地形生态位和扩散过程对暖温带森林群落构建重要性研究
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摘要
森林约覆盖地球陆地的31%,拥有世界陆地物种的三分之二以上,在维持全球生物多样性方面具有重要作用。森林生物多样性是人类衣食住行原材料的重要来源。在全球范围内保护森林生物多样性对人类生存具有重大意义。要使森林生物多样性得到合理有效的保护就需要人们从理论上来研究森林群落构建也即生物多样性维持的有关机制,了解影响群落构建的生态过程和因子,厘清众多因子对群落构建效应的贡献程度。暖温带森林在我国是受人类干扰最早和最为严重的森林类型之一。多年来该区域恢复起来的天然次生林在我国保护森林生物多样性,发挥生态系统功能等方面起着越来越重要的作用,是我国环境恢复和碳排放的重要增值点。研究东灵山地区天然次生林的生物多样性与群落构建机制,不仅有利于了解该地区生物多样性形成的原因,为未来暖温带森林生物多样性保护提供依据,也为解释暖温带森林物种共存与生物多样性维持提供理论支持。
本研究以东灵山地区的暖温带森林为研究对象,通过建立20hm2的暖温带落叶阔叶林动态监测样地,定位每棵树木个体的位置及物种信息,结合地形因素从局域尺度上探讨了暖温带森林群落物种组成与结构,地形因子对群落构建的影响及贡献,扩散过程对群落多样性的影响,以期较为全面和系统地了解暖温带森林群落构建中随机过程与确定性过程的影响及相对贡献。
(1)以20hm2样地内胸径≥1cm的所有木本植物为对象,分析了暖温带森林群落的组成和结构。结果表明,样地内木本植物有58种,隶属于18科33属。独立个体的总数为52,136,包括独立个体分枝的总数为103,284,全部为落叶树种。群落的区系类型以北温带成分居多,同时混有一些亚热带和热带成分,属典型的温带森林类型。群落优势种明显,个体数最多的前5个种的个体数占到总个体数的61%,前20个种占到92%,而其余38个种只占8%。群落成层现象明显,垂直结构由主林层(19个种)、次林层(18个种)和灌木层(21个种)组成。样地所有木本植物个体总径级分布呈倒“J”型,群落更新良好。主林层树种的径级结构近似于双峰或正态分布,而次林层和灌木层树种则表现出倒“J”型或“L”型。几个主要树种的空间分布表现出不同的分布格局,随着径级增大,聚集程度降低。空间分布格局显示主要优势种自身个体在其径级大小的空间分布上互补,不同径级的个体占据了样地内不同的空间位置。
(2)以20hm2样地内的两个优势种群为研究对象,对比分析了样地内两优势种群辽东栎(Quercus wutaishanica)与胡桃楸(Juglans mandshurica)的同物种不同生长阶段,不同物种相同生长阶段群体的地形生境差异性。结果表明,样地内辽东栎种群主要分布在地形坡度大、南坡、西坡、凸出及高海拔的位置上,成年树比幼树更偏好凹地形；胡桃楸种群集中分布在坡度大、北坡、东坡、凹陷及低海拔的地形上,成年树相比幼树偏好分布在南坡、东坡及凹陷的位置。Mann-Whitney U检验显示,辽东栎和胡桃楸各自幼树与成年树的地形因子之间都没有显著差异(P＞0.05),种群内两群体的地形分布范围基本一致；两物种成年树群体在海拔因子的选择上差异显著(P＜0.05),幼树群体除了海拔外,南北坡向上也表现了明显的差异(P＜0.05)。同种群不同生长阶段地形因子的选择显示了物种地形生境的限制及保守性,不同种群同生长阶段地形因子的选择显示了物种间空间分布格局及多样性的维持。这对于理解暖温带森林的物种共存及生物多样性维持具有重要的意义。
(3)以20hm2暖温带天然次生林为研究对象,基于20-ha样地DEM地图,提取海拔、坡度、坡向、凹凸度等地形因子,用GLM模型探讨了地形因子对群落物种丰富度与地上生物量的影响。结果表明,群落地上生物量与物种丰富度分别受不同地形因子的影响,群落地上生物量受Ea,El2,El3,Co3四个非线性因子的显著影响(p＜0.05),而物种丰富度则受Sl,El,El2,El3四个非线性因子的显著作用(p＜0.05),两者受海拔相关因子的影响最大,分别解释了两者3.99%和11.09%的变异。地形因子修正了物种丰富度-地上生物量之间的单峰曲线关系,使其朝着单调与直线化趋势变化。以后在探讨森林群落物种丰富度-生产力关系时,地形因子的修正作用应该受到重视。
(4)利用20hm2温带次生林森林样地植被数据检测了局域尺度上森林发育过程中群落扩散过程与生态位过程的变化,研究了它们在发育过程中的发生特点,区分了2类过程在群落多样性构建中的相对贡献。结果显示,局域尺度扩散过程与生态位过程因群落发育过程不同而有显著差异,其中生态位过程的变化强于扩散过程。扩散过程受森林发育过程的影响表现为前期稳定,后期下降。群落发育过程中扩散过程伴随了冠层稳定的骨架结构及分离的垂直结构,提高了群落物种垂直分布的均匀性,促进了物种共存。局域尺度扩散过程与生态位过程的交互效应是群落构建的主要动力,扩散过程作用要强于环境过滤过程。以上结果暗示除了空间因子等的被动调节外,中性过程与生态位过程对构建群落的相对贡献也受群落发育阶段的影响,群落自身具有主动调节构建的作用。
以上研究结果对于认识自然条件下暖温带森林的群落构建机制具有重要意义,不仅在局域尺度上明确,量化及区分了地形和扩散过程对暖温带森林群落构建的重要性,而且也是全球不同类型森林群落构建机制的有益补充,这对于指导暖温带森林资源和多样性保护及利用,地区环境改善等都具有重要的实践价值。
The forest covers about31%of the Earth land, has more than two-thirds of the world's terrestrial species, and plays an important role in maintaining global biodiversity. Forest biodiversity is an important source of raw materials of the basic necessities of human beings. The conservation of forest biodiversity on a global scale is of great significance for the survival of mankind. In order to be reasonable and effective protection of forest biodiversity, we need to study forest community assembly theory, understand ecological processes and impact factors and clarify their relative contribution on the community assembly. The warm temperate forests are one of the earliest and most serious forest types by human disturbance in China. The natural secondary forests recovered in the region over many years play an increasingly important role in the forest biodiversity conservations and ecosystem function services. They also are the important value-added points of environmental restoration and carbon emissions. The biodiversity and community assembly mechanism researches of the natural secondary forest in Dongling Mountain, are not only conducive to understand the biodiversity formation reasons in the region, and to provide a basis for the warm temperate forest biodiversity protection in the future, but also provide theoretical supports to explain the warm temperate forest species coexistence and biodiversity maintenance.
In this dissertation the warm temperate forests were studied in Dongling Mountain. The warm temperate forest species composition and structure, the impacts and contributions of topographic factors on community assembly, and the impacts of dispersal process on community diversity were explored through the establishment of a20hm2dynamic monitoring temperate deciduous broadleaf forest plots, positioning the locations and species information of each individuals, and combined with the terrain factors at local scale for a more comprehensive and systematic understanding of the impacts and the relative contributions of stochastic processes and deterministic process in the warm temperate forest communities assembly process.
(1) All free-standing woody plants at least1cm in diameter at breast height (DBH,1.3m above ground) as the research object, were mapped, tagged, and identified to species, and their geographic coordinates were recorded in the20-ha Dongling Mountain warm temperate deciduous broad-leaved forest dynamic monitoring plot. We address results on floristic characteristics, community composition, and vertical, size-class, and spatial structure of the Donglingshan forest plot (DLS). These datasets will serve as baseline information accessible to a wide range of future stud-ies. We tagged a total of52,136genotype individuals (103,284individuals including branches of genotype individuals), belonging to58species,33genera and18families. All of these tree species were deciduous. Floristic characteristics of the community suggested a temperate plant flora, including some subtropical and tropical species. There were very obvious dominant species in the plot. Five species comprised61%of all individuals, and20species comprised92%of all individuals, while the other38species comprised only8%of all individuals. Vertical structure was composed of an overstory layer (19species), midstory layer (18species), and a shrub layer (21species). The DBH size-class structure of all species in the plot generally fitted a "reverse J" distribu-tion, thereby indicating good regeneration across the community. The size-class structure of the main species in the overstory layer showed a bimodal or nearly normal distribution, while the most abundant species in the midstory and shrub layers showed "reverse J" or even "L" distributions. Spatial distribution patterns of the dominant species varied with size-class and scale and shifted from closer aggregation to looser aggregation from small to adult or old trees. The size-class spatial distribution patterns of the dominant species showed the different diameter levels of their own individuals occupied different spatial positions in the plot.
(2) A comparative analysis of the terrain habitats differences between the two dominant populations:Quercus wutaishanica and Juglans mandshurica was made according to the same species at different growth stages and the same growth stages of different species in the20hm2warm temperate deciduous broad-leaved forest dynamic monitoring plot in Dongling Mountain. The results showed that, Quercus wutaishanica population mainly distributed at the lager-slope, southern-aspect, western-aspect, more convex and high-altitude locations, and compared with the saplings,the adult trees preferred the concave terrain; Juglans mandshurica population distributed in the steep-slope, northern-aspect, eastern-aspect, more concave and low-altitude terrains, and compared with the saplings, the adult trees preferred at the southern-aspect, eastern-aspect and more concave positions in the plot. The Terrain factors of the sapling groups were not significantly different from the adult groups in each species (P>0.05), namely, their topographical features had the same ranges of variation. The altitude factors of two adult groups were significantly different (P<0.05), however, besides the altitude, the north-south aspects of the sapling groups also showed a significant difference (P<0.05). The topographical selection of the same population at the different growth stages showed the limitations and conservatisms of the species topographic habitats, and the topographical selection of the different populations at the same growth stages showed the spatial distribution patterns and the biodiversity maintenances among species. It is of great significance to understand species coexistence and biodiversity maintenance in the warm temperate forest.
(3) We studied topographic factors such as elevation, slope, aspect, convex effected on community richness and aboveground biomass with the GLM model to clarify the amendment effects of topographical factors on species richness-productivity relationship in the20-ha Donglingshan forest dynamic plot in a warm temperate deciduous broad-leaved secondary forest. The result showed that the terrain factors influencing aboveground biomass are different from those which affected species richness. Aboveground biomass was significantly influenced by four factors:Ea, El2, El3and Co3(p<0.05). Species richness was also influenced by four factors including Sl, El, El2and El3(p<0.05). The altitude-related factors were the biggest influence factors, respectively explaining3.99%and11.09%of the variation of aboveground biomass and species richness. The terrain factor amended the unimodal relationship between species richness and aboveground biomass, and the relationship was adjusted to the monotonous, straight line trend. In exploring the forest species richness and productivity relations in the future, amendments role of topographic factors should be taken seriously.
(4) The impacts of species dispersal and environmental filtering changes on tree richness during the community development scale were tested in the20-ha Dongling Mountain warm temperate deciduous broad-leaved forest dynamic monitoring plot. The relative contributions of species dispersal and environmental filtering to the community diversity were partitioned by using GLM approach. The results showed that, the species dispersal and environmental filtering in community development processes significantly changed, and environmental filtering had the stronger effects than species dispersal on community assembly. Species dispersal index stabilized in the early and declined in late stage of forest development process. The constant species dispersal index was accompanied by stability of the canopy profile structure in early stage, and the divergence of the vertical structure with the community development, which increased the heterogeneity of species vertical distribution and facilitated forest species coexistence. We also found the interactions of species dispersal and environmental filtering were the main driving forces for species diversities, and the species dispersal process was stronger than the environment filtering process to community assembly. This study suggested that in addition to the space factors changing, the relative contributions of neutral and niche processes to the community assembly were different during the community developmental stages.
The above findings are important for the understanding of the warm temperate forest community assembly under the natural conditions. These findings not only quantify and distinguish the relative importances of the terrain and the diffusion process of the warm temperate forest community assembly on a local scale, but also are the useful complements to community assembly mechanisms of the global different type's forest. These conclusions are conductive to better guide the conservation and utilization of warm temperate forest resources and biodiversity, the improvement of the regional environment.
本研究以东灵山地区的暖温带森林为研究对象,通过建立20hm2的暖温带落叶阔叶林动态监测样地,定位每棵树木个体的位置及物种信息,结合地形因素从局域尺度上探讨了暖温带森林群落物种组成与结构,地形因子对群落构建的影响及贡献,扩散过程对群落多样性的影响,以期较为全面和系统地了解暖温带森林群落构建中随机过程与确定性过程的影响及相对贡献。
(1)以20hm2样地内胸径≥1cm的所有木本植物为对象,分析了暖温带森林群落的组成和结构。结果表明,样地内木本植物有58种,隶属于18科33属。独立个体的总数为52,136,包括独立个体分枝的总数为103,284,全部为落叶树种。群落的区系类型以北温带成分居多,同时混有一些亚热带和热带成分,属典型的温带森林类型。群落优势种明显,个体数最多的前5个种的个体数占到总个体数的61%,前20个种占到92%,而其余38个种只占8%。群落成层现象明显,垂直结构由主林层(19个种)、次林层(18个种)和灌木层(21个种)组成。样地所有木本植物个体总径级分布呈倒“J”型,群落更新良好。主林层树种的径级结构近似于双峰或正态分布,而次林层和灌木层树种则表现出倒“J”型或“L”型。几个主要树种的空间分布表现出不同的分布格局,随着径级增大,聚集程度降低。空间分布格局显示主要优势种自身个体在其径级大小的空间分布上互补,不同径级的个体占据了样地内不同的空间位置。
(2)以20hm2样地内的两个优势种群为研究对象,对比分析了样地内两优势种群辽东栎(Quercus wutaishanica)与胡桃楸(Juglans mandshurica)的同物种不同生长阶段,不同物种相同生长阶段群体的地形生境差异性。结果表明,样地内辽东栎种群主要分布在地形坡度大、南坡、西坡、凸出及高海拔的位置上,成年树比幼树更偏好凹地形；胡桃楸种群集中分布在坡度大、北坡、东坡、凹陷及低海拔的地形上,成年树相比幼树偏好分布在南坡、东坡及凹陷的位置。Mann-Whitney U检验显示,辽东栎和胡桃楸各自幼树与成年树的地形因子之间都没有显著差异(P＞0.05),种群内两群体的地形分布范围基本一致；两物种成年树群体在海拔因子的选择上差异显著(P＜0.05),幼树群体除了海拔外,南北坡向上也表现了明显的差异(P＜0.05)。同种群不同生长阶段地形因子的选择显示了物种地形生境的限制及保守性,不同种群同生长阶段地形因子的选择显示了物种间空间分布格局及多样性的维持。这对于理解暖温带森林的物种共存及生物多样性维持具有重要的意义。
(3)以20hm2暖温带天然次生林为研究对象,基于20-ha样地DEM地图,提取海拔、坡度、坡向、凹凸度等地形因子,用GLM模型探讨了地形因子对群落物种丰富度与地上生物量的影响。结果表明,群落地上生物量与物种丰富度分别受不同地形因子的影响,群落地上生物量受Ea,El2,El3,Co3四个非线性因子的显著影响(p＜0.05),而物种丰富度则受Sl,El,El2,El3四个非线性因子的显著作用(p＜0.05),两者受海拔相关因子的影响最大,分别解释了两者3.99%和11.09%的变异。地形因子修正了物种丰富度-地上生物量之间的单峰曲线关系,使其朝着单调与直线化趋势变化。以后在探讨森林群落物种丰富度-生产力关系时,地形因子的修正作用应该受到重视。
(4)利用20hm2温带次生林森林样地植被数据检测了局域尺度上森林发育过程中群落扩散过程与生态位过程的变化,研究了它们在发育过程中的发生特点,区分了2类过程在群落多样性构建中的相对贡献。结果显示,局域尺度扩散过程与生态位过程因群落发育过程不同而有显著差异,其中生态位过程的变化强于扩散过程。扩散过程受森林发育过程的影响表现为前期稳定,后期下降。群落发育过程中扩散过程伴随了冠层稳定的骨架结构及分离的垂直结构,提高了群落物种垂直分布的均匀性,促进了物种共存。局域尺度扩散过程与生态位过程的交互效应是群落构建的主要动力,扩散过程作用要强于环境过滤过程。以上结果暗示除了空间因子等的被动调节外,中性过程与生态位过程对构建群落的相对贡献也受群落发育阶段的影响,群落自身具有主动调节构建的作用。
以上研究结果对于认识自然条件下暖温带森林的群落构建机制具有重要意义,不仅在局域尺度上明确,量化及区分了地形和扩散过程对暖温带森林群落构建的重要性,而且也是全球不同类型森林群落构建机制的有益补充,这对于指导暖温带森林资源和多样性保护及利用,地区环境改善等都具有重要的实践价值。
The forest covers about31%of the Earth land, has more than two-thirds of the world's terrestrial species, and plays an important role in maintaining global biodiversity. Forest biodiversity is an important source of raw materials of the basic necessities of human beings. The conservation of forest biodiversity on a global scale is of great significance for the survival of mankind. In order to be reasonable and effective protection of forest biodiversity, we need to study forest community assembly theory, understand ecological processes and impact factors and clarify their relative contribution on the community assembly. The warm temperate forests are one of the earliest and most serious forest types by human disturbance in China. The natural secondary forests recovered in the region over many years play an increasingly important role in the forest biodiversity conservations and ecosystem function services. They also are the important value-added points of environmental restoration and carbon emissions. The biodiversity and community assembly mechanism researches of the natural secondary forest in Dongling Mountain, are not only conducive to understand the biodiversity formation reasons in the region, and to provide a basis for the warm temperate forest biodiversity protection in the future, but also provide theoretical supports to explain the warm temperate forest species coexistence and biodiversity maintenance.
In this dissertation the warm temperate forests were studied in Dongling Mountain. The warm temperate forest species composition and structure, the impacts and contributions of topographic factors on community assembly, and the impacts of dispersal process on community diversity were explored through the establishment of a20hm2dynamic monitoring temperate deciduous broadleaf forest plots, positioning the locations and species information of each individuals, and combined with the terrain factors at local scale for a more comprehensive and systematic understanding of the impacts and the relative contributions of stochastic processes and deterministic process in the warm temperate forest communities assembly process.
(1) All free-standing woody plants at least1cm in diameter at breast height (DBH,1.3m above ground) as the research object, were mapped, tagged, and identified to species, and their geographic coordinates were recorded in the20-ha Dongling Mountain warm temperate deciduous broad-leaved forest dynamic monitoring plot. We address results on floristic characteristics, community composition, and vertical, size-class, and spatial structure of the Donglingshan forest plot (DLS). These datasets will serve as baseline information accessible to a wide range of future stud-ies. We tagged a total of52,136genotype individuals (103,284individuals including branches of genotype individuals), belonging to58species,33genera and18families. All of these tree species were deciduous. Floristic characteristics of the community suggested a temperate plant flora, including some subtropical and tropical species. There were very obvious dominant species in the plot. Five species comprised61%of all individuals, and20species comprised92%of all individuals, while the other38species comprised only8%of all individuals. Vertical structure was composed of an overstory layer (19species), midstory layer (18species), and a shrub layer (21species). The DBH size-class structure of all species in the plot generally fitted a "reverse J" distribu-tion, thereby indicating good regeneration across the community. The size-class structure of the main species in the overstory layer showed a bimodal or nearly normal distribution, while the most abundant species in the midstory and shrub layers showed "reverse J" or even "L" distributions. Spatial distribution patterns of the dominant species varied with size-class and scale and shifted from closer aggregation to looser aggregation from small to adult or old trees. The size-class spatial distribution patterns of the dominant species showed the different diameter levels of their own individuals occupied different spatial positions in the plot.
(2) A comparative analysis of the terrain habitats differences between the two dominant populations:Quercus wutaishanica and Juglans mandshurica was made according to the same species at different growth stages and the same growth stages of different species in the20hm2warm temperate deciduous broad-leaved forest dynamic monitoring plot in Dongling Mountain. The results showed that, Quercus wutaishanica population mainly distributed at the lager-slope, southern-aspect, western-aspect, more convex and high-altitude locations, and compared with the saplings,the adult trees preferred the concave terrain; Juglans mandshurica population distributed in the steep-slope, northern-aspect, eastern-aspect, more concave and low-altitude terrains, and compared with the saplings, the adult trees preferred at the southern-aspect, eastern-aspect and more concave positions in the plot. The Terrain factors of the sapling groups were not significantly different from the adult groups in each species (P>0.05), namely, their topographical features had the same ranges of variation. The altitude factors of two adult groups were significantly different (P<0.05), however, besides the altitude, the north-south aspects of the sapling groups also showed a significant difference (P<0.05). The topographical selection of the same population at the different growth stages showed the limitations and conservatisms of the species topographic habitats, and the topographical selection of the different populations at the same growth stages showed the spatial distribution patterns and the biodiversity maintenances among species. It is of great significance to understand species coexistence and biodiversity maintenance in the warm temperate forest.
(3) We studied topographic factors such as elevation, slope, aspect, convex effected on community richness and aboveground biomass with the GLM model to clarify the amendment effects of topographical factors on species richness-productivity relationship in the20-ha Donglingshan forest dynamic plot in a warm temperate deciduous broad-leaved secondary forest. The result showed that the terrain factors influencing aboveground biomass are different from those which affected species richness. Aboveground biomass was significantly influenced by four factors:Ea, El2, El3and Co3(p<0.05). Species richness was also influenced by four factors including Sl, El, El2and El3(p<0.05). The altitude-related factors were the biggest influence factors, respectively explaining3.99%and11.09%of the variation of aboveground biomass and species richness. The terrain factor amended the unimodal relationship between species richness and aboveground biomass, and the relationship was adjusted to the monotonous, straight line trend. In exploring the forest species richness and productivity relations in the future, amendments role of topographic factors should be taken seriously.
(4) The impacts of species dispersal and environmental filtering changes on tree richness during the community development scale were tested in the20-ha Dongling Mountain warm temperate deciduous broad-leaved forest dynamic monitoring plot. The relative contributions of species dispersal and environmental filtering to the community diversity were partitioned by using GLM approach. The results showed that, the species dispersal and environmental filtering in community development processes significantly changed, and environmental filtering had the stronger effects than species dispersal on community assembly. Species dispersal index stabilized in the early and declined in late stage of forest development process. The constant species dispersal index was accompanied by stability of the canopy profile structure in early stage, and the divergence of the vertical structure with the community development, which increased the heterogeneity of species vertical distribution and facilitated forest species coexistence. We also found the interactions of species dispersal and environmental filtering were the main driving forces for species diversities, and the species dispersal process was stronger than the environment filtering process to community assembly. This study suggested that in addition to the space factors changing, the relative contributions of neutral and niche processes to the community assembly were different during the community developmental stages.
The above findings are important for the understanding of the warm temperate forest community assembly under the natural conditions. These findings not only quantify and distinguish the relative importances of the terrain and the diffusion process of the warm temperate forest community assembly on a local scale, but also are the useful complements to community assembly mechanisms of the global different type's forest. These conclusions are conductive to better guide the conservation and utilization of warm temperate forest resources and biodiversity, the improvement of the regional environment.
引文
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