多样性排序:方法与实例

多样性已经成为生态学理论与实践中的一个很重要的概念 ,很多多样性指数被引进或设计出来 ,并被广泛地应用于生态系统管理以及环境监测与评价之中。然而 ,很多学者对其也提出了疑议 ,因为当用不同的多样性指数对一组群落进行比较时会得出不同的顺序。Patil和Taillie等人提出用含有一个参数的多样性指数族可以潜在地解决这个问题。本文对这个类型的 1 4种方法做了评述 ,并将其分为 4组。证明了组内各方法间是等价的 ,而不同组的各方法间是不等价的。并认为在进行多样性排序时 ,Patil和Taillie的“右尾和”方法应属首选 ,因为只需计算有限个数值 ,并且该方法判断了群落内多样性的高低。其次 ,Hulbert_Smith_Grassle的方法也值得推荐 ,因为该方法有直观而明确的意义 ,它是一个随机样本中期望包含的物种数目。作为实例 ,还将这些方法应用于北京东灵山地区的 3个森林群落多样性的比较     

An index of divergence from species to life-form diversity based on the notion of intrinsic diversity ordering

In this paper, we propose a measure of divergence from species tolife-form diversity based on the notion of intrinsic diversity ordering. First,species and life-form relative abundances within the analyzed community aredetermined. Next, community intrinsic diversity profile is computed both fromspecies and life-form relative abundances. Finally, the speed of decrease fromspecies to life-form diversity is obtained by comparing the area under thespecies diversity profile with the area under the corresponding life-formdiversity profile. Since life-forms classify plants into ecologically relevantgroups irrespective of their systematics, the proposed measure might be usefulto quantify the ecological similarity of different communities without loosinginformation on traditional taxonomic diversity figures. As an application fordemonstration, a small data set from a garigue community on ultramafic soils ofTuscany (central Italy) is used.     

Unifying and distinguishing diversity ordering methods for comparing communities

Diversity indices have been widely used in ecological research, but they remain problematic in that different indices may rank communities inconsistently. This problem can be solved by using diversity ordering methods, the output of which is a diversity profile in graphical form for each community being compared. In this paper, we demonstrate that existing diversity ordering methods can be classified into four groups and that within-group methods are essentially equivalent, while among-group methods are not. We find that the intrinsic diversity-related methods—i.e., the group containing the right tail-sum method, the logarithmic dominance plot, the majorization method, and the k-dominance plot—provide the most stringent test of diversity ordering, and we recommend the right tail-sum method as the method of preference for practical purposes.     

A Comparative Analysis of Evenness Index Sensitivity

Evenness indices are numerous but the lack of knowledge of their properties is a limitation to their biological usefulness. 15 evenness indices, two of them being recently proposed, were studied in this work. We investigated the sensitivity of each index using (1) 189 macroinvertebrate communities sampled in the field, and (2) a set of communities modified in a controlled way. There is no single way to measure evenness. We demonstrated that a measure should be chosen considering (1) the kind of data analysed and, (2) the index properties wanted by users. Depending on the ecological data set the index should be more sensitive to variations in rare, median or abundant species. For most of macroinvertebrate community analyses, a convenient evenness index requirs to be symmetric, unsensitive to variation on rare taxa, with a large range of variation and can be compared with a diversity index. Depending on diversity measure used, five indices correspond to these criteria: E_Pielou, E_Hurlbert, E_–ln(D), E_1–D, E_MI. Our results were summarized in a table which may help users to select a convenient evenness measure according to their specific data. Concerning index properties, three main features that an evenness index should meet are briefly discussed: dependence with richness, symmetry criteria and variation range. A revue of this controversial subject allowed a best understanding of values obtained with evenness measures depending on their own features.     

The measurement of marine species diversity, with an application to the benthic fauna of the Norwegian continental shelf

Species diversity includes two aspects, the number of species (species richness) and the proportional abundances of the species (heterogeneity diversity). Species richness and heterogeneity diversity can be measured over different scales; a single point, samples, large scales, biogeographical provinces and in assemblages and habitats. In the literature, the terminology of these scales is confused. Here, scales are given a uniform notation. Scales of species richness and heterogeneity diversity are distinguished from turnover (beta) diversity, which is the degree of change in species composition along a gradient. Methods of measurement of the scales of species richness, heterogeneity diversity, turnover diversity and for estimating total species richness are reviewed. Two methods for measuring heterogeneity diversity are recommended Exp H′ (where H′ is the Shannon-Wiener index) and 1/Simpson’s index, together with an equitability index J′. The reviewed methods are then applied to a data set from the Norwegian continental shelf to illustrate the advantages of the recommended methods. Finally, the application of the methods to assessment of effects of disturbance, to studies of gradients of species richness and to conservation issues are discussed.     

Robust estimation of microbial diversity in theory and in practice

Quantifying diversity is of central importance for the study of structure, function and evolution of microbial communities. The estimation of microbial diversity has received renewed attention with the advent of large-scale metagenomic studies. Here, we consider what the diversity observed in a sample tells us about the diversity of the community being sampled. First, we argue that one cannot reliably estimate the absolute and relative number of microbial species present in a community without making unsupported assumptions about species abundance distributions. The reason for this is that sample data do not contain information about the number of rare species in the tail of species abundance distributions. We illustrate the difficulty in comparing species richness estimates by applying Chao''s estimator of species richness to a set of in silico communities: they are ranked incorrectly in the presence of large numbers of rare species. Next, we extend our analysis to a general family of diversity metrics (‘Hill diversities''), and construct lower and upper estimates of diversity values consistent with the sample data. The theory generalizes Chao''s estimator, which we retrieve as the lower estimate of species richness. We show that Shannon and Simpson diversity can be robustly estimated for the in silico communities. We analyze nine metagenomic data sets from a wide range of environments, and show that our findings are relevant for empirically-sampled communities. Hence, we recommend the use of Shannon and Simpson diversity rather than species richness in efforts to quantify and compare microbial diversity.     

A multi-faceted framework of diversity for prioritizing the conservation of fish assemblages

Floodplain waterbodies and their biodiversity are increasingly threatened by human activities. Given the limited resources available to protect them, methods to identify the most valuable areas for biodiversity conservation are urgently needed. In this study, we used freshwater fish assemblages in floodplain waterbodies to propose an innovative method for selecting priority areas based on four aspects of their diversity: taxonomic (i.e. according to species classification), functional (i.e. relationship between species and ecosystem processes), natural heritage (i.e. species threat level), and socio-economic (i.e. species interest to anglers and fishermen) diversity. To quantitatively evaluate those aspects, we selected nine indices derived either from metrics computed at the species level and then combined for each assemblage (species rarity, origin, biodiversity conservation concern, functional uniqueness, functional originality, fishing interest), or from metrics directly computed at the assemblage level (species richness, assemblage rarity, diversity of biological traits). Each of these indices belongs to one of the four aspects of diversity. A synthetic index defined as the sum of the standardized aspects of diversity was used to assess the multi-faceted diversity of fish assemblages. We also investigated whether the two main environmental gradients at the catchment (distance from the sea) and at the floodplain (lateral connectivity of the waterbodies) scales influenced the diversity of fish assemblages, and consequently their potential conservation value. Finally, we propose that the floodplain waterbodies that should be conserved as a priority are those located in the downstream part of the catchment and which have a substantial lateral connectivity with the main channel.     

Rényi多样性指数曲线在评价和监测转基因作物对生物多样性影响中的应用

【背景】多样性比较是评价转基因作物对生物群落影响的标准参数之一。以往用于描述多样性的指数很多,但这些指数各有优缺点,生物学解释不全面。运用现代生物多样性表示方法,能够描述更复杂情况下的多样性关系。本文主要介绍了Rényi多样性指数曲线在生物安全研究中的应用。【方法】Rényi多样性指数曲线与以往常用的一维多样性指数不同,当其等级参数为某些特定值时代表了几个著名的多样性指数,该方法可较为确切地评价复杂的生物群落的多样性。本文在介绍其计算公式后,通过比较河北省2种转Bt基因棉田（孟山都33B和中棉30,均未施农药）以及常规管理（施用农药）和综合治理（IPM,施用农药和释放赤眼蜂相结合）条件下非转基因棉田的蜘蛛群落,证明了该方法的适用性。【结果】Rényi多样性指数曲线分析表明了所有可能的互作类型：1个明确的多样性排序以及2类棉田蜘蛛群落间不同的潜在关系。Rényi指数分析表明,在供试棉田中,中棉30棉田的蜘蛛多样性指数最高,其他3种棉田蜘蛛的多样性高低则难以一概而论。蜘蛛稀有种的多样性以孟山都33B棉田最高,IPM和常规棉田次之;蜘蛛常见种的多样性以IPM棉田最高,常规棉田和孟山都33B棉田次之。【结论与意义】Rényi多样性指数可用于生物群落多样性的综合评价,并可用于评价不同管理措施对生物多样性的影响。该方法弥补了仅用单一数量参数评价多样性的传统方法的缺陷,将不同参数整合成一个具有内在关联的概念框架。     

Tree Diversity in Western Kenya: Using Profiles to Characterise Richness and Evenness

Species diversity is a function of the number of species and the evenness in the abundance of the component species. We calculated diversity and evenness profiles, which allowed comparing the diversity and evenness of communities. We applied the methodology to investigate differences in diversity among the main functions of trees on western Kenyan farms. Many use-groups (all trees and species that provide a specific use) could not be ranked in diversity or evenness. No use-group had perfectly even distributions. Evenness could especially be enhanced for construction materials, fruit, ornamental, firewood, timber and medicine, which included some of the most species-rich groups of the investigated landscape. When considering only the evenness in the distribution of the dominant species, timber, medicine, fruit and beverage ranked lowest (> 60% of trees belonged to the dominant species of these groups). These are also use-groups that are mainly grown by farmers to provide cash through sales. Since not all communities can be ranked in diversity, studies that attempt to order communities in diversity should not base the ordering on a single index, or even a combination of several indices, but use techniques developed for diversity ordering such as the Rényi diversity profile. The rarefaction of diversity profiles described in this article could be used in studies that compare results from surveys with different sample sizes.     

Filling the gap in functional trait databases: use of ecological hypotheses to replace missing data

Functional trait databases are powerful tools in ecology, though most of them contain large amounts of missing values. The goal of this study was to test the effect of imputation methods on the evaluation of trait values at species level and on the subsequent calculation of functional diversity indices at community level using functional trait databases. Two simple imputation methods (average and median), two methods based on ecological hypotheses, and one multiple imputation method were tested using a large plant trait database, together with the influence of the percentage of missing data and differences between functional traits. At community level, the complete‐case approach and three functional diversity indices calculated from grassland plant communities were included. At the species level, one of the methods based on ecological hypothesis was for all traits more accurate than imputation with average or median values, but the multiple imputation method was superior for most of the traits. The method based on functional proximity between species was the best method for traits with an unbalanced distribution, while the method based on the existence of relationships between traits was the best for traits with a balanced distribution. The ranking of the grassland communities for their functional diversity indices was not robust with the complete‐case approach, even for low percentages of missing data. With the imputation methods based on ecological hypotheses, functional diversity indices could be computed with a maximum of 30% of missing data, without affecting the ranking between grassland communities. The multiple imputation method performed well, but not better than single imputation based on ecological hypothesis and adapted to the distribution of the trait values for the functional identity and range of the communities. Ecological studies using functional trait databases have to deal with missing data using imputation methods corresponding to their specific needs and making the most out of the information available in the databases. Within this framework, this study indicates the possibilities and limits of single imputation methods based on ecological hypothesis and concludes that they could be useful when studying the ranking of communities for their functional diversity indices.     

What saplings can tell us about forest expansion over natural grasslands

Questions: 1. Do the species composition, richness and diversity of sapling communities vary significantly in differently sized patches? 2. Do forest patches of different sizes differ in woody plant colonization patterns? Location: São Francisco de Paula, Rio Grande do Sul, Brazil, 29°28'S,50°13'W. Methods: Three woody vegetation types, differing in structural development (patch size) and recovering for 10 years from cattle and burning disturbances, were sampled on grassland. We analysed the composition and complexity of the woody sapling communities, through relative abundance, richness and diversity patterns. We also evaluated recruitment status (residents vs. colonizers) of species in communities occurring in different forest patch size classes. Results : 1. There is a compositional gradient in sapling communities strongly associated with forest patch area. 2. Richness and diversity are positively correlated to patch area, but only in poorly structured patches; large patches present richness and diversity values similar to small patches. 3. Resident to colonizer abundance ratio increases from nurse plants to large patches. The species number proportion between residents and colonizers is similar in small and large patches and did not differ between these patch types. 4. Large patches presented a high number of exclusive species, while nurse plants and small patches did not. Conclusions: Woody plant communities in Araucaria forest patches are associated with patch structure development. Richness and diversity patterns are linked to patch colonization patterns. Generalist species colonize the understorey of nurse plants and small patches; resident species cannot recruit many new individuals. In large patches, sapling recruitment by resident adults precludes the immigration of new species into the patches, limiting richness and diversity.     

A comparative test of phylogenetic diversity indices

Traditional measures of biodiversity, such as species richness, usually treat species as being equal. As this is obviously not the case, measuring diversity in terms of features accumulated over evolutionary history provides additional value to theoretical and applied ecology. Several phylogenetic diversity indices exist, but their behaviour has not yet been tested in a comparative framework. We provide a test of ten commonly used phylogenetic diversity indices based on 40 simulated phylogenies of varying topology. We restrict our analysis to a topological fully resolved tree without information on branch lengths and species lists with presence-absence data. A total of 38,000 artificial communities varying in species richness covering 5-95% of the phylogenies were created by random resampling. The indices were evaluated based on their ability to meet a priori defined requirements. No index meets all requirements, but three indices turned out to be more suitable than others under particular conditions. Average taxonomic distinctness (AvTD) and intensive quadratic entropy (J) are calculated by averaging and are, therefore, unbiased by species richness while reflecting phylogeny per se well. However, averaging leads to the violation of set monotonicity, which requires that species extinction cannot increase the index. Total taxonomic distinctness (TTD) sums up distinctiveness values for particular species across the community. It is therefore strongly linked to species richness and reflects phylogeny per se weakly but satisfies set monotonicity. We suggest that AvTD and J are best applied to studies that compare spatially or temporally rather independent communities that potentially vary strongly in their phylogenetic composition-i.e. where set monotonicity is a more negligible issue, but independence of species richness is desired. In contrast, we suggest that TTD be used in studies that compare rather interdependent communities where changes occur more gradually by species extinction or introduction. Calculating AvTD or TTD, depending on the research question, in addition to species richness is strongly recommended.     

停止人为去除植物功能群后的高寒草甸多样性恢复过程与群落构建

已有大量研究利用功能性状或系统发育来推断群落构建机制, 然而不同过程可能会导致相似的格局。本文基于对甘南高寒草甸植物功能群去除处理后群落恢复过程的跟踪调查, 对比了物种多样性、功能多样性和系统发育多样性的动态变化, 并分析了物种定殖与消失过程对功能多样性和系统发育多样性变化的影响。结果表明: 去除不同数量功能群的群落中: (1)包括物种丰富度(SR)、Shannon-Wiener指数(H°)和Simpson指数(D)在内的传统物种多样性均随时间快速上升并与自然群落趋同, 不同群落的均匀度指数(J)随时间呈增加趋势并趋于相似; 功能多样性(FD)与系统发育多样性(PD)呈现出与物种多样性相似的动态变化趋势, 而平均配对距离(MPD/MPD_a、MFD/MFD_a)则向中等程度聚集。(2)不同群落的功能群和物种组成在短期内均恢复到与自然群落非常相似的程度。(3)物种定殖与消失过程的功能格局是群落恢复过程中趋同效应的主要驱动力。本研究揭示了高寒草甸植物功能群去除停止后群落短期内快速恢复的过程, 说明在小尺度且周边具有大范围未退化草甸的情况下, 无论物种多样性、功能多样性还是系统发育多样性都具有较快的恢复能力, 同时说明了利用群落系统发育多样性格局来推断群落构建机制的局限性。     

Partitioning diversity for conservation analyses

Aim  Differentiation of sites or communities is often measured by partitioning regional or gamma diversity into additive or multiplicative alpha and beta components. The beta component and the ratio of within-group to total diversity (alpha/gamma) are then used to infer the compositional differentiation or similarity of the sites. There is debate about the appropriate measures and partitioning formulas for this purpose. We test the main partitioning methods, using empirical and simulated data, to see if some of these methods lead to false conclusions, and we show how to resolve the problems that we uncover.
Location  South America, Ecuador, Orellana province, Rio Shiripuno.
Methods  We construct sets of real and simulated tropical butterfly communities that can be unambiguously ranked according to their degree of differentiation. We then test whether beta and similarity measures from the different partitioning approaches rank these datasets correctly.
Results  The ratio of within-group diversity to total diversity does not reflect compositional similarity, when the Gini–Simpson index or Shannon entropy are used to measure diversity. Additive beta diversity based on the Gini–Simpson index does not reflect the degree of differentiation between N sites or communities.
Main conclusions  The ratio of within-group to total diversity (alpha/gamma) should not be used to measure the compositional similarity of groups, if diversity is equated with Shannon entropy or the Gini–Simpson index. Conversion of these measures to effective number of species solves these problems. Additive Gini–Simpson beta diversity does not directly reflect the differentiation of N samples or communities. However, when properly transformed onto the unit interval so as to remove the dependence on alpha and N , additive and multiplicative beta measures yield identical normalized measures of relative similarity and differentiation.     

戴云山物种多样性与系统发育多样性海拔梯度分布格局及驱动因子

生物多样性的海拔分布格局是生态学研究的热点。海拔作为综合性因子驱动着植物群落的物种、系统发育与功能多样性的空间分布。以戴云山南坡900-1600 m森林植物群落为研究对象,探讨物种多样性、系统发育指数与环境驱动因子的相互关系以及环境因子在群落构建与多样性维持中的重要意义。结果表明：（1）森林植物群落的系统发育多样性与物种多样性沿海拔均呈现中间高度膨胀格局。（2）物种多样性Margalef指数、Shannon-Wiener指数与系统发育多样性指数呈显著正相关,表明物种多样性越高,系统发育多样性也越高。Shannon-Wiener指数与物种多样性指数（Margalef、Pielou、Simpson指数）、系统发育多样性及系统发育结构都存在显著相关性,一定程度上Shannon-Wiener指数可以代替其他指数。Pielou指数、Simpson指数、Shannon-Wiener指数与系统发育结构NRI （Net relatedness index）指数、NTI （Net nearest taxa index）指数存在显著正相关,表明群落优势度、均匀度与系统发育结构相关性较强。（3）土壤全磷含量是影响系统发育多样性和物种多样性的主要驱动因子,土壤含水量是影响Shannon-Wiener、Pielou、Simpson指数的最显著因子,海拔是影响群落系统发育结构的主要因素。海拔是影响系统发育结构变化的主要环境因子,而土壤因子是影响物种多样性与系统发育多样性的主要因素,进一步验证了物种多样性与系统发育多样性的高度相关,结果旨在揭示物种群落空间分布规律。     

Productivity gradients cause positive diversity–invasibility relationships in microbial communities

Models predict that community invasibility generally declines with species diversity, a prediction confirmed by small‐scale experiments. Large‐scale observations and experiments, however, find that diverse communities tend to be more heavily invaded than simple communities. One hypothesis states that large‐scale environmental heterogeneity, which similarly influences native and invasive species, can cause a positive correlation between diversity and invasibility, overriding the local negative effects of diversity on invasibility. We tested this hypothesis using aquatic microbial communities consisting of protists and rotifers consuming bacteria and nanoflagellates. We constructed a productivity gradient to simulate large‐scale environmental heterogeneity, started communities with the same number of species along this gradient, and subjected equilibrial communities to invasion by non‐resident consumer species. Both invaders and most resident species increased their abundances with resource enrichment, resulting in a positive correlation between diversity and invasibility. Intraspecific interference competition within resident species and the positive effect of enrichment on the number of available resources probably accounted for the higher invasibility with enrichment. Our results provide direct experimental evidence that environmental heterogeneity in productivity can cause a positive diversity–invasibility relationship.     

阿拉善地区不同生境小型兽类群落多样性研究

本文对内蒙古阿拉善地区不同生境中小型兽类群落的物种组成和多样性进行了初步研究。1998 年8～9 月和1999 年8～9 月, 2002 年10 月, 在8 种不同生境类型中共布放54 个样地, 22 606 个有效铗日, 捕获小型兽类标本866 只, 分属2 目6 科, 共20 种。多样性分析结果表明: 8 种不同生境的小型兽类群落Shannon-Wiener 多样性指数在1.186～1.589 之间; Pielou 均匀性指数在0.630 9～0.861 8 之间; Simpson 生态优势度指数在0.234 9～0.415 7 之间; Margalef 丰富度指数在0.931 8～2.020 之间。根据不同生境的小型兽类群落相似性指数聚类结果,可以将8 种生境的小型兽类群落大致划分为4 种类型: 耐旱型(群落Ⅴ和群落Ⅶ)、干旱型(群落Ⅰ、Ⅱ、Ⅲ)、农林交错型(群落Ⅵ和群落Ⅷ) 和森林型(群落Ⅳ) 。生境条件的差异对小型兽类群落物种组成和多样性有一定的限制作用, 而人为干扰有明显影响。     

Species turnover in lake littorals: spatial and temporal variation of benthic macroinvertebrate diversity and community composition

Aims Despite wide consensus that ecological patterns and processes should be studied at multiple spatial scales, the temporal component of diversity variation has remained poorly examined. Specifically, rare species may exhibit patterns of diversity variation profoundly different from those of dominant taxa. Location Southern Finland. Methods We used multiplicative partitioning of true diversities (species richness, Shannon diversity) to identify the most important scale(s) of variation of benthic macroinvertebrate communities across several hierarchical scales, from individual samples to multiple littorals, lakes and years. We also assessed the among‐scale variability of benthic macroinvertebrate community composition by using measures of between‐ and within‐group distances at hierarchical scales. Results On average, a single benthic sample contained 23% of the total regional macroinvertebrate species pool. For both species richness and Shannon diversity, beta‐diversity was clearly the major component of regional diversity, with within‐littoral beta‐diversity (β₁) being the largest component of gamma‐diversity. The interannual component of total diversity was small, being almost negligible for Shannon index. Among‐sample (within‐littoral) diversity was related to variation of substratum heterogeneity at the same scale. By contrast, only a small proportion of rare taxa was found in an average benthic sample. Thus, dominant species among lakes and years were about the same, whereas rare species were mostly detected in a few benthic samples in one lake (or year). For rare species, the temporal component of diversity was more important than spatial turnover at most scales. Main conclusions While individual species occurrences and abundances, particularly those of rare taxa, may vary strongly through space and time, patterns of dominance in lake littoral benthic communities are highly predictable. Consequently, many rare species will be missed in temporally restricted samples of lake littorals. In comprehensive biodiversity surveys, interannual sampling of littoral macroinvertebrate communities is therefore needed.     

竹叶眼子菜种群遗传多样性与群落物种多样性关系的研究

以洪湖中的沉水植物群落及其优势种竹叶眼子菜(Potamogeton wrightii Morong)为研究对象,利用AFLP分子标记结合野外调查的方法,分析保护区(干扰较小)与非保护区(干扰较大,敞水区)中沉水植物群落的物种多样性与竹叶眼子菜遗传多样性之间的相关性。结果显示:洪湖沉水植物群落物种丰富度(S)和物种多样性辛普森指数(D)在保护区和非保护区均未发现显著性差异;竹叶眼子菜的重要值(IV)在保护区(5.2%~23.2%)较非保护区(8.5%~73.3%)稳定。竹叶眼子菜的遗传多样性在两个区未发现显著差异,其遗传多样性与群落物种多样性这两个指标在保护区、非保护区及全湖水平均不相关,说明沉水植物群落的物种多样性和竹叶眼子菜的遗传多样性对不同环境干扰的响应有所差异。     

物种多度和径级尺度对于评价群落系统发育结构的影响: 以尖峰岭热带山地雨林为例

研究不同径级尺度群落系统发育多样性有助于了解不同年龄模式下物种的亲缘关系及其群落系统发育结构; 但是关于物种多度对群落系统发育结构影响的研究较少。以海南尖峰岭热带山地雨林群落为例, 首先在不同径级尺度比较物种多度加权与否分别对4个广泛采用的系统发育指数的影响, 继而利用其中2个经过标准化处理的系统发育多样性指数: 净种间亲缘关系指数(net relatedness index, NRI)和净最近种间亲缘关系指数(nearest taxon index, NTI), 结合群落的生境类型来量度不同局域生境条件下不同径级尺度木本植物系统发育关系。结果发现: (1)未考虑物种多度加权的系统发育平均成对距离(mean pairwise distance, MPD)指数比考虑物种多度加权的MPD指数显著地高估了群落整体系统发育多样性, 且这种现象在小径级尺度(1 cm≤DBH<5 cm)最为明显。因此, 在森林监测样地中对于中、小径级群落系统发育结构研究中建议考虑物种多度信息。(2) 从群落组成整体系统发育结构来看, 尖峰岭热带山地雨林在几乎所有径级尺度和生境下均倾向于系统发育发散, 且随着径级的递增发散程度趋于明显(NRI<0)。(3)从群落组成局部系统发育结构来看, 尖峰岭热带山地雨林在中、小径级倾向于系统发育聚集(NTI>0), 而在大径级(DBH≥15 cm)则倾向于系统发育发散(NTI<0)。总之, 研究群落系统发育结构时应考虑物种多度的影响以及径级尺度效应。