普洱季风常绿阔叶林恢复生态学研究
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摘要
季风常绿阔叶林是云南省南部普洱地区重要的森林类型之一,是热带季雨林、雨林向亚热带常绿阔叶林过渡的植被类型,生物多样性较为丰富。由于过度的森林砍伐和不合理的土地利用导致森林面积的大规模减少,形成不同恢复阶段的退化生态系统。由原始林破坏后发展起来的天然和人工次生林已成为普洱地区重要的森林资源。以往对云南省季风常绿阔叶林的认识局限在原始林的群落结构、物种组成、土壤种子库及生物量上,而对人为干扰过的退化植被则了解甚少。本文以普洱地区不同恢复阶段的季风常绿阔叶林森林砍伐后自然恢复和人工恢复下的群落以及残存的原始林为研究对象,在共计25块样地(2.25hm2)野外群落学调查的基础上,通过比较不同类型群落的组成、结构、多样性、生态位、种间联结、种子库和功能群特征来探讨退化季风常绿阔叶林自然和人工恢复的动态规律,为该地区植被恢复策略的制定提供科学参考。本文的主要结果包括:
1.以野外样方调查数据为基础,对普洱地区季风常绿阔叶林分布海拔范围内的不同恢复群落类型进行数量分类。结果表明:应用TWINSPAN可将季风常绿阔叶林分布海拔范围内恢复群落划分为5个植被类型:季风常绿阔叶林、针阔混交林、针叶林、西南桦林及桉树林,分属原始林、次生林及人工林。恢复方式可分为自然恢复与人工恢复两个恢复序列。自然恢复:恢复15年群落——恢复30年群落——原始林;针叶林——针阔混交林——原始林。从植物区系组成成分看,恢复群落中世界分布科的百分比均高于原始林,由于种源地相同,恢复群落大都处于向季风常绿阔叶林原始林群落的恢复阶段,因此,科、属组成的相似性程度较高,种的组成上既有相似之处,也有一些不同,原始林中国特有种要显著大于恢复群落。原始林与自然恢复群落的生活型谱的相关大于与人工恢复群落,针叶林中一年生植物要多于其它群落类型。原始林的小型叶的比例要小于恢复群落,而叶型则没有反映出群落的恢复程度。
2.通过对云南普洱地区季风常绿阔叶林皆伐后人工和天然恢复15年及30年群落的调查,并以原始林群落为对照,对比分析了恢复方式和恢复时间对季风常绿阔叶林恢复群落的结构和多样性的影响。结果表明:恢复方式对科、属、物种、乔木、灌木、藤本等物种组成和多样性情况无显著影响,而恢复时间则具有显著的影响。科、属、物种、乔木及灌木物种丰富度随着恢复时间的延长呈现先减后增的趋势。所有群落物种丰富度均随径级和高度级的增加而呈现倒“J”型分布,多度则随径级增加呈现偏峰曲线,而随高度级的增加呈现倒“J”型分布。恢复方式和恢复时间与群落结构与多样性变量之间的相关性分析及回归分析表明,恢复时间与多数变量之间具有显著的相关性,而恢复方式则仅与藤本多度具有显著相关性,这说明与恢复方式相比,恢复时间是影响该地区恢复群落结构和多样性的主要因子。
3.通过对云南普洱地区季风常绿阔叶林群落的调查,探讨了不同演替阶段群落物种–面积关系的变化。结果表明:不同演替阶段群落中,取样面积与总物种数、乔木、灌木、藤本物种数均具有极显著相关性,其变化解释率均超过94%;总物种和乔木的物种累积速率(Z )在演替30年的群落中最低,分别为0.334和0.394,灌木(0.437)和藤本(0.326)的Z值则在演替15年的群落中最低;总物种、乔木、灌木、藤本的物种–面积曲线截距在不同演替阶段中无显著变化,但总物种及藤本物种–面积曲线的决定系数(R2)为原始林中显著高于演替15年和30年群落。演替15年群落中,初始乔木及灌木物种丰富度解释了Z值变化的99.97%,但其他演替阶段群落中初始乔木、灌木、藤本及总物种丰富度与Z值均无显著相关性。
4.生态位和种间联结是群落动态研究的焦点之一,物种间相互作用影响着群落的恢复和演替。本文在对云南普洱地区不同恢复阶段(恢复15年、恢复30年和原始林群落)季风常绿阔叶林群落的野外调查基础上,采用生态位宽度、生态位重叠、方差比率(VR)和基于2×2联列表的χ2检验、联结系数(AC)的方法,分析了不同恢复阶段各种群之间的变化趋势。结果表明:3种群落类型中物种的重要值与相对应的生态位宽度之间存在着显著的正相关,恢复15年群落有78.22%的种对之间的生态位重叠值在0.4—0.8之间,说明其群落优势物种存在较强的资源利用性竞争,而恢复30年和原始林群落结构则具有一定的稳定性,多数生态位宽度较大的物种之间的生态位重叠要大于生态位较小的物种,反映出优势物种具有较强的竞争能力。恢复15年和恢复30年群落物种间总体呈不显著正联结,原始林群落则呈不显著负联结。3种群落231种对中,绝大多数种对联结系数未达显著水平,种对间的独立性相对较强,群落中物种间正负联结对数占总种的比例随恢复进程而出现下降趋势。不同恢复阶段群落中联结系数与生态位重叠值之间存在显著正相关,生态位重叠值越大,其联结系数也越大。
5.通过对云南普洱地区不同恢复阶段(恢复15年、30年和原始林群落)季风常绿阔叶林群落的野外调查及土壤种子库的萌发实验,探讨其土壤种子库和幼苗库的基本特征、变化规律及与地上植被的关系。结果表明:在8个样地的土壤种子库中共有2645粒种子(分属37科72属83种)萌发;土壤中种子密度由小到大依次为原始林群落(148±23粒/m~2)<恢复15年群落(362.33±77.05粒/m~2)<恢复30年群落(424±91.53粒/m~2)。原始林群落中种子主要集中在0—2cm层和5—10cm,恢复15年与30年群落在3层间差异不显著;恢复30年群落的Shannon-Wiener指数显著高于恢复15年和原始林群落。3种类型土壤种子库生活型主要以草本植物组成,其中恢复30年群落中多年生草本种子密度显著高于恢复15年和原始林群落,恢复15年群落中一年生草本种子密度要显著高于恢复30年和原始林群落。原始林幼苗库密度要显著大于恢复15年群落,幼苗库密度与土壤种子库种子密度(P<0.05)和林分密度(P<0.01)存在显著负相关,与土壤容重及pH值之间存在显著正相关(P<0.05)。土壤种子库与地上植被的相似性普遍较低,在0.0375—0.1538之间;土壤种子库与草本层的相似性系数要高于土壤种子库与乔木层,幼苗库与乔木层的相似性系数要高于其它类型的相似性系数。
6.藤本植物是森林生态系统的重要组成部分,影响群落的恢复与演替。本文通过对云南普洱地区不同恢复阶段(恢复15年、30年和原始林群落)季风常绿阔叶林群落的野外调查,分析了不同恢复阶段藤本植物的物种丰富度、密度、多度、径级分布、多样性指数及攀援方式,并探讨藤本植物与支持木的关系。结果表明:在0.81hm~2的调查样地中,共发现DBH≥0.1cm的藤本植物1292株(分属34科51属64种)。原始林群落的藤本物种丰富度、密度(DBH<1cm)、胸高断面积和平均胸高断面积都显著高于恢复阶段,原始林和恢复15年群落的密度(DBH≥1cm)、平均胸径和平均长度之间无显著差异,但都显著高于恢复30年。3种群落类型中藤本植物的物种组成和径级分布有显著差异,原始林中藤本植物物种更多,而且大径级(DBH≥10cm)的藤本植物仅出现在原始林及恢复30年的群落。单株藤本攀援的支持木在3种群落类型中均占多数,藤本植物与支持木的胸径存在显著的正相关(P<0.001),原始林群落中DBH≥15cm的支持木更易被攀援,而恢复阶段则相反。茎缠绕藤本植物对原始林的负面影响要显著少于恢复15年及30年群落,而卷须类藤本植物也反映出原始林正处于一个动态变化的阶段,同时根攀缘和搭靠类藤本植物物种组成和多度变化可以反映出干扰后季风常绿阔叶林的恢复程度。森林砍伐是影响不同恢复阶段藤本植物的物种组成和分布的主要因素。
7.附生维管植物是季风常绿阔叶林物种丰富度的重要组成部分。本文通过对云南普洱地区不同恢复阶段(恢复15年、30年和原始林群落)季风常绿阔叶林群落内附生维管植物的调查,分析了不同恢复阶段附生维管植物的物种丰富度、多度、分布和相似性关系,并探讨附生维管植物与宿主的关系。结果表明:(1)在0.81hm~2的调查样地中,共发现附生维管植物3116株(分属9科20属22种);(2)原始林中附生维管植物的物种丰富度(17种)要显著高于恢复30年(7种)和15年(5种)群落;(3)原始林与不同恢复阶段群落中附生维管植物的物种组成有显著不同,恢复15年群落中的附生植物主要由蕨类植物组成,兰科(Orchidaceae)植物是原始林附生植物的主要物种组成,有些物种仅出现在原始林中,可以作为季风常绿阔叶林恢复程度的指示物种;(4)原始林附生维管植物的多度要显著高于恢复阶段;(5)3个群落类型中的附生维管植物在水平方向都为聚集分布,恢复15年群落主要分布在0—5m,原始林中分布可达到20m;(6)原始林中附生维管植物物种丰富度和多度与宿主的胸径之间存在显著正相关,而2个恢复阶段森林中不存在相关性。
The monsoonal broad-leaved evergreen forest is an important vegetation type of the Puer area which located in the Southern Yunnan province, P.R. China. It is a land of enriched biodiversity and considered a transitional vegetation type between the tropical monsoonal rain forest and the subtropical broad-leaved evergreen forest. Due to the extensive deforestation and long-term intensive agricultural landuses, the primary forests have significantly decreased. Thus formed a spectrum of degraded ecosystem of different restoration stages. Therefore, the secondary forests that developed after the natural and artificial restoration by the destruction of primary forests in this region which has become an important forest resources. However, past ecological studies of monsoonal broad-leaves evergreen forest in Yunnan were mainly conducted in community structure, species composition, soil seed bank and biomass of the primary forests, and we still know little about those degraded vegetation after anthropogenic. In this study, field investigations were conducted in totally 25 plots (2.25hm~2) of recovery vegetation and primary forests. The recovery vegetation included different stages of natural restoration and aritficial restoration. On the basis of those field investigations, we analyzed the community composition, structure, diversity, niche, interspecific association, seed banks and the functional group and explored dynamic law of recovery strategy. The goal is to provide science reference for vegetation restoration. The main results are as follows:
1.Analyze the characteristics of the monsoonal broad-leaved evergreen forest and its restoration communities, quantitaitve classification were conducted by TWINSPAN based on field investigation plots. The investigated plots were classified into 7 communities by TWINSPAN, they are primary monsoonal broad-leaved evergreen forest,community of 15 year restoration (15a), community of 30 year restoration (30a), mixed needle broad-leaved forest, needle-leaved forest and eucalyptus forest. recovery types can be divided into natural restoration and artificial restoration: natural recovery: 15a——30a——the primary forest, needle-leaved forest——mixed needle broad-leaved forest——the primary forest. Floristic composition indicated cosmopolitan percentage of restoration communities were higher than the primary forest at familiy level. Because of same species source, restoration communities were at a restoration stage form restoration communities to primary forest. Therefore, the similarity of families and genera were higher degree. the composition of species had some differences. endemic species of the primary forest communities were significantly greater than the recstoration communities. The life-form spectrum bettween restoration communites and primary forest had signicantly correlation, and natural restoration were larger than artificial restoration. Needle-leaved forest had more annuals in all communities. The percentage of microphyll of the primary forest is less than restoration communities, and leaf type did’t reflect restoration degree of community.
2.In the present study, the effect of restoration strategy and time on community structure and diversity were analyzed by investigating species composition, structure, and diversity of different restoration strategy and time community. The results showed that restoration strategy had no effect on species composition and diversity of family, genera, species, tree, shrub, and liana, however, restoration time had significant effect on species composition and diversity. Species richness of family, genera, species, tree, and shrub decreased at the early stage and then increased with community succession. Species richness showed an inverse J-shape with DBH increasing in all community types, while abundance showed single peak curve with DBH increasing and an inverse J-shape with height increasing. The correlation and regression analysis among restoration strategy and time and variables of community structure and diversity showed that there were significant correlation between restoration time and variables of community structure and diversity, and no significant correlation were found between restoration strategy and most of variables of community structure and diversity, which indicated that restoration time was the major factor that affects structure and diversity of community restored.
3.Species-area relationships are one of the most fundamental and most studied problems in all of ecology. However, whether the species-area relationships properties, especially species-area relationships slopes, remain constant through succession time, even though the ecosystem characteristics change on seasonal to evolutionary time scales, is a controversial problem. Focusing on disturbed broad-leaved evergreen forest ecosystems, we analyzed the species-area relationships in different succession stage communities (15a, 30a, and primary forest). Investigation was performed in Pu’er, Yunnan Province. Three plots (30 m×30 m) were located in the 15a, the 30a and the primary forest respectively. Each plot was divided into 36 subplots of 5 m×5 m for the vegetation surveys. Within each subplot all stems of height (H)>1.3 m were counted, measured and identified to the species level. We also recorded environment factors such as crown density, elevation, and slop. Species richness of tree, shrub, liana and total species in the 15a, the 30a and the primary forest were calculated respectively. The species-area relationships, the accumulate rate ( Z ) and the coefficients of determination (R2) of tree, shrub, liana and total species in each succession stage community were studied. We found that the number of total species, trees, shrubs, and lianas had a highly significant correlation with the sample area in different succession stage communities, the area explained over 94% of the total variation in the number of species, trees, shrubs, and lianas. The Z of total species(0.334) and trees(0.394) were the lowest in the 30a, whereas, the Z of shrubs(0.437) and lianas(0.326) were the lowest in the 15a. No significant difference were found for the intercepts of the species-area curve of species, trees, shrubs, and lianas among different succession stage communities, however, the coefficients of determination (R2) of the species-area curve of species and lianas were the highest in the primary forest. A fitted regression line of trees and shrubs explained 99.97% of the variance of Z-value in the 15a, but there were no significant correlation between trees, shrubs, lianas, and total species richness and Z-value in the 30a and the primary forest.
4.Niche and interspecific association is a focus of community dynamics. Interspecific interactions affect restoration and succession of community. Field data collection was biased on 0.81hm~2 plot (including 9 subplots) in different restoration stages (15a, 30a and primary monsoonal broad-leaved evergreen forest) which distributed in Pu'er city,Yunnan province. A series of techniques including niche breadth,niche overlap, the variance ratio (VR),chi-square test,association coefficient (AC) based on 2×2 contingency table were used to analyze change trend of tree population of different restoration stages. The results shown that:There was significant positive correlation (P<0.001) between the importance value and corresponding to niche breadth in 3 communites. 78.22% of species pairs had a niche overlap were between 0.4 and 0.8 in 15a,which indicated a strong resource utilization competition in dominant species. Simultaneously,community structure of 30a and primary forest was stability. The niche breadth increased with niche overlap,and vice versa,which reflected that the dominant species had strong competitiveness. There was a significant positive correlation in total on VR in the 15a and 30a,while the primary forest was no-significant negative association. For chi-square test,Most species pairs did not show significant association in 3 communities,which may show stronger independent distribution. The proportion of positive and negative association decreased from 15a to primary forest. There was a significant positive correlation between association coefficient and niche overlap. Generally speaking, the more niche overlap,the more association coefficient.
5.Soil seed bank and seedling bank is considered an important source for natural restoration in forest ecosystems. Soil seed bank is a dynamic biotic component of plant communities that represents the population’s memory in relation to selective events and seedling plays an important role for numerous tree species. Field data was based on 0.72hm~2 plot (including 8 subplots) in different restoration stages (15a, 30a) and primary monsoon broad-leaved evergreen forest which distribute in Caiyanghe nature reserve, Yixiang township and Xinfang reservoir, Puer city, Yunnan, China. Germination experiments carried out in the greenhouse, which lasted 7 months. On the basis of analyzing soil seed bank and seedling bank density and its relation to the above-ground vegetation. The results were summarized as follows: 2645 seeds germinated in all soil samples, 83 species in72 genera and 37 families were recorded. Total seed density of 30a (424±91.53 seeds/m~2) was significantly higher than primary forest (148±23 seeds/m~2). There was no significant difference between 15a (362.33±77.05 seeds/m~2) and other two types. Seed density of 0-2cm and 5-10cm layer was significantly higer than 2-5cm layer in pirmary forest, while no significant difference among 3 layers in 15a and 30a. Compositae was most abundant species in three community types,because of their wind dispersal way. Shannon-Wiener index of 30a was signifcantly higher than 15a and primary forest, but species richness was not significant difference in three types (including three layer). The seed density of herb was significantly higher than other life form in 3 community types. Seed density of perennial herb of 30a was significantly higher than 15a and primary and seed density of annual herb of 15a was signicantly higer than 30a and primary.and seed density of herb plant and fern plant of 30a was significantly higer than the primary forest. Dicranopteris pedata was most abundant species in restoration stages, especially in 30a. Annual herb offen emerged in early stages of restoration, such as Lindernia nummularifolia, which had short distance to tea park and farm land and had richness seed source. Castanopsis echidnocarpa is dominat species of monsoon broad-leaved evergreen forest, which was most abundant seedling of 30a and primary. Castanopsis calathiformis was most abundant seedling species of 15a. Tree seedlings were more than shrub seedlings. Tree seedling was main composition in all seedling bank, because of survival strategy of seed. Seedling bank density of primary was significantly higher than 15a, which had signifant negative correlation with soil seed bank density (P<0.05)and stand density(P<0.01), had signifantly positive correlation with soil bulk desity and Ph (P<0.05). The similarity coefficient (from 0.0375 to 0.1538) between soil seed bank and above-ground vegetation was generally low in three community types. Similarity coefficient between soil seed bank and herb was higher than soil seed bank and tree layer. Seedling bank was more similar with tree layer than soil seed bank, so forest restoration of monsoon broad-leaved evergreen forest mainly relyed on seed rain of tree layer.
6.Liana is important component of forest ecosystem, which usually affects restoration and succession of the communities. Relationship between liana and host tree has significant implications for forest conservation and sustainable management. Field data collection was based on 0.81hm~2 plot (including 9 subplots) in different restoration stages (15a, 30a and primary monsoonal broad-leaved evergreen forest distribute in Caiyanghe nature reserve, Yixiang Township and Xinfang reservoir, Pu’er city, Yunnan, China. On the basis of analyzing liana species richness, density, abundance, size structure, diversity indices and the climbing mechanism in the different restoration period, the relationship between liana and host tree was discussed. The results shown that: 1292 climbing lianas (DBH≥0.1cm), belonging to 64 species in 51 genera and 34 families were recorded in all plots. The liana richness,density(DBH<0.1cm), basal area at breast height and mean basal area at breast height in primary forest were significantly higher than restoration stages. There was no significant difference of density (DBH≥1cm), average DBH and average length between primary forest and 15a, though both were significantly higher than that of 30a. Species composition and DBH class distribution of lianas varied significantly in the three community types. Mucuna macrocarpa,Celastrus monospermus and Gnetum montanum are most abundant species. These species were found in all restoration stages and primary forest.Mucuna macrocarpa is an early successional species which needs intensive light environment and Gnetum montanum is a late successional species. Species-area curve showed that primary forest had higher species richness than 15a and 30a, which confirmed that primary forest played an important role in biodiversity conservation of lianas. Liana abundance decreased significantly while its DBH class increased. DBH of more than 95% lianas was less than 5cm. The lianas of large diameter class (DBH≥10cm) were found only in primary forest and 30a. One liana per host tree was most abundant in three community types. Host trees were more likely to be infested by lianas when their DBH was larger than 15cm in primary forest and liana-host tree relationship showed different trends in restoration stages. Large tree was more susceptible to be climbed by liana. There was a significant positive correlation (P<0.001) between the liana DBH and host tree DBH. DBH of liana increased with the growth of DBH of host tree in primary and 15a. Stem twining caused more mechanical damage in 4 climbing ways, and the damage for primary forest were significantly lower than that for restoration stages. Density of tendril climber showed that primary forest was undergoing a dynamic process. Species composition and abundance change of root climber and hook climber can be used as an indicator to reflect restoration level of monsoonal broad-leaved evergreen forest of post-disturbance. Primary forest had more Piper flaviflorum than 15a and 30a and Fissistigma acuminatissimum did not appeare in the restoration stages. Deforestation is main factor that affects liana species composition and distribution and reduces lianas abundance and richness significantly. Restoration time has an important effect on regeneration of lianas.
7.Vascular epiphytes are important components of species richness in monsoonal broad-leaved evergreen forest. Field data collection vascular epiphytes was based on 0.81hm~2 plot (including 9 subplots) in different restoration stages (15a, 30a and primary monsoonal broad-leaved evergreen forest distribute in Caiyanghe nature reserve, Yixiang Township and Xinfang reservoir, Pu’er city, Yunnan, China. On the basis of analyzing vascular epiphytes species richness, abundance, distribution, similarity coefficient, the relationship between epiphytes and phorophytes was discussed. The results shown that: (1) 3116 vascular epiphytes, belonging to 22 species in 20 genera and 9 families were recorded in all plots. (2) Diversity of vascular epiphytes was lower in restoration stages (15a, 5 spp. And 30a, 7 spp.) compared to primary forest (17 spp.). (3) Epiphytes species composition in primary forest and restoration stages differed markedly: community of 15a restoration harboured no orchid species but more fern species and primary forest hold more orchid species. Probably the families occurring only in primary forest sites of our study may be useful as bioindicators to determine the degree of restoration in monsoonal broad-leaved evergreen forest. (4) Epiphytes abundance were also lower in restoration stages. The decrease in species numbers and abundance as well as the differences in species composition was mainly due to the less diverse phorophyte structure and less differentiated microclimate in the disturbed and secondary vegetation compared to the primary forest. S?rensen coefficient between 15a and primary forest was higher than between 30a and primary forest. (5) The vascular epiphytes were clumped horizontally in 3 community types. Vertically, vascular epiphytes of 15a were mainly distributed at 0～5m and primary forest was up to 20m. (6) Vascular epiphytes species richness and abundance were both significantly positively correlated with host tree size but restoration stages didn’t have.
1.以野外样方调查数据为基础,对普洱地区季风常绿阔叶林分布海拔范围内的不同恢复群落类型进行数量分类。结果表明:应用TWINSPAN可将季风常绿阔叶林分布海拔范围内恢复群落划分为5个植被类型:季风常绿阔叶林、针阔混交林、针叶林、西南桦林及桉树林,分属原始林、次生林及人工林。恢复方式可分为自然恢复与人工恢复两个恢复序列。自然恢复:恢复15年群落——恢复30年群落——原始林;针叶林——针阔混交林——原始林。从植物区系组成成分看,恢复群落中世界分布科的百分比均高于原始林,由于种源地相同,恢复群落大都处于向季风常绿阔叶林原始林群落的恢复阶段,因此,科、属组成的相似性程度较高,种的组成上既有相似之处,也有一些不同,原始林中国特有种要显著大于恢复群落。原始林与自然恢复群落的生活型谱的相关大于与人工恢复群落,针叶林中一年生植物要多于其它群落类型。原始林的小型叶的比例要小于恢复群落,而叶型则没有反映出群落的恢复程度。
2.通过对云南普洱地区季风常绿阔叶林皆伐后人工和天然恢复15年及30年群落的调查,并以原始林群落为对照,对比分析了恢复方式和恢复时间对季风常绿阔叶林恢复群落的结构和多样性的影响。结果表明:恢复方式对科、属、物种、乔木、灌木、藤本等物种组成和多样性情况无显著影响,而恢复时间则具有显著的影响。科、属、物种、乔木及灌木物种丰富度随着恢复时间的延长呈现先减后增的趋势。所有群落物种丰富度均随径级和高度级的增加而呈现倒“J”型分布,多度则随径级增加呈现偏峰曲线,而随高度级的增加呈现倒“J”型分布。恢复方式和恢复时间与群落结构与多样性变量之间的相关性分析及回归分析表明,恢复时间与多数变量之间具有显著的相关性,而恢复方式则仅与藤本多度具有显著相关性,这说明与恢复方式相比,恢复时间是影响该地区恢复群落结构和多样性的主要因子。
3.通过对云南普洱地区季风常绿阔叶林群落的调查,探讨了不同演替阶段群落物种–面积关系的变化。结果表明:不同演替阶段群落中,取样面积与总物种数、乔木、灌木、藤本物种数均具有极显著相关性,其变化解释率均超过94%;总物种和乔木的物种累积速率(Z )在演替30年的群落中最低,分别为0.334和0.394,灌木(0.437)和藤本(0.326)的Z值则在演替15年的群落中最低;总物种、乔木、灌木、藤本的物种–面积曲线截距在不同演替阶段中无显著变化,但总物种及藤本物种–面积曲线的决定系数(R2)为原始林中显著高于演替15年和30年群落。演替15年群落中,初始乔木及灌木物种丰富度解释了Z值变化的99.97%,但其他演替阶段群落中初始乔木、灌木、藤本及总物种丰富度与Z值均无显著相关性。
4.生态位和种间联结是群落动态研究的焦点之一,物种间相互作用影响着群落的恢复和演替。本文在对云南普洱地区不同恢复阶段(恢复15年、恢复30年和原始林群落)季风常绿阔叶林群落的野外调查基础上,采用生态位宽度、生态位重叠、方差比率(VR)和基于2×2联列表的χ2检验、联结系数(AC)的方法,分析了不同恢复阶段各种群之间的变化趋势。结果表明:3种群落类型中物种的重要值与相对应的生态位宽度之间存在着显著的正相关,恢复15年群落有78.22%的种对之间的生态位重叠值在0.4—0.8之间,说明其群落优势物种存在较强的资源利用性竞争,而恢复30年和原始林群落结构则具有一定的稳定性,多数生态位宽度较大的物种之间的生态位重叠要大于生态位较小的物种,反映出优势物种具有较强的竞争能力。恢复15年和恢复30年群落物种间总体呈不显著正联结,原始林群落则呈不显著负联结。3种群落231种对中,绝大多数种对联结系数未达显著水平,种对间的独立性相对较强,群落中物种间正负联结对数占总种的比例随恢复进程而出现下降趋势。不同恢复阶段群落中联结系数与生态位重叠值之间存在显著正相关,生态位重叠值越大,其联结系数也越大。
5.通过对云南普洱地区不同恢复阶段(恢复15年、30年和原始林群落)季风常绿阔叶林群落的野外调查及土壤种子库的萌发实验,探讨其土壤种子库和幼苗库的基本特征、变化规律及与地上植被的关系。结果表明:在8个样地的土壤种子库中共有2645粒种子(分属37科72属83种)萌发;土壤中种子密度由小到大依次为原始林群落(148±23粒/m~2)<恢复15年群落(362.33±77.05粒/m~2)<恢复30年群落(424±91.53粒/m~2)。原始林群落中种子主要集中在0—2cm层和5—10cm,恢复15年与30年群落在3层间差异不显著;恢复30年群落的Shannon-Wiener指数显著高于恢复15年和原始林群落。3种类型土壤种子库生活型主要以草本植物组成,其中恢复30年群落中多年生草本种子密度显著高于恢复15年和原始林群落,恢复15年群落中一年生草本种子密度要显著高于恢复30年和原始林群落。原始林幼苗库密度要显著大于恢复15年群落,幼苗库密度与土壤种子库种子密度(P<0.05)和林分密度(P<0.01)存在显著负相关,与土壤容重及pH值之间存在显著正相关(P<0.05)。土壤种子库与地上植被的相似性普遍较低,在0.0375—0.1538之间;土壤种子库与草本层的相似性系数要高于土壤种子库与乔木层,幼苗库与乔木层的相似性系数要高于其它类型的相似性系数。
6.藤本植物是森林生态系统的重要组成部分,影响群落的恢复与演替。本文通过对云南普洱地区不同恢复阶段(恢复15年、30年和原始林群落)季风常绿阔叶林群落的野外调查,分析了不同恢复阶段藤本植物的物种丰富度、密度、多度、径级分布、多样性指数及攀援方式,并探讨藤本植物与支持木的关系。结果表明:在0.81hm~2的调查样地中,共发现DBH≥0.1cm的藤本植物1292株(分属34科51属64种)。原始林群落的藤本物种丰富度、密度(DBH<1cm)、胸高断面积和平均胸高断面积都显著高于恢复阶段,原始林和恢复15年群落的密度(DBH≥1cm)、平均胸径和平均长度之间无显著差异,但都显著高于恢复30年。3种群落类型中藤本植物的物种组成和径级分布有显著差异,原始林中藤本植物物种更多,而且大径级(DBH≥10cm)的藤本植物仅出现在原始林及恢复30年的群落。单株藤本攀援的支持木在3种群落类型中均占多数,藤本植物与支持木的胸径存在显著的正相关(P<0.001),原始林群落中DBH≥15cm的支持木更易被攀援,而恢复阶段则相反。茎缠绕藤本植物对原始林的负面影响要显著少于恢复15年及30年群落,而卷须类藤本植物也反映出原始林正处于一个动态变化的阶段,同时根攀缘和搭靠类藤本植物物种组成和多度变化可以反映出干扰后季风常绿阔叶林的恢复程度。森林砍伐是影响不同恢复阶段藤本植物的物种组成和分布的主要因素。
7.附生维管植物是季风常绿阔叶林物种丰富度的重要组成部分。本文通过对云南普洱地区不同恢复阶段(恢复15年、30年和原始林群落)季风常绿阔叶林群落内附生维管植物的调查,分析了不同恢复阶段附生维管植物的物种丰富度、多度、分布和相似性关系,并探讨附生维管植物与宿主的关系。结果表明:(1)在0.81hm~2的调查样地中,共发现附生维管植物3116株(分属9科20属22种);(2)原始林中附生维管植物的物种丰富度(17种)要显著高于恢复30年(7种)和15年(5种)群落;(3)原始林与不同恢复阶段群落中附生维管植物的物种组成有显著不同,恢复15年群落中的附生植物主要由蕨类植物组成,兰科(Orchidaceae)植物是原始林附生植物的主要物种组成,有些物种仅出现在原始林中,可以作为季风常绿阔叶林恢复程度的指示物种;(4)原始林附生维管植物的多度要显著高于恢复阶段;(5)3个群落类型中的附生维管植物在水平方向都为聚集分布,恢复15年群落主要分布在0—5m,原始林中分布可达到20m;(6)原始林中附生维管植物物种丰富度和多度与宿主的胸径之间存在显著正相关,而2个恢复阶段森林中不存在相关性。
The monsoonal broad-leaved evergreen forest is an important vegetation type of the Puer area which located in the Southern Yunnan province, P.R. China. It is a land of enriched biodiversity and considered a transitional vegetation type between the tropical monsoonal rain forest and the subtropical broad-leaved evergreen forest. Due to the extensive deforestation and long-term intensive agricultural landuses, the primary forests have significantly decreased. Thus formed a spectrum of degraded ecosystem of different restoration stages. Therefore, the secondary forests that developed after the natural and artificial restoration by the destruction of primary forests in this region which has become an important forest resources. However, past ecological studies of monsoonal broad-leaves evergreen forest in Yunnan were mainly conducted in community structure, species composition, soil seed bank and biomass of the primary forests, and we still know little about those degraded vegetation after anthropogenic. In this study, field investigations were conducted in totally 25 plots (2.25hm~2) of recovery vegetation and primary forests. The recovery vegetation included different stages of natural restoration and aritficial restoration. On the basis of those field investigations, we analyzed the community composition, structure, diversity, niche, interspecific association, seed banks and the functional group and explored dynamic law of recovery strategy. The goal is to provide science reference for vegetation restoration. The main results are as follows:
1.Analyze the characteristics of the monsoonal broad-leaved evergreen forest and its restoration communities, quantitaitve classification were conducted by TWINSPAN based on field investigation plots. The investigated plots were classified into 7 communities by TWINSPAN, they are primary monsoonal broad-leaved evergreen forest,community of 15 year restoration (15a), community of 30 year restoration (30a), mixed needle broad-leaved forest, needle-leaved forest and eucalyptus forest. recovery types can be divided into natural restoration and artificial restoration: natural recovery: 15a——30a——the primary forest, needle-leaved forest——mixed needle broad-leaved forest——the primary forest. Floristic composition indicated cosmopolitan percentage of restoration communities were higher than the primary forest at familiy level. Because of same species source, restoration communities were at a restoration stage form restoration communities to primary forest. Therefore, the similarity of families and genera were higher degree. the composition of species had some differences. endemic species of the primary forest communities were significantly greater than the recstoration communities. The life-form spectrum bettween restoration communites and primary forest had signicantly correlation, and natural restoration were larger than artificial restoration. Needle-leaved forest had more annuals in all communities. The percentage of microphyll of the primary forest is less than restoration communities, and leaf type did’t reflect restoration degree of community.
2.In the present study, the effect of restoration strategy and time on community structure and diversity were analyzed by investigating species composition, structure, and diversity of different restoration strategy and time community. The results showed that restoration strategy had no effect on species composition and diversity of family, genera, species, tree, shrub, and liana, however, restoration time had significant effect on species composition and diversity. Species richness of family, genera, species, tree, and shrub decreased at the early stage and then increased with community succession. Species richness showed an inverse J-shape with DBH increasing in all community types, while abundance showed single peak curve with DBH increasing and an inverse J-shape with height increasing. The correlation and regression analysis among restoration strategy and time and variables of community structure and diversity showed that there were significant correlation between restoration time and variables of community structure and diversity, and no significant correlation were found between restoration strategy and most of variables of community structure and diversity, which indicated that restoration time was the major factor that affects structure and diversity of community restored.
3.Species-area relationships are one of the most fundamental and most studied problems in all of ecology. However, whether the species-area relationships properties, especially species-area relationships slopes, remain constant through succession time, even though the ecosystem characteristics change on seasonal to evolutionary time scales, is a controversial problem. Focusing on disturbed broad-leaved evergreen forest ecosystems, we analyzed the species-area relationships in different succession stage communities (15a, 30a, and primary forest). Investigation was performed in Pu’er, Yunnan Province. Three plots (30 m×30 m) were located in the 15a, the 30a and the primary forest respectively. Each plot was divided into 36 subplots of 5 m×5 m for the vegetation surveys. Within each subplot all stems of height (H)>1.3 m were counted, measured and identified to the species level. We also recorded environment factors such as crown density, elevation, and slop. Species richness of tree, shrub, liana and total species in the 15a, the 30a and the primary forest were calculated respectively. The species-area relationships, the accumulate rate ( Z ) and the coefficients of determination (R2) of tree, shrub, liana and total species in each succession stage community were studied. We found that the number of total species, trees, shrubs, and lianas had a highly significant correlation with the sample area in different succession stage communities, the area explained over 94% of the total variation in the number of species, trees, shrubs, and lianas. The Z of total species(0.334) and trees(0.394) were the lowest in the 30a, whereas, the Z of shrubs(0.437) and lianas(0.326) were the lowest in the 15a. No significant difference were found for the intercepts of the species-area curve of species, trees, shrubs, and lianas among different succession stage communities, however, the coefficients of determination (R2) of the species-area curve of species and lianas were the highest in the primary forest. A fitted regression line of trees and shrubs explained 99.97% of the variance of Z-value in the 15a, but there were no significant correlation between trees, shrubs, lianas, and total species richness and Z-value in the 30a and the primary forest.
4.Niche and interspecific association is a focus of community dynamics. Interspecific interactions affect restoration and succession of community. Field data collection was biased on 0.81hm~2 plot (including 9 subplots) in different restoration stages (15a, 30a and primary monsoonal broad-leaved evergreen forest) which distributed in Pu'er city,Yunnan province. A series of techniques including niche breadth,niche overlap, the variance ratio (VR),chi-square test,association coefficient (AC) based on 2×2 contingency table were used to analyze change trend of tree population of different restoration stages. The results shown that:There was significant positive correlation (P<0.001) between the importance value and corresponding to niche breadth in 3 communites. 78.22% of species pairs had a niche overlap were between 0.4 and 0.8 in 15a,which indicated a strong resource utilization competition in dominant species. Simultaneously,community structure of 30a and primary forest was stability. The niche breadth increased with niche overlap,and vice versa,which reflected that the dominant species had strong competitiveness. There was a significant positive correlation in total on VR in the 15a and 30a,while the primary forest was no-significant negative association. For chi-square test,Most species pairs did not show significant association in 3 communities,which may show stronger independent distribution. The proportion of positive and negative association decreased from 15a to primary forest. There was a significant positive correlation between association coefficient and niche overlap. Generally speaking, the more niche overlap,the more association coefficient.
5.Soil seed bank and seedling bank is considered an important source for natural restoration in forest ecosystems. Soil seed bank is a dynamic biotic component of plant communities that represents the population’s memory in relation to selective events and seedling plays an important role for numerous tree species. Field data was based on 0.72hm~2 plot (including 8 subplots) in different restoration stages (15a, 30a) and primary monsoon broad-leaved evergreen forest which distribute in Caiyanghe nature reserve, Yixiang township and Xinfang reservoir, Puer city, Yunnan, China. Germination experiments carried out in the greenhouse, which lasted 7 months. On the basis of analyzing soil seed bank and seedling bank density and its relation to the above-ground vegetation. The results were summarized as follows: 2645 seeds germinated in all soil samples, 83 species in72 genera and 37 families were recorded. Total seed density of 30a (424±91.53 seeds/m~2) was significantly higher than primary forest (148±23 seeds/m~2). There was no significant difference between 15a (362.33±77.05 seeds/m~2) and other two types. Seed density of 0-2cm and 5-10cm layer was significantly higer than 2-5cm layer in pirmary forest, while no significant difference among 3 layers in 15a and 30a. Compositae was most abundant species in three community types,because of their wind dispersal way. Shannon-Wiener index of 30a was signifcantly higher than 15a and primary forest, but species richness was not significant difference in three types (including three layer). The seed density of herb was significantly higher than other life form in 3 community types. Seed density of perennial herb of 30a was significantly higher than 15a and primary and seed density of annual herb of 15a was signicantly higer than 30a and primary.and seed density of herb plant and fern plant of 30a was significantly higer than the primary forest. Dicranopteris pedata was most abundant species in restoration stages, especially in 30a. Annual herb offen emerged in early stages of restoration, such as Lindernia nummularifolia, which had short distance to tea park and farm land and had richness seed source. Castanopsis echidnocarpa is dominat species of monsoon broad-leaved evergreen forest, which was most abundant seedling of 30a and primary. Castanopsis calathiformis was most abundant seedling species of 15a. Tree seedlings were more than shrub seedlings. Tree seedling was main composition in all seedling bank, because of survival strategy of seed. Seedling bank density of primary was significantly higher than 15a, which had signifant negative correlation with soil seed bank density (P<0.05)and stand density(P<0.01), had signifantly positive correlation with soil bulk desity and Ph (P<0.05). The similarity coefficient (from 0.0375 to 0.1538) between soil seed bank and above-ground vegetation was generally low in three community types. Similarity coefficient between soil seed bank and herb was higher than soil seed bank and tree layer. Seedling bank was more similar with tree layer than soil seed bank, so forest restoration of monsoon broad-leaved evergreen forest mainly relyed on seed rain of tree layer.
6.Liana is important component of forest ecosystem, which usually affects restoration and succession of the communities. Relationship between liana and host tree has significant implications for forest conservation and sustainable management. Field data collection was based on 0.81hm~2 plot (including 9 subplots) in different restoration stages (15a, 30a and primary monsoonal broad-leaved evergreen forest distribute in Caiyanghe nature reserve, Yixiang Township and Xinfang reservoir, Pu’er city, Yunnan, China. On the basis of analyzing liana species richness, density, abundance, size structure, diversity indices and the climbing mechanism in the different restoration period, the relationship between liana and host tree was discussed. The results shown that: 1292 climbing lianas (DBH≥0.1cm), belonging to 64 species in 51 genera and 34 families were recorded in all plots. The liana richness,density(DBH<0.1cm), basal area at breast height and mean basal area at breast height in primary forest were significantly higher than restoration stages. There was no significant difference of density (DBH≥1cm), average DBH and average length between primary forest and 15a, though both were significantly higher than that of 30a. Species composition and DBH class distribution of lianas varied significantly in the three community types. Mucuna macrocarpa,Celastrus monospermus and Gnetum montanum are most abundant species. These species were found in all restoration stages and primary forest.Mucuna macrocarpa is an early successional species which needs intensive light environment and Gnetum montanum is a late successional species. Species-area curve showed that primary forest had higher species richness than 15a and 30a, which confirmed that primary forest played an important role in biodiversity conservation of lianas. Liana abundance decreased significantly while its DBH class increased. DBH of more than 95% lianas was less than 5cm. The lianas of large diameter class (DBH≥10cm) were found only in primary forest and 30a. One liana per host tree was most abundant in three community types. Host trees were more likely to be infested by lianas when their DBH was larger than 15cm in primary forest and liana-host tree relationship showed different trends in restoration stages. Large tree was more susceptible to be climbed by liana. There was a significant positive correlation (P<0.001) between the liana DBH and host tree DBH. DBH of liana increased with the growth of DBH of host tree in primary and 15a. Stem twining caused more mechanical damage in 4 climbing ways, and the damage for primary forest were significantly lower than that for restoration stages. Density of tendril climber showed that primary forest was undergoing a dynamic process. Species composition and abundance change of root climber and hook climber can be used as an indicator to reflect restoration level of monsoonal broad-leaved evergreen forest of post-disturbance. Primary forest had more Piper flaviflorum than 15a and 30a and Fissistigma acuminatissimum did not appeare in the restoration stages. Deforestation is main factor that affects liana species composition and distribution and reduces lianas abundance and richness significantly. Restoration time has an important effect on regeneration of lianas.
7.Vascular epiphytes are important components of species richness in monsoonal broad-leaved evergreen forest. Field data collection vascular epiphytes was based on 0.81hm~2 plot (including 9 subplots) in different restoration stages (15a, 30a and primary monsoonal broad-leaved evergreen forest distribute in Caiyanghe nature reserve, Yixiang Township and Xinfang reservoir, Pu’er city, Yunnan, China. On the basis of analyzing vascular epiphytes species richness, abundance, distribution, similarity coefficient, the relationship between epiphytes and phorophytes was discussed. The results shown that: (1) 3116 vascular epiphytes, belonging to 22 species in 20 genera and 9 families were recorded in all plots. (2) Diversity of vascular epiphytes was lower in restoration stages (15a, 5 spp. And 30a, 7 spp.) compared to primary forest (17 spp.). (3) Epiphytes species composition in primary forest and restoration stages differed markedly: community of 15a restoration harboured no orchid species but more fern species and primary forest hold more orchid species. Probably the families occurring only in primary forest sites of our study may be useful as bioindicators to determine the degree of restoration in monsoonal broad-leaved evergreen forest. (4) Epiphytes abundance were also lower in restoration stages. The decrease in species numbers and abundance as well as the differences in species composition was mainly due to the less diverse phorophyte structure and less differentiated microclimate in the disturbed and secondary vegetation compared to the primary forest. S?rensen coefficient between 15a and primary forest was higher than between 30a and primary forest. (5) The vascular epiphytes were clumped horizontally in 3 community types. Vertically, vascular epiphytes of 15a were mainly distributed at 0～5m and primary forest was up to 20m. (6) Vascular epiphytes species richness and abundance were both significantly positively correlated with host tree size but restoration stages didn’t have.
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