保安湖—湖湾大型水生植物群落格局的研究

用地质统计学中的半方差和双对数半方差图对保安湖-湖湾(黄风口)的大型水生植物群落格局进行了研究,在不同的尺度上对群落及其主要组成种类的空间异质性进行了定量的描述。结果表明:保安湖黄风口大型水生植物群落的双对数半方差图存在线性区域,群落具有分形特征(自相似性).在不同的尺度下,群落和各组成种类空间格局具有不同的分形维数值,异质性程度存在差异。但其异质性程度不高,建议进行群落调查时,样方尺度应取520m;单一种群调查时的样方尺度取380m.       

Scale-dependent relationships between the spatial distribution of a limiting resource and plant species diversity in an African grassland ecosystem

One cornerstone of ecological theory is that nutrient availability limits the number of species that can inhabit a community. However, the relationship between the spatial distribution of limiting nutrients and species diversity is not well established because there is no single scale appropriate for measuring variation in resource distribution. Instead, the correct scale for analyzing resource variation depends on the range of species sizes within the community. To quantify the relationship between nutrient distribution and plant species diversity, we measured NO₃^- distribution and plant species diversity in 16 paired, modified Whittaker grassland plots in Serengeti National Park, Tanzania. Semivariograms were used to quantify the spatial structure of NO₃^- from scales of 0.4–26 m. Plant species diversity (Shannon-Weiner diversity index; H ) was quantified in 1-m² plots, while plant species richness was measured at multiple spatial scales between 1 and 1,000 m². Small-scale variation in NO₃^- (<0.4 m) was positively correlated with 1-m² H , while 1,000-m² species richness was a log-normal function of average NO₃^- patch size. Nine of the 16 grassland plots had a fractal (self-similar across scales) NO₃^- spatial distribution; of the nine fractal plots, five were adjacent to plots that had a non-fractal distribution of NO₃^-. This finding offered the unique opportunity to test predictions of Ritchie and Olff (1999): when the spatial distribution of limiting resources is fractal, communities should display a left-skewed log-size distribution and a log-normal relationship between net primary production and species richness. These predictions were supported by comparisons of plant size distributions and biomass-richness relationships in paired plots, one with a fractal and one with a non-fractal distribution of NO₃^-. In addition, fractal plots had greater large-scale richness than paired non-fractal plots (1,0–1000 m²), but neither species diversity (H ) nor richness was significantly different at small scales (1 m²). This result is most likely explained by differences in the scale of resource variation among plots: fractal and non-fractal plots had equivalent NO₃^- variation at small scales but differed in NO₃^- variation at large scales (as measured by the fractal dimension). We propose that small-scale variation in NO₃^- is largely due to the direct effects of plants on soil, while patterns of species richness at large scales is controlled by the patch size and fractal dimension of NO₃^- in the landscape. This study provides an important empirical step in understanding the relationship between the spatial distribution of resources and patterns of species diversity across multiple spatial scales.     

毛乌素沙地沙生半灌木群落的空间异质性

在使用 5m× 50 0 0 m样带进行调查获得的植被盖度数据的基础上 ,通过半方差分析和分形分析揭示了毛乌素沙化草地沙生半灌木群落植被盖度的空间变异特点。半方差分析揭示出植被盖度的多尺度变异和等级斑块结构 ,在取样范围内至少存在着 3个等级。分形分析得出毛乌素沙地沙生半木群落植被盖度的分维 D=1 .90 ,接近于 2 ,指示着大部分空间变异发生在较小的尺度上 ;分段分形分析得出不同尺度范围内的分维 ,反映了空间异质性的多尺度变异性。这一结果揭示了荒漠化过程引起的生境破碎化     

地统计学在水生植物群落格局研究中的应用

为了研究水生植物群落的镶嵌分布格局和随着尺度变化的空间特征,应用地统计学中半方差法、半方差理论模型和分形理论,研究了湖北省保安湖沿岸带水生植物群落分布格局,在沿岸带设置了一个160m×160m的方形样方,采用覆盖法将整个样方划分为32×32个格子,记录各格子内出现的物种数,格子内如果有某物种,则将该格子对应的区域化随机变量赋值为1,否则赋值为0。另将群落各格内区域化随机变量赋值为各物种在该采样点随机变量值的总和,运用地统计软件GS+for Windows分析苦草、狐尾藻、金鱼藻和群落的32×32格随机变量表,计算得到它们各向同性模型和不同方向上的分维,从苦草和狐尾藻的各向同性指数模型来看,它们属于聚集性分布,其空间结构比率较高,这表明在小尺度上它们变化较小;从金鱼藻和群落的各向同性线性模型来看,属于中等程度的聚集分布,其空间结构比率很低,这表明它们在小尺度上变化较大。从各向异性模型来看,群落优势种的分维值都较高,说明在调查的群落内,各物种分布比较均匀,根据确定系数判定群落格局的优势方向是西北-东南,与其他群落格局分析方法相比较,该方法在确定群落组分空间依赖范围和相关性大小。以及区分聚集强度上具有一定优势,为定量描述不同尺度下的水生植物群落特征提供了一个新途径。     

山西太岳山兴唐寺红柄白鹃梅群落优势种的空间格局分析

使用点格局分析和分形分析(计盒维数、信息维数、关联维数)方法, 对山西太岳山兴唐寺红柄白鹃梅(Exochorda giraldii)群落进行了空间格局分析。结果表明: 1)红柄白鹃梅群落优势种在较小的尺度上呈现聚集分布的特点, 其中, 杠柳(Periploca sepium)和绣线菊(Spiraea salicifolia)在所有尺度上呈现聚集分布, 红柄白鹃梅和连翘(Forsythia suspensa)随着尺度的增加, 呈现聚集分布—随机分布—均匀分布的规律; 2)红柄白鹃梅群落优势种在不同尺度上表现出聚集分布的特点; 3)红柄白鹃梅群落优势种占据空间能力的大小为: 红柄白鹃梅>连翘>绣线菊>杠柳; 4)红柄白鹃梅群落优势种格局强度的尺度变化程度为: 杠柳>绣线菊>红柄白鹃梅>连翘; 5)红柄白鹃梅群落优势种个体空间相关程度为: 杠柳>红柄白鹃梅>连翘>绣线菊。点格局分析与分形分析结果一致, 揭示了暖温带落叶阔叶林遭到破坏后形成的次生灌丛的空间分布格局。     

Spatial leaf distribution and self-thinning exponent of<Emphasis Type="Italic"> Pinus banksiana</Emphasis> and<Emphasis Type="Italic"> Populus tremuloides</Emphasis>

A hypothesis that the pattern of spatial leaf distribution in forest canopies is numerically related to the exponent of the self-thinning relationship in even-aged monocultures was tested by evaluating the crown fractal dimension of Pinus banksiana (jack pine) and Populus tremuloides (quaking aspen) in Wood Buffalo National Park, Canada. Pure species stands that were considered the most dense for a given mean tree size were measured to establish the empirical self-thinning relationships. The value of the self-thinning exponent was estimated as –1.42 with 95% Confidence Interval (CI) (–1.47, –1.36) for Pinus banksiana, and –1.29 with 95% CI (–1.45, –1.14) for Populus tremuloides. For each species the box dimension of spatial leaf distribution was estimated from unit cylinders described by sequentially lowering in forest canopies, horizontal flaps of one of various diameters attached to the top of a height-measuring pole. The box dimension appeared as 1.95 (1.84, 2.06) for Pinus banksiana, and 2.24 (2.05, 2.43) for Populus tremuloides. By assuming that the box dimension is equivalent to the fractal dimension at the inter-population level, the self-thinning exponent was predicted to be –1.53 (–1.62, –1.45) for Pinus banksiana, and –1.33 (–1.45, –1.23) for Populus tremuloides. The empirical exponent was equivalent to that predicted from the box fractal dimension, as judged by the 95% CI of the dimensions. We conclude that spatial patterns of leaf distribution in forest canopies, as being characterized by the box fractal dimension, are closely related to the value of the self-thinning exponent in the dense monocultures of the species we examined.     

Fractal geometry: a tool for describing spatial patterns of plant communities

Vegetation is a fractal because it exhibits variation over a continuum of scales. The spatial structure of sandrim, bryophyte, pocosin, suburban lawn, forest tree, and forest understory communities was analyzed with a combination of ordination and geostatistical methods. The results either suggest appropriate quadrat sizes and spacings for vegetation research, or they reveal that a sampling design compatible with classical statistics is impossible. The fractal dimensions obtained from these analyses are generally close to 2, implying weak spatial dependence. The fractal dimension is not a constant function of scale, implying that patterns of spatial variation at one scale cannot be extrapolated to other scales.     

Power-law species–area relationships and self-similar species distributions within finite areas

The species–area relationship (SAR) is often expressed as a power law, which indicates scale invariance. It has been claimed that the scale invariance – or self‐similarity at the community level – is not compatible with the self‐similarity at the level of spatial distribution of individual species, because the power law would only emerge if distributions for all species had identical fractal dimensions (FD). Here we show that even if species differ in their FD, the resulting SAR is approximately linear on a log–log scale because observed spatial distributions are inevitably spatially restricted – a phenomenon we term the ‘finite‐area effect’. Using distribution atlases, we demonstrate that the apparent power law of SARs for central European birds is attributable to this finite‐area effect affecting species that indeed reveal self‐similar distributions. We discuss implications of this mechanism producing the SAR.     

Habitat structure and vegetation relationships in central Argentina salt marsh landscapes

Relationships between habitat structure and spatial variations in vegetation composition were determined in catenas of central Argentina salt marsh landscapes. Vegetation was classified following a multi-technique strategy. An analysis of species distributions along an environmental gradient was made and a redundancy analysis was used to relate the environmental variables to vegetation data. The spatial covariation was evaluated through fractal analysis. The vegetation can be divided into four discrete noda that correspond to different topographic positions (summit, backslope, footslope and toeslope, respectively): nodum of Stipa trichotoma + St. tenuissima, nodum of Distichlis scoparia, nodum of Distichlis spicata and nodum of Spartina densiflora. Each of these nodum is characterized by a definite combination of floristic groups. Symmetric aggregation of vegetation borders was observed in all three sites. The existence of vegetation discontinuities along the catenas depended largely on water table depth and salinity which in turn controlled the edaphic salinity. The arrangement of sites by multivariate analysis reflected the influence of a complex gradient representing halomorphic and hydromorphic factors. -diversity was associated with abrupt changes in the physical structure of the catena for a reduced spatial dimension (300-m scale). Absolute diversity and evenness were higher on the summit and declined progressively toward the toeslope. The rate of change was higher on the backslope and the dominant species have different ecological amplitudes overlapping along the gradient. The main operational factors associated with the floristic differences are: (i) the variations in the chemical composition and the seasonal dynamics of the soil solution in the aerated layer of soils, (ii) the salinity and dynamics of groundwater, and (iii) the length of time that the soil is flooded during the rainy season (summer). The fractal dimension was close to 2, implying weak spatial dependence. Fractal dimension varied as a function of scale. The fractogram only revealed a significant spatial dependence on the summit. This spatial dependence was associated with short distances of gradient showing that the organizational pattern of Distichlis spp.- Spartina was related to combinations of underlying environmental factors rather than to a specific position in the catena. The catenas are highly structured spatially, floristic compositions are inextricably linked to this structure. Habitat complexity may directly affect associated vegetation by regulating the hydrohalomorphic conditions in the aerated layer of the soils. The relationships of the habitat-floristic groups are not simple, hydromorphism interacts in a complex way with halomorphism.     

天然黄山松种群格局的分形特征——计盒维数与信息维数

采用计盒维数和信息维数对屏南和寿宁不同群落的天然黄山松(Pinus taiwanensis)种群格局的分形特征进行比较分析.结果表明,天然黄山松种群格局具有分形特征,其计盒维数值在1.299 8～1.862 6之间,不同群落的大小次序为Q3>Q1>Q2>Q4>Q7>Q8>Q5>Q6;信息维数值在1.205 7～1.863 7之间,大小次序是Q3>Q1>Q2>Q4>Q8>Q7>Q5>Q6,屏南天然黄山松近纯林黄山松种群的计盒维数和信息维数均高于寿宁混交林,计盒维数定量描述种群占据水平空间的能力和程度,信息维数揭示种群格局强度的尺度变化及个体分布的非均匀程度,分形维数值的高低与群落环境、种群密度、种群在群落中的优势地位、个体的聚集程度及幼树个体数量等相关.黄山松种群格局分形维数随海拔呈现上下波动变化,1 250～1 270 m是更适生的海拔范围.此外,黄山松种群格局的分形特征存在一定的尺度范围,其拐点尺度是分形范围的下限尺度.     

遗传算法支持下土地利用空间分形特征尺度域的识别

针对土地利用空间分形特征只存在于有限尺度域的现象,采用无标度区内离散点拟合的离差平方和平均值最小作为目标函数,提出了一种基于遗传算法的土地利用空间分形特征尺度域的识别方法,用于准确计算分形维数的有效区间范围。以武汉市武昌区水域空间分形特征为例,利用Quickbird多光谱遥感影像提取土地利用空间信息,重点讨论了基于遗传算法识别土地利用空间分形特征尺度域范围的总体思路、适应度函数和遗传算子等环节;然后分别从测定系数、标准差和无标度区间3个角度,将其同人工判断法、相关系数法以及强化系数法进行对比讨论;并结合研究区域实际的水域空间分布格局,分析不同方法计算所得半径维数的合理性。结果表明,土地利用分形特征尺度域的范围对分形维数计算结果有较大影响;相对于传统计算方法来说,遗传算法在尺度无标度区间识别上具有更高的精度,可以为土地利用空间格局分形特征的研究提供客观指导意见。     

A conceptual framework for the spatial analysis of functional trait diversity

Functional trait diversity is a popular tool in modern ecology, mainly used to infer assembly processes and ecosystem functioning. Patterns of functional trait diversity are shaped by ecological processes such as environmental filtering, species interactions and dispersal that are inherently spatial, and different processes may operate at different spatial scales. Adding a spatial dimension to the analysis of functional trait diversity may thus increase our ability to infer community assembly processes and to predict change in assembly processes following disturbance or land‐use change. Richness, evenness and divergence of functional traits are commonly used indices of functional trait diversity that are known to respond differently to large‐scale filters related to environmental heterogeneity and dispersal and fine‐scale filters related to species interactions (competition). Recent developments in spatial statistics make it possible to separately quantify large‐scale patterns (variation in local means) and fine‐scale patterns (variation around local means) by decomposing overall spatial autocorrelation quantified by Moran's coefficient into its positive and negative components using Moran eigenvector maps (MEM). We thus propose to identify the spatial signature of multiple ecological processes that are potentially acting at different spatial scales by contrasting positive and negative components of spatial autocorrelation for each of the three indices of functional trait diversity. We illustrate this approach with a case study from riparian plant communities, where we test the effects of disturbance on spatial patterns of functional trait diversity. The fine‐scale pattern of all three indices was increased in the disturbed versus control habitat, suggesting an increase in local scale competition and an overall increase in unexplained variance in the post‐disturbance versus control community. Further research using simulation modeling should focus on establishing the proposed link between community assembly rules and spatial patterns of functional trait diversity to maximize our ability to infer multiple processes from spatial community structure.     

广西木论自然保护区铁榄群落主要乔木种群格局的分形特征

为了揭示种群空间分布特征随尺度的变化规律,探讨不同种群对空间资源的占据能力和生态适应能力及分形差异,采用计盒维数、信息维数和关联维数对木论自然保护区铁榄群落主要乔木种群格局特征进行了研究。结果表明,优势种铁榄（Sinosideroxylon pedunculatum）、东女贞（Ligustrum japonicum）、小栾树（Boniodendron minius）、稀有濒危种掌叶木（Handeliodendron bodinieri）、青檀（Pteroceltis tatarinowii）5个种群在相应的尺度具有明显的分形特征;计盒维数和信息维数大小顺序一致,均为：铁榄〉东女贞〉小栾树〉掌叶木〉青檀;在5个种群中,铁榄计盒维数为1．7763最大,接近2,对空间环境的占据能力最强,信息维数（1．7206）也最大,个体聚集强度强,在群落中处于绝对优势地位,青檀计盒维数（0．8316）和信息维数（0．8990）最小,对环境的占据利用能力弱,个体聚集程度整体较弱,在群落中处于伴生地位;在0．5～50m的尺度上,关联维数大小顺序为：铁榄（1．7314）〉东女贞（1．6688）〉小栾树（1．6050）〉青檀（0．7868）,而掌叶木只有在大的尺度上才出现关联,且关联较强。3个分形维数大小的不同组合,能够全面揭示不同种群格局特征和种群在群落中所处地位的差异。     

植被演替过程中种群格局动态的分形分析

该文根据“空间代替时间”的原则和对马尾松(Pinus massoniana)种群的重要值-平均胸径的变动趋势的分析,确定5个森林群落为广东省黑石顶自然保护区森林群落演替的一个时间序列;并在此基础上,运用分形理论的计盒维数和信息维数,对演替过程中马尾松种群空间格局的动态进行了分形分析。结果表明,马尾松种群的计盒维数和信息维数2个参数值均呈递减趋势,与其个体数、重要值变动趋势一致。在群落的演替过程中,马尾松种群的空间占据能力不断下降,种群呈衰退趋势。随其在群落中的优势地位逐渐被其它种群取代,群落将由以马尾松占绝对优势的单优群落演替为多优常绿阔叶林群落。分析结果同时表明,分形分析是群落演替过程中种群空间格局动态研究的有效方法,分形维数则能反映种群格局的尺度变化规律。     

From Hutchinsonian ratios to spatial scaling theory: the interplay among limiting similarity,body size and landscape structure

Spatial scaling theory (SST) relates the physical structure of the environment to species coexistence and community assembly. Although SST is a recognized theory in ecology, few studies have evaluated its predictions, producing contradictory results and frequently failing to meet its assumptions. In addition, the ‘risk predictions’ of SST regarding an increase in species similarity with body size and the dependence of this pattern on the landscape and food fractal dimensions have not been evaluated. This study attempted to account for previous limitations, analyzing these predictions in coleopteran guilds that inhabit 18 temporary ponds. This metacommunity covers a large gradient of environmental variables, including food density, the landscape fractal dimension, the food fractal dimensions and other indicators of pond heterogeneity. Average similarity in carnivorous and herbivorous body sizes systematically increased with guild richness, fulfilling classical predictions of niche theory. Species similarity was associated with body size, but the association reverts from negative to positive as the landscape fractal dimension and heterogeneity increases, a pattern further supported by null model analyses. Several nonexclusive mechanisms may account for this pattern: 1) the body size-dependent landscape perception, through which small animals detect more heterogeneity than larger animals; 2) the reaching of landscape limits by larger species, which prevents them from accessing novel largest clusters; 3) the large differences between the landscape and food fractal dimensions; and 4) the homogenization of the landscape when an integer fractal dimension is reached. These mechanisms may dictate that smaller organisms are more able to capitalize on heterogeneity or available resources than larger organisms, thus promoting increased similarity among smaller species. The presented results support the connection between landscape spatial structure and biodiversity and a mechanistic understanding of this connection from the SST.     

The distribution of the macrobenthos of the White Sea littoral in different spatial scales

Spatial distribution of macrobenthos of middle intertidal zone was studied in scale from centimetres to 30 kilometres along the coastline. The community structure and distribution of the 5 most abundant species (Hydrobia ulvae, Mya arenaria, Macoma baltica. Peloscolex benedeni, Arenicola marina) were considered. Spatial heterogeneity of macrobenthos, estimated as mean dissimilarity between samples, kept constant in scale of centimetres--meters, but increased significantly when enlarged area is considered. Patterns of many species changed with scale from random mosaic to more or less pronounced patchiness, whereas the density of H. ulvae and the structure of the whole community demonstrated fractal (self-similar) patch pattern in wide range of scale from dozens of meters to several kilometres. Spatial correlations between species (the composition of assemblages) and between species and environmental factors were also scale dependent. Some possible effects of scale on the observed spatial distribution of benthos are discussed, and multiscaled analysis of biotic heterogeneity is concluded to be very fruitful.     

莲花山主要森林群落空间格局的分形特征

应用信息维数对莲花山主要森林群落的空间格局进行了研究.结果表明:森林群落的格局强度尺度变化程度由高至低的次序为:岷江冷杉一糙皮桦一白桦群落(Q4)1.84＞云杉-紫果云杉-糙皮桦群落(Q2)1.53＞岷江冷杉群落(Q3)1.14＞紫果云杉-云杉-岷江冷杉群落(Q7)1.10＞华山松群落(Q5)0.99＞岷江冷杉人工林(Q1)0.82＞辽东栎群落(Q6)0.45;群落格局信息维数的差异反映了更新状况的差异,揭示出群落个体分布的非均匀性状况.综合3种分形维数,确定各亚高山寒温性森林群落生长与更新状况的次序为Q4＞Q2＞Q3＞Q7＞Q5＞Q1＞Q6.各群落总体平均记盒维数为1.76,关联维数为1.71,信息维数为1.12;各森林群落生长状况较好,个体聚集程度较低,更新状况一般.分形维数谱将不同森林群落的空间结构特征明确区分开来.     

Fractal properties of forest spatial structure

The definition of fractal dimension of natural objects, which enables to deal with scale dependence of fractal dimension is discussed. Abrupt changes of fractal dimension of spatial structure of terrestrial ecosystems are considered in the context of hierarchical paradigm. On this ground the procedure is proposed for segmentation of a territory, which takes into account the scale dependence of spatial variability of ecological parameters. Using remotely sensed data — normalized difference vegetation index (NDVI) and thermal radiation in the infrared band — fractal dimensions and critical scales are evaluated for different forest types with the help of software, developed for this purpose. The results obtained corroborate the potentialities of fractal approach in ecology. These methods and results can be used for discrimination of remotely sensed data; but further investigations, including detailed comparison of fractal characteristics of remotely sensed forest images with results of on-site field studies are necessary to validate them.     

Structural complexity in mussel beds: the fractal geometry of surface topography

Seafloor topographic complexity is ecologically important because it provides habitat structure and alters boundary-layer flow over the bottom. Despite its importance, there is little agreement on how to define and measure surface complexity. The purpose of this investigation was to utilize fractal geometry of vertical cross-section profiles to characterize the surface topography of the soft-bottom mussel bed (Mytilus edulis L.) at Bob's Cove, ME, USA. Mussels there have been shown previously to have spatially ordered fractal characteristics in the horizontal plane. Two hypotheses were tested. The first was that the bed surface is fractal over the spatial scale of 1.44-200 mm, with fractal dimension less than or equal to 1.26, the value for the Koch curve, our model for bed profiles. The second was that bed surface topography (i.e., in vertical profile) is less complex than the mussel bed spatial pattern (i.e., aerial view in the horizontal plane). Both hypotheses were supported. Cross-sections of plaster casts of the bed produced 88 surface profiles, all of which were fractal over the entire spatial scale of more two orders of magnitude employed in the analysis. Fractal dimension values (D) for individual profiles ranged from 1.031 to 1.310. Fractal dimensions of entire casts ranged up to mean (1.242+/-0.046) and median (1.251) values similar to 1.26, the theoretical value of the Koch curve. The bed surface was less complex than the bed spatial pattern because every profile had D<1.36, the smallest value previously obtained from aerial views of the bed. The investigation demonstrated for the first time that surface topography of a soft-bottom mussel bed was fractal at a spatial scale relevant to hydrodynamic processes and habitat structure important for benthic organisms. The technique of using cross-section profiles from casts of the bed surface avoided possible underestimates of fractal dimension that can result from other profiling methods reported in the literature. The results demonstrate that fractal dimension can be useful in the analysis of habitat space and water flow over any irregular seafloor surface because it incorporates the size, shape, and scale of roughness elements into a simple, numerical metric.     

红树植物木榄种群高度结构的分形特征

 采用计测等值线分形维数的方法,探讨山口国家级红树林自然保护区内的木榄(Bruguiera gymnorrhiza)种群高度结构的特征。结果表明,木榄种群的高度结构具有分形的特征,其计盒维数介于1.61～1.90之间,信息维数介于1.63～1.84之间,分形维数的高低主要与幼树个体的数量、个体的集聚程度和高度层次结构的复杂程度等密切相关。计盒维数定量描述种群占据垂直空间的能力和程度,而信息维数揭示种群高度层次细节的尺度变化强度和个体高度分布的非均匀性程度。分析表明,等值线法能够有效地表征木榄种群高度结构的分