The relationship between species replacement,dissimilarity derived from nestedness,and nestedness

Aim Beta diversity can be partitioned into two components: dissimilarity due to species replacement and dissimilarity due to nestedness ( Baselga, 2010 , Global Ecology and Biogeography, 19 , 134–143). Several contributions have challenged this approach or proposed alternative frameworks. Here, I review the concepts and methods used in these recent contributions, with the aim of clarifying: (1) the rationale behind the partitioning of beta diversity into species replacement and nestedness‐resultant dissimilarity, (2) how, based on this rationale, numerators and denominators of indices have to match, and (3) how nestedness and nestedness‐resultant dissimilarity are related but different concepts. Innovation The rationale behind measures of species replacement (turnover) dictates that the number of species that are replaced between sites (numerator of the index) has to be relativized with respect to the total number of species that could potentially be replaced (denominator). However, a recently proposed partition of Jaccard dissimilarity fails to do this. In consequence, this partition underestimates the contribution of species replacement and overestimates the contribution of richness differences to total dissimilarity. I show how Jaccard dissimilarity can be partitioned into meaningful turnover and nestedness components, and extend these new indices to multiple‐site situations. Finally the concepts of nestedness and nestedness‐resultant dissimilarity are discussed. Main conclusions Nestedness should be assessed using consistent measures that depend both on paired overlap and matrix filling, e.g. NODF, whereas beta‐diversity patterns should be examined using measures that allow the total dissimilarity to be separated into the components of dissimilarity due to species replacement and dissimilarity due to nestedness. In the case of multiple‐site dissimilarity patterns, averaged pairwise indices should never be used because the mean of the pairwise values is unable to accurately reflect the multiple‐site attributes of dissimilarity.     

Testing for differences in beta diversity from plot-to-plot dissimilarities

The central role of beta diversity in describing biodiversity patterns has been investigated in many fields of ecology and biogeography. While a variety of measures of beta diversity have been proposed over the past five decades, the question of how to test for differences in beta diversity among different sets of sampling plots has been addressed only rarely. Here, we describe a simple analytical procedure to test for differences in beta diversity among distinct sets of plots. The advantage of this approach compared to methods that have been previously proposed lies in its randomization procedure. Such a procedure creates a distribution of null values of the test statistic that is compatible with the null hypothesis of no significant difference in multivariate dispersion between the groups. The proposed test was illustrated using a large dataset of plant and water beetle (Coleoptera) assemblages collected from 45 farmland ponds in Ireland.     

A cautionary note on some phylogenetic dissimilarity measures

Aims Measures of plot-to-plot phylogenetic dissimilarity and beta diversity are providing a powerful tool for understanding the complex ecological and evolutionary mechanisms that drive community assembly.Methods Here, we review the properties of some previously published dissimilarity measures that are based on minimum or average phylogenetic dissimilarity between species in different plots.Important findings We first show that some of these measures violate the basic condition that for two identical plots the measures take the value zero. They also violate the condition that the dissimilarity between two identical plots should always be lower than that between two different plots. Such erratic behavior renders these measures unsuitable for measuring plot-to-plot phylogenetic dissimilarity. We next propose a new measure that satisfies these conditions, thus providing a more reasonable way for measuring phylogenetic dissimilarity.     

Incorporating functional dissimilarities into sample‐based rarefaction curves: from taxon resampling to functional resampling

Question: Indices of functional diversity have been seen as the key for integrating information on species richness with measures that focus on those components of community composition related to ecosystem functioning. For comparing species richness among habitats on an equal‐effort basis, so‐called sample‐based rarefaction curves may be used. Given a study area that is sampled for species presence and absence in N plots, sample‐based rarefaction generates the expected number of accumulated species as the number of sampled plots increases from 1 to N. Accordingly, the question for this study is: can we construct a ‘functional rarefaction curve’ that summarizes the expected functional dissimilarity between species when n plots are drawn at random from a larger pool of N plots? Methods: In this paper, we propose a parametric measure of functional diversity that is obtained by combining sample‐based rarefaction techniques that are usually applied to species richness with Rao's quadratic diversity. For a given set of N presence/absence plots, the resulting measure summarizes the expected functional dissimilarity at an increasingly larger cumulative number of plots n (n≤N). Results and Conclusions: Due to its parametric nature, the proposed measure is progressively more sensitive to rare species with increasing plot number, thus rendering this measure adequate for comparing the functional diversity of species assemblages that have been sampled with variable effort.     

Of beta diversity,variance, evenness,and dissimilarity

The amount of variation in species composition among sampling units or beta diversity has become a primary tool for connecting the spatial structure of species assemblages to ecological processes. Many different measures of beta diversity have been developed. Among them, the total variance in the community composition matrix has been proposed as a single‐number estimate of beta diversity. In this study, I first show that this measure summarizes the compositional variation among sampling units after nonlinear transformation of species abundances. Therefore, it is not always adequate for estimating beta diversity. Next, I propose an alternative approach for calculating beta diversity in which variance is substituted by a weighted measure of concentration (i.e., an inverse measure of evenness). The relationship between this new measure of beta diversity and so‐called multiple‐site dissimilarity measures is also discussed.     

Phylogenetic beta diversity of native and alien species in European urban floras

Aim Human activities have weakened biogeographical barriers to dispersal, increasing the rate of introduction of alien plants. However, their impact on beta diversity and floristic homogenization is poorly understood. Our goal is to compare the phylogenetic beta diversity of native species with that of two groups of alien species, archaeophytes and neophytes (introduced before and after ad 1500, respectively), across European urban floras to explore how biological invasions affect phylogenetic turnover at a continental scale. Location Twenty European cities located in six countries between 49 and 53° N latitude in continental Europe and the British Isles. Methods To compare the phylogenetic beta diversity of native and alien species we use the average phylogenetic dissimilarity of individual floras from their group centroid in multivariate space. Differences in phylogenetic beta diversity among different species groups are then assessed using a randomization test for homogeneity of multivariate dispersions. Results Across European urban floras, and when contrasted with natives, archaeophytes are usually associated with lower levels of phylogenetic beta diversity while neophytes tend to increase phylogenetic differentiation. Main conclusions While archaeophytes tend to promote limited homogenization in phylogenetic beta diversity, because of their diverse geographical origin together with short residence times in the invaded regions, neophytes are not promoting biotic homogenization of urban floras across Europe. Therefore, in spite of the increasing rate of alien invasion, an intense phylogenetic homogenization of urban cities is not to be expected soon.     

地理距离及环境差异对云南元江干热河谷植物群落beta多样性的影响

beta多样性反映了群落间物种组成的差异, 是生物多样性研究的热点之一。本研究通过对云南元江干热河谷41个植物群落样方进行调查, 用Jaccard相异系数表征物种beta多样性, 用样方之间的最近谱系距离(mean nearest taxon distance, MNTD)及平均谱系距离(mean pairwise distance, MPD)表征谱系beta多样性, 采用基于距离矩阵的多元回归和方差分解方法, 探讨了该区域干热河谷典型植物群落的物种beta多样性和谱系beta多样性与样方间环境差异(主要是气候)及地理距离之间的关系。结果表明: (1)群落间的地理距离和年平均温度差异对干热河谷植物群落的物种beta多样性和谱系beta多样性有显著影响; (2)地理距离对物种beta多样性和MNTD的影响最大; 地理距离和年平均温度差异对MPD的影响均较大; (3)样方间年平均温度与年平均降水量的差异和地理距离能够解释群落间beta多样性及谱系beta多样性11-13%的变异。以上结果表明, 生态位分化和扩散限制对该地区植物群落的beta多样性均有显著影响, 其中扩散限制的影响可能更大。此外, 人类活动等其他因素也很可能对元江干热河谷的群落组成具有非常重要的影响。     

Measuring fractions of beta diversity and their relationships to nestedness: a theoretical and empirical comparison of novel approaches

Beta diversity and nestedness are central concepts of ecology and biogeography and evaluation of their relationships is in the focus of contemporary ecological and conservation research. Beta diversity patterns are originated from two distinct processes: the replacement (or turnover) of species and the loss (or gain) of species leading to richness differences. Nested distributional patterns are generally thought to have a component deriving from beta diversity which is independent of replacement processes. Quantification of these phenomena is often made by calculating a measure of beta diversity, and the resulting value being subsequently partitioned into a contribution by species replacement plus a fraction shared by beta diversity and nestedness. Three methods have been recently proposed for such partitioning, all of them based on pairwise comparisons of sites. In this paper, the performance of these methods was evaluated on theoretical grounds and tested by a simulation study in which different gradients of dissimilarity, with known degrees of species replacement and species loss, were created. Performance was also tested using empirical data addressing land‐use induced changes in endemic arthropod communities of the Terceira Island in the Azores. We found that the partitioning of β_cc (dissimilarity in terms of the Jaccard index) into two additive fractions, β_‐3 (dissimilarity due to species replacement) plus β_rich (dissimilarity due to richness differences) reflects the species replacement and species loss processes across the simulated gradients in an ecologically and mathematically meaningful way, whilst the other two methods lack mathematical consistency and prove conceptually self‐contradictory. Moreover, the first method identified a selective local extinction process for endemic arthropods, triggered by land‐use changes, while the latter two methods overweighted the replacement component and led to false conclusions. Their basic flaw derives from the fact that the proposed replacement and nestedness components (deemed to account for species loss) are not scaled in the same way as the measure that accounts for the total dissimilarity (Sørensen and Jaccard indices). We therefore recommend the use of β_cc=β_‐3+β_rich, since its components are scaled in the same units and their responses are proportional to the replacement and the gain/loss of species.     

Different roles of elevational and local environmental factors on abundance‐based beta diversity of the soil Enchytraeidae on the Changbai Mountain

The elevational alpha biodiversity gradient in mountain regions is one of the well‐known ecological patterns, but its beta diversity pattern remains poorly known. Examining the beta diversity and its components could enhance the understanding of community assembly mechanism. We studied the beta diversity pattern of the soil enchytraeids along a distinct elevational gradient (705–2,280 m) on the Changbai Mountain, the best‐preserved mountain in northeastern China. The overall abundance‐based community dissimilarity was relatively high (ca. 0.70), largely due to the balanced‐variation component (85%). The overall dissimilarity and its balanced‐variation (substitution) component were related to both local environmental heterogeneity and elevational distance, with the environmental relationships being stronger. In contrast, the abundance‐gradient (subsets) component was not related to the two gradients. The same important spatial and environmental variables were detected in structuring overall dissimilarity and substitution component, different from that in subsets component. Variation partitioning analysis showed that environmental control played a more important role than spatial (vertical and horizontal) factors in structuring the patterns of overall beta diversity and its two components. The predictive power of multivariate analysis was higher for the substitution component (nearly 50%) and overall dissimilarity (35%), but much lower for subsets components (<4%). These findings implied that abundance‐based beta diversity patterns of the soil enchytraeids were the results of different ecological processes (e.g., environmental sorting and dispersal limitation), operating in the two antithetic components. Our study showed the substitution and loss of individuals reflecting different ecological processes and highlights the importance of partitioning beta diversity in assessing biodiversity patterns and their causes.     

Does Ordinal Cover Estimation Offer Reliable Quality Data Structures in Vegetation Ecological Studies?

Using visual estimation of species cover in ordinal interval classes may reduce costs in vegetation studies. In phytosociology, species cover within plots is usually estimated according to the well-known Braun-Blanquet scale and ordinal data from this scale are usually treated using common exploratory analysis tools that are adequate for ratio-scale variables only. This paper addresses whether the visual estimation of ordinal cover data and the treatment of these data with multivariate procedures tailored for ratio-scale data would lead to a significant loss of information with respect to the use of more accurate methods of data collection and analysis. To answer these questions we used three data sets sampled by different authors in different sites of Tuscany (central Italy) in which the species cover is measured with the point quadrat method. For each data set we used a Mantel test to compare the dissimilarity matrices obtained from the original point-quadrat cover data with those obtained from the corresponding ordinal interval classes. The results suggest that the ordinal data are suitable to represent the plot-to-plot dissimilarity structure of all data sets in a reasonable way and that in using such data there is no need to apply dissimilarity coefficients specifically tailored for ordinal scales.     

Effects of freshwater eutrophication on species and functional beta diversity of periphytic algae

Studies about beta diversity and environmental heterogeneity have shown that the strength of the environmental filtering effect may decrease with the increasing scale. These empirical results have related eutrophic aquatic environments to higher values of beta diversity, but never to dissimilarity of species and functional traits of periphytic algae. We tested the hypotheses that periphytic algae have higher dissimilarity of both species and functional traits in eutrophic environments, and that these dissimilarities are related to environmental dissimilarity. To this end, we used richness, density, and four functional traits of periphytic algae and local limnological data from wetlands in the Brazilian savanna (Cerrado). We analyzed the beta diversity and the relationship of species and functional dissimilarities with the environmental dissimilarity and geographic distances. Our hypothesis was confirmed for functional traits dissimilarity and for the importance of the environmental dissimilarity for both species and functional beta diversity. The cultural eutrophication led to a functional homogenization in urban wetlands, which indicates the establishment of species with similar ecological requirements, and consequently, similar ‘roles’ in the ecosystem, and also that sensitive species may have been replaced by tolerant species, leading to declining biodiversity.

     

Multivariate dispersion as a measure of beta diversity

Beta diversity can be defined as the variability in species composition among sampling units for a given area. We propose that it can be measured as the average dissimilarity from individual observation units to their group centroid in multivariate space, using an appropriate dissimilarity measure. Differences in beta diversity among different areas or groups of samples can be tested using this approach. The choice of transformation and dissimilarity measure has important consequences for interpreting results. For kelp holdfast assemblages from New Zealand, variation in species composition was greater in smaller holdfasts, while variation in relative abundances was greater in larger holdasts. Variation in community structure of Norwegian continental shelf macrobenthic fauna increased with increases in environmental heterogeneity, regardless of the measure used. We propose a new dissimilarity measure which allows the relative weight placed on changes in composition vs. abundance to be specified explicitly.     

美洲森林群落beta多样性的纬度梯度性

Beta多样性度量不同时空尺度物种组成的变化,是生物多样性的重要组成部分;理解其地理格局和形成机制已成为当前生物多样性研究的热点问题。基于Alwyn H. Gentry在美洲收集的131个森林样方数据,采用倍性和加性分配方法度量群落beta多样性,检验beta多样性随纬度的变化趋势,并分析其形成机制。研究表明:(1) 美洲森林群落beta多样性随纬度增加显著下降,热带和亚热带地区beta多样性高于温带地区;此格局可由物种分布范围的纬度梯度性和不同粒度(grain)下物种丰富度与纬度回归斜率的差异推论得出;(2) 加性分配方法表明beta多样性对各个温度带森林群落gamma多样性的相对贡献率平均为78.2%,并且随纬度升高而降低;(3) 美洲南半球森林群落beta多样性高于其北半球,这可能反映了区域间物种进化和环境变迁历史的差异。此外,还探讨了不同beta多样性计算方法的适用情景,首次证实了森林生态系统群落水平beta多样性的纬度梯度性,这对研究生物多样性的形成机制和生物多样性保护都具有重要的意义。     

A multiple-site dissimilarity measure for species presence/absence data and its relationship with nestedness and turnover

Multiple-site dissimilarity may be caused by two opposite processes of meta-community organization, such as species nestedness and turnover. Therefore, discriminating among these contributions is necessary for linking multiple-site dissimilarity to ecosystem functioning. This paper introduces a measure of multiple-site dissimilarity or beta diversity for presence/absence data that is based on information on species absences from the species × sites matrix. It is also shown that the newly proposed dissimilarity index can be additively partitioned into species nestedness and turnover.     

Is spectral distance a proxy of beta diversity at different taxonomic ranks? A test using quantile regression

Beta diversity represents a powerful indicator of ecological conditions because of its intrinsic relation with environmental gradients. In this view, remote sensing may be profitably used to derive models characterizing or estimating species turnover over an area. While several examples exist using spectral variability to estimate species diversity at several spatial scales, most of these have relied on standard correlation or regression approaches like the common Ordinary Least Square (OLS) regression which are problematic with noisy data. Moreover, very few attempts were made to derive beta diversity characterization models at different taxonomic ranks. In this paper, we performed quantile regression to test if spectral distance represents a good proxy of beta diversity considering different data thresholds and taxonomic ranks. We used plant distribution data from the North and South Carolina including 146 counties and covering a variety of vegetation formations. The dissimilarity in species composition at different taxonomic ranks (using Sørensen distance) among pairs of counties was compared with their distance in NDVI values derived from 23 yearly MODIS images. Our results indicate that (i) spectral variability represents a good proxy of beta diversity when appropriate statistics are applied and (ii) a lower taxonomic rank is important when changes in species composition are examined spatially using remotely sensed data.     

Contrasting responses of beta diversity components to environmental and host-associated factors in insect ectoparasites

1. The present study investigated whether different components (species replacement and species gains/losses) of compositional and phylogenetic beta diversity of insect ectoparasites responded similarly to environmental and host-associated gradients using a large dataset on distribution of fleas and their rodent hosts in Mongolia. 2. Generalised dissimilarity modelling was applied to investigate whether environmental variables or host dissimilarity was the best predictor of species/lineage replacement and species/lineage gains/losses (= richness difference) components of compositional and phylogenetic flea beta diversity. 3. The total compositional beta diversity of fleas was influenced mainly by the gradient in air temperature and, to a lesser degree, by total host beta diversity, with the former effect being associated with the richness difference component and the latter effect being associated with species replacement component. The total phylogenetic beta diversity of fleas was best explained by the total phylogenetic beta diversity of hosts, with this effect being mainly associated with the lineage replacement component, whereas the lineage richness difference component responded mainly to the temperature gradient. 4. The results of the present indicate that not only multiple beta diversity facets are driven by different factors, but also different components of the same beta diversity facet respond to different environmental (for parasites, including host-associated) gradients. These patterns were masked when only total beta diversity was analysed. 5. This emphasizes the importance of considering the components of insect beta diversity separately. Ignoring the separate components of beta diversity can lead to potentially erroneous inferences about the relative contribution of abiotic and biotic effects on beta diversity.     

Relationship of Productivity to Species Richness in the Xinjiang Temperate Grassland

The relationship between species richness (SR) and aboveground net primary productivity (ANPP) is still a central and debated issue in community ecology. Previous studies have often emphasized the relationship of alpha diversity (number of species identity) to the mean ANPP with respect to the SR-ANPP relationship while neglecting the contribution of beta diversity (dissimilarity in species composition) to the mean ANPP and to the stability of ANPP (coefficient of ANPP: CV of ANPP). In this study, we used alpha and beta diversity, mean ANPP and the CV of ANPP collected from 159 sites and belonging to three vegetation types in the Xinjiang temperate grassland to first examine their trends along climatic factors and among different vegetation types and then test the relationship among alpha (beta) diversity and mean ANPP and the CV of ANPP. Our results showed that in the Xinjiang temperate grasslands, alpha diversity was positively and linearly correlated with MAP but unimodally correlated with MAT. Meanwhile, beta diversity was unimodally correlated with MAP but linearly correlated with MAT. Relative to desert steppe, meadow steppe and typical steppe had the highest alpha and beta diversity, respectively. Except for ANPP exhibiting a quadratic relationship with MAP, no significant relationship was found among ANPP, the CV of ANPP and climatic factors. ANPP and the CV of ANPP also exhibited no apparent patterns in variation among different vegetation types. Our results further showed that mean ANPP was closely associated with alpha diversity. Both linear and unimodal relationships were detected between alpha diversity and mean ANPP, but their particular form was texture-dependent. Meanwhile, the CV of ANPP was positively correlated with beta diversity. Our results indicated that in addition to incorporating alpha diversity and mean ANPP, incorporating beta diversity and the CV of ANPP could expand our understanding of the SR-ANPP relationship.     

Extended dissimilarity: a method of robust estimation of ecological distances from high beta diversity data

It is widely accepted that reliable ordination of ecological data requires a strong linear or ordinal relationship between the dissimilarity of sites, based on species composition, and the ecological distance between them. Certain dissimilarity measures, having the property that they take a fixed maximum value when sites have no species in common, have been shown to be strongly correlated with ecological distance. For ecological gradients of moderate length (moderate beta diversity), such measures, in conjunction with non-metric multidimensional scaling, will reliably yield successful ordinations. However, as beta diversity increases, more sites have no species in common, and such measures invariably under-estimate ecological distance for such sites. Thus ordinations of data with high species turnover (high beta diversity) may fail.Extended dissimilarities are defined using an iterative adaptation of flexible shortest path adjustment applied to the matrix of dissimilarities with fixed maximum values. By means of theoretical argument and simulations, this is shown to lead to far stronger correlations between the adjusted site dissimilarity and ecological distance for ecological gradients of greater length than previously considered. Hence ordinations of extended dissimilarities, by means of either metric or non-metric scaling techniques, are shown to outperform corresponding ordinations of unadjusted dissimilarities, with the difference increasing with increasing beta diversity.     

Substratum simplification reduces beta diversity of stream algal communities

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