长白山针阔混交林种群结构及环境解释
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摘要
原始阔叶红松林是长白山西部地区的地带性顶级植被类型。20世纪以来,长白山原始阔叶红松林资源遭到严重破坏,形成大面积次生林群落。原始阔叶红松林的保护以及退化次生林群落的恢复任务已经迫在眉睫,但我们对该森林生态系统种群及群落结构的认识还十分有限。本文研究了长白山次生杨桦林、次生针阔混交林、原始阔叶红松林中树种空间结构,阐述了树种空间分布与局部生境因子之间的关系;分析了雌雄异株树种种群性比格局及种内、种间竞争作用,揭示了邻体竞争作用对树木径向生长的影响,进一步丰富和扩展了树种共存理论。本文旨在为长白山退化阔叶红松林的恢复和经营提供理论依据。具体研究结果简述如下:
1.长白山针阔混交林树种空间分布及环境解释
次生杨桦林群落:环境因子主要制约幼树和小树的空间分布,对大树影响较小。土壤水分和土壤pH值是影响树种分布的主要环境因子。环境因子(包括环境变量单独解释部分、环境变量与空间变量交互作用部分)对树木种群空间变异解释较差,纯粹空间变量(即空间变量单独解释部分)具有较强的种群空间变异解释能力。环境变量对群落结构空间变异解释量仅为8%。
次生针阔混交林群落:红松大树、蒙古栎大树、蒙古栎小树在空间上呈随机分布,其它树种(不同生长阶段个体)在绝大多数尺度上呈聚集分布。土壤水分及土壤pH是控制群落及树木种群空间结构的主要环境因子。环境因子对簇毛槭、假色槭、鱼鳞松和水曲柳种群空间分布影响较大,对其它树种影响相对较小。环境变量对群落结构空间变异解释量高达21%。
原始阔叶红松林群落:红松、蒙古栎、山荆子、怀槐在研究尺度水平上呈随机分布;水曲柳、紫椴、色木槭等树种在绝大多数研究尺度上为聚集分布。5个优势树种组成的10个种对中,红松与水曲柳、蒙古栎与水曲柳、红松与蒙古栎在空间上彼此独立,其余种对在局部空间尺度上呈正相关关系。5个优势树种中任意3个树种构成的6个种组中,红松-蒙古栎-水曲柳空间独立;其余5个种组(包括红松-蒙古栎-色木槭,红松-蒙古栎-紫椴,蒙古栎-色木槭-水曲柳,蒙古栎-色木槭-紫椴,色木槭-水曲柳-紫椴)在局部空间尺度上正相关。5个优势树种(红松、蒙古栎、紫椴、水曲柳、色木槭)整体上在1-12m呈空间正相关,其余尺度上空间独立。环境因子对树种及群落空间结构的解释均较差。
2.雌雄异株树种种群性比及径向生长格局
次生杨桦林及次生针阔混交林内水曲柳雌、雄植株胸径大小无显著差异:而原始阔叶红松林内雄树胸径显著大于雌树。次生林(次生杨桦林、次生针阔混交林)内性别不显著偏离1∶1;原始阔叶红松林内性比显著偏雄性。次生杨桦林及次生针阔混交林内水曲柳种群的雌树和雄树彼此空间分离;而原始阔叶红松林内雌树和雄树彼此空间独立。次生林内小雌树与大雄树、大雌树与小雄树彼此空间分离。这种性别间的竞争作用限制了某个性别的小植株出现在相对性别大植株周围。
山杨和水曲柳种群性比具有显著的密度依赖性,种群性比随着林分密度的增大而减小。不同林型内山杨和水曲柳种群性比对繁殖植株密度及种群密度反应不同:次生杨桦林样地中,繁殖植株密度及种群密度与山杨种群性比呈显著负相关,与水曲柳种群性比呈显著正相关。次生针阔混交林样地中,繁殖植株密度和种群密度与山杨性比呈显著正相关,与水曲柳种群性比呈显著负相关。
次生杨桦林中水曲柳雄树径向生长速率显著大于雌树。土壤水分、树木大小与雌树和雄树径向生长率显著正相关。雌树比雄树对气候变化响应更加敏感。相邻雌树的竞争作用对雌树径向生长表现为负效应,相邻雄树的竞争作用对雌树径向生长影响不显著。相邻雄树的竞争作用对雄树径向生长表现为负效应,相邻雌树的竞争作用对雄树径向生长影响不显著。因此,同性竞争(雄树-雄树、雌树-雌树)作用是导致雌雄植株径向生长差异的重要因子。
东北红豆杉雄树的径向生长速率显著大于雌树。雌、雄植株径向生长与邻体竞争强度显著负相关,且邻体竞争对雌树径向生长的负效应更大。雌、雄植株对气候因子的响应不同。与雄树相比,雌树径向生长对前一年(8、10月)温度更加敏感。
The original broad-leaved Korean pine forest is the zonal climax vegetation type in the western of Changbai Mountain.Since the 20th century,the resource of original broad-leaved Korean pine forest has been seriously destroyed in Changbai Mountain,which resulting in a large area of secondary forest community.Though the conservation of original broad-leaved Korean pine forest and restoration of degraded secondary forest community are extremely urgent,our knowledge about the structure characteristics of population and community in the forest ecosystem are still limited.Spatial structure of tree species was analyzed in secondary poplar-birch forests,secondary mixed conifer and broad-leaved forest and original broad-leaved Korean pine forest in Changbai Mountain in this paper. In order to enrich and extend the theories of trees coexistence,we studied the relationships between spatial distribution of trees and local habitat factors and the sex ratio pattern of dioecious tree species. And competition between same-sex trees and between opposite-sex trees was analyzed to reveal the effect of inter- and intra-sexual competition on radial growth.This paper aims to provide theories for restoration and management of broad-leaved Korean pine forest in Changbai Mountain.The brief results are as follows:
1.The spatial distribution of tree species and environmental interpretations in the mixed forest.
In secondary poplar-birch forests:environment factors regulated mainly the spatial pattern of saplings and small trees,but there was little effect on the distribution of big trees.Soil moisture and soil pH showed significant influence on the distribution pattern of most tree species,and soil nutrition (including total N,total P,total K) showed a little effect.Environment factors showed a poor interpretation on spatial variation of tree species(including variation interpreted by environment alone and variation interpreted by environment and space in common),but pure spatial variables exhibited strong interpretation power.And only 8%of the variation in community structure could be accounted by environment variables.
In secondary conifer and broadleaved mixed forest:dominant trees of Pinus koraiensis and Quercus mongolica and smaller trees of Q.mongolica tended to random spatial distributions at almost all scales.Soil moisture and soil pH were key environmental factors regulating the spatial structure of the community and individual tree species.Environment factors greatly affected the spatial distribution of Acer barbinerve,A.pseudo-sieboldianum,Picea jezoensis and Fraxinus mandshurica, but there was little effect on other tree species.Altogether 21%of the variation in community structure could be accounted by environment variables.
In original broad-leaved Korean pine forest:P.koraiensis,Q.mongolica,Malus baccata and Maackia amurensis were distributed randomly in the study plot,and A.mono,F.mandshurica and Tilia amurensis were in a clumped pattern at most studied scales.Among 10 species pairs which consisted of five dominant tree species,there were three species pairs(P.koraiensis-F.mandshurica; P.koraiensis-Q.mongolica;Q.mongolica-F.mandshurica) which showed independence.Others showed mutual attraction at several scales.Among six species groups which consisted of five dominant tree species,one species groups(P.koraiensis-Q.mongolica-F.mandshurica) showed independence.Other species groups(P.koraiensis-Q.mongolica-A.mono;P.koraiensis-Q.mongolica -T.amurensis;Q.mongolica-A.mono-F.mandshurica;Q.mongolica-A.mono-T.amurensis; A.mono-F.mandshurica-T.amurensis) showed mutual attraction at several scales.The groups of five tree species showed mutual attraction at 1-12m scales.Environment factors exhibited low explanations for spatial structure of tree populations and community.
2.Sex ratios and radial growth pattern in dioecious tree species
The diameters between genders were not significantly different in the secondary forests,but the diameters of male trees were significantly greater than those of female trees in the old-growth forest. The sex ratio did not significantly deviate from 1:1 in the secondary forests,but was male-biased in the old-growth forest.Spatial segregation between genders within F.mandshurica population was found in secondary forests,but the male trees and female trees were spatially independent in the old-growth forest.Significant segregation between small females versus big males,and big females versus small males was found in the secondary forests,this may also exhibit an intersexual competition effect,which limits the occurrence of small individuals in the vicinity of opposite-sexual big trees in dioecious tree species.
The numbers of males and females increased as increasing sample areas for dioecious tree species,F.mandshurica and Populus davidiana.Sex ratios exhibited significant density-dependent for two dioecious species.Sex ratios decreased with increasing local stand density for F.mandshurica and P.davidiana.Sex ratios showed different responses to density of reproductive individuals and population density in different forest types.The sex ratios of P.davidiana decreased as increasing stem density of reproductive individuals and population density,and those of F.mandshurica increased as increasing stem density of reproductive individuals and population density in secondary poplar-birch forests.But contrary results were found in secondary conifer and broadleaved mixed forest.
Male trees had a faster stem growth rate than female trees in secondary poplar-birch forests. Stem growth rates are positively affected by soil moisture and tree size for both genders.And female trees are more responsive to climate variations than male trees.The growth rates of female trees were negatively affected by neighboring competition from other female trees at all neighborhood distances between 1 and 10m,but were not significantly affected by male trees.The growth rates of male trees were negatively affected by neighboring competition from male trees,but were not significantly affected by female trees.These results suggest that intrasexuai competition(male-male competition and female-female competition) may cause difference in stem growth for both genders.
The radial growth rate of males was significantly larger than that of females in Taxus cuspidata. The radial growth rate associated negatively with neighboring competition for both sex.And female trees were affected more negatively by neighboring competition than males.Different response way to climate factors for males and females was found.Female trees tended to be more sensitive to the variations of temperature in previous August and October.
1.长白山针阔混交林树种空间分布及环境解释
次生杨桦林群落:环境因子主要制约幼树和小树的空间分布,对大树影响较小。土壤水分和土壤pH值是影响树种分布的主要环境因子。环境因子(包括环境变量单独解释部分、环境变量与空间变量交互作用部分)对树木种群空间变异解释较差,纯粹空间变量(即空间变量单独解释部分)具有较强的种群空间变异解释能力。环境变量对群落结构空间变异解释量仅为8%。
次生针阔混交林群落:红松大树、蒙古栎大树、蒙古栎小树在空间上呈随机分布,其它树种(不同生长阶段个体)在绝大多数尺度上呈聚集分布。土壤水分及土壤pH是控制群落及树木种群空间结构的主要环境因子。环境因子对簇毛槭、假色槭、鱼鳞松和水曲柳种群空间分布影响较大,对其它树种影响相对较小。环境变量对群落结构空间变异解释量高达21%。
原始阔叶红松林群落:红松、蒙古栎、山荆子、怀槐在研究尺度水平上呈随机分布;水曲柳、紫椴、色木槭等树种在绝大多数研究尺度上为聚集分布。5个优势树种组成的10个种对中,红松与水曲柳、蒙古栎与水曲柳、红松与蒙古栎在空间上彼此独立,其余种对在局部空间尺度上呈正相关关系。5个优势树种中任意3个树种构成的6个种组中,红松-蒙古栎-水曲柳空间独立;其余5个种组(包括红松-蒙古栎-色木槭,红松-蒙古栎-紫椴,蒙古栎-色木槭-水曲柳,蒙古栎-色木槭-紫椴,色木槭-水曲柳-紫椴)在局部空间尺度上正相关。5个优势树种(红松、蒙古栎、紫椴、水曲柳、色木槭)整体上在1-12m呈空间正相关,其余尺度上空间独立。环境因子对树种及群落空间结构的解释均较差。
2.雌雄异株树种种群性比及径向生长格局
次生杨桦林及次生针阔混交林内水曲柳雌、雄植株胸径大小无显著差异:而原始阔叶红松林内雄树胸径显著大于雌树。次生林(次生杨桦林、次生针阔混交林)内性别不显著偏离1∶1;原始阔叶红松林内性比显著偏雄性。次生杨桦林及次生针阔混交林内水曲柳种群的雌树和雄树彼此空间分离;而原始阔叶红松林内雌树和雄树彼此空间独立。次生林内小雌树与大雄树、大雌树与小雄树彼此空间分离。这种性别间的竞争作用限制了某个性别的小植株出现在相对性别大植株周围。
山杨和水曲柳种群性比具有显著的密度依赖性,种群性比随着林分密度的增大而减小。不同林型内山杨和水曲柳种群性比对繁殖植株密度及种群密度反应不同:次生杨桦林样地中,繁殖植株密度及种群密度与山杨种群性比呈显著负相关,与水曲柳种群性比呈显著正相关。次生针阔混交林样地中,繁殖植株密度和种群密度与山杨性比呈显著正相关,与水曲柳种群性比呈显著负相关。
次生杨桦林中水曲柳雄树径向生长速率显著大于雌树。土壤水分、树木大小与雌树和雄树径向生长率显著正相关。雌树比雄树对气候变化响应更加敏感。相邻雌树的竞争作用对雌树径向生长表现为负效应,相邻雄树的竞争作用对雌树径向生长影响不显著。相邻雄树的竞争作用对雄树径向生长表现为负效应,相邻雌树的竞争作用对雄树径向生长影响不显著。因此,同性竞争(雄树-雄树、雌树-雌树)作用是导致雌雄植株径向生长差异的重要因子。
东北红豆杉雄树的径向生长速率显著大于雌树。雌、雄植株径向生长与邻体竞争强度显著负相关,且邻体竞争对雌树径向生长的负效应更大。雌、雄植株对气候因子的响应不同。与雄树相比,雌树径向生长对前一年(8、10月)温度更加敏感。
The original broad-leaved Korean pine forest is the zonal climax vegetation type in the western of Changbai Mountain.Since the 20th century,the resource of original broad-leaved Korean pine forest has been seriously destroyed in Changbai Mountain,which resulting in a large area of secondary forest community.Though the conservation of original broad-leaved Korean pine forest and restoration of degraded secondary forest community are extremely urgent,our knowledge about the structure characteristics of population and community in the forest ecosystem are still limited.Spatial structure of tree species was analyzed in secondary poplar-birch forests,secondary mixed conifer and broad-leaved forest and original broad-leaved Korean pine forest in Changbai Mountain in this paper. In order to enrich and extend the theories of trees coexistence,we studied the relationships between spatial distribution of trees and local habitat factors and the sex ratio pattern of dioecious tree species. And competition between same-sex trees and between opposite-sex trees was analyzed to reveal the effect of inter- and intra-sexual competition on radial growth.This paper aims to provide theories for restoration and management of broad-leaved Korean pine forest in Changbai Mountain.The brief results are as follows:
1.The spatial distribution of tree species and environmental interpretations in the mixed forest.
In secondary poplar-birch forests:environment factors regulated mainly the spatial pattern of saplings and small trees,but there was little effect on the distribution of big trees.Soil moisture and soil pH showed significant influence on the distribution pattern of most tree species,and soil nutrition (including total N,total P,total K) showed a little effect.Environment factors showed a poor interpretation on spatial variation of tree species(including variation interpreted by environment alone and variation interpreted by environment and space in common),but pure spatial variables exhibited strong interpretation power.And only 8%of the variation in community structure could be accounted by environment variables.
In secondary conifer and broadleaved mixed forest:dominant trees of Pinus koraiensis and Quercus mongolica and smaller trees of Q.mongolica tended to random spatial distributions at almost all scales.Soil moisture and soil pH were key environmental factors regulating the spatial structure of the community and individual tree species.Environment factors greatly affected the spatial distribution of Acer barbinerve,A.pseudo-sieboldianum,Picea jezoensis and Fraxinus mandshurica, but there was little effect on other tree species.Altogether 21%of the variation in community structure could be accounted by environment variables.
In original broad-leaved Korean pine forest:P.koraiensis,Q.mongolica,Malus baccata and Maackia amurensis were distributed randomly in the study plot,and A.mono,F.mandshurica and Tilia amurensis were in a clumped pattern at most studied scales.Among 10 species pairs which consisted of five dominant tree species,there were three species pairs(P.koraiensis-F.mandshurica; P.koraiensis-Q.mongolica;Q.mongolica-F.mandshurica) which showed independence.Others showed mutual attraction at several scales.Among six species groups which consisted of five dominant tree species,one species groups(P.koraiensis-Q.mongolica-F.mandshurica) showed independence.Other species groups(P.koraiensis-Q.mongolica-A.mono;P.koraiensis-Q.mongolica -T.amurensis;Q.mongolica-A.mono-F.mandshurica;Q.mongolica-A.mono-T.amurensis; A.mono-F.mandshurica-T.amurensis) showed mutual attraction at several scales.The groups of five tree species showed mutual attraction at 1-12m scales.Environment factors exhibited low explanations for spatial structure of tree populations and community.
2.Sex ratios and radial growth pattern in dioecious tree species
The diameters between genders were not significantly different in the secondary forests,but the diameters of male trees were significantly greater than those of female trees in the old-growth forest. The sex ratio did not significantly deviate from 1:1 in the secondary forests,but was male-biased in the old-growth forest.Spatial segregation between genders within F.mandshurica population was found in secondary forests,but the male trees and female trees were spatially independent in the old-growth forest.Significant segregation between small females versus big males,and big females versus small males was found in the secondary forests,this may also exhibit an intersexual competition effect,which limits the occurrence of small individuals in the vicinity of opposite-sexual big trees in dioecious tree species.
The numbers of males and females increased as increasing sample areas for dioecious tree species,F.mandshurica and Populus davidiana.Sex ratios exhibited significant density-dependent for two dioecious species.Sex ratios decreased with increasing local stand density for F.mandshurica and P.davidiana.Sex ratios showed different responses to density of reproductive individuals and population density in different forest types.The sex ratios of P.davidiana decreased as increasing stem density of reproductive individuals and population density,and those of F.mandshurica increased as increasing stem density of reproductive individuals and population density in secondary poplar-birch forests.But contrary results were found in secondary conifer and broadleaved mixed forest.
Male trees had a faster stem growth rate than female trees in secondary poplar-birch forests. Stem growth rates are positively affected by soil moisture and tree size for both genders.And female trees are more responsive to climate variations than male trees.The growth rates of female trees were negatively affected by neighboring competition from other female trees at all neighborhood distances between 1 and 10m,but were not significantly affected by male trees.The growth rates of male trees were negatively affected by neighboring competition from male trees,but were not significantly affected by female trees.These results suggest that intrasexuai competition(male-male competition and female-female competition) may cause difference in stem growth for both genders.
The radial growth rate of males was significantly larger than that of females in Taxus cuspidata. The radial growth rate associated negatively with neighboring competition for both sex.And female trees were affected more negatively by neighboring competition than males.Different response way to climate factors for males and females was found.Female trees tended to be more sensitive to the variations of temperature in previous August and October.
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