林分结构多样性研究进展

doi:10.11707/j.1001-7488.20200916

摘要/Abstract

摘要：

林分结构是森林经营与分析中的重要因子，提高林分结构多样性和复杂性是实现森林生态系统生物多样性维持和增加的基础。林分结构多样性表达方法是生态学和林学学科研究的重点议题，本研究从林分结构组成多样性、林木大小分化多样性和林木空间分布多样性3方面综述林分结构多样性研究进展。林分结构组成多样性常以林分组成属性为基本统计单位，以植物群落学中的α物种多样性指数为基础来表达；基于相邻木关系的林分组成结构多样性则通过描述林分中树种的隔离程度来体现。林木大小分化多样性以林木个体胸径、树高和冠幅等因子作为比较指标，大小变异系数、Gini系数、胸径分布偏度、大小分化度等常用来描述林木大小分化程度。林木空间分布多样性的实质是林木在水平空间上的位置分布格局多样性；点格局分析方法能够很好描述林木位置随尺度变化的差异，但对数据要求较高，需要精确的林木定位数据；标记二阶特征可以体现林分中树种和大小随尺度变化的分布情况，对生态学过程和假说具有一定分析和解释能力，但无法表达林分树种组成的丰富程度和林木大小的变异程度，也需要精确的林木定位数据。林分结构复杂性优先于林分结构多样性，二者既有联系又有区别，林分结构多样性常作为林分结构复杂性的替代指标。林分结构是一个多维度概念，现有林分结构多样性表达方法缺乏体现林分结构多维性的指标，探索林分结构多样性和复杂性表达方法仍是一项富有挑战的工作。未来林分结构多样性或复杂性指标的构建需要考虑林分结构多维性，同时要考虑多种属性的权重问题；另外，还要与具体的森林经营活动相结合，能够指导林分结构调整，最终实现增加林分结构多样性、维持和增加森林生态系统多样性的目的。

关键词: 林分结构, 多样性, 组成, 大小分化, 空间分布

Abstract:

Stand structure is an important factor in forest management and analysis. Enhancing the diversity and complexity of stand structure is the basis for maintaining and increasing the biodiversity of forest ecosystems. The method of expressing diversity of stand structure has been a key issue in the study of ecology and forestry disciplines. This paper summarized the research progress of stand structure diversity from the composition diversity of forest stands,the tree size differentiation diversity and the spatial distribution diversity. The composition structural diversity of forest stands is often based on composition attribute and the α species diversity index in plant community science. The diversity of the stand structure based on the neighbourhood relationship reflects the compositional diversity by describing the degree of segregation of the tree species in the stand. The size differentiation diversity of forest trees was compared with the factors such as DBH,tree height and crown width. The indicators of size coefficient,Gini coefficient,DBH distribution and size differentiation were used to describe the size differentiation diversity of stand. The spatial diversity of forest trees is essentially the diversity of spatial distribution patterns of trees in horizontal space. The point pattern analysis method could be used describe the differences of forest position with the change of scale,but it requires accurate tree position data. The mark second-order structural characteristic can reflect the distribution of tree species and the size with the change of scale in the stand and have certain ability to analyze and explain the ecological process and hypothesis,but they cannot express the richness of the tree species and the degree of tree size variation,it also needs accurate tree position data. The complexity of the stand structure takes precedence over the diversity of the stand structure,and the two concepts are related and different. The diversity of the stand structure is often used as an alternative indicator of the complexity of the stand. In the future,the construction of forest structure diversity or complexity indicators needs to take into account the multidimensional characteristic of the forest structure,at the same time,the weighting of multiple attributes must be considered. In addition,the specific forest management activities should be considered in the process of the construction of the forest structure diversity index,and which could be used to guide the stand structural adjustment and finally achieve the goal of increasing the diversity of stand structure and maintaining and increasing the diversity of the forest ecosystem.

Key words: stand structure, diversity, composition, size differentiation, spatial distribution

中图分类号:

S750

赵中华,惠刚盈. 林分结构多样性研究进展[J]. 林业科学, 2020, 56(9): 143-152.

Zhonghua Zhao,Gangying Hui. Advances in Structural Diversity of Stand Structure[J]. Scientia Silvae Sinicae, 2020, 56(9): 143-152.

图/表 4

表1

表2

常用基于相邻木关系的林分组成结构多样性指数"

指数名称Index	计算公式Formula	说明Definition
Pielou分隔指数 Pielou segregation index (Pielou，1961)	$S = 1 - \frac{{N(b + c)}}{{ms + nr}}$	N为样地中林木株数，b、c为不同种相邻木的株数；m、n为作参照树的种A、种B单木数；r、s为作最近相邻木的种A、种B单木数。S越大，林分中树种隔离程度越大N is the total number of trees in plot, b and c is the number of different tree species respectively, m and n is the number of specie A and species B respectively, r and s is the number of nearest neighbor of species A and species B. The larger the value of S, the greater the segregation of the tree species in the stand
简单混交度 Simple mingling (Gadow et al., 1993)	${M_i} = \frac{1}{n}\sum\limits_{j = 1}^n {{v_{ij}}} $	n为相邻木株数，如果第i株林木的第j株相邻木与i为同种，则v_ij=0；否则v_ij=1。M_i越大，由n株相邻木组成的结构单元混交程度越大n is the number of neighbor trees, if i-th tree is the same tree species as j-th neighbor tree, v_ij=0, else, v_ij=1. The larger the value of M_i，the greater the mixed of structural units which consisting of n adjacent woods
林分平均混交度 Stand mean mingling(惠刚盈等，2001)	$\bar M = \frac{1}{N}\sum {{M_i}} $	N为样地中林木株数, M_i为样地内第i株个体的混交度。${\bar M}$越大，林分树种隔离程度越小，树种混交程度越高N is the total number of trees in plot, M_i is the mingling of i-th tree. The larger the value of ${\bar M}$，the smaller the segregation of the tree species, the greater the mixed of stand
树种多样性混交度 Tree species diversity mingling (汤孟平等，2004)	${M_i} = \frac{{{n_i}}}{{{n^2}}}\sum\limits_{j = 1}^n {{v_{ij}}} $	n_i为对象木i的n株相邻木中不同树种个数，n为相邻木数目，如果第i株林木的第j株相邻木与i为同种，则v_ij=0；否则v_ij=1。M_i越大，林分混交程度越高，树种多样性越大n_i is the tree species number in n neighbor trees of object tree i, if i-th tree is the same tree species as j-th neighbor tree, v_ij=0, else, v_ij=1. The larger the value of M_i, the greater the mixed of tree species, the higher the diversity of tree species of the stand
林分修正混交度 Modify stand mingling(惠刚盈等，2007)	${\bar M^\prime } = \frac{1}{{5N}}\sum {\left({{M_i}n_i^\prime } \right)} $	N为样地中林木株数, M_i为样地内第i株个体的混交度，n′_i为第i株树所处结构单元中的树种个数。′越大，林分混交程度越高，树种多样性越大N is the total number of trees in plot, M_i is the mingling of i-th tree.n′_i is tree species number in structure unit of i-th tree. The larger the value of M′, the greater the mixed of tree species, the higher the diversity of tree species of the stand
树种空间多样性指数(TSS) Tree species spatial diversity index (Hui et al., 2011)	${\rm{TSS}} = \sum\limits_{{\rm{sp}} = 1}^n {\left[ {\frac{1}{{5N}}\sum\limits_{i = 1}^{{N_{{\rm{sp}}}}} {({M_i}{S_i}} } \right]} $	N为样地中林木株数, M_i为样地内第i株个体的混交度，n为树种数，N_sp为树种为sp的个体数，S_i为结构单元中的树种数，i为以树种sp为参照树的结构单元数。TSS越大，林分树种空间多样性越高N is the total number of trees in plot, M_i is the mingling of i-th tree. n is the number of tree species, N_sp is the number of species sp, S_i is tree species number in structure unit, i is the number of structure unit which object tree is sp. The higher the value of TSS, it indicates that the tree species spatial diversity of stand is higher
全混交度 Complete mingling (汤孟平等，2012)	${M_{{c_i}}} = \frac{1}{2}\left({{D_i} + \frac{{{c_i}}}{{{n_i}}}} \right){M_i}$	M_i为简单混交度，n_i为最近相邻木株数，c_i为对象木最近相邻木中非同种的个数，D_i为空间结构单元的Simpson指数。M_{c_i}越高，林分混交程度越高M_i is simple mingling, n_i is number of near neighbor trees, c_i is the number of different tree species with object tree, D_i is Simpson index of spatial structure unit. The higher the value of M_{c_i}, the greater the mixed of stand

表2

表3

常用林木大小分化多样性指数(以林木胸径大小为例)"

指数名称Index	计算公式Formula	说明Definition
大小变异系数Size coefficient of variation (Porkress，2004)	${\rm{cv}}\frac{{\sqrt {\frac{{\sum\limits_{i = 1}^n {{{\left({{\rm{DB}}{{\rm{H}}_i} - \overline {{\rm{DBH}}} } \right)}^2}} }}{{n - 1}}} }}{{\overline {{\rm{DBH}}} }}$	n为样地中林木株数, $\overline{{\rm{DBH}}}$为平均胸径DBH的算术平均值。cv越大，林分中树种大小分化程度越大n is the total number of trees in plot, $\overline{{\rm{DBH}}}$ is the arithmetic mean DBH. The higher the value of cv, it indicates that the tree size differentiation of stand is greater
胸径分布偏度Diameter distribution skewness (Porkress，2004)	${\rm{sk}} = \frac{{\sum\limits_{i = 1}^n {{{\left({{\rm{DB}}{{\rm{H}}_i} - \overline {{\rm{DBH}}} } \right)}^3}} }}{{(n - 1) \cdot {\rm{sd}}_{{\rm{DBH}}}^3}}$	n为样地中林木株数, $\overline{{\rm{DBH}}}$为平均胸径DBH的算术平均值，sd为标准差, sk < 0右偏，sk=0对称分布，sk>0左偏。sk绝对值越大，林木大小分化程度越大n is the total number of trees in plot, $\overline{{\rm{DBH}}}$ is the arithmetic mean DBH, sd is standard deviation, sk < 0 means negatively skewed, sk=0 means symmetric, sk>0 means positively skewed. The greater the absolute value of sk, the greater of trees size differentiation
Gini系数 Gini coefficient(Gini，1972)	${\rm{GC}} = \frac{{\sum\limits_{i = 1}^n {(2i - n - 1)} {\rm{B}}{{\rm{A}}_i}}}{{\sum\limits_{i = 1}^n {{\rm{B}}{{\rm{A}}_i}} (n - 1)}}$	n为样地中林木株数，BA_i为林木单株断面积。GC越大，林木大小分化程度越大n is the total number of trees in plot, BA_i is basal area of tree i. The greater the value of GC, the greater of trees size differentiation
大小分化度Diameter differentiation(Gadow et al., 1993)	${T_i} = 1 - \frac{{\min \left({{d_i}, {d_1}} \right)}}{{\max \left({{d_i}, {d_1}} \right)}}$	d_i为参照树直径，d₁为最近1株相邻木直径。T_i越大，林木大小分化程度越大d_i is diameter of object tree, d₁ is diameter of the nearest neighbor tree. The greater the value of T_i, the greater of trees size differentiation
大小比数Neighborhood comparison(惠刚盈等，1999)	${U_i} = \frac{1}{n}\sum\limits_{j = 1}^n {{v_{ij}}} $	n为相邻木株数，如果第i株林木的第j株相邻木较i小，则v_ij=0；否则v_ij=1。U_i越小，由n株相邻木组成结构单元中参照树的优势越大n is number of near neighbor trees, if i-th tree is large than the j-th neighbor tree, v_ij=0, else, v_ij=1. The smaller the value of U_i，the greater of reference tree of structural units which consisting of n adjacent woods
Shannon-Wiener指数 Shannon-Wiener index (Shannon，1949)	${H^\prime } = - \sum\limits_{i = 1}^S {{p_i}} \ln {p_i}$	p_i为第i个径阶林木株数占总林木株数的比例，S为径阶总数。H′越大，林木大小分化程度越高p_i is the percentage of the number of trees of the i-th diameter class in the total number of trees, S is the number of diameter class.The larger the value of H′, the higher of trees size differentiation
Simpson多样性指数 Simpson diversity index (Simpson，1949)	$D = 1 - \sum\limits_1^S {{{\left({{p_i}} \right)}^2}} $	S为径阶总数，p_i为第i个径阶林木株数占总林木株数的比例；D越大，林木大小分化程度越高S is the number of diameter class，p_i is the percentage of the number of trees of the i-th diameter class in the total number of trees. The larger the value of D, the higher of trees size differentiation

表3

表4

常用点格局分析函数"

函数名称Function name	函数形式Function	说明Definition
Ripley’K函数Ripley’Kfunction(Ripley, 1977)	$K(d) = A\sum\limits_{i = 1}^n {\sum\limits_{j = 1}^n {\frac{{{\delta _{ij}}(d)}}{{{n^2}}}} } $	A为样地面积，n为林木个数，d为树i与树j间的距离，当d_ij < d时，δ_ijd=1；当d_ij≥d时，δ_ijd=0 A is area of plot, n is the total number of trees in plot, d is the distance between i and j, when d_ij < d, δ_ijd=1；when d_ij≥d，δ_ijd=0
基Ripley’K函数于L-函数L-function based on Ripley’K function (Besag, 1977)	$\begin{array}{*{20}{c}}{{L_1}(r) = \sqrt {{{\rm{ \mathit{ π} }}^{ - 1}}K(r)} }\\{{L_2}(r) = \sqrt {{{\rm{ \mathit{ π} }}^{ - 1}}K(r)} - r}\end{array}$	K(r)为Ripley’K函数，r为树i与树j间的距离K(r) is Ripley’K function, r is the distance between i and j
双相关函数Pair correlation function	$g(r) = \frac{{{\rho ^{(2)}}(x, y){\rm{d}}x{\rm{d}}y}}{{{\lambda ^2}}}$	λ²为完全随机分布格局中发现被距离r分隔的2个点的概率；ρ⁽²⁾(x, y)dxdy为实际观察窗口中发现2个点的概率λ² is the probability of finding two points separated by r in the complete random distribution pattern; ρ⁽²⁾ (x, y)dxdy is probability of finding two points in actual observation window
O-ring统计O-ring statistics	Or=λg(r)	λ为圆环上点的概率密度，g(r)为双相关函数λ is probability of point on ring, g(r) is pair correlation function
角尺度Uniform angle index (惠刚盈等，1999)	${W_i} = \frac{1}{n}\sum\limits_{j = 1}^n {{v_{ij}}} $	n为相邻木株数，如果相邻木与参照组成的角大于标准角，则v_ij=0；否则v_ij=1。分布格局可用图形、置信区间或显著性检验方法进行判断n is number of near neighbor trees, if the angle composition by adjacent and reference wood is greater than the standard angle, v_ij=0, else, v_ij=1. The distribution pattern can be judged by graph, confidence interval or significant test method

表4

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指数名称Index	计算公式Formula	说明Definition
Shannon-Wiener指数 Shannon-Wiener index (Shannon et al., 1949)	${H^\prime } = - \sum\limits_{i = 1}^S {{p_i}} \ln {p_i}$	p_i为第i个树种株数在林分树木总株数中所占百分比，S为树种数。H′越大，表明林分组成结构多样性越高p_i is the percentage of the i-th tree species in the total number of trees, S is the number of tree species in the stand. The larger the value of H′, the higher the diversity of the components structure of the stand
Pielou均匀度指数 Pielou evenness index (Pielou，1961)	E=H′/lnS	H′为Shannon-Wiener指数，S为树种数。E越大，林分树种分配越均匀H′ is the Shannon-Wiener index, S is the number of tree species. The larger the value of E, the more evenness the distribution of structural components of the forest
Simpson多样性指数 Simpson diversity index (Simpson，1949)	$D = 1 - \sum\limits_1^S {{{\left({{p_i}} \right)}^2}} $	S为树种数，p_i为第i个树种株数在林分树木总株数中所占百分比。D越大，表明林分中树种多样性越高S is the number of tree species, p_i is the percentage of the i-th tree species in the total number of trees. The larger the value of D, the higher the diversity of the components structure of the stand
Margalef丰富度指数 Margalef richness index (Gamito，2010)	R₁=(S-1)/lnN	S为树种数，N为所有树种的个体总数。R₁越大，林分中树种的丰富程度越高S is the number of tree species, N is the total number of trees. The larger the value of R₁, the higher the diversity of the components structure of the stand

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