滇中半湿润常绿阔叶林木本植物的功能特征和功能多样性及其影响因子

doi:10.17520/biods.2023215

生物多样性 ›› 2023, Vol. 31 ›› Issue (11): 23215. DOI: 10.17520/biods.2023215

• 半湿润常绿阔叶林专题 • 上一篇下一篇

滇中半湿润常绿阔叶林木本植物的功能特征和功能多样性及其影响因子

罗彩访¹, 杨涛¹, 张秋雨¹, 王馨培², 沈泽昊¹^,²^,^*()

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, 北京大学地表过程分析与模拟教育部重点实验室, 北京 100871

收稿日期:2023-06-25 接受日期:2023-11-07 出版日期:2023-11-20 发布日期:2023-11-21
通讯作者: * E-mail: shzh@urban.pku.edu.cn
基金资助:
国家自然科学基金(41971228);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)

Plant functional traits, community functional diversity and their environmental determinants of the semi-humid evergreen broad-leaved forest in the Central Yunnan Plateau

Caifang Luo¹, Tao Yang¹, Qiuyu Zhang¹, Xinpei Wang², Zehao Shen¹^,²^,^*()

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 100871s

Received:2023-06-25 Accepted:2023-11-07 Online:2023-11-20 Published:2023-11-21
Contact: * E-mail: shzh@urban.pku.edu.cn

摘要/Abstract

摘要：

半湿润常绿阔叶林是中国常绿阔叶林西部中亚热带类型的代表, 为了解其群落的功能多样性, 本研究对云南滇中地区金光寺、鸡足山、雕翎山、筇竹寺和珠江源5个半湿润常绿阔叶林分布点进行群落调查和木本植物性状测定, 分析不同样地和群落类型的植物功能特征和群落功能多样性, 探究滇中半湿润常绿阔叶林木本植物之间的差异及其环境影响因素。结果表明: (1)不同样地和不同群落类型下功能性状的群落加权平均值大致相当; 灌木层的比叶面积平均值高于乔木层; 乔木层、灌木层的木材平均密度无显著差异(约0.5 g/cm³); (2)基于最大株高、比叶面积和木材密度计算的半湿润常绿阔叶林乔木层功能多样性在不同样地间差异显著(P < 0.05), 其中鸡足山和雕翎山的乔木层功能丰富度、均匀度和冗余度较高, 群落更稳定。不同群落类型所占的功能空间大小不同, 但群落内物种功能分异度、均匀度和冗余度相似。此外, 灌木层的物种丰富度高于乔木层, 功能丰富度却相对较低; (3)功能丰富度和功能冗余度都与物种丰富度显著正相关(P < 0.05), 与其他多样性指数关系不显著。环境因子中气候和土壤因素共同解释群落乔木层、灌木层功能性状和功能多样性的60%以上变异性, 人类活动也会对植物群落功能多样性造成一定影响。本研究揭示了半湿润常绿阔叶林不同群落类型和层次的植物功能性状差异及其区域变化, 为该植被类型的生物多样性和生态系统功能保护与恢复提供科学依据。

关键词: 半湿润常绿阔叶林, 功能性状, 功能多样性, 环境解释

Abstract

Aims: Semi-humid evergreen broad-leaved forest (SEBF) represents a distinct sub-type of evergreen broad-leaved forests within the subtropical region of western China. Despite their ecological significance, the functional diversity of SEBFs remains in adequately understood. This study aims to conduct a comprehensive analysis of plant traits and functional diversity within SEBFs and elucidate the environmental factors influencing their variations.

Methods: This research was conducted across the expanse of the Central Yunnan Plateau, the primary natural habitat for SEBFs. We conducted surveys within five well-protected SEBF sites, namely Jinguangsi (JGS), Jizu Mountains (JZS), Diaoling Mountains (DLS), Qiongzhusi (QZS), and Zhujiangyuan (ZJY). A total of 29 plots measuring 20 m × 20 m were sampled. Key plant traits including the maximum height (H_max), specific leaf area (SLA), and wood density, were recorded for collected samples. Functional diversity indices such as functional richness (FRic), functional divergence (FDiv), functional evenness (FEve), and functional redundancy (Rstar) were calculated based on these traits. Additionally, life-form spectrums and community-weighted mean trait values (CWM) were assessed for different sample sites and community types. Environmental factor analysis, using variance partitioning, was employed to gauge the impact of various ecological factors on the variations in functional diversity within SEBFs.

Results: Within the semi-humid evergreen broad-leaved forests (SEBFs) of the Central Yunnan Plateau, the CWM of key traits, namely H_max, SLA and wood density, exhibited no significant differences among various sample sites and community types. The shrub layer demonstrated higher CWM_SLA values compared to the tree layer, while their CWM_wood_density remained consistent at 0.5 g/cm³. In terms of diversity, both species diversity and functional diversity varied between the tree layer and shrub layer. Notably, JZS and DLS exhibited the highest values for FRic, FEve, and Rstar, indicating greater resilience and stability in these two sample sites. However, different community types of SEBFs, based on various dominant species, displayed similar FDiv, FEve, and Rstar values, differing only in FRic. FRic and Rstar were both significantly positively correlated with species richness (P < 0.05), while their relationships with other functional diversity indices were not significant. Factor analysis revealed that climate and soil nutrient contributed more than 60% of variances in CWM and functional diversity within SBEFs. Additionally, human activities emerged as influential factors in shaping the functional diversity of these forest communities.

Conclusion: Our results shed light on the characteristics of CWM values and functional diversity within the SEBFs, particularly in the context of tree layer and shrub layer. We have observed distinct spatial variations across different sample sites. Interestingly, despite the existence of different community types based on various dominant species, these communities occupy different functional niches yet exhibit a remarkable degree of functional convergence in terms of directional divergence, evenness, and redundancy. These findings provide valuable insights into the functional diversity of SEBFs, which, in turn, can inform and enhance efforts related to forest conservation and biodiversity restoration.

Key words: semi-humid evergreen broad-leaved forest, community-weighted mean trait values, community functional diversity, environmental determinants

罗彩访, 杨涛, 张秋雨, 王馨培, 沈泽昊 (2023) 滇中半湿润常绿阔叶林木本植物的功能特征和功能多样性及其影响因子. 生物多样性, 31, 23215. DOI: 10.17520/biods.2023215.

Caifang Luo, Tao Yang, Qiuyu Zhang, Xinpei Wang, Zehao Shen (2023) Plant functional traits, community functional diversity and their environmental determinants of the semi-humid evergreen broad-leaved forest in the Central Yunnan Plateau. Biodiversity Science, 31, 23215. DOI: 10.17520/biods.2023215.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2023215

https://www.biodiversity-science.net/CN/Y2023/V31/I11/23215

图/表 9

图1 云南省半湿润常绿阔叶林5个调查样地的地理位置及样地照片

Fig. 1 Geographic positions and pictures of the five sampling sites of semi-humid evergreen broad-leaved forests in Yunnan

表1 云南省半湿润常绿阔叶林不同样地的环境因子

Table 1 The environmental factors of semi-humid evergreen broad-leaved forests among different sampling sites in Yunnan

因子 Factors	变量 Variables	金光寺 JGS	鸡足山 JZS	雕翎山 DLS	筇竹寺 QZS	珠江源 ZJY
气候 Climate	年均温 MAT (℃)	11.3	14.7	14.1	15.1	12.5
	年均降水量 MAP (mm)	1,309.3	1,174.0	1,000.2	1,210.5	1,180.0
	湿季均温 MTWQ (℃)	15.4	18.4	18.4	19.5	17.9
	湿季降水量 MPWQ (mm)	636.6	672.8	573.6	692.4	612.8
	干季均温 MTDQ (℃)	5.24	8.71	8.26	9.15	5.95
	干季降水量 MPDQ (mm)	77.8	41.9	40.7	56.3	66.5
	温度季节性 TempS	423.0	422.7	415.5	413.2	465.4
	降水季节性 PrecipS	70.2	90.1	87.8	84.8	79.6
地形 Topography	海拔 Elev (m)	2,504	2,442	2,314	2,197	2,332
土壤 Soil	酸碱度 pH	4.09	5.23	4.70	4.77	3.97
土壤 Soil	全氮 TN (%)	0.31	0.63	0.33	0.57	0.31
	有机碳 OC (mg/g)	9.47	11.2	6.12	5.63	5.04
人类活动 Human activity	距道路最近距离 Dist_road (km)	4.26	0.37	1.47	0.28	0.16
人类活动 Human activity	距居民点最近距离 Dist_resident_. (km)	2.50	1.56	1.85	1.39	1.14
群落类型 Community types		元江栲林 COF	元江栲林 COF 高山栲林 CDF2 黄毛青冈林 CDF	元江栲林 COF 高山栲林 CDF2 白穗柯林 LCF 多变柯林 LVF	元江栲林 COF	元江栲林 COF 灰背栎林 QSF

图2 半湿润常绿阔叶林木本植物中不同生长型物种(灌木、具革质叶片木本植物和具全缘叶片木本植物)重要值占比的比较。(a) (b) (c)为不同样地之间的比较; (d) (e) (f)为不同群落类型之间的比较。不同字母表示差异显著(P < 0.05)。缩写含义见表1。

Fig. 2 Relative proportion of importance values of different growth-form woody species, i.e. shrub species, woody species with leathery leaves, and woody species with entire leaves, within semi-humid evergreen broad-leaved forests. (a), (b), (c) show the multiple comparisons among different sampling sites. (d) (e) (f) show the multiple comparisons among different community types. Different letters denote significant difference of multiple comparisons (P < 0.05). The meanings of abbreviations see Table 1.

图3 半湿润常绿阔叶林植物功能性状(最大株高、比叶面积和木材密度)的群落加权平均值(平均值 ± 标准误差)比较。(a) (b) (c)为不同样地之间的比较; (d) (e) (f)为不同群落类型之间的比较。不同字母表示差异显著(P < 0.05)。缩写含义见表1。

Fig. 3 Community-weighted mean trait values (CWM, mean ± SE) of the maximum height (Hmax), specific leaf area (SLA), and wood density within semi-humid evergreen broad-leaved forests. (a) (b) (c) show the multiple comparisons among different sampling sites. (d) (e) (f) show the multiple comparisons among different community types. Different letters denote significant difference (P < 0.05). The meanings of abbreviations see Table 1.

表2 半湿润常绿阔叶林群落乔木层和灌木层之间功能性状、多样性关系

Table 2 Correlations of community-weighted mean trait values (CWM), species diversity and functional diversity between tree layer and shrub layer within semi-humid evergreen broad-leaved forests. Spearman correlation coefficients (r) showed in this table.

功能性状 CWM	r	物种多样性 Species diversity	r	功能多样性 Functional diversity	r
最大株高 Maximum height (Hmax)	-0.07	物种丰富度 Species richness (SR)	0.76***	功能丰富度 Functional richness (FRic)	0.65***
比叶面积 Specific leaf area (SLA)	-0.17			功能分异度 Functional divergence (FDiv)	0.04
木材密度 Wood density	0.25			功能均匀度 Functional evenness (FEve)	-0.12
				功能冗余度 Functional redundancy (Rstar)	0.11

表3 半湿润常绿阔叶林不同群落类型间多样性差异比较(Wilcoxon秩和检验)。不同字母表示不同群落类型间差异显著(P < 0.05)。

Table 3 Comparison of diversity among different plant community types within semi-humid evergreen broad-leaved forests based on Wilcoxon rank sum test. Different letters denote statistical difference of multiple comparisons (P < 0.05).

群落类型 Community types	样方数 No. of plots	物种丰富度 SR		功能丰富度 FRic		功能分异度 FDiv		功能均匀度 FEve		功能冗余度 Rstar
群落类型 Community types	样方数 No. of plots	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer
灰背栎林 QSF	2	9.3^ab	17.0^ab	7.8^a	5.6^ab	0.86^a	0.71^a	0.61^a	0.62^ab	0.69^a	0.72^ab
黄毛青冈林 CDF	2	13.5^a	12.5^ab	7.9^a	4.2^ab	0.90^a	0.66^a	0.61^a	0.52^ab	0.76^a	0.63^ab
高山栲林 CDF2	4	6.5^b	22.0^a	5.9^ab	7.3^a	0.73^a	0.78^a	0.55^a	0.72^a	0.45^a	0.87^a
白穗柯林 LCF	3	7.5^ab	12.3^ab	7.5^a	3.0^b	0.8^a	0.58^a	0.58^a	0.49^ab	0.63^a	0.58^ab
多变柯林 LVF	2	8.3^ab	10.1^b	7.5^a	2.2^b	0.81^a	0.55^a	0.59^a	0.29^b	0.66^a	0.47^b
元江栲林 COF	16	4.8^b	22.0^a	3.0^b	7.1^a	0.72^a	0.72^a	0.51^a	0.64^ab	0.41^a	0.73^ab

表4 半湿润常绿阔叶林不同样地间群落多样性差异比较(Wilcoxon秩和检验)。不同字母表示不同样地间差异显著(P < 0.05)。

Table 4 Comparison of diversity among different sampling sites within semi-humid evergreen broad-leaved forests based on Wilcoxon rank sum test. Different letters denote statistical difference among different sites (P < 0.05).

样地 Sites	样方数 No. of plots	物种丰富度 SR		功能丰富度 FRic		功能分异度 FDiv		功能均匀度 FEve		功能冗余度 Rstar
样地 Sites	样方数 No. of plots	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer	乔木层 Tree layer	灌木层 Shrub layer
金光寺 JGS	6	2.67^c	17.38^a	0.49^c	9.47^a	0.59^a	0.69^a	0.51^b	0.74^a	0.19^b	0.79^a
鸡足山 JZS	8	10.14^a	10.33^ab	6.93^a	3.80^b	0.81^a	0.66^a	0.83^a	0.55^ab	0.69^a	0.71^a
雕翎山 DLS	7	8.38^ab	7.00^b	6.82^a	2.11^b	0.81^a	0.48^a	0.63^ab	0.47^b	0.64^a	0.33^b
筇竹寺 QZS	2	5.67^bc	16.57^a	4.74^ab	4.64^ab	0.79^a	0.67^a	0.58^b	0.63^ab	0.41^b	0.75^a
珠江源 ZJY	6	3.00^c	10.00^ab	1.74^bc	2.72^b	0.71^a	0.65^a	0.56^b	0.49^ab	0.37^b	0.61^a

表5 半湿润常绿阔叶林群落多样性指数的相关性矩阵(Spearman相关性分析)

Table 5 Correlations among several diversity indices of semi-humid evergreen broad-leaved forests (Spearman correlation analysis)

	多样性指数 Diversity index	乔木层功能多样性 FD of tree layer				灌木层功能多样性 FD of shrub layer
	多样性指数 Diversity index	功能丰富度 FRic	功能分异度 FDiv	功能均匀度 FEve	功能冗余度 Rstar	功能丰富度 FRic	功能分异度 FDiv	功能均匀度 FEve	功能冗余度 Rstar
物种多样性 Species diversity	物种丰富度 SR	0.68***	-0.08	-0.30	0.59**	0.77***	0.10	-0.31	0.60**
功能多样性 Functional diversity (FD)	功能丰富度 FRic	1				1
	功能分异度 FDiv	-0.13	1			0.19	1
	功能均匀度 FEve	-0.34	0.18	1		-0.23	0.36	1
	功能冗余度 Rstar	0.45**	-0.16	-0.23	1	0.40*	0.29	-0.002	1

图4 不同环境因素对半湿润常绿阔叶乔木层(a)和灌木层(b)群落功能性状加权平均值、物种丰富度和功能多样性的贡献。解释量粗体表征随机效应(样地)显著。缩写含义详见图3和表2。

Fig. 4 Contributions of different environmental factors in the CWM of the functional traits, species richness, and functional diversity of tree layer (a) and shrub layer (b) of semi-humid evergreen broad-leaved forests. Different colors indicate different environmental factors, including climate, topography, soil, and human activity. Bold letters of R2 represent the significant effect of random factor (sampling sites). The meanings of abbreviations are shown in Fig. 3 and Table 2.

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滇中半湿润常绿阔叶林木本植物的功能特征和功能多样性及其影响因子

Plant functional traits, community functional diversity and their environmental determinants of the semi-humid evergreen broad-leaved forest in the Central Yunnan Plateau

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