Space-for-time substitutions in climate change ecology and evolution.

In an epoch of rapid environmental change, understanding and predicting how biodiversity will respond to a changing climate is an urgent challenge. Since we seldom have sufficient long-term biological data to use the past to anticipate the future, spatial climate-biotic relationships are often used as a proxy for predicting biotic responses to climate change over time. These 'space-for-time substitutions' (SFTS) have become near ubiquitous in global change biology, but with different subfields largely developing methods in isolation. We review how climate-focussed SFTS are used in four subfields of ecology and evolution, each focussed on a different type of biotic variable - population phenotypes, population genotypes, species' distributions, and ecological communities. We then examine the similarities and differences between subfields in terms of methods, limitations and opportunities. While SFTS are used for a wide range of applications, two main approaches are applied across the four subfields: spatial in situ gradient methods and transplant experiments. We find that SFTS methods share common limitations relating to (i) the causality of identified spatial climate-biotic relationships and (ii) the transferability of these relationships, i.e. whether climate-biotic relationships observed over space are equivalent to those occurring over time. Moreover, despite widespread application of SFTS in climate change research, key assumptions remain largely untested. We highlight opportunities to enhance the robustness of SFTS by addressing key assumptions and limitations, with a particular emphasis on where approaches could be shared between the four subfields.

Abrupt expansion of climate change risks for species globally.

Climate change is already exposing species to dangerous temperatures driving widespread population and geographical contractions. However, little is known about how these risks of thermal exposure will expand across species' existing geographical ranges over time as climate change continues. Here, using geographical data for approximately 36,000 marine and terrestrial species and climate projections to 2100, we show that the area of each species' geographical range at risk of thermal exposure will expand abruptly. On average, more than 50% of the increase in exposure projected for a species will occur in a single decade. This abruptness is partly due to the rapid pace of future projected warming but also because the greater area available at the warm end of thermal gradients constrains species to disproportionately occupy sites close to their upper thermal limit. These geographical constraints on the structure of species ranges operate both on land and in the ocean and mean that, even in the absence of amplifying ecological feedbacks, thermally sensitive species may be inherently vulnerable to sudden warming-driven collapse. With higher levels of warming, the number of species passing these thermal thresholds, and at risk of abrupt and widespread thermal exposure, increases, doubling from less than 15% to more than 30% between 1.5 °C and 2.5 °C of global warming. These results indicate that climate threats to thousands of species are expected to expand abruptly in the coming decades, thereby highlighting the urgency of mitigation and adaptation actions.

Biodiversity: What makes a city bird?

The impacts of urbanisation on biodiversity varies greatly across species. A new study shows how the intrinsic species properties underlying urban tolerance vary globally according to environmental context. This has important implications for conserving biodiversity in a rapidly urbanising world.

Risks to biodiversity from temperature overshoot pathways.

Temperature overshoot pathways entail exceeding a specified global warming level (e.g. 1.5°C or 2°C) followed by a decline in warming, achieved through anthropogenically enhanced CO2 removal from the atmosphere. However, risks to biodiversity from temperature overshoot pathways are poorly described. Here, we explore biodiversity risks from overshoot by synthesizing existing knowledge and quantifying the dynamics of exposure and de-exposure to potentially dangerous temperatures for more than 30 000 species for a 2°C overshoot scenario. Our results suggest that climate risk to biodiversity from temperature overshoot pathways will arrive suddenly, but decrease only gradually. Peak exposure for biodiversity occurs around the same time as peak global warming, but the rate of de-exposure lags behind the temperature decline. While the global overshoot period lasts around 60 years, the duration of elevated exposure of marine and terrestrial biodiversity is substantially longer (around 100 and 130 years, respectively), with some ecological communities never returning to pre-overshoot exposure levels. Key biodiversity impacts may be irreversible and reliance on widespread CO2 removal to reduce warming poses additional risks to biodiversity through altered land use. Avoiding any temperature overshoot must be a priority for reducing biodiversity risks from climate change, followed by limiting the magnitude and duration of any overshoot. More integrated models that include direct and indirect impacts from overshoot are needed to inform policy. This article is part of the theme issue 'Ecological complexity and the biosphere: the next 30 years'.

The macroecology of landscape ecology.

One of landscape ecology's main goals is to unveil how biodiversity is impacted by habitat transformation. However, the discipline suffers from significant context dependency in observed spatial and temporal trends, hindering progress towards understanding the mechanisms driving species declines and preventing the development of accurate estimates of future biodiversity change. Here, we discuss recent evidence that populations' and species' responses to habitat change at the landscape scale are modulated by factors and processes occurring at macroecological scales, such as historical disturbance rates, distance to geographic range edges, and climatic suitability. We suggest that placing landscape ecology studies in a macroecological lens will help to explain seemingly inconsistent results and will ultimately create better predictive models to help mitigate the biodiversity crisis.

A test of Darwin's naturalization conundrum in birds reveals enhanced invasion success in the presence of close relatives.

Biological invasions pose one of the most severe environmental challenges of the twenty-first century. A longstanding idea is that invasion risk is predictable based on the phylogenetic distance - and hence ecological resemblance - between non-native and native species. However, current evidence is contradictory. To explain these mixed results, it has been proposed that the effect is scale-dependent, with invasion inhibited by phylogenetic similarity at small spatial scales but enhanced at larger scales. Analyzing invasion outcomes in a global sample of bird communities, we find no evidence to support this hypothesis. Instead, our results suggest that invaders are locally more successful in the presence of closely related and ecologically similar species, at least in human-altered environments where the majority of invasions have occurred. Functional trait analyses further confirm that the ecological niches of invaders are phylogenetically conserved, supporting the notion that successful invasion in the presence of close relatives is driven by shared adaptations to the types of niches available in novel environments.

AVONET: morphological, ecological and geographical data for all birds.

Functional traits offer a rich quantitative framework for developing and testing theories in evolutionary biology, ecology and ecosystem science. However, the potential of functional traits to drive theoretical advances and refine models of global change can only be fully realised when species-level information is complete. Here we present the AVONET dataset containing comprehensive functional trait data for all birds, including six ecological variables, 11 continuous morphological traits, and information on range size and location. Raw morphological measurements are presented from 90,020 individuals of 11,009 extant bird species sampled from 181 countries. These data are also summarised as species averages in three taxonomic formats, allowing integration with a global phylogeny, geographical range maps, IUCN Red List data and the eBird citizen science database. The AVONET dataset provides the most detailed picture of continuous trait variation for any major radiation of organisms, offering a global template for testing hypotheses and exploring the evolutionary origins, structure and functioning of biodiversity.

Intensive human land uses negatively affect vertebrate functional diversity.

Land-use change is the leading driver of global biodiversity loss thus characterising its impacts on the functional structure of ecological communities is an urgent challenge. Using a database describing vertebrate assemblages in different land uses, we assess how the type and intensity of land use affect the functional diversity of vertebrates globally. We find that human land uses alter local functional structure by driving declines in functional diversity, with the strongest effects in the most disturbed land uses (intensely used urban sites, cropland and pastures), and among amphibians and birds. Both tropical and temperate areas experience important functional losses, which are only partially offset by functional gains. Tropical assemblages are more likely to show decreases in functional diversity that exceed those expected from species loss alone. Our results indicate that land-use change non-randomly reshapes the functional structure of vertebrate assemblages, raising concerns about the continuation of ecological processes sustained by vertebrates.

Loss of functional diversity through anthropogenic extinctions of island birds is not offset by biotic invasions.

Human impacts reshape ecological communities through the extinction and introduction of species. The combined impact of these factors depends on whether non-native species fill the functional roles of extinct species, thus buffering the loss of functional diversity. This question has been difficult to address, because comprehensive information about past extinctions and their traits is generally lacking. We combine detailed information about extinct, extant, and established alien birds to quantify historical changes in functional diversity across nine oceanic archipelagos. We found that alien species often equal or exceed the number of anthropogenic extinctions yet apparently perform a narrower set of functional roles as current island assemblages have undergone a substantial and ubiquitous net loss in functional diversity and increased functional similarity among assemblages. Our results reveal that the introduction of alien species has not prevented anthropogenic extinctions from reducing and homogenizing the functional diversity of native bird assemblages on oceanic archipelagos.

Evolutionary dynamics of the elevational diversity gradient in passerine birds.

Low-elevation regions harbour the majority of the world's species diversity compared to high-elevation areas. This global gradient suggests that lowland species have had more time to diversify, or that net diversification rates have been higher in the lowlands. However, highlands seem to be cradles of diversity as they contain many young endemics, suggesting that their rates of speciation are exceptionally fast. Here we use a phylogenetic diversification model that accounts for the dispersal of species between different elevations to examine the evolutionary dynamics of the elevational diversity gradient in passerine birds, a group that has radiated globally to occupy almost all elevations and latitudes. We find strong support for a model in which passerines diversify at the same rate in the highlands and the lowlands but in which the per-capita rate of dispersal from high to low elevations is more than twice as fast as that in the reverse direction. This suggests that while there is no consistent trend in diversification across elevations, part of the diversity generated by highland regions migrates into the lowlands, thus setting up the observed gradient in passerine diversity. We find that this process drives tropical regions but for temperate areas, the analysis could be hampered by their lower richness. Despite their lower diversity, highland regions are disproportionally important for maintaining diversity in the adjacent lowlands.

Priority effects and the macroevolutionary dynamics of biodiversity.

Priority effects can play a fundamental role in the assembly of ecological communities, but how they shape the dynamics of biodiversity over macroevolutionary timescales remains unclear. Here we develop and analyse a metacommunity model combining local priority effects with niche evolution, speciation and extinction. We show that by promoting the persistence of rare species, local priority effects cause the evolution of higher metacommunity diversity as well as major disparities in richness among evolutionary lineages. However, we also show how classic macroevolutionary patterns of niche incumbency-whereby rates of regional diversification and invasion slow down as ecological niches are filled-do not depend on local priority effects, arising even when invading species continuously displace residents. Together, these results clarify the connection between local priority effects and the filling of ecological niche space, and reveal how the impact of species arrival order on competition fundamentally shapes the generation and maintenance of biodiversity.

Environmental resistance predicts the spread of alien species.

The unabating rise in the number of species introduced outside of their native range makes predicting the spread of alien species an urgent challenge. Most predictions use models of the ecological niche of a species to identify suitable areas for invasion, but these predictions may have limited accuracy. Here, using the global alien avifauna, we demonstrate an alternative approach for predicting alien spread based on the environmental resistance of the landscape. This approach does not require any information on the ecological niche of the invading species and, instead, uses gradients of biotic similarity among native communities in the invaded region to predict the most likely routes of spread. We show that environmental resistance predicts patterns of spread better than a null model of random dispersal or a model based on climate matching to the native range of each species. Applying this approach to simulate future spread reveals major regional differences in projected invasion risk, shaped by proximity to existing invasion hotspots as well as gradients in environmental resistance. Our results show how environmental resistance may provide a general and complementary approach for predicting invasion risk that can be rapidly deployed even when information on the niche or the identity of potential invaders is unknown.

Microbiome and environment explain the absence of correlations between consumers and their diet in Bornean microsnails.

Classical ecological theory posits that species partition resources such that each species occupies a unique resource niche. In general, the availability of more resources allows more species to co-occur. Thus, a strong relationship between communities of consumers and their resources is expected. However, correlations may be influenced by other layers in the food web, or by the environment. Here we show, by studying the relationship between communities of consumers (land snails) and individual diets (from seed plants), that there is in fact no direct, or at most a weak but negative, relationship. However, we found that the diversity of the individual microbiome positively correlates with both consumer community diversity and individual diet diversity in three target species. Moreover, these correlations were affected by various environmental variables, such as anthropogenic activity, habitat island size, and a possibly important nutrient source, guano runoff from nearby caves. Our results suggest that the microbiome and the environment explain the absence of correlations between diet and consumer community diversity. Hence, we advocate that microbiome inventories are routinely added to any community dietary analysis, which our study shows can be done with relatively little extra effort. Our approach presents the tools to quickly obtain an overview of the relationships between consumers and their resources. We anticipate our approach to be useful for ecologists and environmentalists studying different communities in a local food web.

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.

The projected timing of abrupt ecological disruption from climate change.

As anthropogenic climate change continues the risks to biodiversity will increase over time, with future projections indicating that a potentially catastrophic loss of global biodiversity is on the horizon1-3. However, our understanding of when and how abruptly this climate-driven disruption of biodiversity will occur is limited because biodiversity forecasts typically focus on individual snapshots of the future. Here we use annual projections (from 1850 to 2100) of temperature and precipitation across the ranges of more than 30,000 marine and terrestrial species to estimate the timing of their exposure to potentially dangerous climate conditions. We project that future disruption of ecological assemblages as a result of climate change will be abrupt, because within any given ecological assemblage the exposure of most species to climate conditions beyond their realized niche limits occurs almost simultaneously. Under a high-emissions scenario (representative concentration pathway (RCP) 8.5), such abrupt exposure events begin before 2030 in tropical oceans and spread to tropical forests and higher latitudes by 2050. If global warming is kept below 2 °C, less than 2% of assemblages globally are projected to undergo abrupt exposure events of more than 20% of their constituent species; however, the risk accelerates with the magnitude of warming, threatening 15% of assemblages at 4 °C, with similar levels of risk in protected and unprotected areas. These results highlight the impending risk of sudden and severe biodiversity losses from climate change and provide a framework for predicting both when and where these events may occur.

Macroevolutionary convergence connects morphological form to ecological function in birds.

Animals have diversified into a bewildering variety of morphological forms exploiting a complex configuration of trophic niches. Their morphological diversity is widely used as an index of ecosystem function, but the extent to which animal traits predict trophic niches and associated ecological processes is unclear. Here we use the measurements of nine key morphological traits for >99% bird species to show that avian trophic diversity is described by a trait space with four dimensions. The position of species within this space maps with 70-85% accuracy onto major niche axes, including trophic level, dietary resource type and finer-scale variation in foraging behaviour. Phylogenetic analyses reveal that these form-function associations reflect convergence towards predictable trait combinations, indicating that morphological variation is organized into a limited set of dimensions by evolutionary adaptation. Our results establish the minimum dimensionality required for avian functional traits to predict subtle variation in trophic niches and provide a global framework for exploring the origin, function and conservation of bird diversity.

Integrating behaviour and ecology into global biodiversity conservation strategies.

Insights into animal behaviour play an increasingly central role in species-focused conservation practice. However, progress towards incorporating behaviour into regional or global conservation strategies has been more limited, not least because standardized datasets of behavioural traits are generally lacking at wider taxonomic or spatial scales. Here we make use of the recent expansion of global datasets for birds to assess the prospects for including behavioural traits in systematic conservation priority-setting and monitoring programmes. Using International Union for Conservation of Nature Red List classifications for more than 9500 bird species, we show that the incidence of threat can vary substantially across different behavioural categories, and that some types of behaviour-including particular foraging, mating and migration strategies-are significantly more threatened than others. The link between behavioural traits and extinction risk is partly driven by correlations with well-established geographical and ecological factors (e.g. range size, body mass, human population pressure), but our models also reveal that behaviour modifies the effect of these factors, helping to explain broad-scale patterns of extinction risk. Overall, these results suggest that a multi-species approach at the scale of communities, continents and ecosystems can be used to identify and monitor threatened behaviours, and to flag up cases of latent extinction risk, where threatened status may currently be underestimated. Our findings also highlight the importance of comprehensive standardized descriptive data for ecological and behavioural traits, and point the way towards deeper integration of behaviour into quantitative conservation assessments. This article is part of the theme issue 'Linking behaviour to dynamics of populations and communities: application of novel approaches in behavioural ecology to conservation'.

Location-level processes drive the establishment of alien bird populations worldwide.

Human-mediated translocation of species to areas beyond their natural distribution (which results in 'alien' populations1) is a key signature of the Anthropocene2, and is a primary global driver of biodiversity loss and environmental change3. Stemming the tide of invasions requires understanding why some species fail to establish alien populations, and others succeed. To achieve this, we need to integrate the effects of features of the introduction site, the species introduced and the specific introduction event. Determining which, if any, location-level factors affect the success of establishment has proven difficult, owing to the multiple spatial, temporal and phylogenetic axes along which environmental variation may influence population survival. Here we apply Bayesian hierarchical regression analysis to a global spatially and temporally explicit database of introduction events of alien birds4 to show that environmental conditions at the introduction location, notably climatic suitability and the presence of other groups of alien species, are the primary determinants of successful establishment. Species-level traits and the size of the founding population (propagule pressure) exert secondary, but important, effects on success. Thus, current trajectories of anthropogenic environmental change will most probably facilitate future incursions by alien species, but predicting future invasions will require the integration of multiple location-, species- and event-level characteristics.

Dietary niche and the evolution of cranial morphology in birds.

Cranial morphology in birds is thought to be shaped by adaptive evolution for foraging performance. This understanding of ecomorphological evolution is supported by observations of avian island radiations, such as Darwin's finches, which display rapid evolution of skull shape in response to food resource availability and a strong fit between cranial phenotype and trophic ecology. However, a recent analysis of larger clades has suggested that diet is not necessarily a primary driver of cranial shape and that phylogeny and allometry are more significant factors in skull evolution. We use phenome-scale morphometric data across the breadth of extant bird diversity to test the influence of diet and foraging behaviour in shaping cranial evolution. We demonstrate that these trophic characters are significant but very weak predictors of cranial form at this scale. However, dietary groups exhibit significantly different rates of morphological evolution across multiple cranial regions. Granivores and nectarivores exhibit the highest rates of evolution in the face and cranial vault, whereas terrestrial carnivores evolve the slowest. The basisphenoid, occipital, and jaw joint regions have less extreme differences among dietary groups. These patterns demonstrate that dietary niche shapes the tempo and mode of phenotypic evolution in deep time, despite a weaker than expected form-function relationship across large clades.