Random distribution	1-11 (+)
蒙古栎 Quercus mongolica	1-4 (+)	3-12 (+)
水曲柳 Fraxinus mandshurica	随机分布 Random distribution	12-18 (+), 26 (-)
山杨 Populus davidiana	1-5 (+), 9-11 (+), 15-16 (-), 29 (-)	1-3 (+), 7-14 (+)
紫椴 Tilia amurensis		5-10 (+)
红松 Pinus koraiensis		1-15 (+), 17 (+)

图2 东北阔叶红松林不同空间分布类型的物种数随尺度的变化。A, 次生杨桦林小树。B, 次生杨桦林中树。C, 次生杨桦林大树。D, 阔叶红松林小树。E, 阔叶红松林中树。F, 阔叶红松林大树。

Fig. 2 Scale-dependent univariate intraspecific patterns in broadleaf Korean pine forests of Northeastern China. A, Small trees in secondary poplar-birch forests. B, Medium-sized trees in secondary poplar-birch forests. C, Large trees in secondary poplar-birch forests. D, Small trees in broadleaf Korean pine forests. E, Medium-sized trees in broadleaf Korean pine forests. F, Large trees in broadleaf Korean pine forests.

图3 次生杨桦林种间关联随尺度的变化。A, 小树空间关联。B, 中树空间关联。C, 大树空间关联。D, 中树对小树空间关联。E, 大树对小树空间关联。F, 大树对中树空间关联。

Fig. 3 Scale-dependent interspecific association patterns in secondary poplar-birch forests. A, Spatial associations among small trees. B, Spatial associations among medium-sized trees. C, Spatial associations among large trees. D, Spatial associations between medium-sized trees and small trees. E, Spatial associations between large trees and small trees. F, Spatial associations between large trees and medium-sized trees.

图4 阔叶红松林种间关联随尺度的变化。A, 小树空间关联。B, 中树空间关联。C, 大树空间关联。D, 中树对小树空间关联。E, 大树对小树空间关联。F, 大树对中树空间关联。

Fig. 4 Scale-dependent interspecific association patterns in broadleaf Korean pine forests. A, Spatial associations among small trees. B, Spatial associations among medium-sized trees. C, Spatial associations among large trees. D, Spatial associations between medium-sized trees and small trees. E, Spatial associations between large trees and small trees. F, Spatial associations between large trees and medium-sized trees.

图5 不同径级红松的空间分布格局。A, 次生杨桦林红松小树。B, 次生杨桦林红松中树。C, 阔叶红松林红松小树。D, 阔叶红松林红松大树。实线表示单变量成对相关函数g(r)的函数值, 虚线表示单变量成对相关函数g(r)的期望值, 灰色阴影部分表示95%的置信区间。

Fig. 5 Spatial distribution pattern of Pinus koraiensis at different diameter classes. A, Small Pinus koraiensis trees in secondary birch-poplar forest. B, Medium-sized Pinus koraiensis trees in secondary birch-poplar forest. C, Small Pinus koraiensis trees in broadleaf Korean pine forest. D, Large Pinus koraiensis trees in broadleaf Korean pine forest. The solid line indicates the value of the univariate pairwise correlation function g(r), the dashed line indicates the expected value of the univariate pairwise correlation function g(r), and the gray shaded area indicates the 95% confidence interval.

图6 不同径级红松的种内关联。A, 次生杨桦林。B, 阔叶红松林。实线表示双变量成对相关函数g12(r)的函数值, 虚线表示双变量成对相关函数g12(r)的期望值, 灰色阴影部分表示95%的置信区间。

Fig. 6 Intraspecific associations between different diameter classes in Pinus koraiensis. A, Secondary poplar-birch forests. B, Broadleaf Korean pine mixed forests. The solid line indicates the function value of the bivariate pairwise correlation function g12(r), the dashed line indicates the expected value of the bivariate pairwise correlation function g12(r), and the gray shaded area indicates the 95% confidence interval.

图7 不同径级红松与其他优势种的种间关联。A, 次生杨桦林。B, 阔叶红松林。

Fig. 7 Interspecific associations between Pinus koraiensis and other dominant species at different diameter classes. A, Secondary poplar-birch forests. B, Broadleaf Korean pine mixed forests. BP, Betula platyphylla; PD, Populus davidiana; TA, Tilia amurensis.

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东北阔叶红松林不同演替阶段树种分布与空间关联格局

Spatial distribution and association patterns of tree species across successional stages in broadleaf Korean pine forests of Northeast China

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