海南岛热带低地雨林自然恢复过程中木本植物幼苗功能性状及增补动态
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摘要
物种多样性维持机制是群落生态学研究中的一个核心问题。幼苗库是热带森林生态系统的一个基本的组成部分，在群落动态、多样性维持和生态系统运行中发挥着重要作用。然而目前热带林的群落生态学研究主要以胸径(DBH)1cm以上的个体为对象，而对DBH<1cm的幼苗关注较少。幼苗性状对成年树木在生态系统中的地位和作用可能具有决定性的影响，相对于热带树木成年个体，幼苗的功能性状更容易准确获取。本文以海南岛不同恢复阶段的热带低地雨林(刀耕火种后自然恢复30年，60年的次生林和老龄林)为对象，研究了不同生长型木本植物(乔木、灌木和木质藤本)幼苗功能性状随自然恢复过程的变化趋势及其影响因素；通过比较不同恢复阶段幼苗与成年树群落的物种组成及多样性特征，揭示了不同恢复阶段热带低地雨林幼苗的增补动态；通过优势种幼苗移栽试验，分析了幼苗在不同群落中建立的可能性和限制性因子。主要研究结果如下：
1.与老龄林中的优势种幼苗相比，刀耕火种后自然恢复30年和60年次生林群落中的优势种幼苗具有更高的比叶面积、比茎长和比根长，并在两个恢复阶段间没有显著性差异；随着恢复时间的增加，叶干物质比例显著增加，而根干物质比例随恢复逐渐降低，茎干物质比例则在3个恢复阶段没有显著差异；基于幼苗功能性状，应用PCA分析可以将不同恢复阶段的幼苗分成不同的功能群组，老龄林中的物种比次生林中的物种聚集的更为紧密。
2.环境因子随恢复时间的增加而发生变化，土壤含水量在30年次生林中显著低于60年次生林和老龄林，冠层开阔度和土壤养分含量随着恢复时间的增加逐渐降低；幼苗的功能性状与环境因子的相关性随自然恢复过程而发生相应的变化，在恢复30年的次生林中，幼苗的多数功能性状都与土壤含水量密切相关，但到了60年的次生林和老龄林阶段，幼苗的所有功能性状均与其不相关；在恢复30年和60年的次生林中，幼苗的所有功能性状与冠层开阔度均不相关，但在老龄林中比叶面积、叶干物质比例和茎干物质比例都与其显著相关；幼苗功能性状与土壤养分含量在各个恢复阶段中都显著相关。
3.在各恢复阶段的群落中，功能性状随生长型和幼苗生长阶段(高度级)而发生显著变化。幼苗通过调节各功能性状之间的权衡来适应群落环境的变化，乔木和灌木幼苗通过低的比叶面积、低的叶干物质比例和高的茎干物质比例来适应群落下的低光环境，而藤本植物则通过相反的策略实现其在热带低地雨林中的生存。随着幼苗的生长，乔木和灌木植物的幼苗减少对同化器官的投入，增加支撑器官的投入；藤本植物幼苗对同化器官的投入增大，减少了对支撑器官的投入，这符合藤本植物依附于其他植物生长的特性。
4.在各恢复阶段的群落中，幼苗层和成年树(DBH≥7.5cm)的功能性状(比叶面积、叶干物质含量和比茎密度)存在显著差异，幼苗层的叶干物质含量和比茎密度与成年树之间存在线性正相关关系；幼苗层和成年树的比叶面积随恢复阶段呈降低趋势，叶干物质含量和比茎密度均随恢复时间的增加逐渐升高；在次生林中，幼苗层和成年树层功能性状与环境之间的关系基本一致，老龄林中，幼苗层、成年树层功能性状与环境之间的关系有差异。在海南岛热带低地雨林中，虽然幼苗功能性状并不能完全取代成年树的功能性状，但可根据幼苗层的功能性状推测成年树功能性状随自然恢复的变化趋势，在次生林中可以根据幼苗层的功能性状推测成年树功能性状对环境因子的响应规律。
5.随着恢复时间的增加，幼苗的种―面积和种―个体累积速度都在增加，老龄林的累积速度显著高于次生林；群落内幼苗多度及丰富度呈先增大后降低的趋势；次生林与老龄林的幼苗层组成的相似性也逐渐增加；幼苗物种丰富度与不同环境之间的关系随恢复阶段而变化：恢复30年次生林幼苗丰富度与土壤含水量、全氮、全磷含量显著相关，恢复60年次生林幼苗丰富度与冠层开阔度、土壤水分、土壤钾含量显著相关，而老龄林丰富度与冠层开阔度、有效磷、有效钾显著相关；幼苗多度和丰富度随以高度级划分的生长阶段的增加呈逐渐递减的趋势。
6.对不同恢复阶段的幼苗物种组成异质性和环境因子进行冗余分析，结果显示：在30年和60年次生林中，环境因子对幼苗层物种组成异质性的解释方差分别为55.1%和53.5%，而在老龄林中环境因子的解释方差较低，为43.4%。不同恢复阶段幼苗样方分布与环境因子之间的关系不同，恢复30年次生林的幼苗分布主要受土壤含水量和冠层开阔度的影响；恢复60年次生林的幼苗分布主要受土壤养分含量的影响。影响老龄林幼苗分布的环境因子主要是土壤含水量和全磷含量。
7.刀耕火种后自然恢复30年、60年的次生林以及老龄林幼苗的物种丰富度分别占整个群落物种丰富度的63.5%、60.6%和54.9%，说明海南岛热带低地雨林幼苗库是生物多样性中一个重要的组成部分。不同恢复阶段的幼苗(DBH<1cm)、幼树(1cm≤DBH<7.5cm)和成年树(DBH≥7.5cm)之间的补充比例不同，恢复30年次生林分别有71.9%和39.7%物种的幼苗能够进入幼树和成年树层，而恢复60年次生林分别有65.0%和38.0%物种的幼苗能够进入幼树和成年树层，老龄林分别有74.9%和50.9%幼苗进入到幼树和成年树层。
8.以热带低地雨林中分布的4个落叶物种山乌桕(Sapium discolor)、厚皮树(Lanneacoromandelica)、野漆树(Toxicodendron succedaneum)、银珠(Peltophorum tonkinense)和4个常绿物种：米槠(Castanopsis carlesi)、芳槁润楠(Machilus suaveolens)、乌墨(Syzygiumcumini)青梅(Vatica mangachapoi)为研究对象，分别进行4种改变地上和地下竞争的处理(移除地上植被、挖沟、移除地上植被并挖沟和对照)试验，分析了热带低地雨林幼苗更新与生长的影响因素。结果表明：移除地上植被和移除地上植被并挖沟能够显著增加幼苗的相对生长速率，而挖沟对移栽幼苗的相对生长速率影响不显著；落叶树种的幼苗生长速率明显高于常绿树种；幼苗相对生长速率随着光合有效辐射的增大逐渐升高。我们的研究结果表明，光照(地上竞争)是限制海南岛热带低地雨林幼苗生长的最主要因子，随着光合有效辐射的增加，幼苗的相对生长速率显著增加。
Understanding mechanism of species diversity maintenance has been one of the centralissues in community ecology. The seedling bank is a basic component in tropical forestecosystems, which play important roles in community dynamics, diversity maintenance andecosytem functioning.However, the seedlings with a diameter of less than1cm are usuallyneglected. However, seedling functional traits are often strongly related to the performance andfunction of adult plants in an ecosystem. Measurements of seedling traits are easy and accuratecompared to measurements of adult traits.In this thesis,we investigated the diversity andfucntional traits of woody plant(including trees,shrubs and lianas) seedlings in a tropicallowland rain forest of different recovery stages(30years and60years after shifting cultivationand old growth forest) on Hainan island, South China. The variation of seedling functionaltraits of woody plants during the recovery process and its influencing factors were explored.Seedling recruitment dynamics were analyzed by comparing the composition,abundance anddiversity of seedling and tree communities in each recovery stages of the tropical lowlandrainforest. The factors influencing the relative growth rate of8representative speceis of treeseedlings in the tropical lowland rain forest were examined through field experiments. Themain results are as follows.
1. Seedlings in30-year and60-year secondary forests had higher specific leaf area,specific stem length and specific root length. Leaf mass fraction increased significantly, whileroot mass fraction reduced gradually during the recovery process. The seedlings can beaggregated into3functional groups by ordination of Principal compoent analysis The speciesin old growth forest clusted more closely than those in the secondary forests.
2. Environmental factors changed significantly during the recovery process. Soil watercontent were significantly low in30-year forest, while there was no significant difference in60-year forest and old growth forest. Canopy openness and soil nutrient contents decreased withrecovery time. The correlations between seedling functional traits and environmental factors changed correspondingly with the natural recovery process. Most of the seedlings functionaltraits were significantly correlated with soil water content in the30-year forest but nocorrelations existed in the60-year and old growth forests. Seedling functional traits were notcorrelated with canopy openness in the30-year and60-year secondary forests while hadsignificant correaltions in the old growth forest. Seedling functional traits was correlated withsoil nutrient contents in each recovery stage.
3. The functional traits varied significantly with life form and growth stage (height class).Seedlings adapte to the community environment by making the trade-offs between differentfunctional traits. Tree and shrub seedlings adapt to the stressful rainforest understoryenvironment (such as low light) by decreasing specific leaf area and increasing stem massfraction. However, lianas adapt to the environment through the opposite strategy. With thegrowth of seedlings, trees and shrubs reduced the investment to assimilation, and increased theinvestment to support organ. Lianas increased investment to assimilation organs, and reducedto support organs. This is consistent with characteristics of liana attanching to other plants togrowth.
4. Functional traits of seedlings were significantly different from those of the adulttrees(DBH>7.5cm) in each recovery stage. Leaf dry mass content and specific stem density ofseedlings and adult trees were positively correlated. Specific leaf area of both seedlings andadult trees decreased, while leaf dry mass content and specific stem density increased duringthe recovery process. The environmental factors that affected functional traits of both seedlingsand trees were same to a certain extent in secondary forests, but different in old growthforest.These above results suggest that the patterns of change in functional traits and theirrelations with the environmental conditions for adult trees can be indicated in certain degree bythose of the the seedlings in this lowland rain forest.
5. The species-area and species-individual cumulative rates of seedlings increased withthe recovery time. Species accumulation rates in old growth forest were faster than those in thesecondary forests. The seedling abundance and richness first increased and then decreased withrecovery time, which was in line with the intermediate disturbance hypothesis. Species richness of seedlings was correlated with soil water content, total nitrogen and total phosphoruscontent in the30-year forest,while it was correlated with canopy openness, soil water contentand potassium content in the60-year forest; and it was correlated with canopy openness,available phosphorus, and available potassium in the old growth forest. Seedling abundanceand richness showed a gradually decreasing trend with increasing growth stage (height class).
6. The relationships between seedling species composition heterogeneity andenvironmental factors in three successional stages were assessed using redundancy analysis.The variance of environmental factors to interpret heterogeneity of seedlings were55.1%and53.5%respectively in30-year and60-year secondary forests, while lower (43.4%) in oldgrowth forest. The relationship between distribution of seedlings and environmental factors inthree recovery stages were different. Distribution of seedligns in30-year forest was mainlyaffected by soil water content and canopy openness. Distribution of seedligns in60-year forestwas mainly affected by soil nutrient content. Distribution of seedligns in old growth forest wasmainly affected by soil water content and total phosphorus.
7. The species richness of seedlings in30-year forest,60-year forest and old growthforest.accounted for63.5%,60.6%and54.9%of those for the whole coummunity respectively.This result showed that seedling bank is an important component, which should not beneglected in the study of biodiversity in this lowland rain forest. The recruitment rates fromseedlings (DBH<1cm) to saplings (1cm≤DBH<7.5cm) and from seedlings to adult trees(DBH≥7.5cm) were71.9%and39.7%respectively in30-year forest, while those were65.0%and38.0%respectively in60-year forest. In old growth forest,74.9%and50.9%of seedlingspecies grew into saplings and adult trees.
8. Eight representative species (Sapium discolor, Lannea coromandelica, Toxicodendronsuccedaneum, Peltophorum tonkinense, Castanopsis carlesi, Machilus suaveolens, Syzygiumcumini, Vatica mangachapoi) in the tropical lowland rain forest were selected to do thefollowing treatments: removal of the above vegetation (R), trenching (T), removal of the abovevegetation and trenching (R+T), and controlled (C). We measured the relative growth rate ofthese seelings to assess the factors affecting regeneration and growth of the seedlings. We found that the treatments of R and R+T significantly increased the growth rate of seedlings.But the effect of T was not significant. The growth rate of decidous species was higher thanevergreen species. Seedling growth rate increased with the increase of photosyntheticallyactive radiation. The relationship between them could be described by a linear equation. Allthese results showed that light was the most important factor for seedling growth in the tropicallowland rain forest. The below-ground competition had no significant effects on the growth ofseedlings.
1.与老龄林中的优势种幼苗相比，刀耕火种后自然恢复30年和60年次生林群落中的优势种幼苗具有更高的比叶面积、比茎长和比根长，并在两个恢复阶段间没有显著性差异；随着恢复时间的增加，叶干物质比例显著增加，而根干物质比例随恢复逐渐降低，茎干物质比例则在3个恢复阶段没有显著差异；基于幼苗功能性状，应用PCA分析可以将不同恢复阶段的幼苗分成不同的功能群组，老龄林中的物种比次生林中的物种聚集的更为紧密。
2.环境因子随恢复时间的增加而发生变化，土壤含水量在30年次生林中显著低于60年次生林和老龄林，冠层开阔度和土壤养分含量随着恢复时间的增加逐渐降低；幼苗的功能性状与环境因子的相关性随自然恢复过程而发生相应的变化，在恢复30年的次生林中，幼苗的多数功能性状都与土壤含水量密切相关，但到了60年的次生林和老龄林阶段，幼苗的所有功能性状均与其不相关；在恢复30年和60年的次生林中，幼苗的所有功能性状与冠层开阔度均不相关，但在老龄林中比叶面积、叶干物质比例和茎干物质比例都与其显著相关；幼苗功能性状与土壤养分含量在各个恢复阶段中都显著相关。
3.在各恢复阶段的群落中，功能性状随生长型和幼苗生长阶段(高度级)而发生显著变化。幼苗通过调节各功能性状之间的权衡来适应群落环境的变化，乔木和灌木幼苗通过低的比叶面积、低的叶干物质比例和高的茎干物质比例来适应群落下的低光环境，而藤本植物则通过相反的策略实现其在热带低地雨林中的生存。随着幼苗的生长，乔木和灌木植物的幼苗减少对同化器官的投入，增加支撑器官的投入；藤本植物幼苗对同化器官的投入增大，减少了对支撑器官的投入，这符合藤本植物依附于其他植物生长的特性。
4.在各恢复阶段的群落中，幼苗层和成年树(DBH≥7.5cm)的功能性状(比叶面积、叶干物质含量和比茎密度)存在显著差异，幼苗层的叶干物质含量和比茎密度与成年树之间存在线性正相关关系；幼苗层和成年树的比叶面积随恢复阶段呈降低趋势，叶干物质含量和比茎密度均随恢复时间的增加逐渐升高；在次生林中，幼苗层和成年树层功能性状与环境之间的关系基本一致，老龄林中，幼苗层、成年树层功能性状与环境之间的关系有差异。在海南岛热带低地雨林中，虽然幼苗功能性状并不能完全取代成年树的功能性状，但可根据幼苗层的功能性状推测成年树功能性状随自然恢复的变化趋势，在次生林中可以根据幼苗层的功能性状推测成年树功能性状对环境因子的响应规律。
5.随着恢复时间的增加，幼苗的种―面积和种―个体累积速度都在增加，老龄林的累积速度显著高于次生林；群落内幼苗多度及丰富度呈先增大后降低的趋势；次生林与老龄林的幼苗层组成的相似性也逐渐增加；幼苗物种丰富度与不同环境之间的关系随恢复阶段而变化：恢复30年次生林幼苗丰富度与土壤含水量、全氮、全磷含量显著相关，恢复60年次生林幼苗丰富度与冠层开阔度、土壤水分、土壤钾含量显著相关，而老龄林丰富度与冠层开阔度、有效磷、有效钾显著相关；幼苗多度和丰富度随以高度级划分的生长阶段的增加呈逐渐递减的趋势。
6.对不同恢复阶段的幼苗物种组成异质性和环境因子进行冗余分析，结果显示：在30年和60年次生林中，环境因子对幼苗层物种组成异质性的解释方差分别为55.1%和53.5%，而在老龄林中环境因子的解释方差较低，为43.4%。不同恢复阶段幼苗样方分布与环境因子之间的关系不同，恢复30年次生林的幼苗分布主要受土壤含水量和冠层开阔度的影响；恢复60年次生林的幼苗分布主要受土壤养分含量的影响。影响老龄林幼苗分布的环境因子主要是土壤含水量和全磷含量。
7.刀耕火种后自然恢复30年、60年的次生林以及老龄林幼苗的物种丰富度分别占整个群落物种丰富度的63.5%、60.6%和54.9%，说明海南岛热带低地雨林幼苗库是生物多样性中一个重要的组成部分。不同恢复阶段的幼苗(DBH<1cm)、幼树(1cm≤DBH<7.5cm)和成年树(DBH≥7.5cm)之间的补充比例不同，恢复30年次生林分别有71.9%和39.7%物种的幼苗能够进入幼树和成年树层，而恢复60年次生林分别有65.0%和38.0%物种的幼苗能够进入幼树和成年树层，老龄林分别有74.9%和50.9%幼苗进入到幼树和成年树层。
8.以热带低地雨林中分布的4个落叶物种山乌桕(Sapium discolor)、厚皮树(Lanneacoromandelica)、野漆树(Toxicodendron succedaneum)、银珠(Peltophorum tonkinense)和4个常绿物种：米槠(Castanopsis carlesi)、芳槁润楠(Machilus suaveolens)、乌墨(Syzygiumcumini)青梅(Vatica mangachapoi)为研究对象，分别进行4种改变地上和地下竞争的处理(移除地上植被、挖沟、移除地上植被并挖沟和对照)试验，分析了热带低地雨林幼苗更新与生长的影响因素。结果表明：移除地上植被和移除地上植被并挖沟能够显著增加幼苗的相对生长速率，而挖沟对移栽幼苗的相对生长速率影响不显著；落叶树种的幼苗生长速率明显高于常绿树种；幼苗相对生长速率随着光合有效辐射的增大逐渐升高。我们的研究结果表明，光照(地上竞争)是限制海南岛热带低地雨林幼苗生长的最主要因子，随着光合有效辐射的增加，幼苗的相对生长速率显著增加。
Understanding mechanism of species diversity maintenance has been one of the centralissues in community ecology. The seedling bank is a basic component in tropical forestecosystems, which play important roles in community dynamics, diversity maintenance andecosytem functioning.However, the seedlings with a diameter of less than1cm are usuallyneglected. However, seedling functional traits are often strongly related to the performance andfunction of adult plants in an ecosystem. Measurements of seedling traits are easy and accuratecompared to measurements of adult traits.In this thesis,we investigated the diversity andfucntional traits of woody plant(including trees,shrubs and lianas) seedlings in a tropicallowland rain forest of different recovery stages(30years and60years after shifting cultivationand old growth forest) on Hainan island, South China. The variation of seedling functionaltraits of woody plants during the recovery process and its influencing factors were explored.Seedling recruitment dynamics were analyzed by comparing the composition,abundance anddiversity of seedling and tree communities in each recovery stages of the tropical lowlandrainforest. The factors influencing the relative growth rate of8representative speceis of treeseedlings in the tropical lowland rain forest were examined through field experiments. Themain results are as follows.
1. Seedlings in30-year and60-year secondary forests had higher specific leaf area,specific stem length and specific root length. Leaf mass fraction increased significantly, whileroot mass fraction reduced gradually during the recovery process. The seedlings can beaggregated into3functional groups by ordination of Principal compoent analysis The speciesin old growth forest clusted more closely than those in the secondary forests.
2. Environmental factors changed significantly during the recovery process. Soil watercontent were significantly low in30-year forest, while there was no significant difference in60-year forest and old growth forest. Canopy openness and soil nutrient contents decreased withrecovery time. The correlations between seedling functional traits and environmental factors changed correspondingly with the natural recovery process. Most of the seedlings functionaltraits were significantly correlated with soil water content in the30-year forest but nocorrelations existed in the60-year and old growth forests. Seedling functional traits were notcorrelated with canopy openness in the30-year and60-year secondary forests while hadsignificant correaltions in the old growth forest. Seedling functional traits was correlated withsoil nutrient contents in each recovery stage.
3. The functional traits varied significantly with life form and growth stage (height class).Seedlings adapte to the community environment by making the trade-offs between differentfunctional traits. Tree and shrub seedlings adapt to the stressful rainforest understoryenvironment (such as low light) by decreasing specific leaf area and increasing stem massfraction. However, lianas adapt to the environment through the opposite strategy. With thegrowth of seedlings, trees and shrubs reduced the investment to assimilation, and increased theinvestment to support organ. Lianas increased investment to assimilation organs, and reducedto support organs. This is consistent with characteristics of liana attanching to other plants togrowth.
4. Functional traits of seedlings were significantly different from those of the adulttrees(DBH>7.5cm) in each recovery stage. Leaf dry mass content and specific stem density ofseedlings and adult trees were positively correlated. Specific leaf area of both seedlings andadult trees decreased, while leaf dry mass content and specific stem density increased duringthe recovery process. The environmental factors that affected functional traits of both seedlingsand trees were same to a certain extent in secondary forests, but different in old growthforest.These above results suggest that the patterns of change in functional traits and theirrelations with the environmental conditions for adult trees can be indicated in certain degree bythose of the the seedlings in this lowland rain forest.
5. The species-area and species-individual cumulative rates of seedlings increased withthe recovery time. Species accumulation rates in old growth forest were faster than those in thesecondary forests. The seedling abundance and richness first increased and then decreased withrecovery time, which was in line with the intermediate disturbance hypothesis. Species richness of seedlings was correlated with soil water content, total nitrogen and total phosphoruscontent in the30-year forest,while it was correlated with canopy openness, soil water contentand potassium content in the60-year forest; and it was correlated with canopy openness,available phosphorus, and available potassium in the old growth forest. Seedling abundanceand richness showed a gradually decreasing trend with increasing growth stage (height class).
6. The relationships between seedling species composition heterogeneity andenvironmental factors in three successional stages were assessed using redundancy analysis.The variance of environmental factors to interpret heterogeneity of seedlings were55.1%and53.5%respectively in30-year and60-year secondary forests, while lower (43.4%) in oldgrowth forest. The relationship between distribution of seedlings and environmental factors inthree recovery stages were different. Distribution of seedligns in30-year forest was mainlyaffected by soil water content and canopy openness. Distribution of seedligns in60-year forestwas mainly affected by soil nutrient content. Distribution of seedligns in old growth forest wasmainly affected by soil water content and total phosphorus.
7. The species richness of seedlings in30-year forest,60-year forest and old growthforest.accounted for63.5%,60.6%and54.9%of those for the whole coummunity respectively.This result showed that seedling bank is an important component, which should not beneglected in the study of biodiversity in this lowland rain forest. The recruitment rates fromseedlings (DBH<1cm) to saplings (1cm≤DBH<7.5cm) and from seedlings to adult trees(DBH≥7.5cm) were71.9%and39.7%respectively in30-year forest, while those were65.0%and38.0%respectively in60-year forest. In old growth forest,74.9%and50.9%of seedlingspecies grew into saplings and adult trees.
8. Eight representative species (Sapium discolor, Lannea coromandelica, Toxicodendronsuccedaneum, Peltophorum tonkinense, Castanopsis carlesi, Machilus suaveolens, Syzygiumcumini, Vatica mangachapoi) in the tropical lowland rain forest were selected to do thefollowing treatments: removal of the above vegetation (R), trenching (T), removal of the abovevegetation and trenching (R+T), and controlled (C). We measured the relative growth rate ofthese seelings to assess the factors affecting regeneration and growth of the seedlings. We found that the treatments of R and R+T significantly increased the growth rate of seedlings.But the effect of T was not significant. The growth rate of decidous species was higher thanevergreen species. Seedling growth rate increased with the increase of photosyntheticallyactive radiation. The relationship between them could be described by a linear equation. Allthese results showed that light was the most important factor for seedling growth in the tropicallowland rain forest. The below-ground competition had no significant effects on the growth ofseedlings.
引文
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