A link between ecological diversity indices and measures of biodiversity

The practice of environmental planning and protection frequently necessitates the quantification of ecological diversity. Traditional ‘ecological diversity indices’ are based on the abundances of species present. However, such indices are insensitive to taxonomic or similar differences. With equal species abundances they measure the species richness (species number) only. Conversely, so-called ‘biodiversity indices’ are based on species differences, but are insensitive to the abundance conditions. The quadratic entropy index is the only ecological diversity index, the value of which reflects both the differences ‘and’ abundances of the species. When a species list is given without abundance data, then, using the quadratic entropy index and postulating equal abundances, one gets the only biodiversity index derived from a traditional ecological index of diversity. Its extensive form is identical with the sum of differences or distances between the species present. This index trivially satisfies set monotonicity, an important property for biodiversity indices.     

The apportionment of quadratic entropy: a useful alternative for partitioning diversity in ecological data

Many methods that study the diversity within hierarchically structured populations have been developed in genetics. Among them, the analysis of molecular variance (AMOVA) (Excoffier et al., 1992) has the advantage of including evolutionary distances between individuals. AMOVA is a special case of a far more general statistical scheme produced by Rao (1982a; 1986) and called the apportionment of quadratic entropy (APQE). It links diversity and dissimilarity and allows the decomposition of diversity according to a given hierarchy. We apply this framework to ecological data showing that APQE may be very useful for studying diversity at various spatial scales. Moreover, the quadratic entropy has a critical advantage over usual diversity indices because it takes into account differences between species. Finally, the differences that can be incorporated in APQE may be either taxonomic or functional (biological traits), which may be of critical interest for ecologists.     

Quadratic diversity: Its maximization can reduce the richness of species

In recent decades numerous diversity indices have been introduced. Among them the quadratic entropy index Q expresses the mean difference between two individuals chosen from the community at random. Differing from diversity indices habitually employed, Q does not satisfy a property postulated earlier for those measures. Namely, the uniform distribution of species does not necessarily yield the maximal index value. Q is based on the difference matrix of species. For a given matrix one can seek for the vector yielding the maximum quadratic entropy. This task leads to a quadratic programming problem. Using the appropriate program of a program package, we determined the maximum vector for a genetic difference matrix of crane species, as published in the literature. We discovered that some components (frequencies) in the maximum vector are equal to zero. That is, by maximizing the quadratic diversity some species can be eliminated. We discuss briefly the possible implications of this observation. Moreover, even if all elements in the maximum vector are positive, the elements can differ.     

Additive partitioning of Rao's quadratic diversity: a hierarchical approach

Ecologists have long recognized three different components of species diversity: alpha or within-community diversity (α), beta or between-community diversity (β) and gamma or total species diversity in a region (γ). In this framework, β-diversity has been traditionally linked to the other diversity components through a multiplicative model so that it can be expressed as the ratio between γ-diversity and average α-diversity in a set of plots. Yet, more recently, ecologists are starting to partition diversity using the lesser known approach that α- and β-diversities sum to give the γ-diversity. This additive diversity partitioning is based on the decomposition of concave diversity measures for which the total diversity in a pooled set of communities exceeds (or equals) the average diversity within communities. In this paper, first, I shortly revise additive diversity partitioning for traditional diversity measures that are computed from species relative abundances. Next, I show that, under some specific circumstances, the same model can be extended to Rao's quadratic entropy, a measure that combines species relative abundances and pairwise interspecies differences. Finally, in the framework of taxonomic diversity, Rao's quadratic entropy has another decomposition: the sum over the Simpson indices at all the taxonomic levels. Thus, I show that, combining both partitioning models, the contribution of each level in the taxonomic hierarchy to the α- β- and γ-diversity components of Rao's quadratic entropy is made explicit. The proposed diversity decomposition is illustrated with a worked example on data from a plant community on ultramafic soils in Tuscany (central Italy).     

Spatial mismatch between wild bee diversity hotspots and protected areas

Wild bees are critical for multiple ecosystem functions but are currently threatened. Understanding the determinants of the spatial distribution of wild bee diversity is a major research gap for their conservation. We modeled wild bee α and β taxonomic and functional diversity in Switzerland to uncover countrywide diversity patterns and determine the extent to which they provide complementary information, assess the importance of the different drivers structuring wild bee diversity, identify hotspots of wild bee diversity, and determine the overlap between diversity hotspots and the network of protected areas. We used site-level occurrence and trait data from 547 wild bee species across 3343 plots and calculated community attributes, including taxonomic diversity metrics, community mean trait values, and functional diversity metrics. We modeled their distribution with predictors describing gradients of climate, resource availability (vegetation), and anthropogenic influence (i.e., land-use types and beekeeping intensity). Wild bee diversity changed along gradients of climate and resource availability; high-elevation areas had lower functional and taxonomic α diversity, and xeric areas harbored more diverse bee communities. Functional and taxonomic β diversities diverged from this pattern, with high elevations hosting unique species and trait combinations. The proportion of diversity hotspots included in protected areas depended on the biodiversity facet, but most diversity hotspots occurred in unprotected land. Climate and resource availability gradients drove spatial patterns of wild bee diversity, resulting in lower overall diversity at higher elevations, but simultaneously greater taxonomic and functional uniqueness. This spatial mismatch among distinct biodiversity facets and the degree of overlap with protected areas is a challenge to wild bee conservation, especially in the face of global change, and calls for better integrating unprotected land. The application of spatial predictive models represents a valuable tool to aid the future development of protected areas and achieve wild bee conservation goals.     

Scale-dependent variation in coral community similarity across sites, islands, and island groups

Community similarity is the proportion of species richness in a region that is shared on average among communities within that region. The slope of local richness (alpha diversity) regressed on regional richness (gamma diversity) can serve as an index of community similarity across regions with different regional richness. We examined community similarity in corals at three spatial scales (among transects at a site, sites on an island, and islands within an island group) across a 10 000-km longitudinal diversity gradient in the west-central Pacific Ocean. When alpha diversity was regressed on gamma diversity, the slopes, and thus community similarity, increased with scale (0.085, 0.261, and 0.407, respectively) because a greater proportion of gamma diversity was subsumed within alpha diversity as scale increased. Using standard randomization methods, we also examined how community similarity differed between observed and randomized assemblages and how this difference was affected by spatial separation of species within habitat types and specialization of species to three habitat types (reef flats, crests, and slopes). If spatial separation within habitat types and/or habitat specialization (i.e., underdispersion) occurs, fewer species are shared among assemblages than the random expectation. When the locations of individual coral colonies were randomized within and among habitat types, community similarity was 46-47% higher than that for observed assemblages at all three scales. We predicted that spatial separation of coral species within habitat types should increase with scale due to dispersal/extinction dynamics in this insular system, but that specialization of species to different habitat types should not change because habitat differences do not change with scale. However, neither habitat specialization nor spatial separation within habitat types differed among scales. At the two larger scales, each accounted for 22-24% of the difference in community similarity between observed and randomized assemblages. At the smallest scale (transect-site), neither spatial separation within habitat types nor habitat specialization had significant effects on community similarity, probably due to the small size of transect samples. The results suggest that coral species can disperse among islands in an island group as easily as they can among sites on an island over time scales that are relevant to their establishment and persistence on reefs.     

Assessing the Value of the Umbrella‐Species Concept for Conservation Planning with Meta‐Analysis

Abstract: The umbrella‐species concept, which suggests that conservation strategies designed for one species may benefit co‐occurring species, has been promoted as a framework for conservation planning. Nevertheless, there has been considerable variation in the outcome of empirical tests of this concept that has led researchers to question its value, so we used data from 15 published studies in a meta‐analysis to evaluate whether conservation of putative umbrella species also conserves co‐occurring species. We tested the effectiveness of putative umbrella species categorized by taxonomic group, taxonomic similarity to co‐occurring species, body size, generality of resource use, and trophic level to evaluate criteria proposed to guide the selection of umbrella species. We compared species richness and number of individuals (by species and higher taxonomic group) between sites with and without putative umbrella species to test whether more co‐occurring species were present in greater abundances when the area or resource needs of umbrella species were met. Species richness and abundance of co‐occurring species were consistently higher in sites where umbrella species were present than where they were not and for conservation schemes with avian than with mammalian umbrella species. There were no differences in species richness or species abundance with resource generalist or specialist umbrella species or based on taxonomic similarity of umbrella and co‐occurring species. Taxonomic group abundance was higher in across‐taxonomic umbrella species schemes than when umbrella species were of the same taxon as co‐occurring species. Co‐occurring species had similar, or higher, species richness with small‐bodied umbrella species relative to larger‐bodied umbrella species. The only significant difference among umbrella species categorized by trophic level was that species richness was higher with omnivorous than it was with carnivorous avian umbrella species. Our results suggest there is merit to the umbrella‐species concept for conservation, but they do not support the use of the criteria we used to identify umbrella species.     

Does plant species co-occurrence influence soil mite diversity?

Few studies have considered whether plant taxa can be used as predictors of belowground faunal diversity in natural ecosystems. We examined soil mite (Acari) diversity beneath six grass species at the Konza Prairie Biological Station, Kansas, USA. We tested the hypotheses that soil mite species richness, abundance, and taxonomic diversity are greater (1) beneath grasses in dicultures (different species) compared to monocultures (same species), (2) beneath grasses of higher resource quality (lower C:N) compared to lower resource quality, and (3) beneath heterogeneous mixes of grasses (C3 and C4 grasses growing together) compared to homogeneous mixes (C3 or C4 grasses) using natural occurrences of plant species as treatments. This study is the first to examine the interaction between above- and belowground diversity in a natural setting with species-level resolution of a hyper-diverse taxon. Our results indicate that grasses in diculture supported a more species and phylogenetically rich soil mite fauna than was observed for monocultures and that this relationship was significant at depth but not in the upper soil horizon. We noted that mite species richness was not linearly related to grass species richness, which suggests that simple extrapolations of soil faunal diversity based on plant species inventories may underestimate the richness of associated soil mite communities. The distribution of mite size classes in dicultures was considerably different than those for monocultures. There was no difference in soil mite richness between grass combinations of differing resource quality, or resource heterogeneity.     

Effectiveness of terrestrial protected areas for conservation of lake fish communities

Freshwater protected areas are rare even though freshwater ecosystems are among the most imperiled in the world. Conservation actions within terrestrial protected areas (TPAs) such as development or resource extraction regulations may spill over to benefit freshwater ecosystems within their boundaries. Using data from 175 lakes across Ontario, Canada, we compared common indicators of fish‐assemblage status (i.e., species richness, Shannon diversity index, catch per unit effort, and normalized‐length size spectrum slopes) to evaluate whether TPAs benefit lake fish assemblages. Nearest neighbor cluster analysis was used to generate pairs of lakes: inside versus outside, inside versus bordering, and bordering versus outside TPAs based on lake characteristics. The diversity and abundance indicators did not differ significantly across comparisons, but normalized‐length size spectrum slopes (NLSS) were significantly steeper in lakes outside parks. The latter indicated assemblage differences (greater abundances of small‐bodied species) and less‐efficient energy transfer through the trophic levels of assemblages outside parks. Although not significantly different, pollution‐ and turbidity‐tolerant species were more abundant outside parks, whereas 3 of the 4 pollution‐intolerant species were more abundant within parks. Twenty‐one percent of the difference in slopes was related to higher total dissolved solids concentrations and angling pressure. Our results support the hypothesis that TPAs benefit lake fish assemblages and suggest that NLSS slopes are informative indicators for aquatic protected area evaluations because they represent compositional and functional aspects of communities.     

Selection of Multiple Umbrella Species for Functional and Taxonomic Diversity to Represent Urban Biodiversity

Surrogates, such as umbrella species, are commonly used to reduce the complexity of quantifying biodiversity for conservation purposes. The presence of umbrella species is often indicative of high taxonomic diversity; however, functional diversity is now recognized as an important metric for biodiversity and thus should be considered when choosing umbrella species. We identified umbrella species associated with high taxonomic and functional biodiversity in urban areas in Switzerland. We analyzed 39,752 individuals of 574 animal species from 96 study plots and 1397 presences of 262 plant species from 58 plots. Thirty‐one biodiversity measures of 7 taxonomic groups (plants, spiders, bees, ground beetles, lady bugs, weevils and birds) were included in within‐ and across‐taxa analyses. Sixteen measures were taxonomical (species richness and species diversity), whereas 15 were functional (species traits including mobility, resource use, and reproduction). We used indicator value analysis to identify umbrella species associated with single or multiple biodiversity measures. Many umbrella species were indicators of high biodiversity within their own taxonomic group (from 33.3% in weevils to 93.8% in birds), to a lesser extent they were indicators across taxa. Principal component analysis revealed that umbrella species for multiple measures of biodiversity represented different aspects of biodiversity, especially with respect to measures of taxonomic and functional diversity. Thus, even umbrella species for multiple measures of biodiversity were complementary in the biodiversity aspects they represented. Thus, the choice of umbrella species based solely on taxonomic diversity is questionable and may not represent biodiversity comprehensively. Our results suggest that, depending on conservation priorities, managers should choose multiple and complementary umbrella species to assess the state of biodiversity. Selección de Múltiples Especies Paraguas para la Diversidad Funcional y Taxonómica para Representar la Biodiversidad Urbana     

Invertebrate Morphospecies as Surrogates for Species: A Case Study

Environmental monitoring and conservation evaluation in terrestrial habitats may be enhanced by the use of invertebrate inventories, but taxonomic and logistic constraints frequently encountered during conventional taxonomic treatment have greatly restricted their use. To overcome this problem we suggest that nonspecialists may be used to classify invertebrates to morphospecies without compromising scientific accuracy. To test this proposition, large pitfall and litter samples of ants, beetles, and spiders from four forest types were sorted to morphospecies by a nonspecialist and to species by specialists. These data were used to generate morphospecies and species inventories and to estimate richness (α diversity) and turnover (β diversity), information frequently used in the above activities. Our results show that the estimates of richness of ants and spiders varied little between morphospecies and species inventories. Differences between estimates of beetle richness were largely influenced by errors of identification in two families, Curculionidae and Staphylinidae. But morphospecies and species inventories yielded identical ranking of forest type using richness. Turnover was assessed by sample ordination, which revealed similar clusters regardless of the type of inventory. Analysis of similarities of assemblages of ants and beetles showed significant differences between all forest types. Spider assemblages showed a lower level of discrimination. The assessment of turnover was consistent among inventories but different between the major taxa. Our findings suggest that morphospecies may be used as surrogates for species in some environmental monitoring and conservation, in particular when decisions are guided by estimates of richness and the assessment of turnover.     

古田山国家级自然保护区常绿阔叶林类型及其群落物种多样性研究

在群落调查的基础上，分析了古田山自然保护区常绿阔叶林的群落类型和群落物种多样性特征．结果表明：古田山自然保护区常绿阔叶林主要有6种类型(群系)，即甜槠林、栲树林、野含笑-钩栗林、青冈林、虎皮楠-甜槠林、乌冈栎-青冈林，并对各群落类型的结构和物种组成进行了描述．从各群落的外貌、结构和种类组成上看，均具有我国典型常绿阔叶林的基本特征．不同群落类型其物种多样性大小不同，栲树林和野含笑-钩栗林物种多样性较高，虎皮楠-甜槠林和乌冈栎-青冈林物种多样性较低．在群落垂直结构中，灌木层→乔木层→草本层物种多样性依次降低，乔木层与灌木层之间物种多样性差异不显著，草本层的物种丰富度、物种多样性指数均明显小于乔木层和灌木层．与邻近4个山地常绿阔叶林物种多样性相比较，古田山常绿阔叶林物种多样性比纬度位置高的黄山和大别山要大，但比纬度位置低的乌岩岭和缙云山小．     

不同浓度植物根系分泌物微生态效应研究

探讨夹竹桃(Nerium oleander)根系分泌物对土壤微生物量碳、呼吸强度等微生物学特征及土壤微生物群落功能多样性的影响,深入揭示夹竹桃根系分泌物的微生态效应,通过向土壤中添加植物根系分泌物溶液的方法,研究了不同浓度(CK,15 mL去离子水作为对照;LC,5 mL分泌物+10 mL去离子水;MC,10 mL分泌...     

Diversity of ground arthropod community at organic and chemically intensive tea plantation of Darjeeling terai

Tea in Darjeeling foothills and terai are grown conventionally, with application of chemical fertilizers and pesticides, as well as organically without these inputs. Ground level arthropod community was collected from the above two types of tea plots using pitfall traps. Catches from these environments showed variation in the arthropod faunal structure with numerically and taxonomically greater abundance in the organic than that of the conventional plot. Coleopterans were more diverse with largest number of families and Recognizable Taxonomic Units (or morphospecies) in the organic tea plot. The diversity and similarity indices for coleopterans were comparable, in organic and conventional tea plots at species and family levels. The close relationship of the indices suggested that diversity study at family level could be used as surrogate for species level diversity; thus alleviating the laborious and expertise job of taxonomic identification of arthropod species. Faunal diversity study at ground level gave the clue that soil of the organic plantation was healthier than that of the conventional tea plot.     

Measuring diversity: the importance of species similarity

Realistic measures of biodiversity should reflect not only the relative abundances of species, but also the differences between them. We present a natural family of diversity measures taking both factors into account. This is not just another addition to the already long list of diversity indices. Instead, a single formula subsumes many of the most popular indices, including Shannon's, Simpson's, species richness, and Rao's quadratic entropy. These popular indices can then be used and understood in a unified way, and the relationships between them are made plain. The new measures are, moreover, effective numbers, so that percentage changes and ratio comparisons of diversity value are meaningful. We advocate the use of diversity profiles, which provide a faithful graphical representation of the shape of a community; they show how the perceived diversity changes as the emphasis shifts from rare to common species. Communities can usefully be compared by comparing their diversity profiles. We show by example that this is a far more subtle method than any relying on a single statistic. Some ecologists view diversity indices with suspicion, questioning whether they are biologically meaningful. By dropping the naive assumption that distinct species have nothing in common, working with effective numbers, and using diversity profiles, we arrive at a system of diversity measurement that should lay much of this suspicion to rest.     

Quantifying effects of pollution on biodiversity: a case study of highly diverse molluscan assemblages in the Mediterranean

Structured sampling designs are important in the assessment of environmental impacts of variable ecological systems. Recent developments have provided a useful framework extending existing univariate techniques into a multivariate context. Measures of taxonomic relatedness have also been introduced, which complement existing measures of diversity of assemblages. In this study, the potential effects of sewage discharge on spatial patterns of highly diverse molluscan assemblages in a Mediterranean rocky subtidal habitat were investigated. Nine 20 cm×20 cm quadrats were taken from each of three sites (80 m–100 m apart) within a putatively impacted location near a sewage outfall (I) and at each of two control locations (Cs) by destructive sampling by SCUBA divers at a depth of 3 m–4 m. A total of 5507 specimens of 151 species were collected. The average and the variance in total abundance of molluscs were greater, on average, at I than at Cs. Higher abundances at the sewage outfall were largely driven by greater numbers of juvenile molluscs. The Shannon diversity of molluscs (H′) was significantly lower at I, but no difference among locations was detected for the total number of species (S). In addition, the taxonomic distinctness (Δ*) of molluscs was greater at Cs, although it was more variable at I. Multivariate analyses showed that there was a significant difference in the structure of assemblages at I compared with Cs. The location near the outfall was characterized by greater abundances of several species, including especially the gastropods Pusillina philippi, Bittium latreilli, and Bittium reticulatum. There was also greater variability in the structure of assemblages among sites and among quadrats at control locations compared to those near the outfall. Using a suite of univariate and multivariate measures, including diversity indices, detailed information on taxonomic structure and analyses of variability at different spatial scales, provided useful insights into the effects of sewage impacts on these diverse assemblages. These results also highlighted the importance of analysing measures of variance, as well as mean in considering effects of stress in natural communities.     

Priority Wetland Invertebrates as Conservation Surrogates

Abstract: Invertebrates are important functionally in most ecosystems, but seldom appraised as surrogate indicators of biological diversity. Priority species might be good candidates; thus, here we evaluated whether three freshwater invertebrates listed in the U.K. Biodiversity Action Plan indicated the richness, composition, and conservation importance of associated wetland organisms as defined respectively by their alpha diversity, beta diversity, and threat status. Sites occupied by each of the gastropods Segmentina nitida, Anisus vorticulus, and Valvata macrostoma had greater species richness of gastropods and greater conservation importance than other sites. Each also characterized species assemblages associated with significant variations between locations in alpha or beta diversity among other mollusks and aquatic macrophytes. Because of their distinct resource requirements, conserving the three priority species extended the range of wetland types under management for nature conservation by 18% and the associated gastropod niche‐space by around 33%. Although nonpriority species indicated variations in richness, composition, and conservation importance among other organisms as effectively as priority species, none characterized such a wide range of high‐quality wetland types. We conclude that priority invertebrates are no more effective than nonpriority species as indicators of alpha and beta diversity or conservation importance among associated organisms. Nevertheless, conserving priority species can extend the array of distinct environments that are protected for their specialized biodiversity and environmental quality. We suggest that this is a key role for priority species and conservation surrogates more generally, and, on our evidence, can best be delivered through multiple species with contrasting habitat requirements.     

The invasibility of marine algal assemblages: role of functional diversity and identity

The emergence of the biodiversity-ecosystem functioning debate in the last decade has renewed interest in understanding why some communities are more easily invaded than others and how the impact of invasion on recipient communities and ecosystems varies. To date most of the research on invasibility has focused on taxonomic diversity, i.e., species richness. However, functional diversity of the communities should be more relevant for the resistance of the community to invasions, as the extent of functional differences among the species in an assemblage is a major determinant of ecosystem processes. Although coastal marine habitats are among the most heavily invaded ecosystems, studies on community invasibility and vulnerability in these habitats are scarce. We carried out a manipulative field experiment in tide pools of the rocky intertidal to test the hypothesis that increasing functional richness reduces the susceptibility of macroalgal communities to invasion. We selected a priori four functional groups on the basis of previous knowledge of local species characteristics: encrusting, turf, subcanopy, and canopy species. Synthetic assemblages containing one, two, three, or four different functional groups of seaweeds were created, and invasion by native species was monitored over an eight-month period. Cover and resource availability in the assemblages with only one functional group showed different patterns in the use of space and light, confirming true functional differences among our groups. Experimental results showed that the identity of functional groups was more important than functional richness in determining the ability of macroalgal communities to resist invasion and that resistance to invasion was resource-mediated.     

Conserving alpha and beta diversity in wood-production landscapes

International demand for wood and other forest products continues to grow rapidly, and uncertainties remain about how animal communities will respond to intensifying resource extraction associated with woody bioenergy production. We examined changes in alpha and beta diversity of bats, bees, birds, and reptiles across wood production landscapes in the southeastern United States, a biodiversity hotspot that is one of the principal sources of woody biomass globally. We sampled across a spatial gradient of paired forest land-uses (representing pre and postharvest) that allowed us to evaluate biological community changes resulting from several types of biomass harvest. Short-rotation practices and residue removal following clearcuts were associated with reduced alpha diversity (−14.1 and −13.9 species, respectively) and lower beta diversity (i.e., Jaccard dissimilarity) between land-use pairs (0.46 and 0.50, respectively), whereas midrotation thinning increased alpha (+3.5 species) and beta diversity (0.59). Over the course of a stand rotation in a single location, biomass harvesting generally led to less biodiversity. Cross-taxa responses to resource extraction were poorly predicted by alpha diversity: correlations in responses between taxonomic groups were highly variable (−0.2 to 0.4) with large uncertainties. In contrast, beta diversity patterns were highly consistent and predictable across taxa, where correlations in responses between taxonomic groups were all positive (0.05–0.4) with more narrow uncertainties. Beta diversity may, therefore, be a more reliable and information-rich indicator than alpha diversity in understanding animal community response to landscape change. Patterns in beta diversity were primarily driven by turnover instead of species loss or gain, indicating that wood extraction generates habitats that support different biological communities.