云南元谋干热河谷膏桐天然更新种群及其群落特征
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摘要
膏桐(Jatropha curcas Linn.)属大戟科麻疯树属落叶灌木或小乔木。在我国主要分布于南方部分省份,其中以云南省干热河谷地区较为多见。是云南干热河谷地区保持水土、促进植被恢复的树种之一。近年来以其种子油经过加工可制成清洁环保的生物柴油而倍受关注。
本研究以云南元谋干热河谷受人为干扰相对较小的膏桐天然更新种群及其群落为研究对象,在野外调查的基础之上,主要采用数量生态学方法,研究了膏桐种群结构、主要种群分布格局、物种多样性、主要物种生态位、主要物种种间关系、基于群落中主要木本植物的群落排序与聚类。目的在于从种群统计角度分析该种群在此生境下的生存现状及生态对策,推测其生命周期中相对敏感的阶段,分析影响该种群动态的重要因素；揭示该群落物种组成及其数量特征以及主要影响因素、膏桐在群落中的地位及其与主要伴生物种之间的相互关系、主要种群对资源的利用状况、此群落中主要种群的分布特点及其主要影响因子。主要结果和结论如下：
1.总体而言,膏桐种群径级结构呈金字塔形,属于增长型种群。静态生命表分析显示其种群死亡率高峰出现在幼年阶段及中老年过渡阶段。其存活曲线趋于DeeveyⅢ型。生存分析显示该种群的生存状况总体上呈现出前期变化幅度较大、后期变化幅度较小的特点。谱分析显示各周期所起作用随周期的减小而减小。
2.不同大小级及总体膏桐种群均呈集群分布。群落主要木本植物和主要草本植物也大多呈集群分布。各样地的膏桐种群皆呈集群分布。不同样地的不同发育阶段种群的分布格局不尽相同。随着年龄增长,大体上表现为由集群分布向随机分布转变,且这种转变主要发生在中树以及大树这两个阶段。经Greig-Smith格局分析显示不同样地膏桐种群的分布格局规模不尽相同,本研究揭示出的斑块规模出现在50m2和100m2左右,以100m2左右居多。
3.膏桐和扭黄茅分别是此群落木本植物和草本植物的优势种。总体而言,该群落草本植物的物种丰富度指数、多样性指数、均匀度指数高于木本植物与之对应的指数。群落物种丰富度指数、木本植物物种丰富度指数与坡向显著负相关。草本植物的Shannon-Wiener多样性指数与群落郁闭度显著负相关,与土壤pH显著正相关。
4.在群落木本植物中,膏桐、车桑子、戟叶酸模、沙针、华西小石积和帚枝鼠李在群落内分布幅度大,资源利用能力强,竞争能力强；对资源利用的相似性程度较大。而群落中大多数木本种群生态位相似比例值不高、生态位重叠较小,表明该群落的多数木本种群对环境的需求有较大差异。在群落草本植物中,扭黄茅、羽芒菊、芸香草、旱茅、黄细心、蔓草虫豆、假苜蓿的资源利用能力强、对资源利用的相似程度较大。而群落中大多数草本种群生态位相似比例值不高、生态位重叠较小,表明该群落草本植物的多数种群对环境的需求也有较大差异。可见,此群落物种间竞争不强烈,能较为和谐地共存,是该群落处于相对稳定状态的重要原因之一。
5.膏桐群落主要木本种群之间、主要草本种群之间总体上都表现出显著的正关联。Spearman相关分析结果表明,主要木本植物种对和主要草本植物种对中都是少数种对的相关性达到显著或极显著水平。主要木本植物种对中显著正相关、极显著正相关、显著负相关、极显著负相关的种对个数依次为8、5、6、5；主要草本植物种对则依次为2,4,3,3。大多数木本植物种对和大多数草本植物种对的相关都未达到显著水平,种间独立性相对较强,种间关系较为松散。主要木本植物种对和主要草本植物种对中都是全部正相关个数少于全部负相关个数。说明多数物种间的竞争相对较弱,这是群落相对稳定的重要原因之一。
膏桐与多数木本物种负相关,其中与山合欢和毛叶柿之间的负相关达到极显著水平,与车桑子、帚枝鼠李、裸实、千斤拔、野丁香、仙人掌之间正相关,但未达到显著水平。优势草本植物扭黄茅与13种草本物种负相关,其中,扭黄茅与旱茅间显著负相关；与地瓜榕及与鹿藿达到极显著负相关；与其余6种草本植物有微弱的正相关,皆没达到显著水平。
6.CCA分析表明：环境因子与群落分布的相关性由大至小依次为岩石裸露度、坡度、海拔、郁闭度、坡向。多数样地和物种分布于海拔较高、坡度较大、郁闭度较大的地段。建群种膏桐出现于排序轴交叉点附近区域,有较强的适应性。20种主要木本植物与样地的双向聚类分析结果与CCA结果相似。
Jatropha curcas, deciduous shrub or small tree, is classified as Jatropha in Euphobiaceae. In China, it is distributed mainly over some southern provinces. It often grows in Dry Hot Valley of Yuanmou in Yunnan province among those areas. It is one of the good plant species for soil and water conservation, as well as vegetation restoration in Dry Hot Valley in Yunnan. In recent years it has received much attention for its seeds, which can be processed to produce a environment-friendly clean biodiesel fuel.
The natural regeneration population of Jatropha curcas and its community without severe anthropogenic disturbance was studied in Dry Hot Valley of Yuanmou in Yunnan. The population structure, distribution pattern of important species, species diversity, niche of important species, species correlation of important species and the ordination and cluster analysis based on the main woody plant in the community were analyzed by means of mathematic ecology on the basis of the field study. The objectives were to analyze the current survival status and ecological strategy of the population in respect of demography, to infer the comparatively vulnerable demographic stages of the population, to analyze the important factors affecting the population dynamics, to discover the member and the mathematic characteristics of the community as well as their main relevent factors, to discover the status of Jatropha curcas and the relationship between Jatropha curcas and the other important species in the community, to discover the resource utilization characteristic of the important spieces, and to discover the distribution characteristics of the main plant in the community and their main factors. The main results are as follows:
1. The diameter structure diagram of the natural regeneration population of Jatropha curcas belonged to expansive type population pyramid. The static life table analysis indicated that the mortality was high both at the juvenile stage and in the transitional stage from the middle to the old. The shape of the survivorship curve was similar to that of Deevey type III curve. Survival analysis indicated that the survival condition of the population changed considerably in the earlier stage, and relatively slight in the later stage. The spectral analysis showed that the effect of the period was weak with the short of the period.
2. In general, the total Jatropha curcas population followed clumped distribution pattern, so do each developmental stages in the study. Both most of important woody plants and most of important herb in the community followed clumped distribution pattern. Jatropha curcas population in each plot followed clumped distribution pattern. The distribution pattern differed in different developmental stages in different plot. The distribution pattern of the population generally transformed from aggregated distribution into random distribution with age, which often occurred at middle trees or old trees stage for the population. Pattern analysis by Greig-Smith method revealed that the scale of the spatial pattern varied among the populations from different plots, and the identified patch sizes were about50m2and100m2in this study.100m2or so was relatively frequent.
3. Jatropha curcas and Heteropogon contortus was dominant species in woody plant and herb respectively. In general, species richness index, species diversity index and species evenness of the herb are higher than those of the woody plant respectively. There was a significantly negative correlation between species richness index of the community and slope aspect as well as between species richness index of the woody plant and slope aspect. There was a significantly negative correlation between Shannon-Wiener species diversity index of herb and the canopy cover of community. There was a significantly positive correlation between Shannon-Wiener species diversity index of herb and soil pH.
4. In the woody plant of the community, Jatropha curcas, Dodonaea viscose, Rumex hastatus, Osyris wightiana, Osteomeles schwerinae and Rhamnus virgata were widely dispersed over the community. They had great ability to utilize resources and to compete. Any two species in the six woody species were very similar in resource utilization. The niche similarity indexes of the majority of the woody species were low, so were the niche overlap indexes. These indicated that the majority of the wood species differed greatly in the requirement for environment.
In the herb of the community, Heteropogon contortus, Tridax procumbens, Cymbopogon distans, Eremopogon delavayi, Boerhavia diffusa, Atylosia scarabaeoides and Crotalaria medicaginea were widely dispersed over the community. They had great ability to utilize resources. Any two species in the seven herb species were very similar in resource utilization.. The niche similarity indexes of the majority of the herb species were low, so were the niche overlap indexes. These indicated that the majority of the herb species differed greatly in the requirement for environment. The interspecific competition in the community was not fierce. Most species could harmoniously coexist in the community relatively. This is one of the important reasons for the relatively persistence of the community.
5. The overall interspecific correlation analysis showed that significant positive correlation exist both among the20woody species and among the20herb species. Spearman's rank correlation coefficient analysis showed that the minority of the total woody species pairs showed significant correlation or highly significant correlation, so do the total herb species pairs. In the woody plant of the community,8,5,6and8species pairs showed significant positive correlations, highly significant positive correlations, significant negative correlations, and highly significant negative correlations respectively. In the herb of the community,2,4,3and3species pairs showed significant positive correlations, highly significant positive correlations, significant negative correlations, and highly significant negative correlations respectively. Most of the total woody plant species pairs did not show significant correlation, so do the herb species pairs. Most species did not closely correlated with other species. The numbers of species pairs with negative correlation were more than those with positive correlation both in the woody plant and in the herb. This indicated that the interspecific competition in most species was relatively weak in the community, which was one of the important reasons for the relatively persistence of the community.
Most of the woody plant species pairs including Jatropha curcas showed negative correlation. There was a highly significant negative correlation both between Jatropha curcas and Albizia kalkora as well as between Jatropha curcas and Diospyros mollifolia. A small number of woody plant species pairs including Jatropha curcas showed positive correlation, but none of the positive correlation was significant.
Most of the herb species pairs including Heteropogon contortus showed negative correlation. There was a significant negative correlation between Heteropogon contortus and Eremopogon delavayi. There was a highly significant negative correlation both between Heteropogon contortus and Ficus tikoua as well as between Heteropogon contortus and Rhynchosia volubilis. A small number of herb species pairs including Heteropogon contortus showed weak positive correlation, none of the positive correlation was significant.
6. The results of Canonical Correspondence Analysis indicated that the correlations order of the community distribution with the environmental factors were bareness degree of rock, slope gradient, altitude, canopy cover and slope aspect in large to small. The majority of the plots lay in the sites with high altitude, high slope gradient and high canopy cover, so were the main woody plant. Jatropha curcas Linn., the constructive species in the community, appeared in the region near to the intersection of the first two coordinate axes of the CCA ordination plot, which indicated that the species adapted well to the environment. The result of the Two-way Cluster Analysis of the20main woody plant in the community and the plots was similar to that of the Canonical Correspondence Analysis.
本研究以云南元谋干热河谷受人为干扰相对较小的膏桐天然更新种群及其群落为研究对象,在野外调查的基础之上,主要采用数量生态学方法,研究了膏桐种群结构、主要种群分布格局、物种多样性、主要物种生态位、主要物种种间关系、基于群落中主要木本植物的群落排序与聚类。目的在于从种群统计角度分析该种群在此生境下的生存现状及生态对策,推测其生命周期中相对敏感的阶段,分析影响该种群动态的重要因素；揭示该群落物种组成及其数量特征以及主要影响因素、膏桐在群落中的地位及其与主要伴生物种之间的相互关系、主要种群对资源的利用状况、此群落中主要种群的分布特点及其主要影响因子。主要结果和结论如下：
1.总体而言,膏桐种群径级结构呈金字塔形,属于增长型种群。静态生命表分析显示其种群死亡率高峰出现在幼年阶段及中老年过渡阶段。其存活曲线趋于DeeveyⅢ型。生存分析显示该种群的生存状况总体上呈现出前期变化幅度较大、后期变化幅度较小的特点。谱分析显示各周期所起作用随周期的减小而减小。
2.不同大小级及总体膏桐种群均呈集群分布。群落主要木本植物和主要草本植物也大多呈集群分布。各样地的膏桐种群皆呈集群分布。不同样地的不同发育阶段种群的分布格局不尽相同。随着年龄增长,大体上表现为由集群分布向随机分布转变,且这种转变主要发生在中树以及大树这两个阶段。经Greig-Smith格局分析显示不同样地膏桐种群的分布格局规模不尽相同,本研究揭示出的斑块规模出现在50m2和100m2左右,以100m2左右居多。
3.膏桐和扭黄茅分别是此群落木本植物和草本植物的优势种。总体而言,该群落草本植物的物种丰富度指数、多样性指数、均匀度指数高于木本植物与之对应的指数。群落物种丰富度指数、木本植物物种丰富度指数与坡向显著负相关。草本植物的Shannon-Wiener多样性指数与群落郁闭度显著负相关,与土壤pH显著正相关。
4.在群落木本植物中,膏桐、车桑子、戟叶酸模、沙针、华西小石积和帚枝鼠李在群落内分布幅度大,资源利用能力强,竞争能力强；对资源利用的相似性程度较大。而群落中大多数木本种群生态位相似比例值不高、生态位重叠较小,表明该群落的多数木本种群对环境的需求有较大差异。在群落草本植物中,扭黄茅、羽芒菊、芸香草、旱茅、黄细心、蔓草虫豆、假苜蓿的资源利用能力强、对资源利用的相似程度较大。而群落中大多数草本种群生态位相似比例值不高、生态位重叠较小,表明该群落草本植物的多数种群对环境的需求也有较大差异。可见,此群落物种间竞争不强烈,能较为和谐地共存,是该群落处于相对稳定状态的重要原因之一。
5.膏桐群落主要木本种群之间、主要草本种群之间总体上都表现出显著的正关联。Spearman相关分析结果表明,主要木本植物种对和主要草本植物种对中都是少数种对的相关性达到显著或极显著水平。主要木本植物种对中显著正相关、极显著正相关、显著负相关、极显著负相关的种对个数依次为8、5、6、5；主要草本植物种对则依次为2,4,3,3。大多数木本植物种对和大多数草本植物种对的相关都未达到显著水平,种间独立性相对较强,种间关系较为松散。主要木本植物种对和主要草本植物种对中都是全部正相关个数少于全部负相关个数。说明多数物种间的竞争相对较弱,这是群落相对稳定的重要原因之一。
膏桐与多数木本物种负相关,其中与山合欢和毛叶柿之间的负相关达到极显著水平,与车桑子、帚枝鼠李、裸实、千斤拔、野丁香、仙人掌之间正相关,但未达到显著水平。优势草本植物扭黄茅与13种草本物种负相关,其中,扭黄茅与旱茅间显著负相关；与地瓜榕及与鹿藿达到极显著负相关；与其余6种草本植物有微弱的正相关,皆没达到显著水平。
6.CCA分析表明：环境因子与群落分布的相关性由大至小依次为岩石裸露度、坡度、海拔、郁闭度、坡向。多数样地和物种分布于海拔较高、坡度较大、郁闭度较大的地段。建群种膏桐出现于排序轴交叉点附近区域,有较强的适应性。20种主要木本植物与样地的双向聚类分析结果与CCA结果相似。
Jatropha curcas, deciduous shrub or small tree, is classified as Jatropha in Euphobiaceae. In China, it is distributed mainly over some southern provinces. It often grows in Dry Hot Valley of Yuanmou in Yunnan province among those areas. It is one of the good plant species for soil and water conservation, as well as vegetation restoration in Dry Hot Valley in Yunnan. In recent years it has received much attention for its seeds, which can be processed to produce a environment-friendly clean biodiesel fuel.
The natural regeneration population of Jatropha curcas and its community without severe anthropogenic disturbance was studied in Dry Hot Valley of Yuanmou in Yunnan. The population structure, distribution pattern of important species, species diversity, niche of important species, species correlation of important species and the ordination and cluster analysis based on the main woody plant in the community were analyzed by means of mathematic ecology on the basis of the field study. The objectives were to analyze the current survival status and ecological strategy of the population in respect of demography, to infer the comparatively vulnerable demographic stages of the population, to analyze the important factors affecting the population dynamics, to discover the member and the mathematic characteristics of the community as well as their main relevent factors, to discover the status of Jatropha curcas and the relationship between Jatropha curcas and the other important species in the community, to discover the resource utilization characteristic of the important spieces, and to discover the distribution characteristics of the main plant in the community and their main factors. The main results are as follows:
1. The diameter structure diagram of the natural regeneration population of Jatropha curcas belonged to expansive type population pyramid. The static life table analysis indicated that the mortality was high both at the juvenile stage and in the transitional stage from the middle to the old. The shape of the survivorship curve was similar to that of Deevey type III curve. Survival analysis indicated that the survival condition of the population changed considerably in the earlier stage, and relatively slight in the later stage. The spectral analysis showed that the effect of the period was weak with the short of the period.
2. In general, the total Jatropha curcas population followed clumped distribution pattern, so do each developmental stages in the study. Both most of important woody plants and most of important herb in the community followed clumped distribution pattern. Jatropha curcas population in each plot followed clumped distribution pattern. The distribution pattern differed in different developmental stages in different plot. The distribution pattern of the population generally transformed from aggregated distribution into random distribution with age, which often occurred at middle trees or old trees stage for the population. Pattern analysis by Greig-Smith method revealed that the scale of the spatial pattern varied among the populations from different plots, and the identified patch sizes were about50m2and100m2in this study.100m2or so was relatively frequent.
3. Jatropha curcas and Heteropogon contortus was dominant species in woody plant and herb respectively. In general, species richness index, species diversity index and species evenness of the herb are higher than those of the woody plant respectively. There was a significantly negative correlation between species richness index of the community and slope aspect as well as between species richness index of the woody plant and slope aspect. There was a significantly negative correlation between Shannon-Wiener species diversity index of herb and the canopy cover of community. There was a significantly positive correlation between Shannon-Wiener species diversity index of herb and soil pH.
4. In the woody plant of the community, Jatropha curcas, Dodonaea viscose, Rumex hastatus, Osyris wightiana, Osteomeles schwerinae and Rhamnus virgata were widely dispersed over the community. They had great ability to utilize resources and to compete. Any two species in the six woody species were very similar in resource utilization. The niche similarity indexes of the majority of the woody species were low, so were the niche overlap indexes. These indicated that the majority of the wood species differed greatly in the requirement for environment.
In the herb of the community, Heteropogon contortus, Tridax procumbens, Cymbopogon distans, Eremopogon delavayi, Boerhavia diffusa, Atylosia scarabaeoides and Crotalaria medicaginea were widely dispersed over the community. They had great ability to utilize resources. Any two species in the seven herb species were very similar in resource utilization.. The niche similarity indexes of the majority of the herb species were low, so were the niche overlap indexes. These indicated that the majority of the herb species differed greatly in the requirement for environment. The interspecific competition in the community was not fierce. Most species could harmoniously coexist in the community relatively. This is one of the important reasons for the relatively persistence of the community.
5. The overall interspecific correlation analysis showed that significant positive correlation exist both among the20woody species and among the20herb species. Spearman's rank correlation coefficient analysis showed that the minority of the total woody species pairs showed significant correlation or highly significant correlation, so do the total herb species pairs. In the woody plant of the community,8,5,6and8species pairs showed significant positive correlations, highly significant positive correlations, significant negative correlations, and highly significant negative correlations respectively. In the herb of the community,2,4,3and3species pairs showed significant positive correlations, highly significant positive correlations, significant negative correlations, and highly significant negative correlations respectively. Most of the total woody plant species pairs did not show significant correlation, so do the herb species pairs. Most species did not closely correlated with other species. The numbers of species pairs with negative correlation were more than those with positive correlation both in the woody plant and in the herb. This indicated that the interspecific competition in most species was relatively weak in the community, which was one of the important reasons for the relatively persistence of the community.
Most of the woody plant species pairs including Jatropha curcas showed negative correlation. There was a highly significant negative correlation both between Jatropha curcas and Albizia kalkora as well as between Jatropha curcas and Diospyros mollifolia. A small number of woody plant species pairs including Jatropha curcas showed positive correlation, but none of the positive correlation was significant.
Most of the herb species pairs including Heteropogon contortus showed negative correlation. There was a significant negative correlation between Heteropogon contortus and Eremopogon delavayi. There was a highly significant negative correlation both between Heteropogon contortus and Ficus tikoua as well as between Heteropogon contortus and Rhynchosia volubilis. A small number of herb species pairs including Heteropogon contortus showed weak positive correlation, none of the positive correlation was significant.
6. The results of Canonical Correspondence Analysis indicated that the correlations order of the community distribution with the environmental factors were bareness degree of rock, slope gradient, altitude, canopy cover and slope aspect in large to small. The majority of the plots lay in the sites with high altitude, high slope gradient and high canopy cover, so were the main woody plant. Jatropha curcas Linn., the constructive species in the community, appeared in the region near to the intersection of the first two coordinate axes of the CCA ordination plot, which indicated that the species adapted well to the environment. The result of the Two-way Cluster Analysis of the20main woody plant in the community and the plots was similar to that of the Canonical Correspondence Analysis.
引文
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