Evolutionary age and risk of extinction in the global avifauna

Species at high risk of extinction are not distributed at random among higher taxa. Here we demonstrate that there is a positive relationship between the proportion of species in a taxon which are considered to be threatened and the evolutionary age of that taxon, both for the global avifauna and the avifauna of the New World. The potential mechanisms and consequences of the relationship are examined.     

Global distribution and drivers of language extinction risk

Many of the world''s languages face serious risk of extinction. Efforts to prevent this cultural loss are severely constrained by a poor understanding of the geographical patterns and drivers of extinction risk. We quantify the global distribution of language extinction risk—represented by small range and speaker population sizes and rapid declines in the number of speakers—and identify the underlying environmental and socioeconomic drivers. We show that both small range and speaker population sizes are associated with rapid declines in speaker numbers, causing 25% of existing languages to be threatened based on criteria used for species. Language range and population sizes are small in tropical and arctic regions, particularly in areas with high rainfall, high topographic heterogeneity and/or rapidly growing human populations. By contrast, recent speaker declines have mainly occurred at high latitudes and are strongly linked to high economic growth. Threatened languages are numerous in the tropics, the Himalayas and northwestern North America. These results indicate that small-population languages remaining in economically developed regions are seriously threatened by continued speaker declines. However, risks of future language losses are especially high in the tropics and in the Himalayas, as these regions harbour many small-population languages and are undergoing rapid economic growth.     

The evolution of extinction risk: past and present anthropogenic impacts on the primate communities of Madagascar

There are two possible approaches to understanding natural and human-induced changes in the primate communities of Madagascar. One is to begin with present-day and recent historic interactions and work backwards. A second is to begin with paleoecological records of Malagasy primate communities before and immediately following human arrival, and the associated evidence of human and nonhuman primate interactions, and work forwards. On the basis of biological and climatic studies, as well as historic and ethnohistoric records, we are beginning to understand the abiotic and biotic characteristics of Madagascar's habitats, the lemurs' ecological adaptations to these unique habitats, the extent of forest loss, fragmentation and hunting, and the differential vulnerability of extant lemur species to these pressures. On the basis of integrated paleoecological, archaeological and paleontological research, we have begun to construct a detailed chronology for late prehistoric Madagascar. We are beginning to understand the complex sequence of events that led to one of the most dramatic recent megafaunal extinction/extirpation events. Combining the perspectives of the past and the present, we see a complex set of interactions affecting an initially rich but vulnerable fauna. The total evidence refutes any simple, unicausal (e.g. hunting/habitat destruction/climate change) explanation of megafaunal extinctions, yet unequivocally supports a major role--both direct and indirect--for humans as the trigger of the extinction process. It also supports a change over time in the relative importance of hunting versus habitat loss, and in the trophic characteristics of the primate communities in Madagascar.     

Revisiting the impacts of non-random extinction on the tree-of-life

The tree-of-life represents the diversity of living organisms. Species extinction and the concomitant loss of branches from the tree-of-life is therefore a major conservation concern. There is increasing evidence indicating that extinction is phylogenetically non-random, such that if one species is vulnerable to extinction so too are its close relatives. However, the impact of non-random extinctions on the tree-of-life has been a matter of recent debate. Here, we combine simulations with empirical data on extinction risk in mammals. We demonstrate that phylogenetically clustered extinction leads to a disproportionate loss of branches from the tree-of-life, but that the loss of their summed lengths is indistinguishable from random extinction. We argue that under a speciational model of evolution, the number of branches lost might be of equal or greater consequences than the loss of summed branch lengths. We therefore suggest that the impact of non-random extinction on the tree-of-life may have been underestimated.     

Estimating human effects on global extinction

A quantitative technique for estimating extinctions due to clearing of natural ecosystems is described. Applied on a global scale, the method yields preliminary figures on extinctions of flowering plants, butterflies, land birds and land mammals ranging from 5.4 to 15.3% for the period from the beginning of agriculture until the year 1980. Actual numbers of extinctions of mammals and birds to date are much lower, possibly in part due to a tendency for the technique to overestimate species loss at the global scale. However, delayed extinctions of species whose populations have been reduced but not exterminated by habitat destruction are likely, suggesting that human impacts may be more serious than they currently appear.     

Measuring the meltdown: drivers of global amphibian extinction and decline

Habitat loss, climate change, over-exploitation, disease and other factors have been hypothesised in the global decline of amphibian biodiversity. However, the relative importance of and synergies among different drivers are still poorly understood. We present the largest global analysis of roughly 45% of known amphibians (2,583 species) to quantify the influences of life history, climate, human density and habitat loss on declines and extinction risk. Multi-model Bayesian inference reveals that large amphibian species with small geographic range and pronounced seasonality in temperature and precipitation are most likely to be Red-Listed by IUCN. Elevated habitat loss and human densities are also correlated with high threat risk. Range size, habitat loss and more extreme seasonality in precipitation contributed to decline risk in the 2,454 species that declined between 1980 and 2004, compared to species that were stable (n = 1,545) or had increased (n = 28). These empirical results show that amphibian species with restricted ranges should be urgently targeted for conservation.     

Predicting extinction risk of butterflies and moths (Macrolepidoptera) from distribution patterns and species characteristics

An extensive follow-up study of day- and night-active Macrolepidoptera was performed during 2004 at the Kullaberg Nature Reserve located in the south-western part of Sweden. Butterflies were surveyed in an area of 100 km² and night-active moths were trapped in the core area of the reserve. Macrolepidopteran species resident in the area in the 1950s were compared with species resident in the area in 2004. As much as 159 of 597 species (27%) resident in the area in 1950s were not found at all in 2004 and 22 species (4%) had colonised the area. Butterflies exhibited a disastrous decline with a loss of 45% of the fauna, and day-active species had declined more than night-active species had. Species distribution pattern and species characteristics were used to predict the probability that a species had become extinct or colonised the area. Species limited to one or a few food plants, with a short flight-period or restricted to non-forest habitats were all associated with a high extinction risk. Species occurring in fewer European countries and recorded from few provinces in Sweden were all associated with a higher extinction risk compared with ubiquitous species. For expanding species (colonisers), the best predictor was their distribution area in Sweden and colonising species were more likely to be limited to a few provinces in the southernmost part of Sweden. Species extinct from Kullaberg also have decreased in Finland and species that had colonised Kullaberg also have increased in Finland, while the species with no change in Kullaberg are also relatively constant in Finland. Indeed, the macrolepidopteran fauna was severely reduced in the last 50 years and almost 70% of the habitat specialists were lost. For red-listed species the situation was found to be alarming. Almost 70% of red-listed species, resident in the area in the 1950s, was not found in 2004. This study highlights that species composition changes rapidly even in protected nature reserves and that similar changes in the macrolepidopteran fauna seem to occur over large areas.     

Synergies among extinction drivers under global change

If habitat destruction or overexploitation of populations is severe, species loss can occur directly and abruptly. Yet the final descent to extinction is often driven by synergistic processes (amplifying feedbacks) that can be disconnected from the original cause of decline. We review recent observational, experimental and meta-analytic work which together show that owing to interacting and self-reinforcing processes, estimates of extinction risk for most species are more severe than previously recognised. As such, conservation actions which only target single-threat drivers risk being inadequate because of the cascading effects caused by unmanaged synergies. Future work should focus on how climate change will interact with and accelerate ongoing threats to biodiversity, such as habitat degradation, overexploitation and invasive species.     

Sexual selection and the risk of extinction in birds

The relationship between sexual selection and extinction risk has rarely been investigated. This is unfortunate because extinction plays a key role in determining the patterns of species richness seen in extant clades, which form the basis of comparative studies into the role that sexual selection may play in promoting speciation. We investigate the extent to which the perceived risk of extinction relates to four different estimates of sexual selection in 1030 species of birds. We find no evidence that the number of threatened species is distributed unevenly according to a social mating system, and neither of our two measures of pre-mating sexual selection (sexual dimorphism and dichromatism) was related to extinction risk, after controlling for phylogenetic inertia. However, threatened species apparently experience more intense post-mating sexual selection, measured as testis size, than non-threatened species. These results persisted after including body size as a covariate in the analysis, and became even stronger after controlling for clutch size (two known correlates of extinction risk). Sexual selection may therefore be a double-edged process-promoting speciation on one hand but promoting extinction on the other. Furthermore, we suggest that it is post-mating sexual selection, in particular, that is responsible for the negative effect of sexual selection on clade size. Why this might be is unclear, but the mean population fitness of species with high intensities of post-mating sexual selection may be especially low if costs associated with multiple mating are high or if the selection load imposed by post-mating selection is higher relative to that of pre-mating sexual selection.     

Sexual selection and the risk of extinction in mammals

Sexual selection is commonly envisaged as a force working in opposition to natural selection, because extravagant or exaggerated traits could apparently have never evolved via natural selection alone. There is good evidence that a selection load imposed by sexual selection may be eased experimentally by restricting the opportunity for it to operate. Sexual selection could therefore potentially play an important role in influencing the risk of extinction that a population faces, thereby contributing to the apparent selectivity of extinctions. Conversely, recent theory predicts that the likelihood of extinction may decrease when sexual selection is operating because it could accelerate the rate of adaptation in concert with natural selection. So far, comparative evidence (coming mostly from birds) has generally indicated support for the former scenario, but the question remains open. The aim of this study was therefore to examine whether the level of sexual selection (measured as residual testes mass and sexual size dimorphism) was related to the risk of extinction that mammals are currently experiencing. We found no evidence for a relationship between these factors, although our analyses may have been confounded by the possible dominating effect of contemporary anthropogenic factors.     

Fingerprinting the impacts of global change on tropical forests

Recent observations of widespread changes in mature tropical forests such as increasing tree growth, recruitment and mortality rates and increasing above-ground biomass suggest that ''global change'' agents may be causing predictable changes in tropical forests. However, consensus over both the robustness of these changes and the environmental drivers that may be causing them is yet to emerge. This paper focuses on the second part of this debate. We review (i) the evidence that the physical, chemical and biological environment that tropical trees grow in has been altered over recent decades across large areas of the tropics, and (ii) the theoretical, experimental and observational evidence regarding the most likely effects of each of these changes on tropical forests. Ten potential widespread drivers of environmental change were identified: temperature, precipitation, solar radiation, climatic extremes (including El Niño-Southern Oscillation events), atmospheric CO2 concentrations, nutrient deposition, O3/acid depositions, hunting, land-use change and increasing liana numbers. We note that each of these environmental changes is expected to leave a unique ''fingerprint'' in tropical forests, as drivers directly force different processes, have different distributions in space and time and may affect some forests more than others (e.g. depending on soil fertility). Thus, in the third part of the paper we present testable a priori predictions of forest responses to assist ecologists in attributing particular changes in forests to particular causes across multiple datasets. Finally, we discuss how these drivers may change in the future and the possible consequences for tropical forests.     

Genome size and extinction risk in vertebrates

The hypothesis of 'selfish DNA' is tested for the case of animals using the relation between genome size and conservation status of a given species. In contrast to plants, where the larger genome was previously shown to increase the likelihood of extinction, the picture is more complicated in animals. At the within-families and within-orders levels, the larger genome increases the risk of extinction only in reptiles and birds (which have the smallest genomes among tetrapods). In fishes and amphibians, the effect is caused by the higher taxonomic levels (above order). In several phylogenetic lineages of anamniotes, there is a correlation between a higher fraction of threatened species and a lower number of extant species in a lineage with the larger genome. In mammals, no effect was observed at any taxonomic level. The obtained data support the concept of hierarchical selection. It is also shown that, in plants and reptiles, the probability of being threatened increases from less than 10% to more than 80% with the increase in genome size, which can help in establishing conservation priorities.     

Estimation of the extinction risk for high-montane species as a consequence of global warming and assessment of their suitability as cross-taxon indicators

The evidence for climate change is increasing, and global warming could lead to the extinction of some species. Here we estimated the extinction risk of six high-montane species of different taxonomic groups (fern, vascular plant, wood-inhabiting fungus, mollusk, saproxylic beetle, and bird) by modeling their occurrence under two global warming scenarios. We also assessed the cross-taxon indicator suitability of the selected species for monitoring climate change in low-mountain-range forests in southeastern Germany (Bavarian Forest National Park). We tested the influence of temperature and other habitat variables by applying semi-parametric spatial generalized linear models with binomial error. The probability of occurrence for each species under the present conditions and under two conditions of global warming was calculated. To assess the cross-taxon suitability, we tested the predictability of the final generalized linear models for each species using the measured occurrence of the other selected species and a discrimination technique. We identified temperature as the main driver for all selected high-montane species. Our statistical models predict a considerable risk of extinction of these species within the Bavarian Forest National Park as a result of global warming. Our discrimination model indicates that these species have essentially similar relationships with the environment and that five of the six species are suitable as indicators of early signs of global warming. The choice of which indicators to use should involve a consideration of the type of monitoring systems already in place.     

Primate extinction risk and historical patterns of speciation and extinction in relation to body mass

Body mass is thought to influence diversification rates, but previous studies have produced ambiguous results. We investigated patterns of diversification across 100 trees obtained from a new Bayesian inference of primate phylogeny that sampled trees in proportion to their posterior probabilities. First, we used simulations to assess the validity of previous studies that used linear models to investigate the links between IUCN Red List status and body mass. These analyses support the use of linear models for ordinal ranked data on threat status, and phylogenetic generalized linear models revealed a significant positive correlation between current extinction risk and body mass across our tree block. We then investigated historical patterns of speciation and extinction rates using a recently developed maximum-likelihood method. Specifically, we predicted that body mass correlates positively with extinction rate because larger bodied organisms reproduce more slowly, and body mass correlates negatively with speciation rate because smaller bodied organisms are better able to partition niche space. We failed to find evidence that extinction rates covary with body mass across primate phylogeny. Similarly, the speciation rate was generally unrelated to body mass, except in some tests that indicated an increase in the speciation rate with increasing body mass. Importantly, we discovered that our data violated a key assumption of sample randomness with respect to body mass. After correcting for this bias, we found no association between diversification rates and mass.     

Predicting extinction risk in declining species

What biological attributes predispose species to the risk of extinction? There are many hypotheses but so far there has been no systematic analysis for discriminating between them. Using complete phylogenies of contemporary carnivores and primates, we present, to our knowledge, the first comparative test showing that high trophic level, low population density slow life history and, in particular, small geographical range size are all significantly and independently associated with a high extinction risk in declining species. These traits together explain nearly 50% of the total between-species variation in extinction risk. Much of the remaining variation can be accounted for by external anthropogenic factors that affect species irrespective of their biology.     

Comparing risk factors for population extinction

Extinction risk of natural populations of animals and plants is enhanced by many different processes, including habitat size reduction and toxic chemical exposure. We develop a method to evaluate different risk factors in terms of the decrease in the mean extinction time. We choose a population model with logistic growth, environmental and demographic stochasticities with three parameters (intrinsic growth rate r, carrying capacity K, and environmental noise sigma(2)(e)). The reduction in the habitat size decreases carrying capacity K only, whilst toxic chemical exposure decreases survivorship (or fertility) and in effect reduces both r and K. We derived a formula for the reduction in habitat size that decrease the mean extinction time by the same magnitude as a given level of toxic chemical exposure. In a large population (large K) or in a slowly growing population (small r), a small decrease in survivorship can cause the extinction risk increase corresponding to a significant reduction in the habitat size. This conclusion depends also on the nonlinearity of dose-effect relationship. To illustrate the method, we analyse a freshwater fish, Japanese crucian carp (Carassius auratus subsp.) in Lake Biwa.     

Hunting to extinction: biology and regional economy influence extinction risk and the impact of hunting in artiodactyls

Half of all artiodactyls (even-toed hoofed mammals) are threatened with extinction, around double the mammalian average. Here, using a complete species-level phylogeny, we construct a multivariate model to assess for the first time which intrinsic (biological) and extrinsic (anthropogenic and environmental) factors influence variation in extinction risk in artiodactyls. Globally artiodactyls at greatest risk live in economically less developed areas, have older weaning ages and smaller geographical ranges. Our findings suggest that identifying predictors of threat is complicated by interactions between both biological and anthropogenic factors, resulting in differential responses to threatening processes. Artiodactyl species that experience unregulated hunting live in significantly less economically developed areas than those that are not hunted; however, hunted species are more susceptible to extinction if they have slower reproductive rates (older weaning ages). In contrast, risk in non-hunted artiodactyls is unrelated to reproductive rate and more closely associated with the economic development of the region in which they live.     

Interaction strength and extinction risk in a metacommunity

Species interactions and connectivity are both central to explaining the stability of ecological communities and the problem of species extinction. Yet, the role of species interactions for the stability of spatially subdivided communities still eludes ecologists. Ecological models currently address the problem of stability by exploring the role of interaction strength in well mixed habitats, or of connectivity in subdivided communities. Here I propose a unification of interaction strength and connectivity as mechanisms explaining regional community stability. I introduce a metacommunity model based on succession dynamics in coastal ecosystems, incorporating limited dispersal and facilitative interactions. I report a sharp transition in regional stability and extinction probability at intermediate interaction strength, shown to correspond to a phase transition that generates scale-invariant distribution and high regional stability. In contrast with previous studies, stability results from intermediate interaction strength only in subdivided communities, and is associated with large-scale (scale-invariant) synchrony. These results can be generalized to other systems exhibiting phase transitions to show how local interaction strength can be used to resolve the link between regional community stability and pattern formation.     

Climate warming, dispersal inhibition and extinction risk

Global warming impels species to track their shifting habitats or adapt to new conditions. Both processes are critically influenced by individual dispersal. In many animals, dispersal behaviour is plastic, but how organisms with plastic dispersal respond to climate change is basically unknown. Here, we report the analysis of interannual dispersal change from 16 years of monitoring a wild population of the common lizard, and a 12-year manipulation of lizards' diet intended to disentangle the direct effect of temperature rise on dispersal from its effects on resource availability. We show that juvenile dispersal has declined dramatically over the last 16 years, paralleling the rise of spring temperatures during embryogenesis. A mesoscale model of metapopulation dynamics predicts that in general dispersal inhibition will elevate the extinction risk of metapopulations exposed to contrasting effects of climate warming.