An invasive pathogen drives directional niche contractions in amphibians.

Global change is causing an unprecedented restructuring of ecosystems, with the spread of invasive species being a key driver. While population declines of native species due to invasives are well documented, much less is known about whether new biotic interactions reshape niches of native species. Here we quantify geographic range and realized-niche contractions in Australian frog species following the introduction of amphibian chytrid fungus Batrachochytrium dendrobatidis, a pathogen responsible for catastrophic amphibian declines worldwide. We show that chytrid-impacted species experienced proportionately greater contractions in niche breadth than geographic distribution following chytrid emergence. Furthermore, niche contractions were directional, with contemporary distributions of chytrid-impacted species characterized by higher temperatures, lower diurnal temperature range, higher precipitation and lower elevations. Areas with these conditions may enable host persistence with chytrid through lower pathogenicity of the fungus and/or greater demographic resilience. Nevertheless, contraction to a narrower subset of environmental conditions could increase host vulnerability to other threatening processes and should be considered in assessments of extinction risk and during conservation planning. More broadly, our results emphasize that biotic interactions can strongly shape species realized niches and that large-scale niche contractions due to new species interactions-particularly emerging pathogens-could be widespread.

Priorities for conserving the world's terrestrial mammals based on over-the-horizon extinction risk.

Three major axes of global change put the world's mammal biodiversity at risk: climate change, human population growth, and land-use change.1,2,3,4,5,6,7,8,9,10,11,12 In some parts of the world the full effects of these threats on species will only be felt in decades to come, yet conservation emphasizes species currently threatened with extinction, by threats that have already occurred. There have been calls for conservation to become more proactive by anticipating and protecting species that may not yet be threatened, but have a high chance of becoming threatened in the future.3,6,8,10,12,13,14 We refer to this as "over-the-horizon" extinction risk, and we identify such species among the world's nonmarine mammals by considering not only the severity of increase in threats faced by each species, but also the way each species' biology confers sensitivity or robustness to threats. We define four future risk factors based on species' biology and projected exposure to severe change in climate, human population, and land use. We regard species with two or more of these risk factors as especially vulnerable to future extinction risk.10,15,16,17,18,19 Our models predict that by 2100 up to 1,057 (20%) of nonmarine mammal species will have combinations of two or more future risk factors. These species will be particularly concentrated in two future risk hotspots in sub-Saharan Africa and southern/eastern Australia. Proactively targeting species with over-the-horizon extinction risk could help to future-proof global conservation planning and prevent a new wave of mammal species from becoming threatened with extinction by the end of this century.

Global predictors of language endangerment and the future of linguistic diversity.

Language diversity is under threat. While each language is subject to specific social, demographic and political pressures, there may also be common threatening processes. We use an analysis of 6,511 spoken languages with 51 predictor variables spanning aspects of population, documentation, legal recognition, education policy, socioeconomic indicators and environmental features to show that, counter to common perception, contact with other languages per se is not a driver of language loss. However, greater road density, which may encourage population movement, is associated with increased endangerment. Higher average years of schooling is also associated with greater endangerment, evidence that formal education can contribute to loss of language diversity. Without intervention, language loss could triple within 40 years, with at least one language lost per month. To avoid the loss of over 1,500 languages by the end of the century, urgent investment is needed in language documentation, bilingual education programmes and other community-based programmes.

Ecological interactions shape the evolution of flower color in communities across a temperate biodiversity hotspot.

Processes driving the divergence of floral traits may be integral to the extraordinary richness of flowering plants and the assembly of diverse plant communities. Several models of pollinator-mediated floral evolution have been proposed; floral divergence may (i) be directly involved in driving speciation or may occur after speciation driven by (ii) drift or local adaptation in allopatry or (iii) negative interactions between species in sympatry. Here, we generate predictions for patterns of trait divergence and community assembly expected under these three models, and test these predictions in Hakea (Proteaceae), a diverse genus in the Southwest Australian biodiversity hotspot. We quantified functional richness for two key floral traits (pistil length and flower color), as well as phylogenetic distances between species, across ecological communities, and compared these to patterns generated from null models of community assembly. We also estimated the statistical relationship between rates of trait evolution and lineage diversification across the phylogeny. Patterns of community assembly suggest that flower color, but not floral phenology or morphology, or phylogenetic relatedness, is more divergent in communities than expected. Rates of lineage diversification and flower color evolution were negatively correlated across the phylogeny and rates of flower colour evolution were positively related to branching times. These results support a role for diversity-dependent species interactions driving floral divergence during the Hakea radiation, contributing to the development of the extraordinary species richness of southwest Australia.

Clarifying the relationship between body size and extinction risk in amphibians by complete mapping of model space.

In vertebrates, large body size is often a key diagnostic feature of species threatened with extinction. However, in amphibians the link between body size and extinction risk is highly uncertain, with previous studies suggesting positive, negative, u-shaped, or no relationship. Part of the reason for this uncertainty is 'researcher degrees of freedom': the subjectivity and selectivity in choices associated with specifying and fitting models. Here, I clarify the size-threat association in amphibians using Specification Curve Analysis, an analytical approach from the social sciences that attempts to minimize this problem by complete mapping of model space. I find strong support for prevailing negative associations between body size and threat status, the opposite of patterns typical in other vertebrates. This pattern is largely explained by smaller species having smaller geographic ranges, but smaller amphibian species also appear to lack some of the life-history advantages (e.g. higher reproductive output) that are often assumed to 'protect' small species in other taxa. These results highlight the need for a renewed conservation focus on the smallest species of the world's most threatened class of vertebrates, as aquatic habitats become increasingly degraded by human activity.

Macroevolutionary and macroecological approaches to understanding the evolution of stress tolerance in plants.

Environmental stress response in plants has been studied using a wide range of approaches, from lab-based investigation of biochemistry and genetics, to glasshouse studies of physiology and growth rates, to field-based trials and ecological surveys. It is also possible to investigate the evolution of environmental stress responses using macroevolutionary and macroecological analyses, analysing data from many different species, providing a new perspective on the way that environmental stress shapes the evolution and distribution of biodiversity. "Macroevoeco" approaches can produce intriguing results and new ways of looking at old problems. In this review, we focus on studies using phylogenetic analysis to illuminate macroevolutionary patterns in the evolution of environmental stress tolerance in plants. We follow a particular thread from our own research-evolution of salt tolerance-as a case study that illustrates a macroevolutionary way of thinking that opens up a range of broader questions on the evolution of environmental stress tolerances. We consider some potential future applications of macroevolutionary and macroecological analyses to understanding how diverse groups of plants evolve in response to environmental stress, which may allow better prediction of current stress tolerance and a way of predicting the capacity of species to adapt to changing environmental stresses over time.

Spatiophylogenetic modelling of extinction risk reveals evolutionary distinctiveness and brief flowering period as threats in a hotspot plant genus.

Comparative models used to predict species threat status can help identify the diagnostic features of species at risk. Such models often combine variables measured at the species level with spatial variables, causing multiple statistical challenges, including phylogenetic and spatial non-independence. We present a novel Bayesian approach for modelling threat status that simultaneously deals with both forms of non-independence and estimates their relative contribution, and we apply the approach to modelling threat status in the Australian plant genus Hakea. We find that after phylogenetic and spatial effects are accounted for, species with greater evolutionary distinctiveness and a shorter annual flowering period are more likely to be threatened. The model allows us to combine information on evolutionary history, species biology and spatial data, calculate latent extinction risk (potential for non-threatened species to become threatened), estimate the most important drivers of risk for individual species and map spatial patterns in the effects of different predictors on extinction risk. This could be of value for proactive conservation decision-making based on the early identification of species and regions of potential conservation concern.

Equilibrium and non-equilibrium phases in the radiation of Hakea and the drivers of diversity in Mediterranean-type ecosystems.

Mediterranean-type ecosystems (MTEs) contain exceptional plant diversity. Explanations for this diversity are usually classed as either "equilibrium," with elevated MTE diversity resulting from greater ecological carrying capacities, or "non-equilibrium," with MTEs having a greater accumulation of diversity over time than other types of ecosystems. These models have typically been considered as mutually exclusive. Here, we present a trait-based explanatory framework that incorporates both equilibrium and non-equilibrium dynamics. Using a large continental Australian plant radiation (Hakea) as a case study, we identify traits associated with niche partitioning in coexisting species (α-traits) and with environmental filtering (ß-traits), and reconstruct the mode and relative timing of diversification of these traits. Our results point to a radiation with an early non-equilibrium phase marked by divergence of ß-traits as Hakea diversified exponentially and expanded from the southwest Australian MTE into biomes across the Australian continent. This was followed from seven million years ago by an equilibrium phase, marked by diversification of α-traits and a slowdown in lineage diversification as MTE-niches became saturated. These results suggest that processes consistent with both equilibrium and non-equilibrium models have been important during different stages of the radiation of Hakea, and together they provide a richer explanation of present-day diversity patterns.

The ecological drivers of variation in global language diversity.

Language diversity is distributed unevenly over the globe. Intriguingly, patterns of language diversity resemble biodiversity patterns, leading to suggestions that similar mechanisms may underlie both linguistic and biological diversification. Here we present the first global analysis of language diversity that compares the relative importance of two key ecological mechanisms - isolation and ecological risk - after correcting for spatial autocorrelation and phylogenetic non-independence. We find significant effects of climate on language diversity, consistent with the ecological risk hypothesis that areas of high year-round productivity lead to more languages by supporting human cultural groups with smaller distributions. Climate has a much stronger effect on language diversity than landscape features, such as altitudinal range and river density, which might contribute to isolation of cultural groups. The association between biodiversity and language diversity appears to be an incidental effect of their covariation with climate, rather than a causal link between the two.

Reconstructing the Geography of Speciation from Contemporary Biodiversity Data.

Inferring the geographic mode of speciation could help reveal the evolutionary and ecological mechanisms that underlie the generation of biodiversity. Comparative methods have sought to reconstruct the geographic speciation history of clades, using data on phylogeny and species geographic ranges. However, inference from comparative methods has been limited by uncertainty over whether contemporary biodiversity data retain the historic signal of speciation. We constructed a process-based simulation model to determine the influence of speciation mode and postspeciation range evolution on current biodiversity patterns. The simulations suggest that the signal of speciation history remains detectable in species distributions and phylogeny, even when species ranges have evolved substantially through time. We extracted this signal by using a combination of summary statistics that had good power to distinguish speciation modes and then used these statistics to infer the speciation history of 30 plant and animal clades. The results point to broad taxonomic patterns in the modes of speciation, with strongest support for founder speciation in mammals and birds and strongest support for sympatric speciation in plants. Our model and analyses show that broad-scale comparative methods can be a powerful complementary approach to more focused genomic analyses in the study of the patterns and mechanisms of speciation.

Parasites and politics: why cross-cultural studies must control for relatedness, proximity and covariation.

A growing number of studies seek to identify predictors of broad-scale patterns in human cultural diversity, but three sources of non-independence in human cultural variables can bias the results of cross-cultural studies. First, related cultures tend to have many traits in common, regardless of whether those traits are functionally linked. Second, societies in geographical proximity will share many aspects of culture, environment and demography. Third, many cultural traits covary, leading to spurious relationships between traits. Here, we demonstrate tractable methods for dealing with all three sources of bias. We use cross-cultural analyses of proposed associations between human cultural traits and parasite load to illustrate the potential problems of failing to correct for these three forms of statistical non-independence. Associations between parasite stress and sociosexuality, authoritarianism, democracy and language diversity are weak or absent once relatedness and proximity are taken into account, and parasite load has no more power to explain variation in traditionalism, religiosity and collectivism than other measures of biodiversity, climate or population size do. Without correction for statistical non-independence and covariation in cross-cultural analyses, we risk misinterpreting associations between culture and environment.

The phylogeny and biogeography of Hakea (Proteaceae) reveals the role of biome shifts in a continental plant radiation.

The frequency of evolutionary biome shifts during diversification has important implications for our ability to explain geographic patterns of plant diversity. Recent studies present several examples of biome shifts, but whether frequencies of biome shifts closely reflect geographic proximity or environmental similarity of biomes remains poorly known. We explore this question by using phylogenomic methods to estimate the phylogeny of Hakea, a diverse Australian genus occupying a wide range of biomes. Model-based estimation of ancestral regions indicates that Hakea began diversifying in the Mediterranean biome of southern Australia in the Middle Eocene-Early Oligocene, and dispersed repeatedly into other biomes across the continent. We infer around 47 shifts between biomes. Frequencies of shifts between pairs of biomes are usually similar to those expected from their geographic connectedness or climatic similarity, but in some cases are substantially higher or lower than expected, perhaps reflecting how readily key physiological traits can be modified to adapt lineages to new environments. The history of frequent biome-shifting is reflected in the structure of present-day assemblages, which tend to be more phylogenetically diverse than null-model expectations. The case of Hakea demonstrates that the radiation of large plant clades across wide geographic areas need not be constrained by dispersal limitation or conserved adaptations to particular environments.

Environmental niche conservatism explains the accumulation of species richness in Mediterranean-hotspot plant genera.

The causes of exceptionally high plant diversity in Mediterranean-climate biodiversity hotspots are not fully understood. We asked whether a mechanism similar to the tropical niche conservatism hypothesis could explain the diversity of four large genera (Protea, Moraea, Banksia, and Hakea) with distributions within and adjacent to the Greater Cape Floristic Region (South Africa) or the Southwest Floristic Region (Australia). Using phylogenetic and spatial data we estimated the environmental niche of each species, and reconstructed the mode and dynamics of niche evolution, and the geographic history, of each genus. For three genera, there were strong positive relationships between the diversity of clades within a region and their inferred length of occupation of that region. Within genera, there was evidence for strong evolutionary constraint on niche axes associated with climatic seasonality and aridity, with different niche optima for hotspot and nonhotspot clades. Evolutionary transitions away from hotspots were associated with increases in niche breadth and elevated rates of niche evolution. Our results point to a process of "hotspot niche conservatism" whereby the accumulation of plant diversity in Mediterranean-type ecosystems results from longer time for speciation, with dispersal away from hotspots limited by narrow and phylogenetically conserved environmental niches.

Spatial, Phylogenetic, Environmental and Biological Components of Variation in Extinction Risk: A Case Study Using Banksia.

Comparative analyses of extinction risk routinely apply methods that account for phylogenetic non-independence, but few analyses of extinction risk have addressed the possibility of spatial non-independence. We explored patterns of extinction risk in Banksia, a plant genus largely endemic to Australia's southwest biodiversity hotspot, using methods to partition the variance in two response variables (threat status and range size) into phylogenetic, spatial, and independent components. We then estimated the effects of a number of biological and external predictors on extinction risk independently of phylogeny and space. The models explained up to 34.2% of the variation in range size and up to 9.7% of the variation in threat status, nearly all of which was accounted for by the predictors, not by phylogeny or space. In the case of Banksia, therefore, high extinction risk can be clearly linked with biological syndromes (such as a brief flowering period) or geographic indicators of human impact (such as extensive habitat loss), but cannot be predicted from phylogenetic relatedness or geographic proximity.

Detecting Macroevolutionary Self-Destruction from Phylogenies.

Phylogenetic analyses have lent support to the concept of lineage selection: that biological lineages can have heritable traits that influence their capacity to persist and diversify, and thereby affect their representation in biodiversity. While many discussions have focused on "positive" lineage selection, where stably heritable properties of lineages enhance their diversification rate, there are also intriguing examples that seem to represent "negative" lineage selection, where traits reduce the likelihood that a lineage will persist or speciate. In this article, we test whether a particular pattern of negative lineage selection is detectable from the distributions of the trait on a phylogeny. "Self-destructive" traits are those that arise often but then disappear again because they confer either a raised extinction rate or they are prone to a high rate of trait loss. For such a trait, the reconstructed origins will tend to be dispersed across the tips of the phylogeny, rather than defining large clades of related lineages that all share the trait. We examine the utility of four possible measures of "tippiness" as potential indicators of macroevolutionary self-destruction, applying them to phylogenies on which trait evolution has been simulated under different combinations of parameters for speciation, extinction, trait gain, and trait loss. We use an efficient simulation approach that starts with the required number of tips with and without the trait and uses a model to work "backwards" to construct different possible trees that result in that set of tips. We then apply these methods to a number of case studies: salt tolerance in grasses, color polymorphism in birds of prey, and selfing in nightshades. We find that the relative age of species, measured from tip length, can indicate a reduced speciation rate but does not identify traits that increase the extinction rate or the trait loss rate. We show that it is possible to detect cases of macroevolutionary self-destruction by considering the number of tips with the trait that arise from each inferred origin, and the degree to which the trait is scattered across the phylogeny. These metrics, and the methods we present, may be useful for testing macroevolutionary hypotheses from phylogenetic patterns.

Geographic bias in citation rates of conservation research.

We investigated whether the impact of conservation science is greater for research conducted in countries with more pressing conservation problems. We quantified research impact for 231 countries based on 2 citation metrics (mean cites per paper and h index) and fitted models predicting research impact based on number of threatened bird and mammal species (as a measure of conservation importance of a country) and a range of demographic variables. Citation rates of conservation research increased as a country's conservation need increased and as human population, quality of governance, and wealth increased. Even after accounting for these factors, citation rates among regions and countries within regions varied significantly. The conservation research community needs to consider ways to begin addressing the entrenched disadvantages some countries have when it comes to initiating projects and producing high-quality research.

Geographic range shifts do not erase the historic signal of speciation in mammals.

Many evolutionary analyses assume that the positions of species geographic ranges are sufficiently phylogenetically conserved that current ranges reflect ancestral ranges and retain the historic signal of speciation. The validity of this assumption has been challenged, because there is evidence that ranges can shift rapidly and extensively. Here I test the assumption of range conservatism using simulations and empirical tests of phylogenetic signal in geographic positions of ranges within mammal orders, families, and genera. In most taxa, range positions show strong phylogenetic signal, quantified using Pagel's λ, Mantel tests, and a novel method to measure phylogenetic signal near the tips of a phylogeny. Taxa with highly labile range positions are exceptions to the general pattern and include very young groups such as Sciurus that may still be in the early, rapid-expansion phase of adaptive radiation. In two orders containing many species with large distributions (Artiodactyla and Carnivora), temporal patterns of range evolution are consistent with large instantaneous shifts in range position associated with allopatric speciation. In most other taxa, range evolution is better described by models that allow ranges to evolve along branches of the phylogeny. The results point to a common pattern of phylogenetically conserved ranges where the current position of species ranges reflects their position at the time of speciation, modified by gradual drift of range boundaries through time.