基于Ripley <i>L</i>函数的云冷杉针阔混交林优势树种幼树的空间格局分析

许芳泽; 孙海龙; 史景宁; 贺丹妮; 王福增; 向玮

doi:10.12171/j.1000-1522.20230237

基于Ripley L函数的云冷杉针阔混交林优势树种幼树的空间格局分析

1.
北京林业大学森林资源和环境管理国家林草局重点实验室，北京 100083
2.
吉林省长白山森工集团汪清林业分公司，吉林延吉 133200

基金项目: 国家重点研发计划（2022YFD2200502-04），国家自然科学基金项目（32160366）。

详细信息

作者简介:
许芳泽，博士生。主要研究方向：森林生长收获模型。Email：Fangzexu1217@163.com　地址：100083 北京市海淀区北京林业大学林学院

责任作者:
向玮，博士，教授。主要研究方向：森林生长收获模型。Email：wxiang@bjfu.edu.cn　地址：同上。

中图分类号: S791.142；S791.18；S791.247
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出版历程
- 收稿日期: 2023-09-18
- 修回日期: 2023-12-24
- 录用日期: 2024-07-21
- 网络出版日期: 2024-10-11
- 刊出日期: 2024-10-24

Spatial pattern analysis of dominant tree species saplings in spruce-fir coniferous and broadleaved mixed forests based on Ripley L function

1.
State Key Laboratory of Forest Resource and Environmental Management, Beijing Forestry University, Beijing 100083, China
2.
Wangqing Forestry Branch of Changbai Mountain Forest Industry Group, Yanji 133200, Jilin, China

摘要

摘要:
目的
林下幼树的生长和分布通常决定着森林未来林分的结构组成和生长动态，探究优势树种幼树的空间分布格局及种间关联性，以期为森林生态系统的恢复和森林演替动态的预测提供科学依据。
方法
以吉林省汪清县金沟岭林场1 hm²云冷杉针阔混交林样地内的5个优势树种（臭冷杉、鱼鳞云杉、红松、紫椴和五角槭）幼树为研究对象，分别使用一元Ripley L和二元Ripley L 函数对5种幼树的种间空间关联性以及幼树与中树、成树间的种内空间关联性进行分析。
结果
（1）样地内共调查到幼树471株，数量上表现为臭冷杉（357） > 鱼鳞云杉（47） > 紫椴（23） > 五角槭（12） > 红松（14），臭冷杉占据绝对优势。（2）臭冷杉、鱼鳞云杉和紫椴幼树的空间格局在0 ~ 50 m范围内，随尺度增加表现为聚集–均匀–随机的分布模式；五角槭以聚集分布为主，仅在23 ~ 28 m尺度上呈随机分布；红松在14 ~ 21 m、31 ~ 36 m及39 ~ 50 m尺度上呈聚集分布，在其余尺度上均呈随机状态分布。（3）幼树种间关系在中小尺度（0 ~ 36 m）上以显著的空间正相关为主，在大尺度（36 ~ 50 m）以不相关为主，仅臭冷杉幼树在42 ~ 50 m上与其余树种呈显著空间负相关，表明5种幼树间保持着良好的共存的关系。（4）种内空间关联性在0 ~ 50 m尺度上以正相关为主，仅臭冷杉幼树与成树在42 ~ 50 m较大尺度上存在显著空间负相关，表明种内促进作用明显。
结论
在水平结构上，影响该林分幼树空间分布格局及其关联性的主要因素是种子的扩散限制及环境异质性，幼树在小尺度上以聚集分布为主；在垂直结构上，主林层对幼树具有明显的“庇护效应”，5种幼树种内、种间均存在着良好的互利共生的关系。因此，在天然云冷杉针阔混交次生林的恢复中，建议根据林分经营需要，在0 ~ 36 m的小尺度上进行以上幼树的聚集性补植补种，聚集程度为臭冷杉 > 云杉 > 红松，以提高幼树的存活率，促进其天然更新。
- 混交林 /
- 幼树 /
- Ripley L函数 /
- 空间分布格局 /
- 空间关联性
Abstract:
Objective
The structure and dynamics of dominant tree species saplings determine the structure and growth dynamics of future stand. Investigating the spatial distribution patterns and interspecies associations of dominant tree seedlings is of significant theoretical and practical importance for the restoration of forest ecosystems and the prediction of forest succession dynamics.
Method
This study focused on five dominant tree species in a 1-hectare mixed coniferous and broadleaved forest plot in the Jingouling Forest Farm, Wangqing County, Jilin Province of northeastern China. The spatial correlations between five kinds of saplings and the intraspecific spatial correlations between saplings and juveniles and adults were analyzed by univariate Ripley L and bivariate Ripley L functions.
Result
A total of 471 saplings were investigated in the plot, and the quantity size was as follows: Abies nephrolepis (357) > Picea jezoensis var. komarovii (47) > Tilia amurensis (23) > Acer pictum subsp. mono (22) > Pinus koraiensis (14). Abies nephrolepis occupied an absolute advantage. The spatial patterns of Abies nephrolepis, Picea jezoensis var. komarovii and Tilia amurensis saplings with the increase of scale conformed to the aggregation-uniform-random distribution pattern; the scales of aggregated distribution of Pinus koraiensis were 14−21 m, 31−36 m and 39−50 m, and other scales were random. Acer pictum subsp. mono distributed randomly in 0−50 m and distributed aggregately at few scales (23−38 m). The interspecific relationships between saplings were mostly positively correlated with the significant indigenous space (0−36 m), uncorrelated on the 36−42 m scale, and only saplings of Abies nephrolepis were negatively correlated with the significant indigenous space on a large scale (42−50 m), indicating that the five saplings maintained a good relationship of mutual benefit and symbiosis. The intraspecific correlation was mainly spatial positive correlation, and there was a significant negative correlation between saplings and adult trees in large scale (42−50 m).
Conclusion
The spatial distribution and associations of saplings in this forest stand are primarily influenced by seed dispersal limitations and environmental heterogeneity at the horizontal structure level. Saplings predominantly exhibit an aggregated distribution at smaller scales. In the vertical structure, the main forest layer provides a significant “shelter effect” for the saplings. There exists a beneficial symbiotic relationship both within and between the five sapling species. Therefore, in the restoration of natural mixed secondary forest of spruce and fir, it is feasible to conduct seeding of these saplings at a small scale of 0−36 m, based on forest management needs. The degree of aggregation should prioritize fir, followed by spruce, and then pine, to enhance the survival rate of saplings and promote their nature regeneration.
- mixed forests /
- saplings /
- Ripley L function /
- spatial distribution pattern /
- spatial correlation

HTML全文

图 1 优势树种幼树的空间分布格局

实线代表L (r)的实际值，灰色区域代表用零模型进行 99 次模拟形成的置信区间。下同。Solid line stands for L (r) value of spatial pattern，and gray area represents the confidence interval formed after 99 simulations with the zero model. The same below.

Figure 1. Spatial distribution patterns of saplings of dominant tree species

下载: 全尺寸图片幻灯片

图 2 优势树种幼树的种间空间关联性

Figure 2. Interspecific spatial association of saplings of dominant tree species

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图 3 优势树种幼树与中、成树的种内空间关联性

a中树 Medium tree；b成树Adult tree

Figure 3. Intra-specific spatial correlations between saplings, medium and adult trees of dominant tree species

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表 1 树种组成及优势树种幼树的重要值

Table 1 Tree species composition and importance values (IV) of dominant tree species saplings

树种 Tree species	总体数量 Total amount	幼树数量 Number of sapling	乔木层重要值 Important value of tree layer	幼树重要值 Important value of sapling
臭冷杉 Abies nephrolepis	581	357	0.263	0.619
鱼鳞云杉 Picea jezoensis var. komarovii	163	47	0.117	0.112
紫椴 Tilia amurensis	144	23	0.116	0.100
五角槭 Acer pictum subsp. mono	84	22	0.100	0.080
红松 Pinus koraiensis	100	14	0.112	0.055
其他 Others	364	8	0.292	0.034
所有树种 Total tree species	1 436	471	1	1

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表 2 各优势树种不同生长阶段的株数及占比

Table 2 Plant number and proportion at different growth stages of each dominant tree species

树种 Tree species	幼树 Sapling		中树 Medium tree		成树 Adult tree
树种 Tree species	株密度/（株·hm⁻²） Plant density/(tree·ha⁻¹)	百分比 Proportion/%	株密度/（株·hm⁻²） Plant density/(tree·ha⁻¹)	百分比 Proportion/%	株密度/（株·hm⁻²） Plant density/(tree·ha⁻¹)	百分比 Proportion/%
臭冷杉 Abies nephrolepis	357	61.4	193	33.2	31	5.3
鱼鳞云杉 Picea jezoensis var. komarovii	47	28.8	99	60.7	17	10.4
红松 Pinus koraiensis	14	14.6	51	53.1	31	32.3
紫椴 Tilia amurensis	23	16.0	96	66.7	25	17.4
五角槭 Acer pictum subsp. mono	22	26.2	52	61.9	10	11.9
所有树种 Total tree species	463	43.4	491	46.0	114	10.6

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