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人类活动干扰的日益加剧和极端气候变化的异常波动,导致全球生物多样性正以前所未有的速度锐减,影响陆地生态系统的多种生态功能和服务的可持续发展[1-2]。已有研究发现,生物多样性的减少在植物群落中不仅表现为植物物种数量和多样性的减少,还会影响植物功能性状组成、多度和分布范围[3],而植物物种响应外界环境的变化和对植物群落的贡献主要取决于物种所具有的功能性状[4]。因此,生物多样性的减少可能会降低植物功能多样性,从而对植物群落在生态系统功能和服务的作用产生不利影响。
植物功能多样性是指能够影响生态系统功能特征的数值、范围和空间分布[5],它能够将物种功能性状与生态系统功能有机结合起来,通过物种的不同功能性状来解释群落差异,揭示各个物种在生态系统中所起的功能与作用[6]。同时,功能多样性能够反映群落内物种组成、生态位可利用程度以及植物群落中冗余种和种间互补作用,能够更准确地预测植物群落结构的稳定性和抵抗外界干扰的能力[7-8]。因此,开展植物功能多样性的研究对了解生物多样性与生态系统功能之间的关系有着重要意义[9]。
植物生长型代表了植物与生境相关的关键生态策略,体现了植物对外界环境适应性的表征形态[10]。其中,同一生长型的物种,具有表型相似、适应环境条件相似的特点[10]。研究表明,乔木、灌木和藤本植物等生长型与热带森林的结构和功能有着密切的联系[11]。Ali等研究发现乔木层的树冠互补性能够通过对大尺度生态梯度上的光捕获和利用来介导功能多样性对地上生物量的积极影响,从而增加热带天然林的地上生物量[12]。而藤本植物能够调节热带森林的间隙动态、水分关系和冠层间的连通性,并通过与树木竞争各种可利用性资源来影响树木的生长、补充和死亡率,从而影响热带森林的群落结构和物种组成[13-15]。同样,由于灌木矮小耐阴,竞争力较强且死亡率较低,灌木在维持热带雨林多样性和植被动态方面也发挥着重要作用,但在外界环境发生变化时,灌木可能是热带雨林生物多样性中最先受到负面影响的组成部分[16]。
目前,对于植物功能多样性的研究主要集中于高寒草甸[17-18]、温带森林[19]以及亚热带森林[20]等生态系统,而基于热带雨林的研究主要偏重于不同演替阶段植物功能多样性的变化规律[21]。然而,对于热带雨林不同生长型木本植物的功能多样性变化的研究相对较少。
热带雨林是陆地生态系统极为重要的组成部分,仅占全球陆地面积的7%,却维持着世界上超过60%的生物多样性[22-23]。然而,由于气候变化、商业采伐以及自然火灾等因素的影响,热带雨林生境日益破碎化,面积大幅减少。如今,热带雨林已成为高生产力但十分脆弱的生态系统之一[24]。西双版纳热带雨林是我国典型的热带雨林之一,生物多样性丰富,林分结构复杂,在生物多样性保护、全球碳平衡维持和气候调节等方面中发挥着重要作用。自上世纪50年代以来,大规模的天然林被用于橡胶林种植,导致热带雨林面积不断减少,生境破碎化严重[25-26],而西双版纳热带雨林生物多样性的维持机制一直是生态学领域研究的热点,如Asefa等在西双版纳热带森林动态监测样地中利用四种不同的空间点过程模型来估计扩散限制和环境过滤的相对重要性,结果发现环境过滤驱动植物群落远亲物种之间的系统发育和功能多样性,而扩散限制单独和环境过滤共同作用于最邻近的系统发育和功能多样性[27];Yang等在哀牢山和西双版纳这两个森林动态监测样地中使用系统发育和功能多样性方法进行研究,发现环境过滤在哀牢山样地植物群落结构和西双版纳样地生物群落相互作用中发挥了关键作用,其中西双版纳热带雨林的系统发育和功能分散沿地形梯度呈过度分散[28]。周昌艳等以云南西双版纳热带季节雨林20 ha动态监测样地为研究对象,证实了林冠结构驱动局域尺度木本植物功能性状beta多样性格局的形成[29]。这些研究主要是基于热带雨林植物群落整体性或单一生活型木本植物群落,而对于热带雨林不同生活型木本植物功能多样性的研究还鲜有人涉及。因此,本文通过分析西双版纳热带雨林不同生活型木本植物功能性状的差异,揭示热带雨林不同生长型木本植物功能多样性的变化规律以及影响因素,以期为西双版纳热带雨林不同生长型木本植物群落的物种多样性的保护和恢复策略的制定提供理论基础和数据支持。
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利用Pearson相关分析法综合分析了西双版纳热带雨林木本植物功能性状间的相关关系。结果显示,木本植物的大多数功能性状之间存在显著的相关关系(图2)。其中,LC与LDMC呈显著正相关,与SLA、LP呈显著负相关;LDMC与SLA、LP、LN呈显著负相关;WD与LP、LN呈显著负相关;SLA与LP、LN呈显著正相关;LP与LN呈显著正相关。
图 2 西双版纳热带雨林木本植物功能性状间的相关性分析
Figure 2. Correlation analysis of functional traits of woody plants in a tropical rainforest of Xishuangbanna.
不同生长型木本植物的功能性状群落加权平均值之间也存在一定的差异(图3)。其中,乔木的LA、LC、LDMC的群落加权平均值均显著高于灌木和藤本,而LP和SLA的群落加权平均值均显著低于灌木和藤本;灌木的WD的群落加权平均值最高,而LC的群落加权平均值则显著低于乔木和藤本;藤本的LN和SLA的群落加权平均值最高,但LA和WD的群落加权平均值则最低。
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不同生长型木本植物的功能多样性之间存在显著差异。其中,乔木功能丰富度和功能离散度均显著高于灌木和藤本(图4A和B)。灌木和藤本功能均匀度显著高于乔木(图4C)。灌木功能分异度显著高于乔木和藤本(图4D)。
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普通最小二乘法线性回归分析表明,乔木和藤本功能丰富度以及藤本的功能离散度与物种丰富度、多度、系统发育多样性呈正相关(图5A,B,D,M,N和P)。乔木功能均匀度和功能分异度与土壤肥力呈正相关(图5G和K),而藤本功能均匀度与物种丰富度、系统发育多样性呈显著正相关(图5E和H),但藤本功能分异度与物种丰富度、植物系统发育多样性呈负相关(图5I和L),与土壤肥力呈正相关(图5K)。此外,灌木功能离散度与物种丰富度呈正相关(图5M)。
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多元回归分析表明,乔木、灌木和藤本功能多样性与物种丰富度、多度、系统发育多样性和土壤肥力的解释率在0.014~0.486之间变化(表1)。乔木功能丰富度主要与物种丰富度、土壤肥力呈显著正效应,与多度呈显著负效应,而土壤肥力对藤本的功能分异度有显著正效应。灌木功能多样性与物种丰富度、多度、系统发育多样性和土壤肥力均无显著相关性。
表 1 以功能多样性指标作为不同生长型木本植物的响应变量的多元回归分析
Table 1. Multiple regression analysis using functional diversity index as response variable of different growth forms of woody plants
功能丰富度
Functional richness (FRic)功能均匀度
Functional evenness (FEve)功能离散度
Functional dispersion (FDis)功能分异度
Functional divergence (FDiv)乔木
Tree灌木
Shrub藤本
Liana乔木
Tree灌木
Shrub藤本
Liana乔木
Tree灌木
Shrub藤本
Liana乔木
Tree灌木
Shrub藤本
Liana物种丰富度
Species richness1.274** 1.366 −0.098 0.015 0.929 −0.056 0.085 0.537 −0.002 0.581 0.672 −0.036 多度
Abundance−0.362* −0.447 −0.094 0.269 −0.200 −0.160 −0.379 −0.165 −0.063 0.153 0.070 0.171 土壤肥力
Soil fertility0.202* −0.176 −0.003 0.251 0.009 −0.029 −0.042 −0.135 0.046 0.173 0.018 0.307* 植物系统发育
多样性
Plant phylogenetic
diversity−0.350 −1.050 0.611 −0.122 −1.045 0.595 0.107 −0.506 −0.010 −0.758 −0.919 −0.411 R2 0.486 0.110 0.151 0.089 0.088 0.143 0.061 0.081 0.014 0.101 0.065 0.187 注:估计值是预测因子的标准化回归系数。R2,由固定效应解释的方差。显著性水平: *, P<0.05; **,P<0.01; ***, P<0.001
Notes: Estimates are standardized regression coefficients for predictors. R2, the variance explained by fixed effects. Significance level: *, P<0.05; **,P<0.01; ***, P<0.001进一步分析了物种丰富度、多度、系统发育多样性及土壤肥力对不同生长型木本植物功能性状的群落加权平均值(CWM)的影响(图6)。RDA结果显示,乔木植物RDA分析中第一轴和第二轴的解释率分别为78.28%和16.02%,前两轴解释94.30%的差异信息。其中,多度对乔木的功能性状群落加权平均值具有显著负相关(P<0.01),而土壤肥力对乔木的功能性状群落加权平均值具有显著正相关(P<0.05),是影响乔木功能性状的两个主要影响因子(图6A)。在藤本植物RDA分析中第一轴和第二轴的解释率分别为73.74%和11.84%,前两轴解释85.58%的差异信息。其中,藤本的功能性状群落加权平均值与土壤肥力呈显著正相关(P<0.01),而与植物系统发育多样性、物种丰富度呈显著负相关(P<0.05)(图6B)。而灌木植物RDA分析中第一轴和第二轴的解释率分别为60.85%和29.32%,前两轴解释90.17%的差异信息。其中,多度对灌木的功能性状群落加权平均值具有显著负相关(P<0.01),是灌木功能性状的主要影响因子(图6C)。
图 6 西双版纳热带雨林乔木(A)、灌木(B)以及藤本(C)功能性状的群落加权平均值(CWM)与物种丰富度、多度、系统发育多样性和土壤肥力的冗余分析(RDA)
Figure 6. Community weighted mean value (CWM) of functional traits and redundancy analysis (RDA) of species richness, abundance, phylogenetic diversity, and soil fertility of trees(A), shrubs(B), and lianas(C) in a tropical rainforest of Xishuangbanna.
西双版纳热带雨林不同生长型木本植物功能多样性
Functional Diversity in Woody Plants of Different Growth Form in theTropical Rainforest of Xishuangbanna
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摘要:
目的 功能多样性能够预测植物群落结构稳定性和抵抗干扰能力,对维持生态系统功能具有重要作用。 方法 本文基于西双版纳国家级自然保护区勐仑、补蚌和尚勇等片区3块1 hm2动态监测样地的调查数据和木本植物功能性状数据,利用单因素方差分析、主成分分析、Pearson相关分析、冗余分析以及多元线性回归分析等方法,分析西双版纳热带雨林不同生长型木本植物功能多样性的差异,阐明热带雨林木本植物群落的功能特征及主要影响因子。 结果 (1)乔木的叶面积、叶碳含量、叶干物质含量的群落加权平均值均显著高于灌木和藤本,而藤本叶氮含量和比叶面积的群落加权平均值显著高于乔木和灌木。(2)乔木功能丰富度和功能离散度高于灌木和藤本,而灌木功能均匀度和功能分异度均高于乔木和藤本。(3)乔木和藤本功能丰富度以及藤本功能离散度随物种丰富度、多度和系统发育多样性的增加而显著增大;乔木功能均匀度和功能分异度随土壤肥力的增大而显著增大,而灌木功能离散度随物种丰富度的增加而增大。(4)植物多度是影响乔木和灌木功能性状群落加权平均值的主要因素,而系统发育多样性、物种丰富度和土壤肥力是影响藤本功能性状群落加权平均值的主要因素。 结论 本研究表明西双版纳热带雨林不同生长型木本植物群落在功能性状和功能多样性之间存在显著差异性,且植物多度和土壤肥力在热带雨林植物群落功能多样性维持机制中起到重要作用。 Abstract:Aims Functional diversity can predict the stability of plant community structure and the ability to resist disturbance, which plays an important role in sustaining ecosystem functioning. Methods Based on the survey data and the functional traits of woody plants from three 1 hm2 dynamic monitoring plots in Menglun, Bupung, and Shangyong in Xishuangbanna Nature Reserve, China, one-way variance analysis, principal component analysis, Pearson correlation analysis, redundancy analysis, and multiple linear regression analysis were used to analyze the changes and influencing factors of functional diversity among different growth form of woody plants, and to discuss the functional characteristics and main influencing factors of woody plant communities in tropical rainforests in Xishuangbanna. Results (1) The community weighted mean value of leaf area, leaf carbon content and leaf dry matter content of trees were significantly higher than those of shrubs and lianas, while the community weighted mean value of leaf nitrogen content and specific leaf area of lianas were significantly higher than those of trees and shrubs. (2) The functional richness and functional dispersion of trees were higher than those of shrubs and lianas, while the functional evenness and functional divergence of shrubs were higher than those of trees and lianas. (3) The functional richness of trees and lianas and the functional dispersion of lianas increased significantly with increasing plant species richness, abundance and phylogenetic diversity. The functional evenness and functional divergence of trees increased significantly with increasing of soil fertility, while the functional dispersion of shrubs increased with increasing plant species richness. (4) Abundance was the main factor affecting the community weighted mean value of functional traits of trees and shrubs, while phylogenetic diversity, plant species richness and soil fertility were the main factors affecting the community weighted mean value of functional traits of lianas. Conclusion This study reveals that functional traits and functional diversity have significant differences among different growth form of woody plant communities in tropical rainforests in Xishuangbanna. Moreover, woody plant abundance and soil fertility play a crucial role in maintaining the functional diversity of plant communities in tropical rainforests in maintaining plant community functional diversity. -
Key words:
- functional diversity
- / soil fertility
- / abundance
- / tropical rainforest
- / growth form
- / woody plant
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图 6 西双版纳热带雨林乔木(A)、灌木(B)以及藤本(C)功能性状的群落加权平均值(CWM)与物种丰富度、多度、系统发育多样性和土壤肥力的冗余分析(RDA)
Figure 6. Community weighted mean value (CWM) of functional traits and redundancy analysis (RDA) of species richness, abundance, phylogenetic diversity, and soil fertility of trees(A), shrubs(B), and lianas(C) in a tropical rainforest of Xishuangbanna.
表 1 以功能多样性指标作为不同生长型木本植物的响应变量的多元回归分析
Table 1. Multiple regression analysis using functional diversity index as response variable of different growth forms of woody plants
功能丰富度
Functional richness (FRic)功能均匀度
Functional evenness (FEve)功能离散度
Functional dispersion (FDis)功能分异度
Functional divergence (FDiv)乔木
Tree灌木
Shrub藤本
Liana乔木
Tree灌木
Shrub藤本
Liana乔木
Tree灌木
Shrub藤本
Liana乔木
Tree灌木
Shrub藤本
Liana物种丰富度
Species richness1.274** 1.366 −0.098 0.015 0.929 −0.056 0.085 0.537 −0.002 0.581 0.672 −0.036 多度
Abundance−0.362* −0.447 −0.094 0.269 −0.200 −0.160 −0.379 −0.165 −0.063 0.153 0.070 0.171 土壤肥力
Soil fertility0.202* −0.176 −0.003 0.251 0.009 −0.029 −0.042 −0.135 0.046 0.173 0.018 0.307* 植物系统发育
多样性
Plant phylogenetic
diversity−0.350 −1.050 0.611 −0.122 −1.045 0.595 0.107 −0.506 −0.010 −0.758 −0.919 −0.411 R2 0.486 0.110 0.151 0.089 0.088 0.143 0.061 0.081 0.014 0.101 0.065 0.187 注:估计值是预测因子的标准化回归系数。R2,由固定效应解释的方差。显著性水平: *, P<0.05; **,P<0.01; ***, P<0.001
Notes: Estimates are standardized regression coefficients for predictors. R2, the variance explained by fixed effects. Significance level: *, P<0.05; **,P<0.01; ***, P<0.001 -
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