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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.

[Aim] Climate change causes shifts in species ranges globally. Terrestrial plant species often lag behind temperature shifts, and it is unclear to what extent animal-dispersed plants can track climate change. Here, we estimate the ability of bird-dispersed plant species to track future temperature change on a tropical mountain., [Location] Tropical elevational gradient (500–3500 m.a.s.l.) in the Manú biosphere reserve, Peru. [Time period] From 1960–1990 to 2061–2080. [Taxa] Fleshy-fruited plants and avian frugivores. [Methods] Using simulations based on the functional traits of avian frugivores and fruiting plants, we quantified the number of long-distance dispersal (LDD) events that woody plant species would require to track projected temperature shifts on a tropical mountain by the year 2070 under different greenhouse gas emission scenarios [representative concentration pathway (RCP) 2.6, 4.5 and 8.5]. We applied this approach to 343 bird-dispersed woody plant species. [Results] Our simulations revealed that bird-dispersed plants differed in their climate-tracking ability, with large-fruited and canopy plants exhibiting a higher climate-tracking ability. Our simulations also suggested that even under scenarios of strong and intermediate mitigation of greenhouse gas emissions (RCP 2.6 and 4.5), sufficient upslope dispersal would require several LDD events by 2070, which is unlikely for the majority of woody plant species. Furthermore, the ability of plant species to track future changes in temperature increased in simulations with a low degree of trait matching between plants and birds, suggesting that plants in generalized seed-dispersal systems might be more resilient to climate change. [Main conclusion] Our study illustrates how the functional traits of plants and animals can inform predictive models of species dispersal and range shifts under climate change and suggests that the biodiversity of tropical mountain ecosystems is highly vulnerable to future warming. The increasing availability of functional trait data for plants and animals globally will allow parameterization of similar models for many other seed-dispersal systems., Fieldwork at Manú was conducted under the permits 041-2010-AG-DGFFSDGEFFS, 008-2011-AG-DGFFS-DGEFFS, 01-C/C-2010SERNANP-JPNM and 01-2011-SERNANP-PNM-JEF and supported by a scholarship from the German Academic Exchange Service to D.M.D. D.M.D. acknowledges funding from the European Research Council (ERC) under the European Union's Horizon 2020 research and innovation programme (grant number 787638) and the Swiss National Science Foundation (grant number 173342), both awarded to C. H. Graham. W.D.K. acknowledges a Global Ecology grant from the University of Amsterdam Faculty Research Cluster. I.D. was funded by the Alexander von Humboldt Foundation and is now supported by the Balearic Government. S.A.F. was funded by the German Research Foundation (DFG; FR 3246/2-2) and the Leibniz Competition of the Leibniz Association (P52/2017).

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.

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 CO 2 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 CO 2 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’.

AimWe analyse the functional consequences of the changes in species composition resulting from extinctions and introductions on oceanic island bird assemblages. Specifically, we ask if introduced species have compensated the functional loss resulting from species extinctions.LocationSeventy-four oceanic islands (>100 km2) in the Atlantic, Pacific and Indian Oceans.Time periodLate Holocene.Major taxa studiedTerrestrial and freshwater bird species.MethodsWe compiled a species list per island (extinct and extant, native and introduced), and then compiled traits per species. We used single-trait analyses to assess the effects of past species extinctions and introductions on functional composition. Then, we used probabilistic hypervolumes in trait space to calculate functional richness and evenness of original versus present avifaunas of each island (and net change), and to estimate functional originality of extinct and introduced species.ResultsThe net effects of extinctions and introductions were: an increase in average species richness per island (alpha diversity), yet a decline in diversity across all islands (gamma diversity); an average increase in the prevalence of most functional traits (23 out of 35) yet an average decline functional richness and evenness, associated with the fact that extinct species were functionally more original (when compared to extant natives) than introduced species.Main conclusionsIntroduced species are on average offsetting (and even surpassing) the losses of extinct species per island in terms of species richness, and they are increasing the prevalence of most functional traits. However, they are not compensating the loss of functional richness due to extinctions. Current island bird assemblages are becoming functionally poorer, having lost original species and being composed of functionally more homogeneous species. This is likely to have cascading repercussions on the functioning of island ecosystems.

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.

Aim: Urbanization exposes species to novel ecological conditions. Some species thrive in urban areas, whereas many others are excluded from these human-made environments. Previous analyses suggest that the ability to cope with rapid environmental change is associated with long-term patterns of diversification, but whether the suite of traits associated with the ability to colonize urban environments is linked to this process remains poorly understood. Location: World. Time period: Current. Major taxa studied: Passerine birds. Methods: We applied macroevolutionary models to a large dataset of passerine birds to compare the evolutionary history of urban-tolerant species with that of urban-avoidant species. Specifically, we examined models of state-dependent speciation and extinction to assess the macroevolution of urban tolerance as a binary trait, in addition to models of quantitative trait-dependent diversification based on relative urban abundance. We also ran simulation-based model assessments to explore potential sources of bias. Results: We provide evidence that historically, species with traits promoting urban colonization have undergone faster diversification than urban-avoidant species, indicating that urbanization favours clades with a historical tendency towards rapid speciation or reduced extinction. In addition, we find that past transitions towards states that currently impede urban colonization by passerines have been more frequent than in the opposite direction. Furthermore, we find a portion of urban-avoidant passerines to be recent and to undergo fast diversification. All highly supported models give this result consistently. Main conclusions: Urbanization is mainly associated with the loss of lineages that are inherently more vulnerable to extinction over deep time, whereas cities tend to be colonized by less vulnerable lineages, for which urbanization might be neutral or positive in terms of longer-term diversification. Urban avoidance is associated with high rates of recent diversification for some clades occurring in regions with relatively intact natural ecosystems and low current levels of urbanization.

AimThe coexistence and turnover of species along elevation gradients makes tropical mountains hotspots of biodiversity. However, understanding the historical processes through which species arising in geographic isolation (i.e. allopatry) assemble along the same mountain slope (i.e. sympatry) remains a major challenge. Multiple models have been proposed including, (1) the sorting of already elevationally divergent species, (2) the displacement of elevation upon secondary contact, potentially followed by convergence, or (3) elevational conservatism, in which ancestral elevational ranges are retained. However, the relative contribution of these processes to generating patterns of elevational overlap and turnover is unknown.LocationTropical mountains of Central- and South-America.Time periodThe last 12 myr.Major taxa studiedBirds.MethodsWe collate a dataset of 165 avian sister pairs containing estimates of phylogenetic age, geographical and regional elevational range overlap. We develop a framework based on continuous-time Markov models to infer the relative frequency of different historical pathways in explaining present day overlap and turnover of sympatric species along elevational gradients.ResultsWe show that the turnover of closely related bird species across elevation can predominantly be explained by displacement of elevation ranges upon contact (81%) rather than elevational divergence in allopatry (19%). In contrast, overlap along elevation gradients is primarily (88%) explained by conservatism of elevational ranges rather than displacement followed by elevational expansion (12%).Main conclusionsBird communities across tropical elevation gradients are assembled through a mix of processes, including the sorting, displacement and conservatism of species elevation ranges. The dominant role of conservatism in explaining co-occurrence of species on mountain slopes rejects more complex scenarios requiring displacement followed by subsequent expansion. The ability of closely related species to coexist without elevational divergence provides a direct and thus faster pathway to sympatry and may help explain the exceptional species richness of tropical mountains.

Tobias, Joseph A., Sheard, Catherine, Pigot, Alex L., Devenish, Adam J. M., Yang, Jingyi, Sayol, Ferran, Neate‐Clegg, Montague H. C., Alioravainen, Nico, Weeks, Thomas L., Barber, Robert A., Walkden, Patrick A., MacGregor, Hannah E. A., Jones, Samuel E. I., Vincent, Claire, Phillips, Anna G., Marples, Nicola M., Montaño‐Centellas, Flavia A., Leandro‐Silva, Victor, Claramunt, Santiago, Darski, Bianca, Freeman, Benjamin G., Bregman, Tom P., Cooney, Christopher R., Hughes, Emma C., Capp, Elliot J. R., Varley, Zoë K., Friedman, Nicholas R., Korntheuer, Heiko, Corrales‐Vargas, Andrea, Trisos, Christopher H., Weeks, Brian C., Hanz, Dagmar M., Töpfer, Till, Bravo, Gustavo A., Remeš, Vladimír, Nowak, Larissa, Carneiro, Lincoln S., Moncada R., Amilkar J., Matysioková, Beata, Baldassarre, Daniel T., Martínez‐Salinas, Alejandra, Wolfe, Jared D., Chapman, Philip M., Daly, Benjamin G., Sorensen, Marjorie C., Neu, Alexander, Ford, Michael A., Mayhew, Rebekah J., Fabio Silveira, Luis, Kelly, David J., Annorbah, Nathaniel N. D., Pollock, Henry S., Grabowska‐Zhang, Ada M., McEntee, Jay P., Carlos T. Gonzalez, Juan, Meneses, Camila G., Muñoz, Marcia C., Powell, Luke L., Jamie, Gabriel A., Matthews, Thomas J., Johnson, Oscar, Brito, Guilherme R. R., Zyskowski, Kristof, Crates, Ross, Harvey, Michael G., Jurado Zevallos, Maura, Hosner, Peter A., Bradfer‐Lawrence, Tom, Maley, James M., Stiles, F. Gary, Lima, Hevana S., Provost, Kaiya L., Chibesa, Moses, Mashao, Mmatjie, Howard, Jeffrey T., Mlamba, Edson, Chua, Marcus A. H., Li, Bicheng, Gómez, M. Isabel, García, Natalia C., Päckert, Martin, Fuchs, Jérôme, Ali, Jarome R., Derryberry, Elizabeth P., Carlson, Monica L., Urriza, Rolly C., Brzeski, Kristin E., Prawiradilaga, Dewi M., Rayner, Matt J., Miller, Eliot T., Bowie, Rauri C. K., Lafontaine, René‐Marie, Scofield, R. Paul, Lou, Yingqiang, Somarathna, Lankani, Lepage, Denis, Illif, Marshall, Neuschulz, Eike Lena, Templin, Mathias, Dehling, D. Matthias, Cooper, Jacob C., Pauwels, Olivier S. G., Analuddin, Kangkuso, Fjeldså, Jon, Seddon, Nathalie, Sweet, Paul R., DeClerck, Fabrice A. J., Naka, Luciano N., Brawn, Jeffrey D., Aleixo, Alexandre, Böhning‐Gaese, Katrin, Rahbek, Carsten, Fritz, Susanne A., Thomas, Gavin H., Schleuning, Matthias (2022): AVONET: morphological, ecological and geographical data for all birds. Ecology Letters 25 (3): 581-597, DOI: 10.1111/ele.13898, URL: http://dx.doi.org/10.1111/ele.13898

The origin, distribution, and function of biological diversity are fundamental themes of ecology and evolutionary biology. Research on birds has played a major role in the history and development of these ideas, yet progress was for many decades limited by a focus on patterns of current diversity, often restricted to particular clades or regions. Deeper insight is now emerging from a recent wave of integrative studies combining comprehensive phylogenetic, environmental, and functional trait data at unprecedented scales. We review these empirical advances and describe how they are reshaping our understanding of global patterns of bird diversity and the processes by which it arises, with implications for avian biogeography and functional ecology. Further expansion and integration of data sets may help to resolve longstanding debates about the evolutionary origins of biodiversity and offer a framework for understanding and predicting the response of ecosystems to environmental change.

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.

Description, Anthropogenic extinctions and alien establishments cause an overall loss of functional diversity of island bird communities., 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.

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.

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.

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. Bayesian hierarchical regression analysis of a global database of bird introduction events reveals the environmental, climatic and biotic factors that are the primary determinants of the successful establishment of populations of alien species.

A key question in invasion biology is why some regions have more alien species than others. Here, we provide a general framework to answer this. We model alien species richness as a function of the number of species introduced (colonization pressure) and the probability that each species establishes, which is a function of propagule pressure (the number of introduction events and the number of individuals per event) and the probability that a founding individual leaves a surviving lineage (lineage survival probability). With this model we show that: (a) alien species richness is most sensitive to variation in colonization pressure; (b) heterogeneity in lineage survival probability lowers the probability of population establishment, implying alien richness should be lower in more spatially or temporally variable environments; (c) heterogeneity in lineage survival probability leads to higher alien richness when a given propagule pressure is divided into more introduction events, each involving fewer individuals; and (d) we cannot quantify how specific components, such as lineage survival probability, influence alien species richness without data on other components and knowledge of how these covary. Overall, the model provides novel insights into the factors influencing alien species richness, and shows why we expect anthropogenic effects to be critical to this.

Despite biological invasions are one of the main environmental problems of the twenty-first century, there is still no theoretical or empirical agreement on whether a high phylogenetic relatedness between exotic and native species positively or negatively affect invasion success. To resolve this conundrum, it has been proposed that the effect might be scale-dependent, being negative at smaller spatial scales and positive at larger scales. Here we show that this scale-dependent pattern may be a sampling artefact associated with species-area effects and a non-random pattern of species introductions. We support this conclusion with simulations and empirical data on invaded and non-invaded avian communities in regions from five continents. We further show that at smaller-scales —where these artifacts are negligible— invasion success generally increases with the presence of closely-related species, but that predictive accuracy largely depends on considering the influence of human-related disturbances in facilitating invasions.

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.

Low elevation regions harbor the majority of the world’s species diversity compared to high elevation areas. This global elevational diversity gradient, suggests that lowland species have had more time to diversify, or that net diversification rates have been higher in the lowlands (either due to higher ecological limits or intrinsically higher diversification rates). 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 where passerines diversify at the same rate in the highlands and the lowlands but where the rate of dispersal from high to low elevations is more than twice as fast as in the reverse direction. This suggests that while there is no consistent trend in diversification across elevations, highland regions act as species pumps because the diversity they generate migrates into the lowlands, thus setting up the observed gradient in passerine diversity. This species pump is particularly strong in the tropics, where the inferred rate of speciation is 1.4 times faster than in the temperate zone. We conclude that despite their lower diversity, highland regions are disproportionally important for maintaining diversity in the adjacent lowlands. The extinction of species in the tropical highlands due to rapid climate change this century could thus have major and long-lasting impacts on global passerine diversity.

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.

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.

Aim Accelerating rates of anthropogenic introductions are leading to a dramatic restructuring of species distributions globally. However, the extent to which invasions alter the imprint of evolutionary history in species geographical ranges remains unclear. Here, we provide a global assessment of how the introduction, establishment and spread of alien species alters the phylogenetic signal in geographical range size using birds as a model system. Location Global. Time period Contemporaneous. Taxa Birds. Methods We compare the phylogenetic signal in alien range size with that of native distributions of species globally (n = 9,993) and across different stages in the invasion pathway, from introduced (n = 965) to established species (n = 359). Using stochastic simulations, we test whether differences in phylogenetic signal arise from nonrandom patterns of species introduction, establishment or spread. Results Geographical range size in birds exhibits an intermediate phylogenetic signal, driven by the spatial clustering of closely related species. Nonrandom introductions, biased towards wide‐ranging species from particular clades and regions, produce an anomalously strong phylogenetic signal in the native range size of introduced species. In contrast, the phylogenetic signal in alien range size is substantially weaker than for native distributions. This weak phylogenetic signal cannot be explained by a lack of time for dispersal but is instead regulated by phylogenetic correlations across species in the location and number of introduction events. Main conclusions We demonstrate that the effects of anthropogenic introductions on the phylogenetic signal in range size vary across different stages in the invasion pathway. The process of transport and introduction amplifies the phylogenetic signal in the pool of potential invaders, whereas the subsequent pattern of spread decouples variation in alien range size from phylogenetic ancestry. Together, our findings suggest that evolutionary relatedness is likely to be a relatively weak predictor of the spread of invasive species.

Aim: To identify the factors that influence the availability of data on the negative impacts of alien bird species, in order to understand why more than 70% are currently classified as Data Deficient (DD) by the Environmental Impact Classification of Alien Taxa (EICAT) protocol. / Location: Global. / Methods: Information on factors hypothesised to influence the availability of impact data were collated for 344 alien bird species (107 with impact data and 237 DD). These data were analysed using mixed effects models accounting for phylogenetic non-independence of species (MCMCglmm). / Results: Data deficiency in the negative impacts of alien birds is not randomly distributed. Residence time, relative brain size and alien range size were found to be strongly related to the availability of data on impacts. / Main conclusions: The availability of data on the negative impacts of alien birds is mainly influenced by the spatial and temporal extents of their alien ranges. The results of this study suggest that the impacts of some DD alien birds are likely to be minor (e.g. species with comparatively long residence times as aliens, such as the common waxbill (Estrilda astrild) and the Java sparrow (Padda oryzivora)). However, the results also suggest that some DD alien birds may have damaging impacts (e.g. species from orders of alien birds known for their impacts to biodiversity but with comparatively small alien ranges, such as the New Caledonian crow (Corvus moneduloides)). This implies that at least some DD alien birds may have impacts that are being overlooked. Studies examining the traits that influence the severity of alien bird impacts are needed to help to predict which DD species are more likely to impact upon biodiversity.

1.Identifying the factors that determine the success of biological invasions has major consequences for both ecological theory and conservation decision-making. Reliably inferring these factors depends on adequately accounting for the known effects of propagule pressure on establishment success, but detailed information on the size and number of introduced populations is often lacking. 2.Here, we conduct simulations to explore the effects of incomplete knowledge of propagule pressure on inferences regarding the correlates of establishment success. We compare situations where we have complete information on propagule number and propagule size across species, allowing success to be modelled at the population scale (population-level analysis), with those where data for propagule size (species-level analysis) and both propagule size and number (location-level analysis) are unavailable. We assess the ability to correctly infer the effects of a covariate on establishment success when this covariate exhibits varying degrees of correlation with propagule pressure. 3.We show that when establishment success is modelled at the level of species, rather than populations, this leads to an elevated tendency to incorrectly infer the effects of species level traits on establishment success (higher Type-1 error and lower power), particular when traits in question are strongly associated with propagule pressure. These biases are magnified when using proxies for propagule pressure, such as the number of locations where species have been introduced, and are magnified further when these proxies are converted to binary variables. 4.Our results validate the current best practice for the analysis of establishment success in alien species at the population-level, when the effects of propagule size and a covariate predicting establishment probability can both be reliably inferred. However, given the growing interest in understanding correlates of biological invasions, they strongly urge caution in interpreting results based on incomplete knowledge of propagule pressure. This article is protected by copyright. All rights reserved.

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.

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 for99% 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.

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.

Slow-downs in lineage accumulation in phylogenies suggest that speciation rates decline as diversity increases. Likelihood methods have been developed to detect such diversity-dependence. However, a thorough test of whether such approaches correctly infer diversity-dependence is lacking.Here we simulate phylogenetic branching under linear negative diversity-dependent and diversity-independent models and estimate from the simulated phylogenies the maximum likelihood parameters for three different conditionings – on survival of the birth-death process given the crown age, on tree size (N), and on tree size given the crown age. We report the accuracy of recovering the simulation parameters and the reliability of the model selection based on the χ2 likelihood ratio test.Parameter estimate accuracyConditioning on survival given the crown age yields a severe bias of the carrying capacity K toward N, and an upward bias of the speciation rate, particularly in clades where diversity-dependent feedbacks are still weak (N « K). Conditioning on N yields an overestimate of K and an underestimate of speciation rate, particularly when saturation has been reached. Dual conditioning yields relatively unbiased parameter estimates on average, but the deviation from the true value for any single estimate may be large.Model selection reliabilityThe frequency of incorrectly rejecting a diversity-independent model when the simulation was diversity-independent (Type I error) differs substantially from the significance level α used in the likelihood ratio test, rendering the likelihood ratio test inappropriate. The frequency of correctly rejecting the diversity-independent model when the simulation was diversity-dependent (power) is larger when the clade is closer to equilibrium, and for conditioning on crown age.We conclude that conditioning on crown age has the best statistical properties overall, but caution that parameter estimates may be biased. To assess parameter uncertainty in future studies of diversity-dependence on real data, we recommend parametric bootstrapping, examination of the likelihood surface and comparison of estimates across the types of conditioning. To assess model selection reliability we discourage the use of the χ2 likelihood ratio test or AIC (which are equivalent in this case), but recommend a likelihood ratio test based on parametric bootstrap. We illustrate this method for the diversification of Dendroica warblers.

The role of ecological limits in regulating the distribution and diversification of species remains controversial. Although such limits must ultimately arise from constraints on local species coexistence, this spatial context is missing from most macroevolutionary models. Here, we develop a stochastic, spatially explicit model of species diversification to explore the phylogenetic and biogeographic patterns expected when local diversity is bounded. We show how local ecological limits, by regulating opportunities for range expansion and thus rates of speciation and extinction, lead to temporal slowdowns in diversification and predictable differences in equilibrium diversity between regions. However, our models also show that even when regions have identical diversity limits, the dynamics of diversification and total number of species supported at equilibrium can vary dramatically depending on the relative size of geographic and local ecological niche space. Our model predicts that small regions with higher local ecological limits support a higher standing diversity and more balanced phylogenetic trees than large geographic areas with more stringent constraints on local coexistence. Our findings highlight how considering the spatial context of diversification can provide new insights into the role of ecological limits in driving variation in biodiversity across space, time, and clades.

This research was funded by the Netherlands Organisation for Scientific Research VENI grant 863.13.003 (to A.L.P.), NASA Biodiversity grant NNX11AP72G and NSF grants NSF DBI 1262600, DBI 0960550 and DEB 1026764 (to W.J.), the Oxford Clarendon Fund and US-UK Fulbright Commission (to C.S.), and the John Fell Fund and NERC grant NE/I028068/1 (to J.A.T.). Ecological communities are assembled from the overlapping of species in geographic space, but the mechanisms facilitating or limiting such overlaps are difficult to resolve. Here, we combine phylogenetic, morphological and environmental data to model how multiple processes regulate the origin and maintenance of geographic range overlap across 1,115 pairs of avian sister species globally. We show that coexistence cannot be adequately predicted by either dispersal-assembly (that is, biogeographic) models or niche-assembly models alone. Instead, our results overwhelmingly support an integrated model with different assembly processes dominating at different stages of coexistence. The initial attainment of narrow geographic overlap is dictated by intrinsic dispersal ability and the time available for dispersal, whereas wider coexistence is largely dependent on niche availability, increasing with ecosystem productivity and divergence in niche-related traits, and apparently declining as communities become saturated with species. Furthermore, although coexistence of any individual pair of species is highly stochastic, we find that integrating assembly processes allows broad variation in the incidence and extent of coexistence to be predicted with reasonable accuracy. Our findings demonstrate how phylogenetic data coupled with environmental factors and functional traits can begin to clarify the multi-layered processes shaping the distribution of biodiversity at large spatial scales. Postprint

Quantifying the role of biodiversity in ecosystems not only requires understanding the links between species and the ecological functions and services they provide, but also how these factors relate to measurable indices, such as functional traits and phylogenetic diversity. However, these relationships remain poorly understood, especially for heterotrophic organisms within complex ecological networks. Here, we assemble data on avian traits across a global sample of mutualistic plant–frugivore networks to critically assess how the functional roles of frugivores are associated with their intrinsic traits, as well as their evolutionary and functional distinctiveness. We find strong evidence for niche complementarity, with phenotypically and phylogenetically distinct birds interacting with more unique sets of plants. However, interaction strengths—the number of plant species dependent on a frugivore—were unrelated to evolutionary or functional distinctiveness, largely because distinct frugivores tend to be locally rare, and thus have fewer connections across the network. Instead, interaction strengths were better predicted by intrinsic traits, including body size, gape width and dietary specialization. Our analysis provides general support for the use of traits in quantifying species ecological functions, but also highlights the need to go beyond simple metrics of functional or phylogenetic diversity to consider the multiple pathways through which traits may determine ecological processes.

International audience; Meeting the ever-increasing needs of the Earth's human population without excessively reducing biological diversity is one of the greatest challenges facing humanity, suggesting that newapproaches to biodiversity conservation are required. One idea rapidly gaining momentum-as well as opposition-is to incorporate the values of biodiversity into decision-making using economic methods. Here, we develop several lines of argument for how biodiversity might be valued, building on recent developments in natural science, economics and science-policy processes. Then we provide a synoptic guide to the papers in this special feature, summarizing recent research advances relevant to biodiversity valuation and management. Current evidence suggests that more biodiverse systems have greater stability and resilience, and that by maximizing key components of biodiversity we maximize an ecosystem's long-term value. Moreover, many services and values arising from biodiversity are interdependent, and often poorly captured by standard economic models. We conclude that economic valuation approaches to biodiversity conservation should (i) account for interdependency and (ii) complement rather than replace traditional approaches. To identify possible solutions, we present a framework for understanding the foundational role of hard-to-quantify ` biodiversity services' in sustaining the value of ecosystems to humanity, and then use this framework to highlight new directions for pure and applied research. In most cases, clarifying the links between biodiversity and ecosystem services, and developing effective policy and practice for managing biodiversity, will require a genuinely interdisciplinary approach.

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’.

Whether biotic interactions limit geographic ranges has long been controversial, and traditional analyses of static distribution patterns have made little progress towards resolving this debate. Here, we use a novel phylogenetic approach to test whether biotic interactions constrain the transition to secondary sympatry following speciation. Applying this temporal framework to a diverse clade of passerine birds (Furnariidae), we reject models of geographic range overlap limited purely by dispersal or environmental constraints, and instead show that rates of secondary sympatry are positively associated with both the phylogenetic and morphological distance between species. Thus, transition rates to sympatry increase with time since divergence and accelerate as the ecological differences between species accumulate. Taken together, these results provide strong empirical evidence that biotic interactions - and primarily ecological competition - limit species distributions across large spatial and temporal scales. They also offer phylogenetic and trait-based metrics by which these interactions can be incorporated into ecological forecasting models. © 2012 Blackwell Publishing Ltd/CNRS.

The association between species richness and ecosystem energy availability is one of the major geographic trends in biodiversity. It is often explained in terms of energetic constraints, such that coexistence among competing species is limited in low productivity environments. However, it has proven challenging to reject alternative views, including the null hypothesis that species richness has simply had more time to accumulate in productive regions, and thus the role of energetic constraints in limiting coexistence remains largely unknown. We use the phylogenetic relationships and geographic ranges of sister species (pairs of lineages who are each other’s closest extant relatives) to examine the association between energy availability and coexistence across an entire vertebrate class (Aves). We show that the incidence of coexistence among sister species increases with overall species richness and is elevated in more productive ecosystems, even when accounting for differences in the evolutionary time available for coexistence to occur. Our results indicate that energy availability promotes species coexistence in closely related lineages, providing a key step toward a more mechanistic understanding of the productivity–richness relationship underlying global gradients in biodiversity., An analysis of phylogenetic and geographic data across the world's bird species reveals that the probability of species coexistence is constrained by the availability of energy in the environment., Author Summary The increase in the number of species with the availability of energy in the environment is one of the most general but least understood patterns in global biodiversity. The finite amount of energy flowing through an ecosystem has long been suspected to place a fundamental constraint on the ability of species to subdivide ecological resources, with greater energy availability—and, thus, ecosystem productivity—in the tropics, potentially facilitating increased levels of coexistence. However, empirical support for this hypothesis has been lacking, raising the possibility that richness is higher in productive ecosystems for largely historical reasons, including the greater geological age and area of tropical biomes, which increases the period of time available for diversity to accumulate. By combining phylogenetic and geographic data from across the world’s bird species, we show that greater ecosystem productivity is associated with an increased probability of coexistence among closely related lineages and that this pattern contributes to the higher species richness in the tropics. Our results confirm that contemporary gradients in species richness are fundamentally shaped by energetic constraints on coexistence.

Phylogenetic trees often depart from the expectations of stochastic models, exhibiting imbalance in diversification among lineages and slowdowns in the rate of lineage accumulation through time. Such departures have led to a widespread perception that ecological differences among species or adaptation and subsequent niche filling are required to explain patterns of diversification. However, a key element missing from models of diversification is the geographical context of speciation and extinction. In this study, we develop a spatially explicit model of geographic range evolution and cladogenesis, where speciation arises via vicariance or peripatry, and explore the effects of these processes on patterns of diversification. We compare the results with those observed in 41 reconstructed avian trees. Our model shows that nonconstant rates of speciation and extinction are emergent properties of the apportioning of geographic ranges that accompanies speciation. The dynamics of diversification exhibit wide variation, depending on the mode of speciation, tendency for range expansion, and rate of range evolution. By varying these parameters, the model is able to capture many, but not all, of the features exhibited by birth-death trees and extant bird clades. Under scenarios with relatively stable geographic ranges, strong slowdowns in diversification rates are produced, with faster rates of range dynamics leading to constant or accelerating rates of apparent diversification. A peripatric model of speciation with stable ranges also generates highly unbalanced trees typical of bird phylogenies but fails to produce realistic range size distributions among the extant species. Results most similar to those of a birth-death process are reached under a peripatric speciation scenario with highly volatile range dynamics. Taken together, our results demonstrate that considering the geographical context of speciation and extinction provides a more conservative null model of diversification and offers a very different perspective on the phylogenetic patterns expected in the absence of ecology.

The environmental factors limiting species' ranges across broad geographic and taxonomic scales are central to questions regarding the geographic variation in biodiversity and impacts of environmental change. However, our understanding remains relatively limited owing to the small-scale, correlative nature of most previous analyses. In this study, we provide a global test of the environmental determinants of range limits in birds, using both stochastic and environmentally deterministic models of range expansion to simulate spatial patterns in the shape of species' distributions. We show that spatial variation in range shape can be well explained by the action of a few key climatic variables in limiting range expansion. Furthermore, we demonstrate that these patterns cannot be explained by random processes alone. In addition, our study also identifies important differences in the factors constraining range expansion between biogeographic realms and between widespread and rare species. Although temperature was the principal climatic determinant of range shape at a global scale, its effects were driven primarily by widespread species, with the ranges of rare taxa better predicted by gradients in precipitation or topography. These key results were robust with respect to the index used to quantify range shape. Our analysis helps resolve long running debates regarding the role of environmentally deterministic and stochastic processes in structuring biodiversity and extends our understanding of the factors limiting species' ranges and the response of broad scale biodiversity patterns to environmental change.