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

doi:10.17521/cjpe.2025.0365

Abstract

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

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[J]. Chin J Plant Ecol, 2026, 50(4): 833-845.

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URL: https://www.plant-ecology.com/EN/10.17521/cjpe.2025.0365

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

Figures/Tables 10

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.

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 (+)

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 (+)

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 (+)

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.

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.

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.

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.

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.

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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