Loss of species and functions in a deforested megadiverse tropical forest.

Tropical species richness is threatened by habitat degradation associated with land-use conversion, yet the consequences for functional diversity remain little understood. Progress has been hindered by difficulties in obtaining comprehensive species-level trait information to characterize entire assemblages and insufficient appreciation that increasing land-cover heterogeneity potentially compensates for species loss. We examined the impacts of tropical deforestation associated with land-use heterogeneity on bird species richness, functional redundancy, functional diversity, and associated components (i.e., alpha diversity, species dissimilarity, and interaction strength of the relationship between abundance and functional dissimilarity). We analyzed over 200 georeferenced bird assemblages in the Atlantic Forest of Brazil. We characterized the functional role of the species of each assemblage and modeled biodiversity metrics as a function of forest cover and land-cover heterogeneity. Replacement of native Atlantic Forest with a mosaic of land uses (e.g., agriculture, pastures, and urbanization) reduced bird species richness in a nonrandom way. Core forest species, or species considered sensitive to edges, tended to be absent in communities in heterogenous environments. Overall, functional diversity and functional redundancy of bird species were not affected by forest loss. However, birds in highly heterogenous habitats were functionally distinct from birds in forest, suggesting a shift in community composition toward mosaic-exclusive species led by land-cover heterogeneity. Threatened species of the Atlantic Forest did not seem to tolerate degraded and heterogeneous environments; they remained primarily in areas with large forest tracts. Our results shed light on the complex effects of native forest transformation to mosaics of anthropogenic landscapes and emphasize the importance of considering the effects of deforestation and land-use heterogeneity when assessing deforestation effects on Neotropical biodiversity.

Pérdida de especies y funciones en un bosque tropical megadiverso deforestado Resumen La riqueza de especies tropicales está amenazada por la degradación asociada con la conversión del uso de suelo, y aun así entendemos muy poco de las consecuencias que esto tiene para la diversidad funcional. El progreso está obstaculizado por las dificultades para obtener información completa de los rasgos a nivel de especie para caracterizar ensamblajes completos y la apreciación insuficiente de que la heterogeneidad creciente de la cobertura del suelo tiene el potencial para compensar la pérdida de especies. Analizamos el impacto que tiene la deforestación tropical asociada con la heterogeneidad del uso de suelo sobre la riqueza de especies de aves, la redundancia funcional, la diversidad funcional y sus componentes asociados (es decir, diversidad alfa, disimilitud de especies y fuerza de interacción de la relación entre la abundancia y la disimilitud funcional). Analizamos más de 200 ensamblajes georreferenciados de aves en el Bosque Atlántico de Brasil. Caracterizamos el papel funcional de las especies de cada ensamblaje y modelamos las medidas de biodiversidad como función de la cobertura forestal y de la heterogeneidad del uso de suelo. La sustitución del Bosque Atlántico nativo con un mosaico de usos de suelo (p. ej.: agricultura, pastura y urbanización) redujo la riqueza de especies de una manera no aleatoria. Las especies nucleares del bosque, o las especies consideradas como sensibles a los bordes, tendieron a estar ausentes en las comunidades de los ambientes heterogéneas. En general, la diversidad y la redundancia funcionales de las especies de aves no se vieron afectadas por la pérdida del bosque. Sin embargo, las aves en los hábitats con alta heterogeneidad eran funcionalmente distintas a las aves de los bosques, lo que sugiere un cambio en la composición x de la comunidad hacia especies exclusivas de mosaicos llevadas por la heterogeneidad de la cobertura del suelo. Las especies amenazadas del Bosque Atlántico no parecieron tolerar el ambiente degradado y heterogéneo pues permanecieron principalmente en las áreas con grandes extensiones de bosque. Nuestros resultados arrojan luz sobre los efectos complejos de la transformación de los bosques nativos en mosaicos de paisajes antropogénicos y recalcan la importancia de considerar los efectos de la deforestación y la heterogeneidad del uso de suelo cuando se evalúan los efectos de la deforestación sobre la biodiversidad neotropical.

Disturbed habitats locally reduce the signal of deep evolutionary history in functional traits of plants.

The functioning of present ecosystems reflects deep evolutionary history of locally cooccurring species if their functional traits show high phylogenetic signal (PS). However, we do not understand what drives local PS. We hypothesize that local PS is high in undisturbed and stressful habitats, either due to ongoing local assembly of species that maintained ancestral traits, or to past evolutionary maintenance of ancestral traits within habitat species-pools, or to both. We quantified PS and diversity of 10 traits within 6704 local plant communities across 38 Dutch habitat types differing in disturbance or stress. Mean local PS varied 50-fold among habitat types, often independently of phylogenetic or trait diversity. Mean local PS decreased with disturbance but showed no consistent relationship to stress. Mean local PS exceeded species-pool PS, reflecting nonrandom subsampling from the pool. Disturbance or stress related more strongly to mean local than to species-pool PS. Disturbed habitats harbour species with evolutionary divergent trait values, probably driven by ongoing, local assembly of species: environmental fluctuations might maintain different trait values within lineages through an evolutionary storage effect. If functional traits do not reflect phylogeny, ecosystem functioning might not be contingent on the presence of particular lineages, and lineages might establish evolutionarily novel interactions.

Revisiting species and areas of interest for conserving global mammalian phylogenetic diversity.

Various prioritisation strategies have been developed to cope with accelerating biodiversity loss and limited conservation resources. These strategies could become more engaging for decision-makers if they reflected the positive effects conservation can have on future projected biodiversity, by targeting net positive outcomes in future projected biodiversity, rather than reflecting the negative consequences of further biodiversity losses only. Hoping to inform the post-2020 biodiversity framework, we here apply this approach of targeting net positive outcomes in future projected biodiversity to phylogenetic diversity (PD) to re-identify species and areas of interest for conserving global mammalian PD. We identify priority species/areas as those whose protection would maximise gains in future projected PD. We also identify loss-significant species/areas as those whose/where extinction(s) would maximise losses in future projected PD. We show that our priority species/areas differ from loss-significant species/areas. While our priority species are mostly similar to those identified by the EDGE of Existence Programme, our priority areas generally differ from previously-identified ones for global mammal conservation. We further highlight that these newly-identified species/areas of interest currently lack protection and offer some guidance for their future management.

Biomass of slow life history species increases as local bottom trawl effort decreases in the Celtic sea.

Due to its selective removal, fishing pressure has long influenced the dynamics of species based on their life history traits. Sensitivity to fishing increases along a "fast-to-slow" gradient of life history strategies, and the "slow" species (large, long-lived, late-maturing, giving birth to few large offspring) require the most time to recover from fishing. In the North East Atlantic, after having reached extreme levels, fishing pressure has decreased since the 1980's due to management measures such as total allowable catch (TAC) or area closure. An effect on the distribution of species as well as a potential recovery could be expected. However, temporal patterns of life history strategies are rarely linked to management measures. In addition, a larger emphasis is often put on exploited or emblematic sensitive species but rarely on assembly processes at the ecosystem scale (both commercial and non-commercial species). Based on a 17-year time series of 101 taxa (fishes, elasmobranchs, bivalves, cephalopods and crustaceans), we observed a negative relationship between the biomass of taxa sensitive to fishing and bottom trawling pressure, as well as an increase in their total biomass in the Celtic Sea. Over the whole area, stochasticity appeared as the dominant assembly process. Deterministic assembly processes were at play in the centre of the area where significant overdispersion (caused by the presence of both slow and fast taxa) were observed. The absence of sensitive taxa from the rest of the Celtic Sea appeared to be caused mainly by a historical effect of environmental filtering when fishing was high. At the local scale, we related the decrease in fishing pressure to the increase in biomass of five of the most sensitive taxa. This local decrease in fishing pressure, resulting from the implementation of an area closure, highlights the positive effect of such management measures in less than two decades.

Assembly rules of helminth parasite communities in grey mullets: combining components of diversity.

Organisms aggregate in ecological communities. It has been widely debated whether these associations are explained by deterministic or, in contrast, random processes. The answer may vary, depending on the level of an organisational scale (α, ß and γ) and the facet of diversity considered: taxonomic, functional and phylogenetic. Diversity at the level of a sampling unit (i.e. host individual) is the α diversity; ß diversity represents the extent of dissimilarity in diversity among sampling units (within a level of an organisational scale, ß1; between levels of an organisational scale, ß2); and the total diversity of a system is γ diversity. Thus, the combination of facets and levels of a scale may be useful to disentangle the mechanisms driving the composition of a parasite community. Using helminth parasite taxonomic, functional, and a proxy for phylogenetic diversity of three species of grey mullets (Teleostei: Mugilidae) from the Mediterranean Sea, we show that random and deterministic processes of different nature explain the assemblage of parasite communities. The parasite community at a host individual (α) was invariably a random subset of the total diversity in the community for the three facets of diversity. At the ß1 level, taxonomic diversity was lower than expected by chance, whereas functional diversity and the proxy for phylogenetic diversity were random. At the ß2 level, diversity patterns suggested environmental filtering of the parasite assemblage: species, trait, and phylogenetic compositions of parasite communities seemed to depend primarily on the species of host, but also on the locality and season. Our study shows that parasite communities are not totally understood if any of the components (i.e. facets and levels) of diversity is neglected.

The worldwide impact of urbanisation on avian functional diversity.

Urbanisation is driving rapid declines in species richness and abundance worldwide, but the general implications for ecosystem function and services remain poorly understood. Here, we integrate global data on bird communities with comprehensive information on traits associated with ecological processes to show that assemblages in highly urbanised environments have substantially different functional composition and 20% less functional diversity on average than surrounding natural habitats. These changes occur without significant decreases in functional dissimilarity between species; instead, they are caused by a decrease in species richness and abundance evenness, leading to declines in functional redundancy. The reconfiguration and decline of native functional diversity in cities are not compensated by the presence of exotic species but are less severe under moderate levels of urbanisation. Thus, urbanisation has substantial negative impacts on functional diversity, potentially resulting in impaired provision of ecosystem services, but these impacts can be reduced by less intensive urbanisation practices.

Environment outweighs the effects of fishing in regulating demersal community structure in an exploited marine ecosystem.

Global climate change has already caused bottom temperatures of coastal marine ecosystems to increase worldwide. These ecosystems face many pressures, of which fishing is one of the most important. While consequences of global warming on commercial species are studied extensively, the importance of the increase in bottom temperature and of variation in fishing effort is more rarely considered together in these exploited ecosystems. Using a 17 year time series from an international bottom trawl survey, we investigated covariations of an entire demersal ecosystem (101 taxa) with the environment in the Celtic Sea. Our results showed that over the past two decades, biotic communities in the Celtic Sea were likely controlled more by environmental variables than fisheries, probably due to its long history of exploitation. At the scale of the entire zone, relations between taxa and the environment remained stable over the years, but at a local scale, in the center of the Celtic Sea, dynamics were probably driven by interannual variation in temperature. Fishing was an important factor structuring species assemblages at the beginning of the time series (2000) but decreased in importance after 2009. This was most likely caused by a change in spatial distribution of fishing effort, following a change in targeted taxa from nephrops to deeper water anglerfish that did not covary with fishing effort. Increasing bottom temperatures could induce additional changes in the coming years, notably in the cold-water commercial species cod, hake, nephrops, and American plaice. We showed that analyzing covariation is an effective way to screen a large number of taxa and highlight those that may be most susceptible to future simultaneous increases in temperature and changes in exploitation pattern by fisheries. This information can be particularly relevant for ecosystem assessments.

Explosive breeding in tropical anurans: environmental triggers, community composition and acoustic structure.

BACKGROUND: Anurans largely rely on acoustic communication for sexual selection and reproduction. While multiple studies have focused on the calling activity patterns of prolonged breeding assemblages, species that concentrate their reproduction in short-time windows, explosive breeders, are still largely unknown, probably because of their ephemeral nature. In tropical regions, multiple species of explosive breeders may simultaneously aggregate leading to massive, mixed and dynamic choruses. To understand the environmental triggers, the phenology and composition of these choruses, we collected acoustic and environmental data at five ponds in French Guiana during a rainy season, assessing acoustic communities before and during explosive breeding events. RESULTS: We detected in each pond two explosive breeding events, lasting between 24 and 70 h. The rainfall during the previous 48 h was the most important factor predicting the emergence of these events. During explosive breeding events, we identified a temporal factor that clearly distinguished pre- and mid-explosive communities. A common pool of explosive breeders co-occurred in most of the events, namely Chiasmocleis shudikarensis, Trachycephalus coriaceus and Ceratophrys cornuta. Nevertheless, the species composition was remarkably variable between ponds and for each pond between the first and the second events. The acoustic structure of explosive breeding communities had outlying levels of amplitude and unexpected low acoustic diversity, significantly lower than the communities preceding explosive breeding events. CONCLUSIONS: Explosive breeding communities were tightly linked with specific rainfall patterns. With climate change increasing rainfall variability in tropical regions, such communities may experience significant shifts in their timing, distribution and composition. In structurally similar habitats, located in the same region without obvious barriers, our results highlight the variation in composition across explosive breeding events. The characteristic acoustic structure of explosive breeding events stands out from the circadian acoustic environment being easily detected at long distance, probably reflecting behavioural singularities and conveying heterospecific information announcing the availability of short-lived breeding sites. Our data provides a baseline against which future changes, possibly linked to climate change, can be measured, contributing to a better understanding on the causes, patterns and consequences of these unique assemblages.

Reconciling the concepts and measures of diversity, rarity and originality in ecology and evolution.

The concept of biological diversity, or biodiversity, is at the core of evolutionary and ecological studies. Many indices of biodiversity have been developed in the last four decades, with species being one of the central units of these indices. However, evolutionary and ecological studies need a precise description of species' characteristics to best quantify inter-species diversity, as species are not equivalent and exchangeable. One of the first concepts characterizing species in biodiversity studies was abundance-based rarity. Abundance-based rarity was then complemented by trait- and phylo-based rarity, called species' trait-based and phylogenetic originalities, respectively. Originality, which is a property of an individual species, represents a species' contribution to the overall diversity of a reference set of species. Originality can also be defined as the rarity of a species' characteristics such as the state of a functional trait, which is often assumed to be represented by the position of the species on a phylogenetic tree. We review and compare various approaches for measuring originality, rarity and diversity and demonstrate that (i) even if attempts to bridge these concepts do exist, only a few ecological and evolutionary studies have tried to combine them all in the past two decades; (ii) phylo- and trait-based diversity indices can be written as a function of species rarity and originality measures in several ways; and (iii) there is a need for the joint use of these three types of indices to understand community assembly processes and species' roles in ecosystem functioning in order to protect biodiversity efficiently.

Mammal extinctions and the increasing isolation of humans on the tree of life.

A sixth great mass extinction is ongoing due to the direct and indirect effects of human pressures. However, not all lineages are affected equally. From an anthropocentric perspective, it is often purported that humans hold a unique place on Earth. Here, we show that our current impacts on the natural world risk realizing that expectation. We simulated species loss on the mammalian phylogenetic tree, informed by species current extinction risks. We explored how Homo sapiens could become isolated in the tree if species currently threatened with extinction disappeared. We analyzed correlates of mammal extinctions risks that may drive this isolation pattern. We show that, within mammals, and more particularly within primates, extinction risks increase with the number of known threat types, and decrease with geographic range size. Extinctions increase with species body mass, trophic level, and the median longitudinal extent of each species range in mammals but not within primates. The risks of extinction are frequently high among H. sapiens close relatives. Pruning threatened primates, including apes (Hominidae, Hylobatidae), from the tree of life will lead to our species being among those with the fewest close relatives. If no action is taken, we will thus not only lose crucial biodiversity for the preservation of Earth ecosystems, but also a key living reference to what makes us human.

An ordination approach to explore similarities among communities.

Analysis of similarities among communities can help to decipher the biogeographical, evolutionary, and ecological factors that drive local diversity. Recent indices of similarity among communities incorporate not only information on species presence and abundance but also information on how similar species are in their traits and how closely related they are in terms of taxonomy or phylogeny. Towards this aim, trait-based, taxonomic or phylogenetic similarities among species have been defined and bounded between 0 (species are maximally distinct) and 1 (species are similar). A required property for an index of similarity between two communities is that it must provide minimum similarity (0) where communities have maximally distinct species, as well as maximum similarity (1) where communities are equivalent in their trait, taxonomic or phylogenetic compositions. Here, I developed a new ordination methodology that conforms to the requirement: double similarity principal component analysis (DSPCA). DSPCA summarizes multidimensional trait-based, taxonomic or phylogenetic similarities among communities into orthogonal axes. The species that drive each similarity pattern can be identified together with their traits or with their taxonomic or phylogenetic positions. I applied this methodology to theoretical examples and to empirical data sets on bird and bat communities to illustrate key properties of DSPCA. I compared the results obtained with DSPCA with those provided by related approaches. Theoretical and empirical case studies highlight the following additional properties of DSPCA: (i) axes are orthogonal and identify independent (dis)similarity patterns between communities; (ii) the more functionally, taxonomically or phylogenetically similar communities are, the closer they are on an axis; (iii) the coordinate of a species on an axis expresses how representative the species is of the pattern identified by the axis; and (iv) a species is representative of x communities if the functional, taxonomic or phylogenetic characteristics of this species are very common within each of these x communities. DSPCA is an efficient approach to visualize functional, taxonomic and phylogenetic similarities between communities. It is also a useful alternative to recent methods dedicated to phylogenetic diversity patterns. It will be an asset for all studies that aim to compare functional, taxonomic, genetic and phylogenetic diversity.

Trait and phylogenetic diversity provide insights into community assembly of reef-associated shrimps (Palaemonidae) at different spatial scales across the Chagos Archipelago.

Coral reefs are the most biodiverse marine ecosystem and one of the most threatened by global climate change impacts. The vast majority of diversity on reefs is comprised of small invertebrates that live within the reef structure, termed the cryptofauna. This component of biodiversity is hugely understudied, and many species remain undescribed. This study represents a rare analysis of assembly processes structuring a distinct group of cryptofauna, the Palaemonidae, in the Chagos Archipelago, a reef ecosystem under minimal direct human impacts in the central Indian Ocean. The Palaemonidae are a diverse group of Caridae (infraorder of shrimps) that inhabit many different niches on coral reefs and are of particular interest because of their varied habitat associations. Phylogenetic and trait diversity and phylogenetic signal were used to infer likely drivers of community structure. The mechanisms driving palaemonid community assembly and maintenance in the Chagos Archipelago showed distinct spatial patterns. At local scales, among coral colonies and among reefs fringing individual atolls, significant trait, and phylogenetic clustering patterns suggest environmental filtering may be a dominant ecological process driving Palaemonidae community structure, although local competition through equalizing mechanisms may also play a role in shaping the local community structure. Importantly, we also tested the robustness of phylogenetic diversity to changes in evolutionary information as multi-gene phylogenies are resource intensive and for large families, such as the Palaemonidae, are often incomplete. These tests demonstrated a very modest impact on phylogenetic community structure, with only one of the four genes (PEPCK gene) in the phylogeny affecting phylogenetic diversity patterns, which provides useful information for future studies on large families with incomplete phylogenies. These findings contribute to our limited knowledge of this component of biodiversity in a marine locality as close to undisturbed by humans as can be found. It also provides a rare evaluation of phylogenetic diversity methods.

Ecological versatility and the assembly of multiple competitors: cautionary notes for assembly inferences.

The role of niche differences and competition is invoked when one finds coexisting species to be more dissimilar in trait composition than expected at random in community assembly studies. This approach has been questioned as competition has been hypothesized to either lead to communities assembled by similar or dissimilar species, depending on whether species similarity reflects fitness or niche differences, respectively. A current problem is that the arguments used to draw relationships between competition and species similarity are based on pairwise theoretical examples, while in nature competition can occurs among a constellation of species with different levels of versatility in resources used. By versatility we mean the documented ability of some species to escape competition for commonly used resources by changing for marginal and unused resources. Thus, a versatile species will have the ability to decrease niche overlap with all other species when facing strong competitors. When these species are embedded in multiple interactions the role of pairwise niche and fitness differences could be reduced due to indirect effects and thus competition would not be detectable. Here we developed a coexistence theory where competition occurs simultaneously among multiple species with different levels of versatility and then used it in a simulation to unravel patterns of species similarity during community assembly. We found that simulated communities can be assembled by species with more, less or equal similarity compared to a null model when using a mean distance based metric (SES.MPD). However, contrasting these varied results, we consistently found species overdispersion using a nearest neighbor-based metric (SES.MNTD), even when species differences reflected more directly their competitive abilities than their niche differences. Strong tendency to overdispersion emerged when high ecological versatility promoted large niche differences and enabled coexistence. This is because versatility to use marginal resources compensates possible fitness differences among species. Our findings provide mounting evidence of the important role of minimum niche differences and versatility in resource consumption for species embedded in multiple direct and indirect interactions.

A new method for quantifying the phylogenetic redundancy of biological communities.

The increasing use of phylogenetic methods in community ecology recognizes that accumulated evolutionary differences among species mirror, to some extent, ecological processes. The scope of this work is thus to propose a new method for the measurement of community-level phylogenetic redundancy, which takes into account the branching pattern of the underlying phylogeny. Like for functional redundancy, a measure of phylogenetic redundancy can be described as a normalized measure in the range (0-1) that relates the observed community-level phylogenetic diversity to the value of a hypothetical assemblage with the same abundance distribution of the focal community in which all species had independent evolution. Therefore, phylogenetic redundancy can be interpreted as the diversity decrease that is obtained by taking into account the evolutionary relationships among species in the calculation of diversity. The behavior of the proposed method, for which we provide a simple R function called 'phyloredundancy', was evaluated with published data on Alpine plant communities along a primary succession on a glacier foreland in northern Italy. As shown by our results, the method accounts for the length of shared branches in the phylogeny, producing a coherent framework for describing the evolutionary relationships within a species assemblage. Being based on classical diversity measures, which have been used in ecology for decades, it also has a great potential for future research in phylogenetic community ecology.

Ancestrality and evolution of trait syndromes in finches (Fringillidae).

Species traits have been hypothesized by one of us (Ponge, 2013) to evolve in a correlated manner as species colonize stable, undisturbed habitats, shifting from "ancestral" to "derived" strategies. We predicted that generalism, r-selection, sexual monomorphism, and migration/gregariousness are the ancestral states (collectively called strategy A) and evolved correlatively toward specialism, K-selection, sexual dimorphism, and residence/territoriality as habitat stabilized (collectively called B strategy). We analyzed the correlated evolution of four syndromes, summarizing the covariation between 53 traits, respectively, involved in ecological specialization, r-K gradient, sexual selection, and dispersal/social behaviors in 81 species representative of Fringillidae, a bird family with available natural history information and that shows variability for all these traits. The ancestrality of strategy A was supported for three of the four syndromes, the ancestrality of generalism having a weaker support, except for the core group Carduelinae (69 species). It appeared that two different B-strategies evolved from the ancestral state A, both associated with highly predictable environments: one in poorly seasonal environments, called B1, with species living permanently in lowland tropics, with "slow pace of life" and weak sexual dimorphism, and one in highly seasonal environments, called B2, with species breeding out-of-the-tropics, migratory, with a "fast pace of life" and high sexual dimorphism.

Urbanisation and the loss of phylogenetic diversity in birds.

Despite the recognised conservation value of phylogenetic diversity, little is known about how it is affected by the urbanisation process. Combining a complete avian phylogeny with surveys along urbanisation gradients from five continents, we show that highly urbanised environments supported on average 450 million fewer years of evolutionary history than the surrounding natural environments. This loss was primarily caused by species loss and could have been higher had not been partially compensated by the addition of urban exploiters and some exotic species. Highly urbanised environments also supported fewer evolutionary distinctive species, implying a disproportionate loss of evolutionary history. Compared with highly urbanised environments, changes in phylogenetic richness and evolutionary distinctiveness were less substantial in moderately urbanised environments. Protecting pristine environments is therefore essential for maintaining phylogenetic diversity, but moderate levels of urbanisation still preserve much of the original diversity.

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.

Predicting loss of evolutionary history: Where are we?

The Earth's evolutionary history is threatened by species loss in the current sixth mass extinction event in Earth's history. Such extinction events not only eliminate species but also their unique evolutionary histories. Here we review the expected loss of Earth's evolutionary history quantified by phylogenetic diversity (PD) and evolutionary distinctiveness (ED) at risk. Due to the general paucity of data, global evolutionary history losses have been predicted for only a few groups, such as mammals, birds, amphibians, plants, corals and fishes. Among these groups, there is now empirical support that extinction threats are clustered on the phylogeny; however this is not always a sufficient condition to cause higher loss of phylogenetic diversity in comparison to a scenario of random extinctions. Extinctions of the most evolutionarily distinct species and the shape of phylogenetic trees are additional factors that can elevate losses of evolutionary history. Consequently, impacts of species extinctions differ among groups and regions, and even if global losses are low within large groups, losses can be high among subgroups or within some regions. Further, we show that PD and ED are poorly protected by current conservation practices. While evolutionary history can be indirectly protected by current conservation schemes, optimizing its preservation requires integrating phylogenetic indices with those that capture rarity and extinction risk. Measures based on PD and ED could bring solutions to conservation issues, however they are still rarely used in practice, probably because the reasons to protect evolutionary history are not clear for practitioners or due to a lack of data. However, important advances have been made in the availability of phylogenetic trees and methods for their construction, as well as assessments of extinction risk. Some challenges remain, and looking forward, research should prioritize the assessment of expected PD and ED loss for more taxonomic groups and test the assumption that preserving ED and PD also protects rare species and ecosystem services. Such research will be useful to inform and guide the conservation of Earth's biodiversity and the services it provides.

Integrating data-deficient species in analyses of evolutionary history loss.

There is an increasing interest in measuring loss of phylogenetic diversity and evolutionary distinctiveness which together depict the evolutionary history of conservation interest. Those losses are assessed through the evolutionary relationships between species and species threat status or extinction probabilities. Yet, available information is not always sufficient to quantify the threat status of species that are then classified as data deficient. Data-deficient species are a crucial issue as they cause incomplete assessments of the loss of phylogenetic diversity and evolutionary distinctiveness. We aimed to explore the potential bias caused by data-deficient species in estimating four widely used indices: HEDGE, EDGE, PDloss, and Expected PDloss. Second, we tested four different widely applicable and multitaxa imputation methods and their potential to minimize the bias for those four indices. Two methods are based on a best- vs. worst-case extinction scenarios, one is based on the frequency distribution of threat status within a taxonomic group and one is based on correlates of extinction risks. We showed that data-deficient species led to important bias in predictions of evolutionary history loss (especially high underestimation when they were removed). This issue was particularly important when data-deficient species tended to be clustered in the tree of life. The imputation method based on correlates of extinction risks, especially geographic range size, had the best performance and enabled us to improve risk assessments. Solving threat status of DD species can fundamentally change our understanding of loss of phylogenetic diversity. We found that this loss could be substantially higher than previously found in amphibians, squamate reptiles, and carnivores. We also identified species that are of high priority for the conservation of evolutionary distinctiveness.

The Evolutionary Legacy of Diversification Predicts Ecosystem Function.

Theory suggests that the structure of evolutionary history represented in a species community may affect its functioning, but phylogenetic diversity metrics do not allow for the identification of major differences in this structure. Here we propose a new metric, ELDERness (for Evolutionary Legacy of DivERsity) to estimate evolutionary branching patterns within communities by fitting a polynomial function to lineage-through-time (LTT) plots. We illustrate how real and simulated community branching patterns can be more correctly described by ELDERness and can successfully predict ecosystem functioning. In particular, the evolutionary history of branching patterns can be encapsulated by the parameters of third-order polynomial functions and further measured through only two parameters, the "ELDERness surfaces." These parameters captured variation in productivity of a grassland community better than existing phylogenetic diversity or diversification metrics and independent of species richness or presence of nitrogen fixers. Specifically, communities with small ELDERness surfaces (constant accumulation of lineages through time in LTT plots) were more productive, consistent with increased productivity resulting from complementary lineages combined with niche filling within lineages. Overall, while existing phylogenetic diversity metrics remain useful in many contexts, we suggest that our ELDERness approach better enables testing hypotheses that relate complex patterns of macroevolutionary history represented in local communities to ecosystem functioning.