云南鸡足山半湿润常绿阔叶林群落木本植物多样性格局与环境解释

doi:10.17520/biods.2023217

生物多样性 ›› 2023, Vol. 31 ›› Issue (11): 23217. DOI: 10.17520/biods.2023217

• 半湿润常绿阔叶林专题 • 上一篇下一篇

云南鸡足山半湿润常绿阔叶林群落木本植物多样性格局与环境解释

王晓凤¹, 饶杰生¹, 杨涛¹, 刘文聪¹, 田希¹, 陈稀¹, 刘其明², 徐衍潇², 张秋雨¹, 张洪强³, 张旭³, 欧晓昆¹, 沈泽昊²^,¹^,^*()

1.云南大学生态与环境学院, 昆明 650091
2.北京大学城市与环境学院, 地表过程分析与模拟教育部重点实验室, 北京 100871
3.云南佑图生物科技有限公司, 昆明 650599

收稿日期:2023-06-25 接受日期:2023-08-29 出版日期:2023-11-20 发布日期:2023-12-23
通讯作者: * E-mail: shzh@urban.pku.edu.cn
基金资助:
云南省科技厅-云南大学双一流学科建设重大专项(2018FY001-002);云南省基础研究计划重大项目课题(202101BC07 0002)

Spatial variation and determinants of woody plant species diversity in a semi-humid evergreen broad-leaved forest in the Jizu Mountains, Yunnan

Xiaofeng Wang¹, Jiesheng Rao¹, Tao Yang¹, Wencong Liu¹, Xi Tian¹, Xi Chen¹, Qiming Liu², Yanxiao Xu², Qiuyu Zhang¹, Hongqiang Zhang³, Xu Zhang³, Xiaokun Ou¹, Zehao Shen²^,¹^,^*()

1 School of Ecology and Environmental Science, Yunnan University, Kunming 650091
2 Key Laboratory of MOE for Earth Surface Processes, College of Urban and Environmental Sciences, Peking University, Beijing 100871
3 Yunnan Youtu Biotechnology Co., LTD, Kunming 650599

Received:2023-06-25 Accepted:2023-08-29 Online:2023-11-20 Published:2023-12-23
Contact: * E-mail: shzh@urban.pku.edu.cn

摘要/Abstract

摘要：

生物多样性格局是生物群落构建过程的反映, 有助于理解生态系统的物种共存机制。半湿润常绿阔叶林是亚热带西部半湿润气候下的地带性植被类型, 对其群落内物种多样性的空间异质格局及影响因子的认识还比较缺乏, 基于大样地的研究还未见报道。本文基于鸡足山20.16 ha森林动态监测样地504个样方的木本植物调查数据和TWINSPAN群落分类结果, 分析了样方尺度上α多样性和β多样性的空间分布格局及其环境影响因子。结果表明: (1)样地内云南松(Pinus yunnanensis)群落及其乔木和灌木的物种丰富度均高于高山栲(Castanopsis delavayi)群落和元江栲(C. orthacantha)群落; 高山栲群落及其灌木物种丰富度低于元江栲群落, 乔木物种丰富度则高于元江栲群落。相反, 元江栲群落的Shannon-Wiener多样性指数最高, 其次为高山栲群落, 云南松群落的最低。基于净种间亲缘关系指数, 元江栲群落、高山栲群落、云南松群落的谱系结构分别为聚集、随机、发散。(2)对物种丰富度存在普遍性影响的因子有土壤总氮、pH值、相对海拔高度、木本植物的胸高断面积之和; 导致乔木种丰富度变化的主要因子有土壤总氮、pH值、胸高断面积之和; 导致灌木种丰富度变化的主要因子是相对海拔高度、土壤总氮、坡度和地形湿度指数。(3)物种β多样性格局显示, 随着空间距离和环境距离的增大, 物种差异性和物种周转率增大, 即环境越相似, 群落越相似。上述结果表明, 鸡足山样地中先锋性的云南松群落与稳定的元江栲群落和高山栲群落在物种多样性和构成上存在显著差异, 指示了半湿润常绿阔叶林异质斑块所对应的物种共存和物种多样性维持特征, 为进一步的机理性研究提供了有效证据。

关键词: 鸡足山, α多样性, β多样性, 环境因子, 地理加权回归模型, Mantel检验

Abstract

Aims: Spatial patterns of biodiversity can reflect the community assembly processes and the mechanisms of species coexistence within ecosystems. The semi-humid evergreen broad-leaved forest is a type of vegetation zone located in the semi-humid climate of the western subtropical region of China. To date, there have been many studies investigating the species diversity of this type of vegetation, but studies into the patterns of spatial heterogeneity of species diversity and its determinants are still relatively lacking, and research utilizing large dynamics plot has not yet been reported. To better understand the maintenance mechanism of species diversity, we explored the characteristics of spatial heterogeneity and species diversity and their main environmental determinants in a 20.16 ha forest dynamics plot located in the Jizu Mountains.

Methods: Based on data collected from woody plants and TWINSPAN classification results from 504 quadrats 20 m × 20 m in the 20.16 ha forest dynamics plot in the Jizu Mountains, this article utilized regression analysis and the Mantel test to analyze the distribution patterns of α and β diversity, and their correlation with respective environmental factors.

Results (1) In the plot, we found that the species richness of the Pinus yunnanensis community and its trees and shrubs were all higher than that of the Castanopsis delavayi community and C. orthacantha community; the C. delavayi community and its shrubs species richness were both higher than that of the C. orthacantha community, while its tree species richness was higher than C. orthacantha community. On the contrary, the Shannon-Wiener value of the C. orthacantha community was found to be the highest among all species, followed by the C. delavayi community, while the value of the P. yunnanensis community was the lowest. Based on the net interspecific relationship index (NRI), the phylogenetic structures of the C. orthacantha community, C. delavayi community, and P. yunnanensis community were clustering, random, and overdispersion, respectively. (2) The factors that demonstrated a universal impact on species richness included the total nitrogen and pH, the relative altitude, and the sum of basal area of woody plants. The main factors affecting the richness of tree diversity included total nitrogen level and pH and the sum of basal area. The dominant factor influencing shrub species richness was the relative altitude, followed by total nitrogen, slope, and the topographic wetness index. (3) The species β diversity pattern showed that differences in species composition and turnover rate increased with increasing spatial and environmental distances, indicating that the more similar the environment, the more similar the community.

Conclusion: This study reflects the significant differences in species diversity and composition between the pioneer P. yunnanensis community and the stable C. orthacantha community and C. delavayi community in the Jizu Mountains forest dynamics plot, which describes the characteristics of species coexistence and species diversity maintenance corresponding to the heterogeneous patches of semi-humid evergreen broad-leaved forests, and provides a basis for further mechanistic investigations.

Key words: Jizu Mountains, α diversity, β diversity, environmental factors, geographically weighted regression model, Mantel test

王晓凤, 饶杰生, 杨涛, 刘文聪, 田希, 陈稀, 刘其明, 徐衍潇, 张秋雨, 张洪强, 张旭, 欧晓昆, 沈泽昊 (2023) 云南鸡足山半湿润常绿阔叶林群落木本植物多样性格局与环境解释. 生物多样性, 31, 23217. DOI: 10.17520/biods.2023217.

Xiaofeng Wang, Jiesheng Rao, Tao Yang, Wencong Liu, Xi Tian, Xi Chen, Qiming Liu, Yanxiao Xu, Qiuyu Zhang, Hongqiang Zhang, Xu Zhang, Xiaokun Ou, Zehao Shen (2023) Spatial variation and determinants of woody plant species diversity in a semi-humid evergreen broad-leaved forest in the Jizu Mountains, Yunnan. Biodiversity Science, 31, 23217. DOI: 10.17520/biods.2023217.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2023217

https://www.biodiversity-science.net/CN/Y2023/V31/I11/23217

图/表 8

表1 鸡足山半湿润常绿阔叶林动态监测样地3个群落的物种数量统计

Table 1 Statistics of species numbers of three communities in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains

群落类型 Community type	样方数量 Number of quadrats	面积 Area (ha)	群落 Community		乔木 Trees		灌木 Shrubs
群落类型 Community type	样方数量 Number of quadrats	面积 Area (ha)	物种数量 Species number	物种密度 Species density (species/ha)	物种数量 Species number	物种密度 Species density (species/ha)	物种数量 Species number	物种密度 Species dendity (species/ha)
云南松群落 Pinus yunnanensis community	67	2.68	71	26.49	26	9.70	45	16.79
高山栲群落 Castanopsis delavayi community	203	8.12	72	8.87	31	3.82	41	5.05
元江栲群落 C. orthacantha community	234	9.36	82	8.76	31	3.31	51	5.45
总计 Total	504	20.16	103	5.11	38	1.88	65	3.22

图1 鸡足山半湿润常绿阔叶林动态监测样地3个群落的种-面积曲线(a, b, c)和种-多度曲线(d, e)。(a)木本植物; (b)乔木; (c)灌木; (d)木本植物; (e)常绿阔叶种和落叶阔叶种。图中虚线部分为实际种-面积对数函数模型的拟合延长线。

Fig. 1 Species-area curves (a, b, c) and species-abundance curves (d, e) of the three communities in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains. (a) Woody plant; (b) Trees; (c) Shrubs; (d) Woody plant; (e) Evergreen and deciduous broad-leaved species. The dashed line in the figure represents the fitting extension of the actual species-area logarithmic function model.

图2 鸡足山半湿润常绿阔叶林动态监测样地504个样方的α多样性和谱系多样性空间格局。(a)物种丰富度(SR); (b) Shannon-Wiener多样性指数(H′); (c)净种间亲缘关系指数(NRI)。

Fig. 2 Spatial variation of α diversity and lineage diversity among 504 quadrats in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains. (a) Species richness (SR); (b) Shannon-Wiener diversity index (H′); (c) Net interspecific relationship index (NRI).

表2 鸡足山半湿润常绿阔叶林动态监测样地3个群落木本植物、乔木、灌木的Shannon-Wiener多样性指数、物种丰富度格局的普通最小二乘(OLS)回归模型和地理加权回归(GWR)模型拟合优度(R2)

Table 2 Fitting degrees (R2) of ordinary least squares (OLS) regression model and geographically weighted regression (GWR) model on patterns of Shannon-Wiener diversity index and species richness of woody plants, trees and shrubs of the three communities in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains

尺度 Scale	多样性指数 Diversity index	云南松群落 Pinus yunnanensis community		高山栲群落 Castanopsis delavayi community		元江栲群落 C. orthacantha community
尺度 Scale	多样性指数 Diversity index	OLS	GWR	OLS	GWR	OLS	GWR
木本植物 Woody plant	Shannon-Wiener多样性指数 Shannon-Wiener diversity index	0.661	0.798	0.358	0.777	0.254	0.637
木本植物 Woody plant	物种丰富度 Species richness	0.25	0.43	0.303	0.533	0.314	0.474
乔木 Trees	Shannon-Wiener多样性指数 Shannon-Wiener diversity index	0.557	0.734	0.396	0.811	0.282	0.819
乔木 Trees	物种丰富度 Species richness	0.248	0.336	0.179	0.465	0.185	0.546
灌木 Shrubs	Shannon-Wiener多样性指数 Shannon-Wiener diversity index	0.537	0.664	0.310	0.655	0.294	0.566
灌木 Shrubs	物种丰富度 Species richness	0.157	0.341	0.363	0.583	0.409	0.468

图3 基于普通最小二乘(OLS)回归模型(a)和地理加权回归(GWR)模型(b)解释鸡足山半湿润常绿阔叶林动态监测样地3个群落的木本植物、乔木、灌木的物种丰富度格局。Age: 群落年龄; BasA: 胸高断面积之和; Clay: 黏粒含量; Silt: 粉粒含量; Sand: 砂粒含量; OrgC: 有机碳; TotN: 总氮; AvaP: 有效磷; pH: pH值; Alti: 相对海拔高度; Slop: 坡度; Aspe: 坡向; Curv: 曲率; Roug: 粗糙度; Twi: 地形湿度指数。

Fig. 3 Explanation of species richness patterns of woody plant, trees, and shrubs of the three communities in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains based on ordinary least squares (OLS) regression model (a) and geographically weighted regression (GWR) (b) model. Age, Community age; BasA, Sum of basal area; Clay, Clay content; Silt, Silt content; Sand, Sand content; OrgC, Organic carbon; TotN, Total nitrogen; AvaP, Available phosphorus; pH, pH value; Alti, Relative altitude; Slop, Slope; Aspe, Aspect; Curv, Curvature; Roug, Roughness; Twi, Topographic wetness index.

图4 鸡足山半湿润常绿阔叶林动态监测样地在20 m粒度的Whittaker指数和物种周转率与环境距离和空间距离的关系。Age: 群落年龄; BasA: 胸高断面积之和; Clay: 黏粒含量; Silt: 粉粒含量; Sand: 砂粒含量; OrgC: 有机碳; TotN: 总氮; AvaP: 有效磷; pH: pH值; Alti: 相对海拔高度; Slop: 坡度; Aspe: 坡向; Curv: 曲率; Roug: 粗糙度; Twi: 地形湿度指数; Spa: 空间距离。

Fig. 4 Relationship between Whittaker’s index and species turnover rate with environmental distance and spatial distance at 20 m grain, estimated within the forest dynamics plot of the semi-humid evergreen broad-leaved forests in the Jizu Mountains. Age, Community age; BasA, Sum of basal area; Clay, Clay content; Silt, Silt content; Sand, Sand content; OrgC, Organic carbon; TotN, Total nitrogen; AvaP, Available phosphorus; pH, pH value; Alti, Relative altitude; Slop, Slope; Aspe, Aspect; Curv, Curvature; Roug, Roughness; Twi, Topographic wetness index; Spa, Spatial distance.

表3 鸡足山半湿润常绿阔叶林动态监测样地在20 m粒度的Whittaker指数与环境距离和空间距离的Mantel (对角线上方)和偏Mantel (对角线下方)检验结果

Table 3 Mantel (above the diagonal) and partial Mantel (below the diagonal) tests for the relationships between Whittaker’s index and environmental and spatial distances at 20 m grain in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains

	Whittaker指数 β_W	环境距离 Env	空间距离 Spa
Whittaker指数 Whittaker’s index (β_W)	-	0.325, P = 0.001	0.456, P = 0.001
环境距离 Environment distance (Env)	0.253, P = 0.001	-	0.233, P = 0.001
空间距离 Spatial distance (Spa)	0.413, P = 0.001	0.101, P = 0.001	-

表4 鸡足山半湿润常绿阔叶林动态监测样地在20 m粒度的物种周转率与环境距离和空间距离的Mantel (对角线上方)和偏Mantel (对角线下方)检验结果

Table 4 Mantel (above the diagonal) and partial Mantel (below the diagonal) tests for the relationships between species turnover rate and environmental and spatial distances at 20 m grain in the forest dynamics plot in a semi-humid evergreen broad-leaved forest in the Jizu Mountains

	物种周转率 β_tu	环境距离 Env	空间距离 Spa
物种周转率 Species turnover rate (β_tu)	-	0.317, P = 0.001	0.431, P = 0.001
环境距离 Environment distance (Env)	0.247, P = 0.001	-	0.233, P = 0.001
空间距离 Spatial distance (Spa)	0.387, P = 0.001	0.113, P = 0.001	-

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云南鸡足山半湿润常绿阔叶林群落木本植物多样性格局与环境解释

Spatial variation and determinants of woody plant species diversity in a semi-humid evergreen broad-leaved forest in the Jizu Mountains, Yunnan

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