长白山阔叶红松林植物多样性及其群落空间结构研究
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摘要
森林是陆地生态系统的主体，是人类赖以生存的重要自然资源，对调节生态平衡具有不可替代的作用。长白山阔叶红松林与世界其它地区的森林一样，在遭受长期的采伐干扰后，形成了小面积的原始林与大面积的次生林并存的局面。如何恢复林分质量偏低的次生林是森林可持续经营面临的迫切任务。为此，本文从阔叶红松林及其次生林的时间格局变化和群落空间结构变化两个方面研究了次生林与原始林在组织、结构方面的差异和差距，并探讨了生态位互补作用对维持原始林稳定性的重要作用，为以生态学知识为基础的可持续森林经营提供了理论依据。兹分述如下：
在长白山自然保护区内及长白山白河林业局光明林场，用Topcon全站仪共设置了四块永久样地。样地类型及面积大小依次为：阔叶红松林样地(100m×100m)；近熟林样地(260m×200m)；中龄林样地(260m×200m)，成熟林样地(160m×100m)。对各样地内所有的乔木、对阔叶红松林的全部灌木及中龄林样地的部分灌木进行了每木相对坐标定位及每木调查，对阔叶红松林、中龄林及成熟林内的草本按机械布点设置(1m×1m)样方并在生长季节内按月份调查其多样性变化。
调查表明：阔叶红松林带共有维管植物265种，其中乔木47种，灌木31种，草本187种。
阔叶红松林、近熟林、成熟林和中龄林中的树种数分别为20、29、34和40种。各林型的乔木多样性高低与它们的物种丰富度一致。中龄林阳性先锋树种优势度明显，演替层树种组成最丰富。未来顶级群落中的优势种群在次生林中的相对多度高于原始林，相对优势度低于原始林，相对频度变化较小，其中水曲柳(Fraxinus mandshurica Rupr.)在各个林型中的各项指标变化最大，色木槭(Acer mono Maxim.)在各个林型中的表现最稳定。
阔叶红松林与中龄林的灌木分别为18种和30种。中龄林林下灌木种类、平均密度及生物多样性远远高于阔叶红松林。
中龄林、成熟林及阔叶红松林林下草本植物种数依次为158种、137种和103种。林下草本层多样性在生长季节内都经历了由低到高，再由高到低的过程。遗传距离检验证明，同一个群落，草本层在各个月份之间存在显著差异，其遗传距离差异从大到小依次为：5月与6月；6月与7月；8月与9月；7月与8月。各个样地间草本层在5月没有显著差异，在其它各月间差异显著。
各种类型的次生林在乔木层、灌木层以及草本层的多样性均高于阔叶红松林，主要是因为在次生林的各个层次喜光植物的比例都有不同程度的增加。
Ripley'(K)点格局分析表明：阔叶红松林中，红松(Pinus koraiensis Sieb. et Zucc.)在0—13m、水曲柳(F. mandshurica Rupr.)在7-9m、糠椴(Tilia mandshurica Rupr. et Maxim.)在0—47m、紫椴(T. amurensis Rupr.)在0—7m及9—30m的尺度内为聚集分布，其余尺度为随机分布，蒙古栎(Quercus mongolica Fish. ex Turcz.)在所有尺度上呈随机分布。上层乔木优势种群关联不显著的种对包括：紫椴—水曲柳，蒙古栎—红松；在小尺度内负关联的种对包括：紫椴—蒙古栎，紫椴—水曲柳；在较大尺度内显著负关联的种对是：糠椴—紫椴。亲缘关系接近的树种负关联的尺度范围大，生态习性接近的优势种在小尺度上负关联，林冠层优势种总体上呈随机分布的格局，物种之间由于生态位分化使关联不显著。色木槭(A. mono Maxim.)及白牛槭(A. mandshuricum Maxim.)在所有尺度上呈负相关关系。色木槭和白牛槭在0—7m范围内呈显著负关联。青楷槭(A. tegmentosum Maxim.)、假色槭(A. pseudo-sieboldianum(Pax.)Kom.)和簇毛槭(A. barbinerve Maxim.)在较大尺度内都呈聚集分布，且都在较小尺度内达到了最大聚集强度，这些物种在群落中存在小尺度聚集分布，多个小尺度斑块复合形成更大尺度的聚集分布的特点。槭属类树种相互间都存在较大尺度内的空间负关联关系。林冠上层乔木和色木槭之间的关联度不显著，上层乔木和白牛槭之间呈显著的负相关。上层乔木和下层乔木之间，中层乔木与下层乔木之间，整体上呈现先负关联后随机的相互关系。差别只在于关联的尺度大小。灌木层优势种在较大尺度内均为聚集分布。暴马丁香(Syringa reticulata(Blume)Hara var. mandshurica(Maxim.)Hara)和毛榛子(Corylus mandshurica Maxim. et Rupr.)在0—12m内显著负关联，东北山梅花(Philadelphus schrenkii Rupr.)与东北溲疏(Beutzia amurensis(Regel)Airy-Shaw)在0—15m尺度内显著负关联；东北山梅花与毛榛子在0—37m内显著负关联；暴马丁香与东北山梅花仅在0—6m尺度范围内显著负关联。
角尺度分析表明：阔叶红松林及其次生林整体上呈聚集分布的格局。
中龄林、成熟林、近熟林以及阔叶红松林混交度大小依次为：0.54、0.63、0.64、0.53。研究表明：在同一个群落内，上层乔木的混交度水平高，下层乔木的混交度水平低；种子繁殖的树种的混交度水平高，萌蘖繁殖的混交度水平低，两种繁殖方式兼有的树种混交度水平较高。在次生林中，先锋树种的混交度水平高，未来顶级群落的优势树种的混交度总体偏低，在原始林中，林冠上层的优势树种的混交度高。
相邻木结构单元大小比数分析表明：在同一个群落，上层乔木在生长上占优势，下层乔木在生长上处于被压状态；在次生演替的相对早期阶段，先锋树种在生长上处于优势，未来顶级群落的树种暂时处于被压状态；在次生演替相对中期阶段，阳性树种和未来顶级群落的优势树种共同占优势；在顶级群落中，先锋树种被淘汰，优势树种在生长上占有绝对优势。
对阔叶红松林及其次生林的生物多样性变化、优势种群的点格局分析以及相邻木结构单元的空间结构分析，定量的阐明了在次生林中，先锋树种在小尺度内同种个体共存的现象很少见，先锋树种在生长上处于优势，未来顶级群落的优势树种由于遭到采伐等干扰，又由于林下有不同规模的幼苗和幼树存在，混交度水平较低，生长上不占优势，随着次生林的恢复，先锋树种及先锋草本植物逐渐被淘汰，生物多样性下降，未来顶级群落中的优势树种在生长上逐渐处于优势地位，相对优势度逐渐升高，又由于自疏作用，上层乔木的混交度水平得到提高，同层树种之间以及不同层树种之间生态位逐渐分化，种群格局趋于随机分布，群落趋于稳定。
The forest is the principal part of the terrestrial ecosystem, an important natural resource for the people to live on, which functions indispensably on the adjustment of the eco-balance. The situation is that the phenomenon of the coexistence of the small-area virgin forest and the large-area secondary forest in the region of Mount Changbai is similar to that of any other region, because of cutting and other mankind disturbance. So, how to recover the low-quality secondary forest is the stringent problem faced with the sustainable development of forest for us to solve. Therefore, the paper focuses on the difference of the organization, structure between the virgin forest and secondary forest, with the example of the change of temporal pattern and dimensional structure on the broad-leaved korean pine forest and its secondary forest. The paper also discussed the reciprocal function of niche is of great importance for the maintenance of the stability of the virgin forest. The detail is as follows:
In the region of the Guangming Forest Centre of the Baihe Forest Bureau and the natural reserve of Mount Changbai, four fixed-plots have been set up with Topcon total station. The type and size of the plots is respectively as:
Broad-leaved korean pine forest plot (100m×100m) Closely mature forest plot (260m × 200m ) Middle-year forest plot (260m × 200m ) Secondary mature forest plot (160m × 100m)
The coordinate positioning and research of each single tree are carried out on the whole of the shrubs of the broad-leaved korean pine forest and part of the shrubs of the middle-year forest. The research of the diversity change in the growing seasons is also been performed with the sample setting of 1m × 1m of broad-leaved korean pine forest, middle-year forest and secondary mature forest.
The research shows that there are vascular plants of 265 species, in which there are 47 tree species, 31 shrub species and 187 herb species.
The number of tree species is 20、 29、 34 and 40 in the broad-leaved korean pine forest, closely mature forest, secondary mature forest, middle-year forest respectively. The abundance of the tree diversity of each type of forests above is consistent with that of their species. The dominance of the pioneer species is obvious in the middle-year forest. And the species in the succession layer is most abundant. The relative abundance of the dominant plant population in the future climax community is more than that in the virgin forest, while its relative dominance is less and the change of the relative frequency is minor. Fraxinus mandshurica Rupr.is one of them, all of whose indexes changes most. And the indexes of Acer mono Maxim stays most stable.
The species of shrubs are 18 and 30 in the broad-leaved korean pine forest and middle-year forest respectively. The species of the shrubs under the trees, its average density and biodiversity of the middle-year forest are far higher than that of the broad-leaved korean pine forest
The species of herbs are 158, 137 and 103 under the middle-year forest, secondary mature forest and broad-leaved korean pine forest respectively. The diversity of the herb layer experiences the process from the lower class to higher class, then vice versa in the growing season. The test of the hereditary distances demonstrates that there are obvious differences of herb layer between months in the same community. The detailed figures of the difference of the hereditary distances arranged from the longest to the lowest are as follows: May and June, June and July, August and September, July and August. What's more, the herb layers of different plots have no obvious differences in May, while with great difference in other months.
The diversity of tree, shrub and herb layers is much more abundant in the types of secondary forests than that in the broad-leaved korean pine forest. The main cause is that there is some increase of the propotion of the sun plants in each of these layers in the secondary forests.
Ripley's K (d) function shows that Pinus koraiensis is in the aggregated distribution in the range of 0-13m, F. mandshurica 7-9m, Tilia mandshurica 0-47m and T. amurensis 0-7m and 9-30m.They are in the random distribution in other ranges. Quercus mongolica is in the random distribution of all the ranges. The pairs of the dominant species in the upper stratum whose relativity is minor include: T. amurensis - F. mandshurica, and Q. mongolica - P. koraiensis; The pairs that are negative association in a narrow rage include: T. amurensis - Q. mongolica, and T. amurensis - F. mandshurica; While in a wide range includes T. mandshurica - T. amurensis. The species who have close relationship with each other is more negatively associated in a wide range, and the dominant species whose ecological behavior are more close to each other shows negative association in a narrow rage. The dominant species of the crown layer demonstrate a pattern of random distribution, and the interrelationship of species is not obvious because of the polarization of the niches. A. mono shows obvious negative association in all levels of range. So does with A. mandshuricum. The two are obviously negatively associated with each other in the range from 0 to 7m. A. tegmentosum, A. pseudo-sieboldianum and A. barbinerve are in the aggregated distribution in a wider range, and they have reached the maximal aggregated intensity in the minor range. There exists narrow-ranged aggregated distribution among these species, and then many narrow-ranged patches constitute a wider-ranged aggregated distribution. The species of Q. mongolica have a wider-ranged spatial negative association each other. There is no significant relationship between the upper stratum in the crown layer and A. mono, while the upper stratum is negatively associated with A, mandshuricum obviously. There exists the interrelationship that first negative association and then random distribution in the whole in the example of the upper stratum and the lower stratum, the middle stratum and the lower stratum. The only difference lies in the range size of the interrelationship. Syringa reticulata(Blume)Hara. var.mandshurica and Corylus mandshurica only shows obvious negative association in the scale from 0 to 12m., Philadelphus schrenkii and Deutzia amurensis from 0 to 15m, C. mandshurica and P. schrenkii from 0 to 37m, and S. reticulata(Blume)Hara var.mandshurica and P. schrenkii only from 0 to 6m.
The analysis of the stand neighborhood pattern demonstrates that the broad-leaved korean pine forest and its secondary forests show the aggregated distribution in a whole.
The magnitude of the minglings of the middle-year forest, secondary mature forest, closely mature forest and the broad-leaved korean pine forest shows as follows : 0.54,0.63,0.64,0.53.
The research shows that in the same community, the higher the level of the minglings in the upper stratum, the lower the level of the minglings in the lower stratum; the higher the level of the minglings in the species of the seed-breeding, the lower the level of the mingling the stump-breeding. When a species has the both breeding ways , the level of mingling will be higher. In the secondary forests, the level of the mingling of eh pioneer species is higher, while the dominant species of the future climax community lower. In the virgin forests, the dominant species of the upper crown layers have higher level of mingling.
The analysis of the neighborhood comparison of the nearest neighborhood stand structure unit shows that in a community, the upper stratum take advantage over the lower stratum in growth, which accords with the practice; in the relatively early phase of the secondary succession, pioneer species dominate in growth, and the species of the future climax community are restrained temporarily; in the relatively middle phase, pioneer species and dominant plants of the future climax community share the advantage; in the climax community, pioneer species is first to be eliminated, while the dominant species take the sheer advantage.
The research includes the analysis of the biodiversity change of the broad-leaved korean pine forest and its secondary forests, Ripley's K (d) function of the dominant plant population and the spatial structure of the neighborhood stand structure unit. Those findings clarify quantificationally that in the secondary forests, co-existence of the same types of the pioneer species is rare, while the pioneer species take advantage in growth. Therefore, the level of mingling of the dominant species in the future climax community is low because of the cutting and the existence of the young trees, and it is alse inferior to others in growth. With the recovery of the secondary forests, the pioneer species and herbs are been eliminating, and there comes the decline in biodiversity. The dominant species of the future climax community is beginning to take the advantage in growth and its relative dominance increases. But due to "self-thinning", the level of mingling of the upper trees is growing, while the niche between the species of the same layers and the different layers gradually differentiates, the population pattern become randomly distributed and the community grows to be stable.
在长白山自然保护区内及长白山白河林业局光明林场，用Topcon全站仪共设置了四块永久样地。样地类型及面积大小依次为：阔叶红松林样地(100m×100m)；近熟林样地(260m×200m)；中龄林样地(260m×200m)，成熟林样地(160m×100m)。对各样地内所有的乔木、对阔叶红松林的全部灌木及中龄林样地的部分灌木进行了每木相对坐标定位及每木调查，对阔叶红松林、中龄林及成熟林内的草本按机械布点设置(1m×1m)样方并在生长季节内按月份调查其多样性变化。
调查表明：阔叶红松林带共有维管植物265种，其中乔木47种，灌木31种，草本187种。
阔叶红松林、近熟林、成熟林和中龄林中的树种数分别为20、29、34和40种。各林型的乔木多样性高低与它们的物种丰富度一致。中龄林阳性先锋树种优势度明显，演替层树种组成最丰富。未来顶级群落中的优势种群在次生林中的相对多度高于原始林，相对优势度低于原始林，相对频度变化较小，其中水曲柳(Fraxinus mandshurica Rupr.)在各个林型中的各项指标变化最大，色木槭(Acer mono Maxim.)在各个林型中的表现最稳定。
阔叶红松林与中龄林的灌木分别为18种和30种。中龄林林下灌木种类、平均密度及生物多样性远远高于阔叶红松林。
中龄林、成熟林及阔叶红松林林下草本植物种数依次为158种、137种和103种。林下草本层多样性在生长季节内都经历了由低到高，再由高到低的过程。遗传距离检验证明，同一个群落，草本层在各个月份之间存在显著差异，其遗传距离差异从大到小依次为：5月与6月；6月与7月；8月与9月；7月与8月。各个样地间草本层在5月没有显著差异，在其它各月间差异显著。
各种类型的次生林在乔木层、灌木层以及草本层的多样性均高于阔叶红松林，主要是因为在次生林的各个层次喜光植物的比例都有不同程度的增加。
Ripley'(K)点格局分析表明：阔叶红松林中，红松(Pinus koraiensis Sieb. et Zucc.)在0—13m、水曲柳(F. mandshurica Rupr.)在7-9m、糠椴(Tilia mandshurica Rupr. et Maxim.)在0—47m、紫椴(T. amurensis Rupr.)在0—7m及9—30m的尺度内为聚集分布，其余尺度为随机分布，蒙古栎(Quercus mongolica Fish. ex Turcz.)在所有尺度上呈随机分布。上层乔木优势种群关联不显著的种对包括：紫椴—水曲柳，蒙古栎—红松；在小尺度内负关联的种对包括：紫椴—蒙古栎，紫椴—水曲柳；在较大尺度内显著负关联的种对是：糠椴—紫椴。亲缘关系接近的树种负关联的尺度范围大，生态习性接近的优势种在小尺度上负关联，林冠层优势种总体上呈随机分布的格局，物种之间由于生态位分化使关联不显著。色木槭(A. mono Maxim.)及白牛槭(A. mandshuricum Maxim.)在所有尺度上呈负相关关系。色木槭和白牛槭在0—7m范围内呈显著负关联。青楷槭(A. tegmentosum Maxim.)、假色槭(A. pseudo-sieboldianum(Pax.)Kom.)和簇毛槭(A. barbinerve Maxim.)在较大尺度内都呈聚集分布，且都在较小尺度内达到了最大聚集强度，这些物种在群落中存在小尺度聚集分布，多个小尺度斑块复合形成更大尺度的聚集分布的特点。槭属类树种相互间都存在较大尺度内的空间负关联关系。林冠上层乔木和色木槭之间的关联度不显著，上层乔木和白牛槭之间呈显著的负相关。上层乔木和下层乔木之间，中层乔木与下层乔木之间，整体上呈现先负关联后随机的相互关系。差别只在于关联的尺度大小。灌木层优势种在较大尺度内均为聚集分布。暴马丁香(Syringa reticulata(Blume)Hara var. mandshurica(Maxim.)Hara)和毛榛子(Corylus mandshurica Maxim. et Rupr.)在0—12m内显著负关联，东北山梅花(Philadelphus schrenkii Rupr.)与东北溲疏(Beutzia amurensis(Regel)Airy-Shaw)在0—15m尺度内显著负关联；东北山梅花与毛榛子在0—37m内显著负关联；暴马丁香与东北山梅花仅在0—6m尺度范围内显著负关联。
角尺度分析表明：阔叶红松林及其次生林整体上呈聚集分布的格局。
中龄林、成熟林、近熟林以及阔叶红松林混交度大小依次为：0.54、0.63、0.64、0.53。研究表明：在同一个群落内，上层乔木的混交度水平高，下层乔木的混交度水平低；种子繁殖的树种的混交度水平高，萌蘖繁殖的混交度水平低，两种繁殖方式兼有的树种混交度水平较高。在次生林中，先锋树种的混交度水平高，未来顶级群落的优势树种的混交度总体偏低，在原始林中，林冠上层的优势树种的混交度高。
相邻木结构单元大小比数分析表明：在同一个群落，上层乔木在生长上占优势，下层乔木在生长上处于被压状态；在次生演替的相对早期阶段，先锋树种在生长上处于优势，未来顶级群落的树种暂时处于被压状态；在次生演替相对中期阶段，阳性树种和未来顶级群落的优势树种共同占优势；在顶级群落中，先锋树种被淘汰，优势树种在生长上占有绝对优势。
对阔叶红松林及其次生林的生物多样性变化、优势种群的点格局分析以及相邻木结构单元的空间结构分析，定量的阐明了在次生林中，先锋树种在小尺度内同种个体共存的现象很少见，先锋树种在生长上处于优势，未来顶级群落的优势树种由于遭到采伐等干扰，又由于林下有不同规模的幼苗和幼树存在，混交度水平较低，生长上不占优势，随着次生林的恢复，先锋树种及先锋草本植物逐渐被淘汰，生物多样性下降，未来顶级群落中的优势树种在生长上逐渐处于优势地位，相对优势度逐渐升高，又由于自疏作用，上层乔木的混交度水平得到提高，同层树种之间以及不同层树种之间生态位逐渐分化，种群格局趋于随机分布，群落趋于稳定。
The forest is the principal part of the terrestrial ecosystem, an important natural resource for the people to live on, which functions indispensably on the adjustment of the eco-balance. The situation is that the phenomenon of the coexistence of the small-area virgin forest and the large-area secondary forest in the region of Mount Changbai is similar to that of any other region, because of cutting and other mankind disturbance. So, how to recover the low-quality secondary forest is the stringent problem faced with the sustainable development of forest for us to solve. Therefore, the paper focuses on the difference of the organization, structure between the virgin forest and secondary forest, with the example of the change of temporal pattern and dimensional structure on the broad-leaved korean pine forest and its secondary forest. The paper also discussed the reciprocal function of niche is of great importance for the maintenance of the stability of the virgin forest. The detail is as follows:
In the region of the Guangming Forest Centre of the Baihe Forest Bureau and the natural reserve of Mount Changbai, four fixed-plots have been set up with Topcon total station. The type and size of the plots is respectively as:
Broad-leaved korean pine forest plot (100m×100m) Closely mature forest plot (260m × 200m ) Middle-year forest plot (260m × 200m ) Secondary mature forest plot (160m × 100m)
The coordinate positioning and research of each single tree are carried out on the whole of the shrubs of the broad-leaved korean pine forest and part of the shrubs of the middle-year forest. The research of the diversity change in the growing seasons is also been performed with the sample setting of 1m × 1m of broad-leaved korean pine forest, middle-year forest and secondary mature forest.
The research shows that there are vascular plants of 265 species, in which there are 47 tree species, 31 shrub species and 187 herb species.
The number of tree species is 20、 29、 34 and 40 in the broad-leaved korean pine forest, closely mature forest, secondary mature forest, middle-year forest respectively. The abundance of the tree diversity of each type of forests above is consistent with that of their species. The dominance of the pioneer species is obvious in the middle-year forest. And the species in the succession layer is most abundant. The relative abundance of the dominant plant population in the future climax community is more than that in the virgin forest, while its relative dominance is less and the change of the relative frequency is minor. Fraxinus mandshurica Rupr.is one of them, all of whose indexes changes most. And the indexes of Acer mono Maxim stays most stable.
The species of shrubs are 18 and 30 in the broad-leaved korean pine forest and middle-year forest respectively. The species of the shrubs under the trees, its average density and biodiversity of the middle-year forest are far higher than that of the broad-leaved korean pine forest
The species of herbs are 158, 137 and 103 under the middle-year forest, secondary mature forest and broad-leaved korean pine forest respectively. The diversity of the herb layer experiences the process from the lower class to higher class, then vice versa in the growing season. The test of the hereditary distances demonstrates that there are obvious differences of herb layer between months in the same community. The detailed figures of the difference of the hereditary distances arranged from the longest to the lowest are as follows: May and June, June and July, August and September, July and August. What's more, the herb layers of different plots have no obvious differences in May, while with great difference in other months.
The diversity of tree, shrub and herb layers is much more abundant in the types of secondary forests than that in the broad-leaved korean pine forest. The main cause is that there is some increase of the propotion of the sun plants in each of these layers in the secondary forests.
Ripley's K (d) function shows that Pinus koraiensis is in the aggregated distribution in the range of 0-13m, F. mandshurica 7-9m, Tilia mandshurica 0-47m and T. amurensis 0-7m and 9-30m.They are in the random distribution in other ranges. Quercus mongolica is in the random distribution of all the ranges. The pairs of the dominant species in the upper stratum whose relativity is minor include: T. amurensis - F. mandshurica, and Q. mongolica - P. koraiensis; The pairs that are negative association in a narrow rage include: T. amurensis - Q. mongolica, and T. amurensis - F. mandshurica; While in a wide range includes T. mandshurica - T. amurensis. The species who have close relationship with each other is more negatively associated in a wide range, and the dominant species whose ecological behavior are more close to each other shows negative association in a narrow rage. The dominant species of the crown layer demonstrate a pattern of random distribution, and the interrelationship of species is not obvious because of the polarization of the niches. A. mono shows obvious negative association in all levels of range. So does with A. mandshuricum. The two are obviously negatively associated with each other in the range from 0 to 7m. A. tegmentosum, A. pseudo-sieboldianum and A. barbinerve are in the aggregated distribution in a wider range, and they have reached the maximal aggregated intensity in the minor range. There exists narrow-ranged aggregated distribution among these species, and then many narrow-ranged patches constitute a wider-ranged aggregated distribution. The species of Q. mongolica have a wider-ranged spatial negative association each other. There is no significant relationship between the upper stratum in the crown layer and A. mono, while the upper stratum is negatively associated with A, mandshuricum obviously. There exists the interrelationship that first negative association and then random distribution in the whole in the example of the upper stratum and the lower stratum, the middle stratum and the lower stratum. The only difference lies in the range size of the interrelationship. Syringa reticulata(Blume)Hara. var.mandshurica and Corylus mandshurica only shows obvious negative association in the scale from 0 to 12m., Philadelphus schrenkii and Deutzia amurensis from 0 to 15m, C. mandshurica and P. schrenkii from 0 to 37m, and S. reticulata(Blume)Hara var.mandshurica and P. schrenkii only from 0 to 6m.
The analysis of the stand neighborhood pattern demonstrates that the broad-leaved korean pine forest and its secondary forests show the aggregated distribution in a whole.
The magnitude of the minglings of the middle-year forest, secondary mature forest, closely mature forest and the broad-leaved korean pine forest shows as follows : 0.54,0.63,0.64,0.53.
The research shows that in the same community, the higher the level of the minglings in the upper stratum, the lower the level of the minglings in the lower stratum; the higher the level of the minglings in the species of the seed-breeding, the lower the level of the mingling the stump-breeding. When a species has the both breeding ways , the level of mingling will be higher. In the secondary forests, the level of the mingling of eh pioneer species is higher, while the dominant species of the future climax community lower. In the virgin forests, the dominant species of the upper crown layers have higher level of mingling.
The analysis of the neighborhood comparison of the nearest neighborhood stand structure unit shows that in a community, the upper stratum take advantage over the lower stratum in growth, which accords with the practice; in the relatively early phase of the secondary succession, pioneer species dominate in growth, and the species of the future climax community are restrained temporarily; in the relatively middle phase, pioneer species and dominant plants of the future climax community share the advantage; in the climax community, pioneer species is first to be eliminated, while the dominant species take the sheer advantage.
The research includes the analysis of the biodiversity change of the broad-leaved korean pine forest and its secondary forests, Ripley's K (d) function of the dominant plant population and the spatial structure of the neighborhood stand structure unit. Those findings clarify quantificationally that in the secondary forests, co-existence of the same types of the pioneer species is rare, while the pioneer species take advantage in growth. Therefore, the level of mingling of the dominant species in the future climax community is low because of the cutting and the existence of the young trees, and it is alse inferior to others in growth. With the recovery of the secondary forests, the pioneer species and herbs are been eliminating, and there comes the decline in biodiversity. The dominant species of the future climax community is beginning to take the advantage in growth and its relative dominance increases. But due to "self-thinning", the level of mingling of the upper trees is growing, while the niche between the species of the same layers and the different layers gradually differentiates, the population pattern become randomly distributed and the community grows to be stable.
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