A metric of biodiversity that integrates abundance,phylogeny, and function

A metric of biodiversity is proposed that combines three of its key components: abundance, phylogeny, and ecological function. This metric is an expansion of the current abundance‐based metric that uses Hill numbers, the effective number of types in a sample if all types had the same mean proportional abundance. I define analogous proportional measures of phylogenetic divergence and functional distinctiveness. Phylogenetic divergence is measured as the sum of the proportional share of each species of a given branch of a phylogeny. Functional distinctiveness can be measured in two ways, as the proportional share of each species of a specified ecological function, or as the relative distance of each species based on functional trait values. Because all three aspects of biodiversity are measured in the same fashion (relative proportions) in similar units (effective numbers of species), an integrated metric can be defined. The combined metric provides understanding of covariation among the components and how management for one component may trade off against others. The metric can be partitioned into components of richness and evenness, and into subsets and variation among subsets, all of which can be related through a simple multiplicative framework. This metric is a complement to, rather than a replacement of, current metrics of phylogenetic and functional diversity. More work is needed to link this new metric to ecological theory, determine its error structure, and devise methods for its effective assessment.     

Functional and phylogenetic dimensions are more important than the taxonomic dimension for capturing variation in stream fish communities

Biodiversity is inherently multidimensional in nature, differences in evolutionary history, attributes of species, taxonomic composition constitutes a small fraction of whole variation present in this multidimensional space. Despite its multidimensional characteristic, biodiversity has been traditionally measured by assessing its dimensions separately through metrics of diversity. However, assessing multiple dimensions in a common framework opens the possibility of answering interesting questions that, until now, are poorly understood, such as: What dimensions capture most of the variation present in biodiversity among communities? We assess this question by extending the framework of Importance Values (IVs) to three dimensions of variation in biodiversity, functional, taxonomic and phylogenetic, and evaluate which of these captures the most variation in biodiversity space. To address this question we used data from stream fish communities of the Ivinhema River Basin in Brazil. We found that functional and phylogenetic dimensions are more important than the taxonomic dimension (represented by richness) in capturing variation in the biodiversity space formed by these three dimensions together. Furthermore, the IVs of these three dimensions were similar along an altitudinal gradient, indicating similar contributions by a given dimension in different environmental conditions. We highlight the importance of adopting a multidimensional approach when describing biodiversity, since richness (the proxy for taxonomic dimension), despite being the most commonly used, is an incomplete surrogate to capture the variation present in the biodiversity space of stream fish communities.     

Contrasting changes in the abundance and diversity of North American bird assemblages from 1971 to 2010

Although it is generally recognized that global biodiversity is declining, few studies have examined long‐term changes in multiple biodiversity dimensions simultaneously. In this study, we quantified and compared temporal changes in the abundance, taxonomic diversity, functional diversity, and phylogenetic diversity of bird assemblages, using roadside monitoring data of the North American Breeding Bird Survey from 1971 to 2010. We calculated 12 abundance and diversity metrics based on 5‐year average abundances of 519 species for each of 768 monitoring routes. We did this for all bird species together as well as for four subgroups based on breeding habitat affinity (grassland, woodland, wetland, and shrubland breeders). The majority of the biodiversity metrics increased or remained constant over the study period, whereas the overall abundance of birds showed a pronounced decrease, primarily driven by declines of the most abundant species. These results highlight how stable or even increasing metrics of taxonomic, functional, or phylogenetic diversity may occur in parallel with substantial losses of individuals. We further found that patterns of change differed among the species subgroups, with both abundance and diversity increasing for woodland birds and decreasing for grassland breeders. The contrasting changes between abundance and diversity and among the breeding habitat groups underscore the relevance of a multifaceted approach to measuring biodiversity change. Our findings further stress the importance of monitoring the overall abundance of individuals in addition to metrics of taxonomic, functional, or phylogenetic diversity, thus confirming the importance of population abundance as an essential biodiversity variable.     

How to choose a biodiversity indicator – Redundancy and complementarity of biodiversity metrics in a freshwater ecosystem

A range of biodiversity metrics are available to assess the ecological integrity of aquatic ecosystems. However, performance varies considerably among different types of metrics and provides different information regarding ecosystem conditions, thus making difficult the selection of appropriate metrics for biomonitoring. The present study evaluated the robustness of six biodiversity metrics to assess environmental change and determine their utility as relevant indicators of ecosystem biodiversity and functionality. Traditional metrics such as species richness and Shannon diversity were considered along with alternative metrics such as functional diversity, size diversity and taxonomic distinctness. To that end, invertebrate assemblages in a river floodplain were used as a case study to evaluate the performance of metrics using Generalized Additive Models (GAM). GAM explained between eight and 49% of the variability in biodiversity. The regression models exhibited differences in the response of biodiversity indicators to environmental factors, suggesting that intermediate levels of turbidity and low salinity are conditions favouring increased biodiversity in the study area. Based on correlations among metrics and responses to primary environmental factors, it is concluded that Shannon and functional diversity, and rarefied species richness generated similar information regarding ecosystem conditions (i.e., the metrics were redundant); while size diversity and distinctness provided useful additional data characterizing ecosystem quality (i.e., the metrics were complementary). Functional diversity indicated not only number and dominance of species, but also each species functional role in the community, and was therefore the most informative biodiversity metric. Nevertheless, the use of a combination of metrics, for example functional and size diversity, and variation in taxonomic distinctness, provides complementary data that will serve to achieve a more thorough understanding of ecosystem structure and function, and response to primary environmental influences.     

A guide to phylogenetic metrics for conservation,community ecology and macroecology

The use of phylogenies in ecology is increasingly common and has broadened our understanding of biological diversity. Ecological sub‐disciplines, particularly conservation, community ecology and macroecology, all recognize the value of evolutionary relationships but the resulting development of phylogenetic approaches has led to a proliferation of phylogenetic diversity metrics. The use of many metrics across the sub‐disciplines hampers potential meta‐analyses, syntheses, and generalizations of existing results. Further, there is no guide for selecting the appropriate metric for a given question, and different metrics are frequently used to address similar questions. To improve the choice, application, and interpretation of phylo‐diversity metrics, we organize existing metrics by expanding on a unifying framework for phylogenetic information. Generally, questions about phylogenetic relationships within or between assemblages tend to ask three types of question: how much; how different; or how regular? We show that these questions reflect three dimensions of a phylogenetic tree: richness, divergence, and regularity. We classify 70 existing phylo‐diversity metrics based on their mathematical form within these three dimensions and identify ‘anchor’ representatives: for α‐diversity metrics these are PD (Faith's phylogenetic diversity), MPD (mean pairwise distance), and VPD (variation of pairwise distances). By analysing mathematical formulae and using simulations, we use this framework to identify metrics that mix dimensions, and we provide a guide to choosing and using the most appropriate metrics. We show that metric choice requires connecting the research question with the correct dimension of the framework and that there are logical approaches to selecting and interpreting metrics. The guide outlined herein will help researchers navigate the current jungle of indices.     

Stronger Tests of Mechanisms Underlying Geographic Gradients of Biodiversity: Insights from the Dimensionality of Biodiversity

Inference involving diversity gradients typically is gathered by mechanistic tests involving single dimensions of biodiversity such as species richness. Nonetheless, because traits such as geographic range size, trophic status or phenotypic characteristics are tied to a particular species, mechanistic effects driving broad diversity patterns should manifest across numerous dimensions of biodiversity. We develop an approach of stronger inference based on numerous dimensions of biodiversity and apply it to evaluate one such putative mechanism: the mid-domain effect (MDE). Species composition of 10,000-km² grid cells was determined by overlaying geographic range maps of 133 noctilionoid bat taxa. We determined empirical diversity gradients in the Neotropics by calculating species richness and three indices each of phylogenetic, functional and phenetic diversity for each grid cell. We also created 1,000 simulated gradients of each examined metric of biodiversity based on a MDE model to estimate patterns expected if species distributions were randomly placed within the Neotropics. For each simulation run, we regressed the observed gradient onto the MDE-expected gradient. If a MDE drives empirical gradients, then coefficients of determination from such an analysis should be high, the intercept no different from zero and the slope no different than unity. Species richness gradients predicted by the MDE fit empirical patterns. The MDE produced strong spatially structured gradients of taxonomic, phylogenetic, functional and phenetic diversity. Nonetheless, expected values generated from the MDE for most dimensions of biodiversity exhibited poor fit to most empirical patterns. The MDE cannot account for most empirical patterns of biodiversity. Fuller understanding of latitudinal gradients will come from simultaneous examination of relative effects of random, environmental and historical mechanisms to better understand distribution and abundance of the current biota.     

Taxonomic,functional, and phylogenetic dimensions of rodent biodiversity along an extensive tropical elevational gradient

Relationships among taxonomic, functional, and phylogenetic dimensions of biodiversity provide insight about the relative contributions of ecological and evolutionary processes in structuring local assemblages. We used data for rodent species distributions from an extensive tropical elevational gradient to 1) describe elevational gradients for each of three dimensions of biodiversity, 2) evaluate the sufficiency of species richness as a surrogate for other dimensions, and 3) quantify the relative support for mechanisms that increase or decrease phylogenetic or functional dispersion. Taxonomic biodiversity was quantified by species richness, as well as by richness, evenness, diversity, dominance, and rarity at generic and familial levels. Morphological and categorical traits were used to estimate functional biodiversity, and an ultrametric mammalian supertree was used as the basis for estimating phylogenetic biodiversity. Elevational gradients of each dimension of biodiversity were strong, with significant linear and non‐linear components based on orthogonal polynomial regression. Empirical linear and non‐linear regression components were consistently different than those expected based on species richness for generic, familial, and phylogenetic biodiversity, but not for functional biodiversity. Nevertheless, the congruence of dimensions of biodiversity based on correlation analyses indicated that any one dimension is a useful surrogate for the other dimensions for rodents at Manu. Given variation in species richness, assemblages from lowland rainforests comprised more biodiversity than expected, whereas assemblages from cloud and elfin forests represented less biodiversity than expected. Warm temperatures, vertical complexity of the vegetation, and high productivity likely facilitate niche differentiation in rainforests, whereas cricetid rodents are competitively superior to other clades in the less structurally complex, less productive, and colder, high elevation habitats.     

Evidence of neotropical anuran community disruption on rice crops: a multidimensional evaluation

Agricultural expansion is a major driver of biodiversity loss, especially in the megadiverse tropics. Rice is among the world’s most important food crops, invariably affecting biodiversity worldwide. Although the effects of habitat conversion to rice crops on biodiversity are not completely understood, landscape modification often creates conditions that benefit some species and excludes others. We conducted an integrative evaluation of the effects that habitat conversion to irrigated rice crops has on anuran communities from a Cerrado-Amazon ecotone. We adopted a multidimensional approach to compare anuran communities from agricultural and pristine environments considering (i) taxonomic metrics; (ii) functional and phylogenetic diversity; (iii) selected and excluded traits and (iv) body condition indices. When compared to their pristine counterparts, agricultural waterbodies showed increased functional divergence and decreased species diversity and functional richness. Furthermore, agricultural anuran communities exhibited lower phylogenetic diversity. Nonetheless, taxonomic diversity did not vary significantly, suggesting that it should not be used without complementary metrics. Species with small range, habitat specialization, small clutches and large body size were excluded from rice crops. Furthermore, frogs showed lower body condition in crops than in pristine areas. Understanding how species traits correlate with specific responses to agriculture will allow better predictions of the functional effects of anthropogenic land-use. Maintaining high diversity in anthropogenic environments is important for ecosystem resilience because diverse communities are more likely to hold multiple species capable of contributing to ecological functions. Our results show that converting natural vegetation to irrigated rice crops drives many species to local extinction, and resilient species to exhibit lower body condition.     

Old and new challenges in using species diversity for assessing biodiversity

Although the maintenance of diversity of living systems is critical for ecosystem functioning, the accelerating pace of global change is threatening its preservation. Standardized methods for biodiversity assessment and monitoring are needed. Species diversity is one of the most widely adopted metrics for assessing patterns and processes of biodiversity, at both ecological and biogeographic scales. However, those perspectives differ because of the types of data that can be feasibly collected, resulting in differences in the questions that can be addressed. Despite a theoretical consensus on diversity metrics, standardized methods for its measurement are lacking, especially at the scales needed to monitor biodiversity for conservation and management purposes. We review the conceptual framework for species diversity, examine common metrics, and explore their use for biodiversity conservation and management. Key differences in diversity measures at ecological and biogeographic scales are the completeness of species lists and the ability to include information on species abundances. We analyse the major pitfalls and problems with quantitative measurement of species diversity, look at the use of weighting measures by phylogenetic distance, discuss potential solutions and propose a research agenda to solve the major existing problems.     

Taxonomic and functional diversity covary in rock pool microalgal communities despite their different drivers

Local biodiversity has traditionally been estimated with taxonomic diversity metrics such as species richness. Recently, the concept of biodiversity has been extended beyond species identity by ecological traits determining the functional role of a species in a community. This interspecific functional diversity typically responds more strongly to local environmental variation compared with taxonomic diversity, while taxonomic diversity may mirror more strongly dispersal processes compared with functional metrics. Several trait‐based indices have been developed to measure functional diversity for various organisms and habitat types, but studies of their applicability on aquatic microbial communities have been underrepresented. We examined the drivers and covariance of taxonomic and functional diversity among diatom rock pool communities on the Baltic Sea coast. We quantified three taxonomic (species richness, Shannon''s diversity, and Pielou''s evenness) and three functional (functional richness, evenness, and divergence) diversity indices and determined abiotic factors best explaining variation in these indices by generalized linear mixed models. The six diversity indices were highly collinear except functional evenness, which merely correlated significantly with taxonomic evenness. All diversity indices were always explained by water conductivity and temperature–sampling month interaction. Taxonomic diversity was further consistently explained by pool distance to the sea, and functional richness and divergence by pool location. The explained variance in regression models did not markedly differ between taxonomic and functional metrics. Our findings do not clearly support the superiority of neither set of diversity indices in explaining coastal microbial diversity, but rather highlight the general overlap among the indices. However, as individual metrics may be driven by different factors, the greatest advantage in assessing biodiversity is nevertheless probably achieved with a simultaneous application of the taxonomic and functional diversity metrics.     

Multiple facets of stream macroinvertebrate alpha diversity are driven by different ecological factors across an extensive altitudinal gradient

Environmental filtering and spatial structuring are important ecological processes for the generation and maintenance of biodiversity. However, the relative importance of these ecological drivers for multiple facets of diversity is still poorly understood in highland streams. Here, we examined the responses of three facets of stream macroinvertebrate alpha diversity to local environmental, landscape‐climate and spatial factors in a near‐pristine highland riverine ecosystem. Taxonomic (species richness, Shannon diversity, and evenness), functional (functional richness, evenness, divergence, and Rao's Quadratic entropy), and a proxy of phylogenetic alpha diversity (taxonomic distinctness and variation in taxonomic distinctness) were calculated for macroinvertebrate assemblages in 55 stream sites. Then Pearson correlation coefficient was used to explore congruence of indices within and across the three diversity facets. Finally, multiple linear regression models and variation partitioning were employed to identify the relative importance of different ecological drivers of biodiversity. We found most correlations between the diversity indices within the same facet, and between functional richness and species richness were relatively strong. The two phylogenetic diversity indices were quite independent from taxonomic diversity but correlated with functional diversity indices to some extent. Taxonomic and functional diversity were more strongly determined by environmental variables, while phylogenetic diversity was better explained by spatial factors. In terms of environmental variables, habitat‐scale variables describing habitat complexity and water physical features played the primary role in determining the diversity patterns of all three facets, whereas landscape factors appeared less influential. Our findings indicated that both environmental and spatial factors are important ecological drivers for biodiversity patterns of macroinvertebrates in Tibetan streams, although their relative importance was contingent on different facets of diversity. Such findings verified the complementary roles of taxonomic, functional and phylogenetic diversity, and highlighted the importance of comprehensively considering multiple ecological drivers for different facets of diversity in biodiversity assessment.     

Scale‐dependent complementarity of climatic velocity and environmental diversity for identifying priority areas for conservation under climate change

As most regions of the earth transition to altered climatic conditions, new methods are needed to identify refugia and other areas whose conservation would facilitate persistence of biodiversity under climate change. We compared several common approaches to conservation planning focused on climate resilience over a broad range of ecological settings across North America and evaluated how commonalities in the priority areas identified by different methods varied with regional context and spatial scale. Our results indicate that priority areas based on different environmental diversity metrics differed substantially from each other and from priorities based on spatiotemporal metrics such as climatic velocity. Refugia identified by diversity or velocity metrics were not strongly associated with the current protected area system, suggesting the need for additional conservation measures including protection of refugia. Despite the inherent uncertainties in predicting future climate, we found that variation among climatic velocities derived from different general circulation models and emissions pathways was less than the variation among the suite of environmental diversity metrics. To address uncertainty created by this variation, planners can combine priorities identified by alternative metrics at a single resolution and downweight areas of high variation between metrics. Alternately, coarse‐resolution velocity metrics can be combined with fine‐resolution diversity metrics in order to leverage the respective strengths of the two groups of metrics as tools for identification of potential macro‐ and microrefugia that in combination maximize both transient and long‐term resilience to climate change. Planners should compare and integrate approaches that span a range of model complexity and spatial scale to match the range of ecological and physical processes influencing persistence of biodiversity and identify a conservation network resilient to threats operating at multiple scales.     

Species pool size and rainfall account for the relationship between biodiversity and biomass production in natural forests of China

The strength of biodiversity–biomass production relationships increases with increasing environmental stress and time. However, we know little about the effects of abiotic (e.g., climate) and biotic (e.g., species pool and community composition) factors on this trend. Whether variation in biomass production is best explained by phylogenetic diversity metrics or traditional measures of species richness also remains elusive. We compiled estimates of community composition and biomass production for tree species in 111 permanent quadrats spanning three natural forests (tropical, subtropical, and temperate) in China. Based on ~10 years of data, we compared temperature, rainfall, species pool size, and community composition in each forest each year. We estimated species richness and phylogenetic diversity in each quadrat each year; the latter metric was based on the sum of branch lengths of a phylogeny that connects species in each quadrat each year. Using generalized linear mixed‐effect models, we found that top‐ranked models included the interaction between forest and biodiversity and the interaction between forest and year for both biodiversity metrics. Variation in biomass production was best explained by phylogenetic diversity; biomass production generally increased with phylogenetic diversity, and the relationship was stronger in subtropical and temperate forests. Increasing species pool size, temperature, and rainfall and decreasing inter‐quadrat dissimilarity range shifted the relationship between biomass production and phylogenetic diversity from positive to neutral. When considered alone, species pool size had the strongest influence on biomass production, while species pool size, rainfall, and their interaction with phylogenetic diversity constituted the top‐ranked model. Our study highlights the importance of species pool size and rainfall on the relationship between phylogenetic diversity and biomass production in natural forest ecosystems.     

Selecting plant species for practical restoration of degraded lands using a multiple‐trait approach

Ecological restoration is essential in rehabilitating degraded areas and safeguarding biodiversity, ecosystem services and human welfare. Using functional traits to plan restoration strategies has been suggested as they are the main ecological attributes that underlie ecosystem processes and services. However, few studies have translated ecological theory into actual restoration practices that can be easily used by different stakeholders. In this article, we applied a multiple‐trait approach to select plant species for the restoration of degraded lands inside the Brazilian Amazon Forests. We selected 10 traits encompassing ease of management, geographical distribution and interactions with animals and other ecosystem services and scored these traits using 118 native species. Then, we ranked all species according to the total number of traits that they exhibited to obtain a list of 53 highly ranked species. In addition, we employed non‐metric multidimensional scaling (NMDS) to assess the variation in these traits across the entire group of species. Based on the results, we selected a subset of species that maximizes functional diversity (high variability). We performed a sparse linear discriminant analysis (SLDA) to highlight a minimum set of traits to effectively discriminate botanical families. The final list of species and their traits highlight the importance of preserving not only the historical reference of a focused ecosystem but also its functional diversity to restore the interaction with local fauna, enrich the food chain and guarantee ecosystem services for local communities.     

Multifaceted Analysis of Patch-Level Plant Diversity in Response to Landscape Spatial Pattern and History on Mediterranean Dunes

Landscape structure is known to critically affect biodiversity. However, although the multi-facetted character of biodiversity is widely recognized, few studies have linked landscape spatial pattern and history simultaneously to multiple facets (taxonomic, functional, and phylogenetic) and spatial components (α, β, and γ) of plant diversity. We set out to reveal whether landscape parameters have specific effects on the separate diversity facets and components of plant diversity at a patch scale on coastal dune landscapes of Central Italy. For each landscape patch, we computed a set of patch-based metrics relying on multi-temporal land-cover maps. Based on a database of plant community plots, on functional traits from field measurements and on a dated phylogenetic tree, we calculated taxonomic (TD), functional (FD), and phylogenetic diversity (PD) within each patch at α, β, and γ level. Diversity measures were then related to the landscape metrics via linear mixed-effect models. Landscape pattern and transformations affected TD only moderately in coastal dune ecosystems. We found much stronger and contrasted effects on FD and PD. FD increased in patches surrounded by human-dominated habitats; PD was higher in fragmented patches, particularly in the Mediterranean macchia. Moreover, landscape pattern affected differently the single communities, the turnover among communities and the pool of species within the patch (α, β, and γ components). Our results call for the combined inclusion of FD and PD and their partitions into ecological analyses, being TD too crude to capture the comprehensive and contrasted response of plant diversity to landscape spatial pattern.     

Phylogenetic community structure metrics and null models: a review with new methods and software

Competitive exclusion and habitat filtering influence community assembly, but ecologists and evolutionary biologists have not reached consensus on how to quantify patterns that would reveal the action of these processes. Currently, at least 22 α‐diversity and 10 β‐diversity metrics of community phylogenetic structure can be combined with nine null models (eight for β‐diversity metrics), providing 278 potentially distinct approaches to test for phylogenetic clustering and overdispersion. Selecting the appropriate approach for a study is daunting. First, we describe similarities among metrics and null models across variance in phylogeny size and shape, species abundance, and species richness. Second, we develop spatially explicit, individual‐based simulations of neutral, competitive exclusion, or habitat filtering community assembly, and quantify the performance (type I and II error rates) of all 278 metric and null model combinations against each assembly process. Many α‐diversity metrics and null models are at least functionally equivalent, reducing the number of truly unique metrics to 12 and the number of unique metric + null model combinations to 72. An even smaller subset of metric and null model combinations showed robust statistical performance. For α‐diversity metrics, phylogenetic diversity and mean nearest taxon distance were best able to detect habitat filtering, while mean pairwise phylogenetic distance‐based metrics were best able to detect competitive exclusion. Overall, β‐diversity metrics tended to have greater power to detect habitat filtering and competitive exclusion than α‐diversity metrics, but had higher type 1 error in some cases. Across both α‐ and β‐diversity metrics, null model selection affected type I error rates more than metric selection. A null model that maintained species richness, and approximately maintained species occurrence frequency and abundance across sites, exhibited low type I and II error rates. This regional null model simulates neutral dispersal of individuals into local communities by sampling from a regional species pool. We introduce a flexible new R package, metricTester, to facilitate robust analyses of method performance.     

Amphibian Species Contribute Similarly to Taxonomic,but not Functional and Phylogenetic Diversity: Inferences from Amphibian Biodiversity on Emei Mountain

Understanding the relationships between species,communities,and biodiversity are important challenges in conservation ecology.Current biodiversity conservation activities usually focus on species that are rare,endemic,distinctive,or at risk of extinction.However,empirical studies of whether such species contribute more to aspects of biodiversity than common species are still relatively rare.The aim of the present study was to assess the contribution of individual amphibian species to different facets of biodiversity,and to test whether species of conservation interest contribute more to taxonomic,functional,and phylogenetic diversity than do species without special conservation status.To answer these questions,19 000 simulated random communities with a gradient of species richness were created by shuffling the regional pool of species inhabiting Emei Mountain.Differences of diversity values were then computed before and after removing individual species in these random communities.Our results indicated that although individual species contributed similarly to taxonomic diversity,their contribution to functional and phylogenetic diversity was more idiosyncratic.This was primarily driven by the diverse functional attributes of species and the differences in phylogenetic relationships among species.Additionally,species of conservation interest did not show a significantly higher contribution to any facet of biodiversity.Our results support the claims that the usefulness of metrics based only on species richness is limited.Instead,assemblages that include species with functional and phylogenetic diversity should be protected to maintain biodiversity.     

The components of biodiversity,with a particular focus on phylogenetic information

We present a framework for biodiversity metrics that organizes the growing panoply of metrics. Our framework distinguishes metrics based on the type of information–abundance, phylogeny, function–and two common properties–magnitude and variability. Our new metrics of phylogenetic diversity are based on a partition of the total branch lengths of a cladogram into the proportional share of each species, including: a measure of divergence which standardizes the amount of evolutionary divergence by species richness and time depth of the cladogram; a measure of regularity which is maximal when the tree is perfectly symmetrical so that all species have the same proportional branch lengths; a measure that combines information on the magnitude and variability of abundance with phylogenetic variability, and a measure of phylogenetically weighted effective mean abundance; and indicate how those metrics can be decomposed into α and β components. We illustrate the utility of these new metrics using empirical data on the bat fauna of Manu, Peru. Divergence was greatest in lowland rainforest and at the transition between cloud and elfin forests, and least in upper elfin forests and in cloud forests. In contrast, regularity was greatest in lowland rainforest, dipping to its smallest values in mid‐elevation cloud forests, and then increasing in high elevation elfin forests. These patterns indicate that the first species to drop out with increasing elevation are ones that are closely related to other species in the metacommunity. Measures of the effective number of phylogenetically independent or distinct species decreased very rapidly with elevation, and β‐diversity was larger. In contrast, a comparison of feeding guilds shows a different effect of phylogenetic patterning. Along the elevational gradient, each guild generally loses some species from each clade–rather than entire clades–explaining the maintenance of functional diversity as phylogenetic diversity decreases.