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.     

野生人参自然分布的典型原始阔叶红松林群落及其特征研究

本文通过物种多样性指数、生态优势度、群落均匀度、重要值等指标综合分析了野生人参自然分布的典型椴树阔叶红松林的群落特征．结果认为椴树阔叶红松林内物种数及其个体数量较多,多样性指数及群落的均匀度都较高,物种分布均匀并具有明显的层次,生态优势度较低,具有较稳定的群落结构．这种群落特征给人参的生活和生长创造了适宜的生态条件,是人参这一古老遗植物在这种群落中得以长期保存和生长繁衍的主要因素．     

Deterministic diversity changes in freshwater phytoplankton in the Yunnan–Guizhou Plateau lakes in China

Diversity measures reflect different aspects of a community, which are determined by different ecological processes. However, information is still limited on the ecological processes that are represented by different measures of species diversity. In this study, the primary driving factors for richness and diversity indices were tested. The possible ecological processes represented by each index were analyzed. First, the type of ecological process that governed the phytoplankton community in the Yunnan–Guizhou Plateau lakes, either deterministic or stochastic, was identified by Caswell's neutral model. The results indicate that a deterministic process governs the phytoplankton community. Second, the driving factors of richness and diversity indices were screened with mixed models. The results suggest that the variation of phytoplankton richness in different lakes or sites was primarily related to bottom-up factors. The variations in evenness and other measures based on the relative abundance were driven by both top-down and bottom-up factors, such as zooplankton biomass, and pH and mean light, respectively. Finally, although the different measures of diversity may respond to specific bottom-up or top-down processes, the responses to the two processes were not independent of each other. These findings will increase our understanding of the relationships between ecological processes and diversity measures for freshwater phytoplankton.     

Diversity patterns in grasslands along a landscape gradient in northwestern France

Abstract. Several measures of biodiversity were calculated (species richness SR, species diversity H', species evenness J', mean similarity, mosaic diversity and factorial diversity) in vascular plant communities along a landscape gradient in the Seine valley, Normandy, France. For these communities, we also recorded environmental and management data. Species and environmental data were analysed simultaneously by Canonical Correspondence Analysis (CCA) in order to study their relationships. CCA identifies one main landscape gradient linked to a set of highly linked ecological factors. Three community types were identified along this gradient: calcicolous communities on chalk slopes, mesophilous communities on colluvium and hydrophilous communities on alluviums. The measures of biodiversity between these groups and their variations along the landscape gradient indicate similar patterns for H', J' and SR. Between‐community biodiversity measures allow consideration of the distribution of species among communities in the landscape. Factorial diversity accounts for the organisation of the communities with reference to the basic mechanisms of species coexistence. Affinity analysis (similarity and mosaic diversity) measures the compositional pattern diversity, which is the function of the variation in species richness. We discuss the indicative versus the predictive value of these measures of biodiversity as regards ecological factors and processes and their application for conservation purposes.     

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.     

On the information-theoretical meaning of Hill's parametric evenness

The degree to which abundances are divided equitably among community species or evenness is a basic property of any biological community. Several evenness indices have been proposed to summarize community structure. However, despite their potential applicability in ecological research, none seems to be generally preferred. In this paper we show that, unlike other evenness indices without any clear information-theoretical meaning, Hill's parametric diversity measure E _,0 has an immediate relation to Rényi's generalized information. Therefore, E _,0 might be adequate for summarizing community structure within the context of a general theoretical framework of diversity analysis based on information theory.     

Exploring the Phylogenetic Structure of Ecological Communities: An Example for Rain Forest Trees

Because of the correlation expected between the phylogenetic relatedness of two taxa and their net ecological similarity, a measure of the overall phylogenetic relatedness of a community of interacting organisms can be used to investigate the contemporary ecological processes that structure community composition. I describe two indices that use the number of nodes that separate taxa on a phylogeny as a measure of their phylogenetic relatedness. As an example of the use of these indices in community analysis, I compared the mean observed net relatedness of trees (>/=10 cm diameter at breast height) in each of 28 plots (each 0.16 ha) in a Bornean rain forest with the net relatedness expected if species were drawn randomly from the species pool (of the 324 species in the 28 plots), using a supertree that I assembled from published sources. I found that the species in plots were more phylogenetically related than expected by chance, a result that was insensitive to various modifications to the basic methodology. I tentatively infer that variation in habitat among plots causes ecologically more similar species to co-occur within plots. Finally, I suggest a range of applications for phylogenetic relatedness measures in community analysis.     

The decoupling of abundance and species richness in lizard communities

1. Patterns of species richness often correlate strongly with measures of energy. The more individuals hypothesis (MIH) proposes that this relationship is facilitated by greater resources supporting larger populations, which are less likely to become extinct. Hence, the MIH predicts that community abundance and species richness will be positively related. 2. Recently, Buckley & Jetz (2010, Journal of Animal Ecology, 79, 358-365) documented a decoupling of community abundance and species richness in lizard communities in south-west United States, such that richer communities did not contain more individuals. They predicted, as a consequence of the mechanisms driving the decoupling, a more even distribution of species abundances in species-rich communities, evidenced by a positive relationship between species evenness and species richness. 3. We found a similar decoupling of the relationship between abundance and species richness for lizard communities in semi-arid south-eastern Australia. However, we note that a positive relationship between evenness and richness is expected because of the nature of the indices used. We illustrate this mathematically and empirically using data from both sets of lizard communities. When we used a measure of evenness, which is robust to species richness, there was no relationship between evenness and richness in either data set. 4. For lizard communities in both Australia and the United States, species dominance decreased as species richness increased. Further, with the iterative removal of the first, second and third most dominant species from each community, the relationship between abundance and species richness became increasingly more positive. 5. Our data support the contention that species richness in lizard communities is not directly related to the number of individuals an environment can support. We propose an alternative hypothesis regarding how the decoupling of abundance and richness is accommodated; namely, an inverse relationship between species dominance and species richness, possibly because of ecological release.     

An updated consumer’s guide to evenness and related indices

Ecologists widely agree that species diversity consists of two components, richness (the number of species) and evenness (a measure of the equitability of the proportional abundances of those species). However, no consensus on an exact definition of evenness (or equitability) has emerged. Instead, numerous equitability indices have been used in the ecological literature, as different researchers have preferred indices with different mathematical properties. In this paper, I show that the phrase ‘species diversity consists of two independent components, richness and evenness’ logically leads to one particular definition of evenness (Evenness = Diversity/Richness). To facilitate accurate communication, I propose that the term ‘evenness’ be used only to refer to this phenomenon, and that other terms be used for the equitability indices that measure other things. Here I provide a review of popular equitability indices, explain what each measures in practice, and show how they relate to each other and to evenness itself. I also explore how the partitioning of diversity into richness and evenness components is related to the partitioning of diversity into alpha and beta components. Dissecting the indices makes it easier to see the conceptual differences among them. Such understanding is necessary to ensure that an appropriate index is chosen for the questions at hand, as well as to interpret the index values correctly and to assess when index values can and when they cannot be considered comparable.     

广东森林群落的组成结构数量特征

本文探索森林群落组成结构水平的定量描述方法。通过用Shannon—Wiener多样性指数、Simpson生态优势度指标以及群落均匀度指标,测定广东范围内22个森林群落的物种多样性、生态优势度和群落均匀度。结果表明这三种指标值能有效地表征群落的组成结构特征;广东亚热带常绿阔叶林群落的物种多样性指数为4—5左右,群落均匀度为0.7—0.8左右,生态优势度为0.08—0.12左右。文章还对测定结果进行分析,进一步探讨了三种指标值的相互关系,三种指标值在群落学研究中的意义,以及在林业实践上的作用等。     

Structural analysis of fouling community development in the damariscotta river estuary,maine

The development of the marine fouling community at a station in the Damariscotta River Estuary was studied in respect to current theories of ecological succession. This community may be divided into two subcomponents, the primary fouling assemblage, which attaches directly to the substratum, and the secondary foulers which live on and among the primary fouling assemblage. Structure of the total fouling community was examined using measures of diversity, evenness, and dominance, as well as the numbers of species and abundance.Multiple regression analysis was used to establish a hierarchical list of the factors influencing secondary fouling community structure. A definite successional trend was observed, although the length of the study precluded any observation of a climax community.     

A semantic taxonomy for diversity measures

Community diversity has been studied extensively in relation to its effects on ecosystem functioning. Testing the consequences of diversity on ecosystem processes will require measures to be available based on a rigorous conceptualization of their very meaning. In the last decades, literally dozens of measures of diversity have been proposed. However, rather than using unrelated metrics, we need to identify their separate components so that possible links between them and ecosystem functioning can be examined using an agreed-upon language. In this paper, first, a short overview on new and old measures of community diversity is presented. Next, I propose a general framework in which most of the existing measures of diversity are sorted into four interrelated semantic classes: richness, abundance-weighted diversity, evenness and divergence. In this view, this paper constitutes an attempt to organize the very large number of existing diversity measures avoiding ambiguities in the meaning of the different facets of community diversity.     

Integrated ecosystem health assessment based on eco-exergy theory: A case study of the Jiangsu coastal area

The main objective of ecosystem health management is to preserve the capacity of ecosystems to respond to disturbances and future changes. We proposed a set of ecological indicators for coastal ecosystem health assessment using physical stressors such as total suspended matter, chemical stressors including nutrients and heavy metal pollutants, community structure metrics including species richness, diversity and evenness, and ecosystem level eco-exergy indicators. The results of our case study indicate that the health status of the Jiangsu coastal ecosystem is limited by environmental stressors and factors that affect the community species diversity. The health status of nektonic and benthic communities is reflected by water quality and sediment physicochemical properties, respectively. The results of our case study demonstrate that the integrated ecological health indicator system can provide a comprehensive assessment that corresponds with the current health of coastal ecosystems and a reliable theoretical basis for regional coastal management.     

The ecology of differences: assessing community assembly with trait and evolutionary distances

Species enter and persist in local communities because of their ecological fit to local conditions, and recently, ecologists have moved from measuring diversity as species richness and evenness, to using measures that reflect species ecological differences. There are two principal approaches for quantifying species ecological differences: functional (trait‐based) and phylogenetic pairwise distances between species. Both approaches have produced new ecological insights, yet at the same time methodological issues and assumptions limit them. Traits and phylogeny may provide different, and perhaps complementary, information about species' differences. To adequately test assembly hypotheses, a framework integrating the information provided by traits and phylogenies is required. We propose an intuitive measure for combining functional and phylogenetic pairwise distances, which provides a useful way to assess how functional and phylogenetic distances contribute to understanding patterns of community assembly. Here, we show that both traits and phylogeny inform community assembly patterns in alpine plant communities across an elevation gradient, because they represent complementary information. Differences in historical selection pressures have produced variation in the strength of the trait‐phylogeny correlation, and as such, integrating traits and phylogeny can enhance the ability to detect assembly patterns across habitats or environmental gradients.     

浙江天童山区灌丛群落的物种多样性及其与演替的关系

根据浙江天童山区灌丛群落的样地调查数据,采用Shannon－Wiener多样性指数、Simpson指数和群落均匀度指数研究了灌丛群落的物种多样性,并与当地的常绿阔叶林进行了比较。结果表明：这3种指标能够有效地表征亚热带灌丛群落的组成结构特征。天童灌丛群落灌木层的物种多样性指数 SW 高于常绿阔叶林中乔木层,而低于常绿阔叶林中灌木层。灌丛群落灌木层的生态优势度 SN 低于常绿阔叶林中乔木层,但高于常绿阔叶林中灌木层。灌丛群落灌木层的群落均匀度 PW 比常绿阔叶林中乔木层的和灌水层的都低。此外,文中还就本区灌丛群落物种多样性的特点,讨论了加速其进展演替的恢复措施。     

Cyanobacteria dominance influences resource use efficiency and community turnover in phytoplankton and zooplankton communities

Freshwater biodiversity loss potentially disrupts ecosystem services related to water quality and may negatively impact ecosystem functioning and temporal community turnover. We analysed a data set containing phytoplankton and zooplankton community data from 131 lakes through 9 years in an agricultural region to test predictions that plankton communities with low biodiversity are less efficient in their use of limiting resources and display greater community turnover (measured as community dissimilarity). Phytoplankton resource use efficiency (RUE = biomass per unit resource) was negatively related to phytoplankton evenness (measured as Pielou's evenness), whereas zooplankton RUE was positively related to phytoplankton evenness. Phytoplankton and zooplankton RUE were high and low, respectively, when Cyanobacteria, especially Microcystis sp., dominated. Phytoplankton communities displayed slower community turnover rates when dominated by few genera. Our findings, which counter findings of many terrestrial studies, suggest that Cyanobacteria dominance may play important roles in ecosystem functioning and community turnover in nutrient‐enriched lakes.     

Species interactions mediate phylogenetic community structure in a hyperdiverse lizard assemblage from arid Australia

Evolutionary history can exert a profound influence on ecological communities, but few generalities have emerged concerning the relationships among phylogeny, community membership, and niche evolution. We compared phylogenetic community structure and niche evolution in three lizard clades (Ctenotus skinks, agamids, and diplodactyline geckos) from arid Australia. We surveyed lizard communities at 32 sites in the northwestern Great Victoria Desert and generated complete species-level molecular phylogenies for regional representatives of the three clades. We document a striking pattern of phylogenetic evenness within local communities for all groups: pairwise correlations in species abundance across sites are negatively related to phylogenetic similarity. By modeling site suitability on the basis of species' habitat preferences, we demonstrate that phylogenetic evenness generally persists even after controlling for habitat filtering among species. This phylogenetic evenness is coupled with evolutionary lability of habitat-associated traits, to the extent that closely related species are more divergent in habitat use than distantly related species. In contrast, lizard diets are phylogenetically conserved, and pairwise dietary overlap between species is negatively related to phylogenetic distance in two of the three clades. Our results suggest that contemporary and historical species interactions have led to similar patterns of community structure across multiple clades in one of the world's most diverse lizard communities.