亚热带森林中环境和物种多样性对灌木存活的影响

doi:10.17520/biods.2022443

生物多样性 ›› 2023, Vol. 31 ›› Issue (3): 22443. DOI: 10.17520/biods.2022443

• 研究报告: 植物多样性 • 上一篇下一篇

亚热带森林中环境和物种多样性对灌木存活的影响

薛玉洁¹^,², 程安鹏², 李珊², 刘晓娟², 李景文¹^,^*()

1.北京林业大学生态与自然保护学院, 北京 100083
2.中国科学院植物研究所浙江钱江源森林生物多样性国家野外科学观测研究站, 北京 100093

收稿日期:2022-08-03 接受日期:2022-09-20 出版日期:2023-03-20 发布日期:2022-11-11
通讯作者: 李景文
作者简介:* E-mail: lijingwen@bjfu.edu.cn
基金资助:
国家重点研发计划(2022YFF0802300);中国科学院先导科技专项(B类)(XDB31000000)

The effects of environment and species diversity on shrub survival in subtropical forests

Yujie Xue¹^,², Anpeng Cheng², Shan Li², Xiaojuan Liu², Jingwen Li¹^,^*()

1 School of Ecology and Nature Conservation, Beijing Forestry University, Beijing 100083
2 Zhejiang Qianjiangyuan Forest Biodiversity National Observation and Research Station, Institute of Botany, Chinese Academy of Sciences, Beijing 100093

Received:2022-08-03 Accepted:2022-09-20 Online:2023-03-20 Published:2022-11-11
Contact: Jingwen Li

摘要/Abstract

摘要：

灌木是森林生态系统的重要组成部分, 在维持物种多样性、促进养分循环、保护幼苗更新以及促进营养级相互作用等方面具有重要作用, 其存活情况对森林群落组成和生态系统功能具有重要意义。但亚热带森林中灌木的存活情况及其影响因素仍有待探究。本研究依托于中国亚热带森林生物多样性与生态系统功能实验研究平台(BEF-China), 利用105个不同乔木与灌木多样性相嵌套样方中2012年和2019年的灌木个体存活数据, 以及样方水平的土壤养分数据、地形数据、物种多样性数据和物种水平的功能性状数据, 通过构建一系列广义线型混合模型(GLMM)探讨了非生物因素、生物因素以及灌木功能性状对灌木存活的影响。结果表明: 不同灌木物种的存活率具有显著差异。坡度较小的山谷、山鞍部位以及土壤碳氮比较高的环境更有利于灌木存活; 乔木和灌木多样性对灌木存活影响有限, 但乔木多样性的增加可通过形成更郁闭的林冠提高灌木存活; 灌木功能性状显著影响灌木存活, 且倾向于保守型生活策略(即较小的比叶面积、叶片氮含量、气孔大小, 较高的叶干物质含量、木质密度)的灌木物种具有更易于存活。通过模型比较发现, 包含所有因素的全模型对灌木存活的解释度最高, 不同影响因素的贡献率由大到小依次为非生物因素、灌木功能性状以及生物因素。本研究表明灌木存活受到多方面因素的共同影响, 尽管土壤和地形等非生物因素的变化影响了灌木的存活, 但不同灌木由于功能性状间的差异同样形成了不同的存活表现, 而物种多样性的直接影响可能需要更长时间的监测才能更加显著。

关键词: BEF-China, 功能性状, 灌木存活, 亚热带森林, 乔木多样性

Abstract

Aims: Shrubs are a vital component of forest ecosystems, playing an important role in maintaining species diversity, promoting nutrient cycling, protecting seedling regeneration, and fostering multi-trophic interactions. Shrub survival is an integral factor in forest community composition and function. However, previous studies lack insights on factors influencing shrub survival. Here, we examed the influence of different factors on shrub survival in order to understand the mechanisms underlying shrub survival.

Methods: The study examined 105 plots with varying tree and shrub species diversity in the Biodiversity-Ecosystem Functioning Experiment China platform (BEF-China). By collecting shrub survival in 2012 and 2019, topographical variables, soil variables, tree and shrub species richness in plot level, associated with shrub functional traits in species level, we analyzed the effects of abiotic factors, biotic factors and functional traits on shrub survival using the generalized linear mixed model (GLMM).

Results: We found significant differences in the survival rates among 10 shrub species. Shrubs tended to have higher survival rates in valleys or gently sloping saddles, and in plots with high soil C/N ratio. Neither tree nor shrub diversity had a significant effect on shrub survival, but tree diversity could indirectly improve shrub survival by forming a denser canopy. Shrub functional traits generally significantly affected their survival. For example, shrub species with conservative traits like low specific leaf area, low leaf nitrogen content, small stomatal area, high leaf dry matter content, and high wood density survived better. Abiotic factors had the largest impact on shrub survival, followed by shrub functional traits and then biotic factors. However, the best model for shrub survival included all the factors studied.

Conclusion: The survival of shrubs is the joint result of many factors. Abiotic factors play an important role and differing functional traits also influence shrub survival. However, the impact of species diversity on shrub survival may need a longer window of observation.

Key words: BEF-China, functional trait, shrub survival, subtropical forest, tree species diversity

薛玉洁, 程安鹏, 李珊, 刘晓娟, 李景文 (2023) 亚热带森林中环境和物种多样性对灌木存活的影响. 生物多样性, 31, 22443. DOI: 10.17520/biods.2022443.

Yujie Xue, Anpeng Cheng, Shan Li, Xiaojuan Liu, Jingwen Li (2023) The effects of environment and species diversity on shrub survival in subtropical forests. Biodiversity Science, 31, 22443. DOI: 10.17520/biods.2022443.

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链接本文: https://www.biodiversity-science.net/CN/10.17520/biods.2022443

https://www.biodiversity-science.net/CN/Y2023/V31/I3/22443

图/表 6

图1 BEF-China样地A中灌木种植设计示意图。(a)样地中灌木种植样方分布, 共计35个4亩大样方; (b)每个4亩大样方内分别设置灌木多样性为2、4、8的1亩样方; (c) 1亩样方中灌木和乔木种植位置示意图; (d)不同灌木多样性样方的林下景观。

Fig. 1 Design of shrub planting in BEF-China Site A. (a) Overview of shrub plots in Site A, in total 35 super-plots; (b) In each super-plot, shrubs were planted into the three 1-mu plots with richness levels of 2, 4 and 8 shrub species, respectively; (c) Scheme of shrub planting positions in relation to the tree planting positions; (d) Different views in plots with different shrub diversity.

图2 不同物种(a)或演替阶段(b)灌木存活率的变化。不同字母表示不同灌木物种间存活率差异显著(P < 0.05, Tukey检验)。EuMy: 大果卫矛; ArCr: 朱砂根; LoCh: 檵木; RhIn: 石斑木; EuMu: 格药柃; SyBu: 赤楠; CaCh: 浙江红山茶; GaJa: 栀子; DiMy: 杨梅叶蚊母树; DiBu: 小叶蚊母树; E表示在自然演替群落中, 该灌木出现于演替早期群落; E/L表示在自然演替中该灌木从早期到后期一直存在; I/L表示该灌木在演替中期到演替后期出现; L表示该灌木仅在演替后期出现。

Fig. 2 Survival rates of different shrub species (a) or different successional stages (b). Different letters show statistically significant differences according to a Tukey test at 0.05 level. EuMy, Euonymus myrianthus; ArCr, Ardisia crenata; LoCh, Loropetalum chinense; RhIn, Rhaphiolepis indica; EuMu, Eurya muricata; SyBu, Syzygium buxifolium; CaCh, Camellia chekiangoleosa; GaJa, Gardenia jasminoides; DiMy, Distylium myricoides; DiBu, D. buxifolium. E, Shrub occurs in early-successional stages; E/L, Shrub occurs in all successional stages; I/L, Shrub occurs in intermediate- and late-successional stages; L, Shrub occurs in late-successional stages.

图3 非生物因子(a)、生物因子(b)以及功能性状(c)对灌木存活的相对影响。Soil pH: 土壤pH值; Soil N: 土壤氮含量; Soil C: 土壤碳含量; Soil C/N: 土壤碳氮比; topo.PC1-4: 地形因素第一至第四主成分; logTR: 乔木多样性(以2为底的对数); logSR: 灌木多样性(以2为底的对数); logTR:logSR: logTR和logSR的交互效应; Canopy cover: 林冠盖度; trait.PC1-4: 功能性状第一至第四主成分。• P < 0.1; * P < 0.05; *** P < 0.001; 空心圆代表效应不显著(P > 0.1)。

Fig. 3 Relative effect of abiotic factors (a), biotic factors (b) and shrub functional traits (c) on shrub survival. Soil N, Soil nitrogen content; Soil C, Soil carbon content; Soil C/N, Soil carbon to nitrogen ratio; topo.PC1-4, The first, second, third, and fourth principal component of topography, respectively; logTR, Tree richness (log2-transformed contrast); logSR, Shrub richness (log2-transformed contrast); logTR:logSR, Interaction of logTR and logSR; Canopy cover, The canopy cover degree of tree; trait.PC1-4, The first, second, third, and fourth principal component of traits, respectively. • P < 0.1; * P < 0.05; *** P < 0.001. Unfilled circles indicate insignificant effects (P > 0.1).

图4 灌木功能性状主成分分析(PCA)结果。SA: 气孔面积; LA: 叶面积; Chl: 叶绿素含量; Thickness: 叶片厚度; LDMC: 叶干物质含量; Height: 株高; Crown area: 冠幅; WD: 木质密度; SD: 气孔密度; LC: 叶片碳含量; SLA: 比叶面积; LN: 叶片氮含量; EuMy、ArCr、LoCh、RhIn、EuMu、SyBu、CaCh、GaJa、DiMy、DiBu含义见图2。

Fig. 4 Principal component analysis (PCA) of shrub functional traits. SA, Stomatal area; LA, Leaf area; Chl, Leaf chlorophyll content; Thickness, Leaf thickness; LDMC, Leaf dry matter content; Height, Shrub height; Crown area, Shrub crown area; WD, Wood density; SD, Stomatal density; LC, Leaf carbon content; SLA, Specific leaf area; LN, Leaf nitrogen content; EuMy, ArCr, LoCh, RhIn, EuMu, SyBu, CaCh, GaJa, DiMy, and DiBu see Fig. 2.

表1 不同灌木存活模型的AIC值以及R2

Table 1 AIC values and R2 of different models for shrub survival

模型 Models	AIC	ΔAIC	边际决定系数 Marginal R²	条件决定系数 Conditional R²
非生物模型 Abiotic model	44,022.1	19.9	0.028	0.424
生物模型 Biotic model	44,051.1	48.9	0.005	0.426
功能性状模型 Trait model	44,041.2	39	0.261	0.409
全模型 Full model	44,002.2	0	0.306	0.408

图5 全模型中各类因子对灌木存活的相对影响。biotic: 生物因素; abiotic: 环境因素; trait: 功能性状; trait.PC1、trait.PC2、trait.PC3、trait.PC4、Soil pH、Soil C、Soil N、Soil C/N、topo.PC1、topo.PC2、topo.PC3、topo.PC4、logTR、logSR、 logTR:logSR、Canopy cover含义见图3。? P < 0.1; * P < 0.05; ** P < 0.01; *** P < 0.001。空心圆代表效应不显著(P > 0.1)。

Fig. 5 Parameter estimates (standardized regression coefficients) and relative importance of fixed factors. trait, Functional traits; abiotic, Abiotic factors; biotic, Biotic factors; trait.PC1, trait.PC2, trait.PC3, trait.PC4, Soil pH, Soil C, Soil N, Soil C/N, topo.PC1, topo.PC2, topo.PC3, topo.PC4, logTR, logSR, logTR:logSR and Canopy cover see Fig. 3. ? P < 0.1; * P < 0.05; ** P < 0.01; *** P < 0.001. Unfilled circles indicate insignificant effects (P > 0.1).

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亚热带森林中环境和物种多样性对灌木存活的影响

The effects of environment and species diversity on shrub survival in subtropical forests

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