云南香格里拉亚高山寒温性针叶林优势种空间分布格局及种内种间关联性

doi:10.17521/cjpe.2024.0066

植物生态学报 ›› 2025, Vol. 49 ›› Issue (2): 268-281.DOI: 10.17521/cjpe.2024.0066 cstr: 32100.14.cjpe.2024.0066

云南香格里拉亚高山寒温性针叶林优势种空间分布格局及种内种间关联性

万嘉敏¹, 张彩彩¹^,^*(), 邓云²^,³^,^*(), 顾荣², 斯那取宗⁴, 吴俊华⁴, 娄启妍⁴, 陈梅⁴, 张志明⁵, 林露湘²^,³

¹大理大学东喜玛拉雅研究院, 云南大理 671003
²中国科学院西双版纳热带植物园热带森林生态学重点实验室, 云南勐腊 666303
³云南西双版纳森林生态系统国家野外科学观测研究站, 云南勐腊 666303
⁴香格里拉普达措国家公园管理局, 云南迪庆 674499
⁵云南大学生态与环境学院西南跨境生态安全教育部重点实验室, 昆明 650091

收稿日期:2024-03-09 接受日期:2024-06-20 出版日期:2025-02-20 发布日期:2025-02-20
通讯作者: ^*张彩彩: (zhangcc@eastern-himalaya.cn);
邓云: (dy@xtbg.org.cn)
基金资助:
中央财政林草生态保护恢复资金(WNLY-2022-06-018);国家自然科学基金-云南联合基金(U1902203);国家自然科学基金(32160268);云南省基础研究专项(202101BC070002);云南省重点研发计划(202303AC100009)

Spatial distribution patterns and intraspecific and interspecific associations of dominant species in subalpine cold-temperate coniferous forests of Shangri-La, Yunnan, China

WAN Jia-Min¹, ZHANG Cai-Cai¹^,^*(), DENG Yun²^,³^,^*(), GU Rong², SINA Qu-Zong⁴, WU Jun-Hua⁴, LOU Qi-Yan⁴, CHEN Mei⁴, ZHANG Zhi-Ming⁵, LIN Lu-Xiang²^,³

¹Institute of Eastern-Himalaya Biodiversity Research, Dali University, Dali, Yunnan 671003, China
²CAS Key Laboratory of Tropical Forest Ecology, Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Mengla, Yunnan 666303, China
³National Forest Ecosystem Research Station at Xishuangbanna, Mengla, Yunnan 666303, China
⁴Potatso National Park Authority in Shangri-La, Dêqên, Yunnan 674499, China
⁵Key Laboratory for Transboundary Ecosecurity of Southwest China of Ministry of Education, School of Ecology and Environmental Sciencé, Yunnan University, Kunming 650091, China

Received:2024-03-09 Accepted:2024-06-20 Online:2025-02-20 Published:2025-02-20
Supported by:
Central Government Finance for Forest and Grassland Ecological Protection and Restoration(WNLY-2022-06-018);Joint Fund of the National Natural Science Foundation of China-Yunnan Province(U1902203);National Natural Science Foundation of China(32160268);Basic Research Special Project of Yunnan Province(202101BC070002);Key Research and Development Plan of Yunnan Province(202303AC100009)

摘要/Abstract

摘要： 植物种群空间分布格局是散布限制和环境过滤等多种生态过程综合作用的结果。分布在高山树线交错带的植物因其特殊的生境, 对气候变化表现出高度的敏感性。因此, 研究这些植物的空间分布格局及其相互关系, 对理解和预测高山林线森林群落的动态和发展趋势至关重要。该研究基于云南香格里拉亚高山寒温性针叶林20 hm²动态监测样地的调查数据, 以样地内优势种长苞冷杉(Abies georgei)、亚乔木层优势种红棕杜鹃(Rhododendron rubiginosum)和西南花楸(Sorbus rehderiana)、灌木层优势种唐古特忍冬(Lonicera tangutica)和云南双盾木(Dipelta yunnanensis)为研究对象, 采用空间点格局方法分析各优势种的空间分布格局、长苞冷杉不同发育阶段间的种内关联性、长苞冷杉与其他优势种间的种间关联性, 以及其他优势种种间关联性, 并使用Torus-translation方法检验这些植物与地形因子的关联性。结果表明: (1)长苞冷杉的幼树和中树均呈现聚集分布, 这主要由散布限制和生境异质性驱动; 而成树主要呈随机分布, 表明密度依赖性的竞争对大径级个体分布的主导作用。亚乔木层和灌木层的优势种均呈聚集分布, 但剔除环境异质性后部分优势种转变为随机分布, 说明环境过滤驱动了树种空间分布模式。(2)长苞冷杉的幼树与中树呈正关联, 可能是小径级个体通过集群作用来提高抵御外界环境胁迫的能力。幼树和中树与成树呈负关联, 这主要受由密度制约引起的专一性病原菌和植食性昆虫的侵害以及大个体对小个体的不对称竞争的影响。(3)长苞冷杉的幼树与亚乔木层和灌木层的优势种分别呈正关联和负关联; 中树与其他优势种大多表现为负关联, 而成树则多表现为正关联; 乔木层和灌木层优势种之间多表现为正关联。说明亚高山寒温性针叶林优势种之间存在复杂的动态平衡。各优势种通过独特的生存策略和资源利用方式来实现长期共存, 最终形成以长苞冷杉为主导的相对稳定的顶极群落。(4)坡度与长苞冷杉的幼树和中树的密度显著负相关, 与红棕杜鹃和云南双盾木显著正相关, 说明长苞冷杉与其他优势种发生了坡度生态位的分化。此外, 由于冬季积雪时间较长等不利因素, 凹凸度也对优势种的分布具有显著的影响。总体而言, 地形驱动的生境过滤可能是维持亚高山寒温性针叶林群落构建的主要驱动力。

关键词: 长苞冷杉, 点格局, 空间关联性, 生境关联

Abstract:

Aims The spatial distribution patterns of plant populations result from the combined effects of multiple ecological processes, such as dispersal limitation and environmental filtering. The plants distributed in alpine treeline ecotones are highly sensitive to climate change due to their unique habitats. Therefore, studying the spatial distribution patterns of these plants and their correlations is critical for understanding and predicting the dynamics and trends of forest communities in alpine treelines.

Methods This study is based on the inventory data collected from a 20 hm²dynamics plot of a subalpine cold-temperate coniferous forest in Shangri-La, Yunnan, China. The dominant tree species identified were Abies georgei, Lonicera tangutica, Dipelta yunnanensis, Rhododendron rubiginosum, and Sorbus rehderiana. The spatial point pattern method was used to analyze the spatial distribution pattern of each dominant species, the intraspecific association of A. georgei at different developmental stages, the interspecific association between A. georgei and the other dominant species, and the interspecific association among the other dominant species. Additionally, the Torus-translation method was applied to test the associations between these plants and topographic factors.

Important findings (1) Sapling and juvenile trees of A. georgei demonstrated aggregated distributions, primarily driven by dispersal limitation and habitat heterogeneity. In contrast, adult trees exhibited a predominantly random distribution, suggesting that density-dependent competition may be the primary factor influencing the distribution of individuals in large-diameter classes. The dominant species in both the subtree layer and shrub layer also demonstrated aggregated distribution. However, the posterior partial advantage of the environmental heterogeneity transformed into a random distribution, indicating that environmental filtering might be responsible for driving the spatial distribution pattern of these tree species. (2) Positive associations were observed between sapling and juvenile trees of A. georgei indicating that small-diameter individuals tend to congregate due to an enhanced capacity to cope with external environmental stresses. Conversely, saplings and juvenile trees were negatively correlated with adult trees. This was mainly due to the infestation of specific pathogens and phytophagous insects caused by density constraints and asymmetric competition of large individuals against smaller ones. (3) There were positive and negative correlations between the saplings and the dominant species in the subtree layer and the shrub layer, respectively. The juvenile trees and other dominant species revealed predominantly negative correlation, while the adult trees showed predominantly positive correlation. The majority of the dominant species in the tree layer and shrub layer exhibited positive correlation, indicating a complex dynamic balance within the dominant species in the subalpine cold-temperate coniferous forest. The long-term coexistence of each dominant species in the plot is achieved through their unique survival strategies and resource utilization, and ultimately leading to the formation of a relatively stable successional climax community dominated by A. georgei. (4) Slope was found to be significantly negatively correlated with sapling and juvenile trees of A. georgei, and significantly positively related to R. rubiginosum and D. yunnanensis. This suggests that the slope ecological niche differentiation occurred between A. georgei and other dominant species. Additionally, convexity was found to exert a significant effect on the distribution of dominant species due to adverse conditions such as prolonged snowpack in winter. In conclusion, the habitat filtering driven by topography is the main driver that maintains community assembly in subalpine cold-temperate coniferous forests.

Key words: Abies georgei, spatial point pattern, spatial association, habitat association

万嘉敏, 张彩彩, 邓云, 顾荣, 斯那取宗, 吴俊华, 娄启妍, 陈梅, 张志明, 林露湘. 云南香格里拉亚高山寒温性针叶林优势种空间分布格局及种内种间关联性. 植物生态学报, 2025, 49(2): 268-281. DOI: 10.17521/cjpe.2024.0066

WAN Jia-Min, ZHANG Cai-Cai, DENG Yun, GU Rong, SINA Qu-Zong, WU Jun-Hua, LOU Qi-Yan, CHEN Mei, ZHANG Zhi-Ming, LIN Lu-Xiang. Spatial distribution patterns and intraspecific and interspecific associations of dominant species in subalpine cold-temperate coniferous forests of Shangri-La, Yunnan, China. Chinese Journal of Plant Ecology, 2025, 49(2): 268-281. DOI: 10.17521/cjpe.2024.0066

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链接本文: https://www.plant-ecology.com/CN/10.17521/cjpe.2024.0066

https://www.plant-ecology.com/CN/Y2025/V49/I2/268

图/表 6

图1 云南香格里拉亚高山寒温性针叶林20 hm2动态监测样地中长苞冷杉种群不同发育阶段的空间分布格局。A, 幼树。B, 中树。C, 成树。黑色实线表示单变量成对相关函数g(r)的函数值, 红色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99%的置信区间。地图中绿色表示低海拔, 红色表示高海拔。p值为拟合优度检验结果。

Fig. 1 Spatial distribution pattern of Abies georgei at different developmental stages in the 20 hm2 dynamics plot of subalpine cold-temperate coniferous forest in Shangri-La, Yunnan. A, Saplings. B, Juvenile trees. C, Adult trees. Solid black lines represents the value of the univariate pair-correlation g(r) function, the dashed red line represents the expected value of the univariate pair-correlation g(r) function, and the gray shaded part represents the 99% confidence interval. Green on the map indicates low altitudes and red indicates high altitudes. The p-value is the goodness of fit test result.

图2 云南香格里拉亚高山寒温性针叶林20 hm2动态监测样地中亚乔木层和灌木层优势种的空间分布格局。A, 红棕杜鹃。B, 西南花楸。C, 唐古特忍冬。D, 云南双盾木。黑色实线表示单变量成对相关函数g(r)的函数值, 红色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99%的置信区间。地图中绿色表示低海拔, 红色表示高海拔。p值为拟合优度检验结果。

Fig. 2 Spatial distribution pattern of dominant species in the subtree layer and shrub layer in the 20 hm2 dynamics plot of subalpine cold-temperate coniferous forest in Shangri-La, Yunnan. A, Rhododendron rubiginosum. B, Sorbus rehderiana. C, Lonicera tangutica. D, Dipelta yunnanensis. Solid black lines represent the value of the univariate pair-correlation g(r) function, the dashed red lines represent the expected value of the univariate pair-correlation g(r) function, and the gray shaded parts represent the 99% confidence interval. Green on the map indicates low altitudes and red indicates high altitudes. The p-value is the goodness of fit test result.

图3 云南香格里拉亚高山寒温性针叶林20 hm2动态监测样地中长苞冷杉种群不同发育阶段的种内关联性。A, 幼树和中树。B, 幼树和成树。C, 中树和成树。黑色曲线表示双变量成对相关函数g12(r)的函数值, 黑色直线表示双变量成对相关函数g12(r)的期望值, 灰色虚线表示99%的置信区间。p值为拟合优度检验结果。

Fig. 3 Intraspecific correlation of Abies georgei population at different developmental stages in the 20 hm2 dynamics plot of subalpine cold-temperate coniferous forest in Shangri-La, Yunnan. A, Sapling and juvenile trees. B, Sapling and adult trees. C, Juvenile and adult trees. The black curve represents the function value of the bivariate pair-correlation function g12(r), and the black straight line represents the expected value of the bivariate pair-correlation g12(r) function. The dashed gray line indicates a 99% confidence interval. The p-value is the goodness of fit test result.

图4 云南香格里拉亚高山寒温性针叶林20 hm2动态监测样地中长苞冷杉种群不同发育阶段与其他优势种的种间关联性。A, 长苞冷杉与亚乔木层优势种之间的关联性。B, 长苞冷杉与灌木层优势种之间的关联性。p值为拟合优度检验结果。

Fig. 4 Interspecific relationship between Abies georgei at different developmental stages and other dominant species in the 20 hm2 dynamics plot of subalpine cold-temperate coniferous forest in Shangri-La, Yunnan. A, Relationship between Rhododendron rubiginosum and dominant species in subtree layers. B, Relationship between Rhododendron rubiginosum and dominant species in shrub layers. DY, Dipelta yunnanensis; LT, Lonicera tangutica; RR, Rhododendron rubiginosum; SR, Sorbus rehderiana. The p-value is the goodness of fit test result.

图5 云南香格里拉亚高山寒温性针叶林20 hm2动态监测样地中亚乔木层和灌木层优势种的种间关联性。p值为拟合优度检验结果。

Fig. 5 Interspecific relationship of dominant species in subtree layer and shrub layer in 20 hm2 dynamics plot of subalpine cold-temperate coniferous forest in Shangri-La, Yunnan. DY, Dipelta yunnanensis; LT, Lonicera tangutica; RR, Rhododendron rubiginosum; SR, Sorbus rehderiana. The p-value is the goodness of fit test result.

表1 云南香格里拉亚高山寒温性针叶林20 hm2动态监测样地长苞冷杉种群及其他优势种与地形因子的Torus-translation检验及物种密度与地形因子的Spearman相关系数

Table 1 Torus-translation test and Spearman correlation coefficient of species density and topographic factors of Abies georgei and other dominant species in the 20 hm2 dynamics plot of subalpine cold-temperate coniferous forest in Shangri-La, Yunnan

发育阶段/物种 Development stage/species	海拔 Altitude	坡度 Slope	凹凸度 Convex	正弦坡向 Sin (aspect)	余弦坡向 Cos (aspect)
幼树 Sapling	0.16	-0.41^*	0.08	-0.13	-0.13
中树 Juvenile	0.07	-0.30^*	-0.08	0.06	0.08
成树 Adult	0.27^*	0.04	0.34^*	0.07	0.08
红棕杜鹃 Rhododendron rubiginosum	-0.17	0.39^*	0.28^*	-0.07	-0.07
西南花楸 Sorbus rehderiana	-0.26^*	0.15	-0.08	-0.07	-0.10
唐古特忍冬 Lonicera tangutica	0.50^*	-0.04	0.17^*	0.35^*	0.38^*
云南双盾木Dipelta yunnanensis	0.05	0.49^*	0.08	0.50^*	0.51^*

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云南香格里拉亚高山寒温性针叶林优势种空间分布格局及种内种间关联性

Spatial distribution patterns and intraspecific and interspecific associations of dominant species in subalpine cold-temperate coniferous forests of Shangri-La, Yunnan, China

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