影响广东黑石顶树附生苔藓分布的环境因子

通过对广东省黑石顶自然保护区内5个2500 m2样地内树附生苔藓的调查及有关环境因子的测定,研究了树附生苔藓的分布格局及其与环境因子的关系.树附生苔藓在不同高度的分布存在一定梯度,20 cm高处树附生苔藓的种类数与盖度均大于60 cm及更高处,且其群落优势种的数量组成与后者存在较大差异.不同树种附生苔藓盖度和种数差异较大.基于附生苔藓植物的盖度进行DCA排序及聚类分析将树种分成四组,马尾松(Pinus massoniana)因具有两种特有的网藓(巴西网藓Syrrhopodon prolifer和鞘刺网藓S.armatus)单独一组,福建青冈(Quercus chungii)亦与其余各种的差异均较大,形成一组,其余阔叶树种根据其所处的森林类型分成两组,针阔叶混交林内的阔叶树种和次生阔叶林内的阔叶树种各形成一组.对环境因子及树皮含水量和pH的分析显示,垂直梯度上空气湿度的差异可能是造成附生苔藓在不同高度分布差异的主要影响因子之一,不同树种附生苔藓的差异在一定程度上受树皮pH的影响,而与树皮含水量无关.同一树种上树附生苔藓的分布又在一定程度上受森林类型的影响.     

Protective management of trees against debarking by deer negatively impacts bryophyte diversity

When wildlife populations become too large, they impact other flora and fauna within the ecosystems that they inhabit. For example, the recent rise in population numbers of sika deer in Japan has led to the stripping of bark from tree overstories in forested areas. This has led to protective management actions, such as wrapping the trunks of trees in wire mesh. The present study investigates the impact of this management action on epiphytic diversity at Mt. Ohdaigahara, which is one of the hotspots for bryophyte diversity in Japan. The correlation between the diversity of epiphytic bryophytes and environmental variables was examined, including the presence/absence of wire mesh protection. A generalized linear model showed that species richness and bryophyte cover was significantly correlated with both tree diameter (at 1.5 m height) and tree density (P < 0.01), but negatively correlated with wire mesh protection. Inductively coupled plasma-mass spectrometry analysis showed a significant 3- to 6-fold higher concentration of zinc in bryophytes occupying tree bark under wire mesh protection than for those without wire mesh. Hence, the high sensitivity of bryophytes to zinc accumulation, as a result of toxicity caused by galvanized iron mesh, has led to the loss of species richness and bryophyte cover on tree trunks. Furthermore, other heavy metals found in wire mesh may also contribute to the negative effect on bryophytes. Therefore, to establish best practices for biodiversity conservation that include bryophytes, materials that are free of heavy metals should be preferentially used for tree protection.     

Epiphytic bryophyte diversity and range distributions along an elevational gradient in Marojejy,Madagascar

Describing spatial variation in species richness and understanding its links to ecological mechanisms are complementary approaches for explaining geographical patterns of richness. The study of elevational gradients holds enormous potential for understanding the factors underlying global diversity. This paper investigates the pattern of species richness and range-size distribution of epiphytic bryophytes along an elevational gradient in Marojejy National Park, northeast Madagascar. The main objectives are to describe bryophyte species composition and endemism in Marojejy National Park, to describe the species richness and distribution patterns of epiphytic bryophytes along an elevational gradient from 250 m to 2050 m and to evaluate the explanatory value of environmental variables for the observed patterns. Bryophyte samples were collected following a nested design with four hierarchical levels: elevational belts, plots, quadrats, and microplots. In total, 254 epiphytic bryophyte species were recorded, comprising 157 liverworts and 97 mosses. Twenty-three of these are endemic to Madagascar. Species richness exhibits a hump-shaped pattern along the elevational gradient, peaking at 1,250 m. Eighty-seven percent of the total recorded species have a range distribution lower than 1,000 m, at which point 36% are restricted to these single elevations. Our results suggest that mean temperature, relative humidity, and vapor pressure deficit play important roles in shaping the richness pattern observed in this study. While the liverwort richness pattern did not correlate to vapor pressure deficit and responded only weakly to relative humidity, the richness pattern shown by mosses correlates well with mean temperature, relative humidity, and vapor pressure deficit.     

The effects of stand structure on ground-floor bryophyte assemblages in temperate mixed forests

The effect of tree species composition, stand structure characteristics and substrate availability on ground-floor bryophyte assemblages was studied in mixed deciduous forests of Western Hungary. Species composition, species richness and cover of bryophytes occurring on the soil and logs were analysed as dependent variables. The whole assemblage and functional groups defined on the basis of substrate preference were investigated separately. Substrate availability (open soil, logs) was the most prominent factor in determining species composition, cover and diversity positively, while the litter of deciduous trees had a negative effect on the occurrence of forest floor bryophytes. Besides, bryophyte species richness increased with tree species and stand structural diversity, and for specialist epiphytic and epixylic species log volume was essential. Sapling density and light heterogeneity were influential on bryophyte cover, especially for the dominant terricolous species. Many variables of the forest floor bryophyte community can be estimated efficiently by examining stand structure in the studied region. Selective cutting increasing tree species diversity, stand structural heterogeneity and dead wood volume can maintain higher bryophyte diversity in this region than the shelter-wood system producing even-aged, monodominant, structurally homogenous stands.     

武汉市城区苔藓植物多样性和分布及与环境因子的关系

对武汉市7个中心城区和部分远郊城区共26个样点73个样地的苔藓群落进行调查,采集苔藓植物样本共431份。经鉴定调查区内共有苔藓植物20科35属91种,其中狭叶小羽藓（Haplocladium angustifolium（Hampe et C.Muell.）Broth.）和钝叶绢藓（Entodon obtusatus Broth.）等为优势种,分布于其中的24个样点。多样性指数分析结果显示,位于青龙山国家森林公园的α多样性Patrick和Shannon-Wiener指数最高,说明该样点苔藓植物群落的复杂程度高,群落所含信息量大。Patrick指数与环境因子的Pearson相关性分析结果显示,土壤含水率和与主要干道距离是显著影响地面生苔藓植物多样性的环境因子,而显著影响树附生苔藓植物多样性的环境因子仅与距主要干道距离相关。用典范对应分析法（CCA）研究26个样点中苔藓植物的盖度与主要环境因子的关系,结果发现人为干扰程度、草本盖度、与城市主要干道的距离等对地面生和树附生苔藓植物的分布都有显著影响。其中地面生苔藓植物的分布受人为干扰和草本盖度影响最为显著;而树附生苔藓植物的分布受附生树干胸径及与主要干道距离影响最为显著。研究结果表明武汉市区苔藓植物的多样性和分布受人为干扰较大,树附生苔藓对汽车尾气等因素更为敏感。     

Epiphytes of Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis>) plantations in Ireland and the effects of open spaces

The epiphytes of the trunks and branches of mature Sitka spruce (Picea sitchensis) trees were studied in twelve plantations containing open spaces (glades, rides and roads) in the east and southwest of Ireland. A pair of trees was studied at each site: one tree at the south-facing edge of an open space and one in the forest interior. Spruce trees were found to support a moderately diverse range of bryophytes and lichens, including two relatively rare bryophyte species. Clear patterns in vertical distribution were identified, with bryophyte richness and cover decreasing and lichen richness and cover increasing from the tree base to the upper trunk. The open spaces themselves did not appear to affect overall epiphyte diversity, with no significant differences in any of the diversity measures between edge and interior trees. The main effect of open spaces was on the epiphyte cover of the edge trees. This was related to increased light levels combined with the presence of live branches from close to ground level on the south sides of the edge trees, which produced optimum conditions for bryophytes at the tree base and lichens in the upper plots. However, this dense side-canopy negatively affected epiphyte diversity on the north sides of the edge trees. Further research is required to assess the effects of open spaces within forestry plantations on epiphyte diversity.     

Epiphytes of Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis>) plantations in Ireland and the effects of open spaces

The epiphytes of the trunks and branches of mature Sitka spruce (Picea sitchensis) trees were studied in twelve plantations containing open spaces (glades, rides and roads) in the east and southwest of Ireland. A pair of trees was studied at each site: one tree at the south-facing edge of an open space and one in the forest interior. Spruce trees were found to support a moderately diverse range of bryophytes and lichens, including two relatively rare bryophyte species. Clear patterns in vertical distribution were identified, with bryophyte richness and cover decreasing and lichen richness and cover increasing from the tree base to the upper trunk. The open spaces themselves did not appear to affect overall epiphyte diversity, with no significant differences in any of the diversity measures between edge and interior trees. The main effect of open spaces was on the epiphyte cover of the edge trees. This was related to increased light levels combined with the presence of live branches from close to ground level on the south sides of the edge trees, which produced optimum conditions for bryophytes at the tree base and lichens in the upper plots. However, this dense side-canopy negatively affected epiphyte diversity on the north sides of the edge trees. Further research is required to assess the effects of open spaces within forestry plantations on epiphyte diversity.     

Bole epiphytic bryophytes on <Emphasis Type="Italic">Lithocarpus xylocarpus</Emphasis> (Kurz) Markgr. in the Ailao Mountains,SW China

Epiphytic bryophytes growing on Lithocarpus xylocarpus (Kurz) Markgr. trunks of different diameter classes in primary (132 plots) and secondary (84 plots) Lithocarpus forests in the Ailao Mountains, SW China, were surveyed and analyzed to determine species composition and richness, and to identify environmental variables that may affect it. Among the 65 species (belonging to 32 genera, 19 families) found, 28 occurred in both forests, with Syrrhopodon gardneri (Hook.) Schwaegr. predominanting. Species richness and total coverage in primary forest were significantly higher than in secondary forest. We suggest that a period of perhaps much more than 110 years is necessary for the recovery of epiphytic bryoflora in montane forest of SW China. Fan, turf, and smooth mat are the most important life forms, with high occurrences in both forests. The life form composition of epiphytic bryophytes is determined mainly by microhabitat and host age. Tree age, the presence of primary forest, bark pH, and plot exposure are the environmental variables that have significantly influenced the composition of epiphytic bryophytes. Tree age explained most variations in epiphytic bryoflora. Bark pH is another important parameter that significantly influenced the epiphytic bryophyte community, but seemed indirectly correlated with tree age. Primary forest is a favorable habitat for epiphytes, due mainly to its diversified canopy structure and the presence of large diameter hosts. Moisture-laden southwest trade winds and forest structure could differentiate microclimate and impel a distinct composition of epiphytes in windward and leeward exposures.     

广东黑石顶森林苔藓群落特征初探

通过对5个面积为2500m2的样地中树附生和地面苔藓植物的定量调查,初步研究了广东黑石顶森林中苔藓植物多样性特征。本次调查共发现苔藓植物50种,包括30种苔类和20种藓类。样地中树附生苔藓植物40种,地面苔藓植物24种,前者远高于后者。各样地中地面苔藓植物的种类为8-13种,盖度0.59%-1.12%,坡度及小生境的多样性对地面苔藓植物物种多样性及盖度均有很大影响;树附生苔藓植物12-20种,盖度0.63%-1.63%。结果显示30a的次生林中苔藓植物的物种丰富度及盖度均可恢复到与成熟阔叶林接近的水平。相似性分析表明,成熟阔叶林内苔藓植物种类组成与针阔叶混交林及30a的次生阔叶林差异不明显,但前者的苔藓植物群落结构与后者的差异较大。     

Factors influencing epiphytic bryophyte and lichen species richness at different spatial scales in managed temperate forests

The effect of management related factors on species richness of epiphytic bryophytes and lichens was studied in managed deciduous-coniferous mixed forests in Western-Hungary. At the stand level, the potential explanatory variables were tree species composition, stand structure, microclimate and light conditions, landscape and historical variables; while at tree level host tree species, tree size and light were studied. Species richness of the two epiphyte groups was positively correlated. Both for lichen and bryophyte plot level richness, the composition and diversity of tree species and the abundance of shrub layer were the most influential positive factors. Besides, for bryophytes the presence of large trees, while for lichens amount and heterogeneity of light were important. Tree level richness was mainly determined by host tree species for both groups. For bryophytes oaks, while for lichens oaks and hornbeam turned out the most favourable hosts. Tree size generally increased tree level species richness, except on pine for bryophytes and on hornbeam for lichens. The key variables for epiphytic diversity of the region were directly influenced by recent forest management; historical and landscape variables were not influential. Forest management oriented to the conservation of epiphytes should focus on: (i) the maintenance of tree species diversity in mixed stands; (ii) increment the proportion of deciduous trees (mainly oaks); (iii) conserving large trees within the stands; (iv) providing the presence of shrub and regeneration layer; (v) creating heterogeneous light conditions. For these purposes tree selection and selective cutting management seem more appropriate than shelterwood system.     

Vertical distribution of epiphytic bryophytes in an Indonesian rainforest

We studied species richness, composition and vertical distribution of epiphytic bryophytes in submontane rainforest of Central Sulawesi. Bryophytes were sampled on eight canopy trees and on eight trees in the forest understorey. Microclimate was measured at trunk bases and at crown bases. The total recorded number of 146 epiphytic bryophyte species is among the highest ever reported for tropical forests and underlines the importance of the Malesian region as a global biodiversity hotspot. Species composition differed significantly between understorey trees and canopy tree trunks on the one hand, and the forest canopy on the other. Fourty-five percent of the bryophyte species were restricted to canopy tree crowns, 12% to the understorey. Dendroid and fan-like species mainly occurred in the forest understorey whereas tufts were most species rich in the tree crowns. The findings reflect the different microclimatic regimes and substrates found in the understorey and in the forest canopy. The results indicate that assessments of the bryophyte diversity of tropical forests are inadequate when understorey trees and tree crowns are excluded.     

辽东山地古石河冰缘地貌森林生态系统苔藓物种多样性研究

基于9个20 m×30 m森林群落样地的调查数据,采用物种丰富度、α和β多样性指数,对辽东山地古石河冰缘地貌不同林型石生、树生苔藓植物物种多样性进行定量研究,采用皮尔逊相关分析方法对其影响因素进行分析。结果显示,古石河冰缘地貌苔藓植物共有26科46属59种;不同林型石生、树生苔藓植物物种丰富度和α多样性指数均为:暗针叶林针阔混交林落叶阔叶林;石生苔藓植物β多样性指数最高为落叶阔叶林-针阔混交林间(0.44),最低为落叶阔叶林-暗针叶林间(0.33);树生苔藓植物β多样性指数最高为针阔混交林-暗针叶林间(0.40),最低为落叶阔叶林-暗针叶林间(0.25);分析表明,林冠层郁闭度、海拔高度是影响辽东山地古石河冰缘地貌森林生态系统苔藓物种多样性的重要因子。     

Bryophyte colonisation in experimental microcosms: the role of nutrients, defoliation and vascular vegetation

A three-year multi-factorial microcosm experiment simulating fertilisation, defoliation and the composition of vascular vegetation in a dry grassland succession was used to test four hypotheses concerning the establishment and survival of bryophytes in grassland vegetation. H₁: bryophyte cover may be used to predict bryophyte species richness. H₂: bryophyte richness is suppressed at high nutrient levels and promoted by defoliation of vascular plants. H₃: species richness of bryophytes is influenced by the species composition of the vascular vegetation. H₄: bryophyte species richness is negatively correlated with vascular plant biomass.
The relationship between bryophyte richness and bryophyte cover was found to follow the classical species-area richness curve. Bryophyte species richness responded positively to defoliation and negatively to fertilisation. The species composition of vascular vegetation had no significant effect on bryophyte richness. Bryophyte species richness was lower at high vascular plant biomass and vascular plant dry weight above 400 g m⁻² appeared fatal to bryophytes. At high nutrient levels, defoliation increased bryophyte richness, but defoliation did not fully compensate for the negative effect of fertilisation. The study reinforces the concern for short lived shuttle bryophytes in the agricultural landscape.     

Endozoochory by slugs can increase bryophyte establishment and species richness

Herbivory can affect plant community composition and diversity by removing biomass and reducing light competition. Herbivory may particularly benefit low growing species such as bryophytes, which are frequently limited by light competition. Gastropods are important herbivores of seed plants and cryptogams, furthermore, they can disperse propagules such as seeds and spores via endozoochory. However, whether gastropod herbivory can reduce the dominance of vascular plants and thereby promote the germination and establishment of endozoochorously dispersed bryophyte spores has never been tested experimentally. Moreover, it is unclear whether these possible interacting effects can influence bryophyte species richness. Here, we tested for endozoochorous spore dispersal by slugs, in combination with sowing of vascular plants, in a fully factorial common garden experiment. Enclosures contained either slugs previously fed with bryophyte sporophytes, control slugs, or no slugs. After 21 days the bryophyte cover was on average 2.8 times higher (3.9% versus 1.4%) and after eight months the bryophyte species richness 2.6 times higher (5.8 versus 2.2) in enclosures containing slugs previously fed with bryophyte sporophytes than in the other treatments. Furthermore, after eight months high vascular plant cover reduced bryophyte diversity. On average enclosures without seed sowing harboured 1.6 times more bryophyte species than the ones with seed sowing (4.2 versus 2.6), indicating competitive effects of vascular plants on bryophytes. Our findings suggest that slugs are important dispersal vectors for bryophytes and that they can increase bryophyte populations and maintain bryophyte diversity by reducing the dominance of vascular plants.     

Bryophyte Species Richness and Composition along an Altitudinal Gradient in Gongga Mountain,China

An investigation of terrestrial bryophyte species diversity and community structure along an altitudinal gradient from 2,001 to 4,221 m a.s.l. in Gongga Mountain in Sichuan, China was carried out in June 2010. Factors which might affect bryophyte species composition and diversity, including climate, elevation, slope, depth of litter, vegetation type, soil pH and soil Eh, were examined to understand the altitudinal feature of bryophyte distribution. A total of 14 representative elevations were chosen along an altitudinal gradient, with study sites at each elevation chosen according to habitat type (forests, grasslands) and accessibility. At each elevation, three 100 m × 2 m transects that are 50 m apart were set along the contour line, and three 50 cm × 50 cm quadrats were set along each transect at an interval of 30 m. Species diversity, cover, biomass, and thickness of terrestrial bryophytes were examined. A total of 165 species, including 42 liverworts and 123 mosses, are recorded in Gongga mountain. Ground bryophyte species richness does not show any clear elevation trend. The terrestrial bryophyte cover increases with elevation. The terrestrial bryophyte biomass and thickness display a clear humped relationship with the elevation, with the maximum around 3,758 m. At this altitude, biomass is 700.3 g m⁻² and the maximum thickness is 8 cm. Bryophyte distribution is primarily associated with the depth of litter, the air temperature and the precipitation. Further studies are necessary to include other epiphytes types and vascular vegetation in a larger altitudinal range.     

Tree age‐dependent changes among epiphytic bryophyte communities in Mediterranean environments. A case study from Sicily (Italy)

Abstract

The epiphytic bryophytes inhabiting trees of different size/age in a Quercus ilex wood from Madonie Mountains (northern Sicily, Italy) were studied with the purpose of describing the changes that take place in the bryophyte stratum during the tree lifespan. Results indicate an increase of bryophyte cover combined with a progressive decrease of epiphytic lichens and the existence of an active process of species and community substitution. The way these processes take place corresponds to a succession sequence, which is characterised by a high number of pioneer species on the youngest trees and a sharp decline of species number on middle‐aged and old trees due to the great spread of Leptodon smithii and a few other pleurocarpous mosses. The comparison of these traits to those from central Spain evidences differences in patterns of tree colonisation by epiphyte bryophytes between the two areas and suggests that climatic conditions play a chief role in the epiphytic bryophyte establishment and succession in Mediterranean areas.     

Hidden crown jewels: the role of tree crowns for bryophyte and lichen species richness in sycamore maple wooded pastures

Tree crowns typically cover the vast majority of the surface area of trees, but they are rarely considered in diversity surveys of epiphytic bryophytes and lichens, especially in temperate Europe. Usually only stems are sampled. We assessed the number of bryophyte and lichen species on stems and in crowns of 80 solitary sycamore maple trees (Acer pseudoplatanus) at six sites in wooded pastures in the northern Alps. The total number of species detected per tree ranged from 13 to 60 for bryophytes, from 25 to 67 for lichens, and from 42 to 104 for bryophytes and lichens considered together. At the tree level, 29 % of bryophyte and 61 % of lichen species were recorded only in the crown. Considering all sampled trees together, only 4 % of bryophyte, compared to 34 % of lichen species, were never recorded on the stem. Five out of 10 red-listed bryophyte species and 29 out of 39 red-listed lichen species were more frequent in crowns. The species richness detected per tree was unexpectedly high, whereas the proportion of exclusive crown species was similar to studies from forest trees. For bryophytes, in contrast to lichens, sampling several stems can give a good estimation of the species present at a site. However, frequency estimates may be highly biased for lichens and bryophytes if crowns are not considered. Our study demonstrates that tree crowns need to be considered in research on these taxa, especially in biodiversity surveys and in conservation tasks involving lichens and to a lesser degree also bryophytes.     

Predicting bryophyte diversity in grassland and eucalypt-dominated remnants in subhumid Tasmania

Aim To determine the environmental factors associated with bryophyte diversity in remnants in a fragmented, agricultural landscape. Location Eighty‐two remnants of tussock grassland, eucalypt woodland and eucalypt forest in the subhumid Midlands region of Tasmania, Australia. Methods Remnants were surveyed for bryophytes and predictor variables, such as vascular plant cover, climate, and topography. Management histories for each remnant were compiled using both site observation and landowner surveys. Bryophyte cover, richness, and composition were related to the independent variables using simple correlation and general linear models. Results We found weak relationships between the dependent variables and the fragmentation variables (remnant area, remnant perimeter to area ratio, distance to nearest remnant, distance to nearest larger remnant, and remnant age). Instead, climatic variables were important in predicting bryophytes, in particular those affecting humidity (minimum temperature of the coldest month, precipitation). Despite extensive sheep grazing in this landscape, grazing was not correlated with bryophyte diversity. Bryophyte diversity was not explained by vascular plant richness and was only weakly explained by composition, but was predicted by the cover of vascular plants. There was greater bryophyte cover and richness and different composition where the cover of native vascular plants was lower. Main conclusions The implications of our results are that all remnants, regardless of area, age and isolation, appear to be valuable for bryophyte conservation in this highly altered landscape. Our results also suggest that the cover of the vascular plant community, rather than its diversity, holds promise as a guide to bryophyte diversity. Bryophyte composition was similar between sites and a focus on the most species‐rich sites may be the best conservation strategy in this ecosystem.     

Effects of management and adjacent forest on the heathland bryophyte layer

Heathlands are severely threatened by decreasing habitat quantity and quality. In these habitats, bryophytes form an important component of plant diversity. Therefore, the aims of this study were to investigate the effects of management and adjacent forest land use on the heathland bryophyte layer. Furthermore, effects of these factors on invasive bryophyte species were studied.

Bryophyte communities of 11 dry heathland patches bordering forest were surveyed. In these heathlands, management turned out to be an important factor explaining community composition of bryophytes. Furthermore, significant effects of adjacent land use on bryophyte community composition and species richness were found. However, these effects were rather limited in extent, occurring only close to habitat borders, i.e., up to maximum 5 m into the heathland patch. Also for individual bryophyte species, effects of adjacent forest could only be observed within maximum 5 m from the edge. One remarkable edge effect was the increased dominance of the invasive species Campylopus introflexus near the forest edge, especially at grazed sites. Effects of adjacent land use seemed to predominate management effects. Consequently, management cannot serve as a tool to mitigate edge effects on bryophyte species in heathland patches.     

浙江西天目山主要森林类型的苔藓多样性比较

苔藓是森林的重要组分, 是森林保护区的重要保护对象, 在物种资源和生态系统功能维护中有重要作用。该研究以浙江西天目山国家自然保护区内7种主要森林类型(落叶矮林、落叶阔叶林、常绿-落叶阔叶混交林、常绿阔叶林、针阔混交林、针叶林和竹林)内的苔藓植物为对象, 调查了32个10 m × 10 m的样地, 记录地面生苔藓植物盖度和树附生苔藓植物多度, 采用重要值、相似性系数、多样性指数分析了森林类型间的苔藓植物多样性差异。共采集969份标本, 隶属41科84属142种, 其中苔类植物13科18属33种, 藓类植物28科66属109种, 优势科为灰藓科、青藓科和羽藓科。2种混交林(常绿-落叶阔叶混交林和针阔混交林)的物种丰富度和多样性指数均高于其余5种森林, 其中物种丰富度以针阔混交林最高, 苔藓植物多样性则以常绿-落叶阔叶混交林最高, 竹林两者均为最低。海拔等环境因子较为接近的植被类型的苔藓植物多样性相似性较高, 常绿阔叶林与针叶林相似性最高, 而落叶矮林和竹林相似性最小。