Climate-land-use interactions shape tropical mountain biodiversity and ecosystem functions.

Agriculture and the exploitation of natural resources have transformed tropical mountain ecosystems across the world, and the consequences of these transformations for biodiversity and ecosystem functioning are largely unknown1-3. Conclusions that are derived from studies in non-mountainous areas are not suitable for predicting the effects of land-use changes on tropical mountains because the climatic environment rapidly changes with elevation, which may mitigate or amplify the effects of land use4,5. It is of key importance to understand how the interplay of climate and land use constrains biodiversity and ecosystem functions to determine the consequences of global change for mountain ecosystems. Here we show that the interacting effects of climate and land use reshape elevational trends in biodiversity and ecosystem functions on Africa's largest mountain, Mount Kilimanjaro (Tanzania). We find that increasing land-use intensity causes larger losses of plant and animal species richness in the arid lowlands than in humid submontane and montane zones. Increases in land-use intensity are associated with significant changes in the composition of plant, animal and microorganism communities; stronger modifications of plant and animal communities occur in arid and humid ecosystems, respectively. Temperature, precipitation and land use jointly modulate soil properties, nutrient turnover, greenhouse gas emissions, plant biomass and productivity, as well as animal interactions. Our data suggest that the response of ecosystem functions to land-use intensity depends strongly on climate; more-severe changes in ecosystem functioning occur in the arid lowlands and the cold montane zone. Interactions between climate and land use explained-on average-54% of the variation in species richness, species composition and ecosystem functions, whereas only 30% of variation was related to single drivers. Our study reveals that climate can modulate the effects of land use on biodiversity and ecosystem functioning, and points to a lowered resistance of ecosystems in climatically challenging environments to ongoing land-use changes in tropical mountainous regions.

Projected climate change impacts on the phylogenetic diversity of the world's terrestrial birds: more than species numbers.

Ongoing climate change is a major threat to biodiversity. As abiotic tolerances and dispersal abilities vary, species-specific responses have the potential to further amplify or ameliorate the ensuing impacts on species assemblages. Here, we investigate the effects of climate change on species distributions across non-marine birds, quantifying its projected impact on species richness (SR) as well as on different aspects of phylogenetic diversity globally. Going beyond previous work, we disentangle the potential impacts of species gains versus losses on assemblage-level phylogenetic diversity under climate change and compare the projected impacts to randomized assemblage changes. We show that beyond its effects on SR, climate change could have profound impacts on assemblage-level phylogenetic diversity and composition, which differ significantly from random changes and among regions. Though marked species losses are most frequent in tropical and subtropical areas in our projections, phylogenetic restructuring of species communities is likely to occur all across the globe. Furthermore, our results indicate that the most severe changes to the phylogenetic diversity of local assemblages are likely to be caused by species range shifts and local species gains rather than range reductions and extinctions. Our findings highlight the importance of considering diverse measures in climate impact assessments.

A research framework for projecting ecosystem change in highly diverse tropical mountain ecosystems.

Tropical mountain ecosystems are threatened by climate and land-use changes. Their diversity and complexity make projections how they respond to environmental changes challenging. A suitable way are trait-based approaches, by distinguishing between response traits that determine the resistance of species to environmental changes and effect traits that are relevant for species' interactions, biotic processes, and ecosystem functions. The combination of those approaches with land surface models (LSM) linking the functional community composition to ecosystem functions provides new ways to project the response of ecosystems to environmental changes. With the interdisciplinary project RESPECT, we propose a research framework that uses a trait-based response-effect-framework (REF) to quantify relationships between abiotic conditions, the diversity of functional traits in communities, and associated biotic processes, informing a biodiversity-LSM. We apply the framework to a megadiverse tropical mountain forest. We use a plot design along an elevation and a land-use gradient to collect data on abiotic drivers, functional traits, and biotic processes. We integrate these data to build the biodiversity-LSM and illustrate how to test the model. REF results show that aboveground biomass production is not directly related to changing climatic conditions, but indirectly through associated changes in functional traits. Herbivory is directly related to changing abiotic conditions. The biodiversity-LSM informed by local functional trait and soil data improved the simulation of biomass production substantially. We conclude that local data, also derived from previous projects (platform Ecuador), are key elements of the research framework. We specify essential datasets to apply this framework to other mountain ecosystems.

Bioenergy cropland expansion may offset positive effects of climate change mitigation for global vertebrate diversity.

Climate and land-use change interactively affect biodiversity. Large-scale expansions of bioenergy have been suggested as an important component for climate change mitigation. Here we use harmonized climate and land-use projections to investigate their potential combined impacts on global vertebrate diversity under a low- and a high-level emission scenario. We combine climate-based species distribution models for the world's amphibians, birds, and mammals with land-use change simulations and identify areas threatened by both climate and land-use change in the future. The combined projected effects of climate and land-use change on vertebrate diversity are similar under the two scenarios, with land-use change effects being stronger under the low- and climate change effects under the high-emission scenario. Under the low-emission scenario, increases in bioenergy cropland may cause severe impacts in biodiversity that are not compensated by lower climate change impacts. Under this low-emission scenario, larger proportions of species distributions and a higher number of small-range species may become impacted by the combination of land-use and climate change than under the high-emission scenario, largely a result of bioenergy cropland expansion. Our findings highlight the need to carefully consider both climate and land-use change when projecting biodiversity impacts. We show that biodiversity is likely to suffer severely if bioenergy cropland expansion remains a major component of climate change mitigation strategies. Our study calls for an immediate and significant reduction in energy consumption for the benefit of both biodiversity and to achieve the goals of the Paris Agreement.

Long-term declines of European insectivorous bird populations and potential causes.

Evidence of declines in insect populations has recently received considerable scientific and societal attention. However, the lack of long-term insect monitoring makes it difficult to assess whether declines are geographically widespread. By contrast, bird populations are well monitored and often used as indicators of environmental change. We compared the population trends of European insectivorous birds with those of other birds to assess whether patterns in bird population trends were consistent with declines of insects. We further examined whether declines were evident for insectivores with different habitats, foraging strata, and other ecological preferences. Bird population trends were estimated for Europe (1990-2015) and Denmark (1990-2016). On average, insectivores declined over the study period (13% across Europe and 28% in Denmark), whereas omnivores had stable populations. Seedeaters also declined (28% across Europe; 34% in Denmark), but this assessment was based on fewer species than for other groups. The effects of insectivory were stronger for farmland species (especially grassland species), for ground feeders, and for cold-adapted species. Insectivory was associated with long-distance migration, which was also linked to population declines. However, many insectivores had stable populations, especially habitat generalists. Our findings suggest that the decline of insectivores is primarily associated with agricultural intensification and loss of grassland habitat. The loss of both seed and insect specialists indicates an overall trend toward bird communities dominated by diet generalists.

Declinaciones a Largo Plazo de Poblaciones de Aves Insectívoras en Europa y las Causas Probables Resumen La evidencia de las declinaciones poblacionales de insectos ha recibido recientemente una atención considerable por parte de la comunidad científica y la sociedad. Sin embargo, la falta de un monitoreo prolongado de los insectos complica valorar si estas declinaciones tienen una distribución extensa geográficamente. Como contraste, las poblaciones de aves tienen un monitoreo constante y con frecuencia se usan como indicadores del cambio climático. Comparamos las tendencias poblacionales de las aves insectívoras de Europa con las de otras aves para valorar si los patrones en las tendencias poblacionales de aves son consistentes con las declinaciones de insectos. Además examinamos si las declinaciones eran evidentes para aves insectívoras con diferentes hábitats, estratos de alimentación, y otras preferencias ecológicas. Las tendencias poblacionales de las aves se estimaron para Europa (1990 - 2015) y para Dinamarca (1990 - 2016). En promedio, las aves insectívoras declinaron a lo largo del periodo de estudio (13% en Europa y 28% en Dinamarca) mientras que las aves omnívoras tuvieron poblaciones estables. Las poblaciones de aves que se alimentan de semillas también declinaron (28% en Europa; 34% en Dinamarca), pero esta valoración se basó en menos especies que para los otros grupos. Los efectos de la insectivoría fueron más evidentes para las especies de tierras agrícolas (especialmente las especies de pastizales), para las especies que se alimentan sobre el suelo y para las especies adaptadas al frío. La insectivoría estuvo asociada con la migración de larga distancia, la cual también estuvo ligada a las declinaciones poblacionales. Sin embargo, muchas aves insectívoras tuvieron poblaciones estables, especialmente aquellas generalistas de hábitat. Nuestros hallazgos sugieren que la declinación de las aves insectívoras está asociada principalmente con la intensificación agrícola y la pérdida de pastizales. La pérdida de aves cuya alimentación es especialista en insectos o en semillas indica una tendencia general hacia comunidades de aves dominadas por aquellas con dietas generalistas.

Different responses of taxonomic and functional bird diversity to forest fragmentation across an elevational gradient.

Many studies have investigated how habitat fragmentation affects the taxonomic and functional diversity of species assemblages. However, the joint effects of habitat fragmentation and environmental conditions on taxonomic and functional diversity, for instance across elevational gradients, have largely been neglected so far. In this study, we compare whether taxonomic and functional indicators show similar or distinct responses to forest fragmentation across an elevational gradient. We based our analysis on a comprehensive data set of species-rich bird assemblages from tropical montane forest in the Southern Andes of Ecuador. We monitored birds over 2 years in two habitat types (continuous and fragmented forest) at three elevations (i.e., 1000, 2000, and 3000 m a.s.l) and measured nine morphological traits for each bird species on museum specimens. Bird species richness and abundance were significantly higher in fragmented compared to continuous forests and decreased towards high elevations. In contrast, functional diversity was significantly reduced in fragmented compared to continuous forests at low elevations, but fragmentation effects on functional diversity tended to be reversed at high elevations. Our results demonstrate that taxonomic and functional indicators can show decoupled responses to forest fragmentation and that these effects are highly variable across elevations. Our findings reveal that functional homogenization in bird communities in response to fragmentation can be masked by apparent increases in taxonomic diversity, particularly in diverse communities at low elevations.

Direct and indirect effects of plant and frugivore diversity on structural and functional components of fruit removal by birds.

Seed dispersal is an important ecosystem function, but it is contentious how structural and functional diversity of plant and bird communities are associated with seed-dispersal functions. We used structural equation models to test how structural (i.e., abundance, species richness) and functional diversity (i.e., functional dispersion and community-weighted means of functional traits) of fruiting plants and frugivorous birds directly and indirectly influence the respective components of fruit removal. We recorded plant and bird diversity in point counts and observed plant-frugivore interactions in a tropical mountain forest in Ecuador. We also recorded plant and bird morphological traits to calculate measures of functional diversity. We found that fruit abundance had a positive direct effect on bird abundance, which directly and indirectly mediated the abundance of removed fruits. Plant and bird species richness were only directly related to the richness of the removed fruits. Functional dispersion of the plant community was positively associated to that of the bird community and to that of the removed fruits. Consistently, we found positive associations between community-weighted means of plant and bird traits and between community-weighted means of plant traits and that of plants with removed fruits. In contrast, community-weighted means of the bird community were unrelated to that of the removed fruits. Overall, our results suggest that plant abundance directly and indirectly influences fruit removal, likely because of avian fruit tracking. However, we did not find strong links between the functional diversity of the frugivore community and removed fruits, suggesting that other factors in addition to plant-animal trait matching might be important for the functional diversity of removed fruits. Our findings highlight the importance of frugivore abundance for maintaining seed dispersal by animals in tropical forests.

Twenty-million-year relationship between mammalian diversity and primary productivity.

At global and regional scales, primary productivity strongly correlates with richness patterns of extant animals across space, suggesting that resource availability and climatic conditions drive patterns of diversity. However, the existence and consistency of such diversity-productivity relationships through geological history is unclear. Here we provide a comprehensive quantitative test of the diversity-productivity relationship for terrestrial large mammals through time across broad temporal and spatial scales. We combine >14,000 occurrences for 690 fossil genera through the Neogene (23-1.8 Mya) with regional estimates of primary productivity from fossil plant communities in North America and Europe. We show a significant positive diversity-productivity relationship through the 20-million-year record, providing evidence on unprecedented spatial and temporal scales that this relationship is a general pattern in the ecology and paleo-ecology of our planet. Further, we discover that genus richness today does not match the fossil relationship, suggesting that a combination of human impacts and Pleistocene climate variability has modified the 20-million-year ecological relationship by strongly reducing primary productivity and driving many mammalian species into decline or to extinction.

Disentangling the effects of multiple environmental drivers on population changes within communities.

The effects of different environmental drivers on the changes in species' population abundances can be difficult to disentangle as they often act simultaneously. Researchers have built statistical models that include environmental variables (such as annual temperature) or species attributes (such as a species' temperature preference), which are assumed to detect the impacts of specific drivers (such as climate change). However, these approaches are often applied separately or, if combined, not explicitly compared. We show the complementary insights gained by applying both these approaches to a community dataset on Danish terrestrial birds. We use our analysis to compare the relative importance of climate change and agricultural land-use change for the abundance changes within the community between 1983 and 2013. Population models were fitted to the community data of species' annual abundances with predictors comprising: species attributes (species' temperature and habitat preferences), environmental variables (climatic and agricultural land-use change variables) or both. Relationships between species' abundances and environmental variables were used to identify the drivers associated with average abundance changes of species in the community. Relationships between species' abundances and their attributes were used to understand the drivers causing interspecific variation in abundance changes. Warmer winters were positively associated with community-level abundances, and warm-adapted species had more positive abundance changes than cold-adapted ones. Agricultural land-use area was negatively associated with community-level abundances, and birds using a high proportion of meadow and habitat specialists had more negative abundance changes than birds using other habitats and habitat generalists. Effect sizes of environmental variables were larger for agricultural land-use change while those of species attributes were larger for climate change. The environmental data approach suggested that agricultural land-use change has decreased the average abundances of species in the community, affecting total community size while the species attribute-based approach suggested that climate change has caused variation in abundance among species, affecting community composition. Environmental variables and species attributes that are hypothesized to link to specific drivers can be used together to provide complementary information on the impacts of different drivers on communities.

An estimate of the number of tropical tree species.

The high species richness of tropical forests has long been recognized, yet there remains substantial uncertainty regarding the actual number of tropical tree species. Using a pantropical tree inventory database from closed canopy forests, consisting of 657,630 trees belonging to 11,371 species, we use a fitted value of Fisher's alpha and an approximate pantropical stem total to estimate the minimum number of tropical forest tree species to fall between ∼ 40,000 and ∼ 53,000, i.e., at the high end of previous estimates. Contrary to common assumption, the Indo-Pacific region was found to be as species-rich as the Neotropics, with both regions having a minimum of ∼ 19,000-25,000 tree species. Continental Africa is relatively depauperate with a minimum of ∼ 4,500-6,000 tree species. Very few species are shared among the African, American, and the Indo-Pacific regions. We provide a methodological framework for estimating species richness in trees that may help refine species richness estimates of tree-dependent taxa.

Cross-taxa generalities in the relationship between population abundance and ambient temperatures.

Identifying patterns in the effects of temperature on species' population abundances could help develop a general framework for predicting the consequences of climate change across different communities and realms. We used long-term population time series data from terrestrial, freshwater, and marine species communities within central Europe to compare the effects of temperature on abundance across a broad range of taxonomic groups. We asked whether there was an average relationship between temperatures in different seasons and annual abundances of species in a community, and whether species attributes (temperature range of distribution, range size, habitat breadth, dispersal ability, body size, and lifespan) explained interspecific variation in the relationship between temperature and abundance. We found that, on average, warmer winter temperatures were associated with greater abundances in terrestrial communities (ground beetles, spiders, and birds) but not always in aquatic communities (freshwater and marine invertebrates and fish). The abundances of species with large geographical ranges, larger body sizes, and longer lifespans tended to be less related to temperature. Our results suggest that climate change may have, in general, positive effects on species' abundances within many terrestrial communities in central Europe while the effects are less predictable in aquatic communities.

Functionally specialised birds respond flexibly to seasonal changes in fruit availability.

Interactions between resource and consumer species result in complex ecological networks. The overall structure of these networks is often stable in space and time, but little is known about the temporal stability of the functional roles of consumer species in these networks. We used a trait-based approach to investigate whether consumers (frugivorous birds) show similar degrees of functional specialisation on resources (plants) in ecological networks across seasons. We additionally tested whether closely related bird species have similar degrees of functional specialisation and whether birds that are functionally specialised on specific resource types within a season are flexible in switching to other resource types in other seasons. We analysed four seasonal replicates of two species-rich plant-frugivore networks from the tropical Andes. To quantify fruit preferences of frugivorous birds, we projected their interactions with plants into a multidimensional plant trait space. To measure functional specialisation of birds, we calculated a species' functional niche breadth (the extent of seasonal plant trait space utilised by a particular bird) and functional originality (the extent to which a bird species' fruit preference functionally differs from those of other species in a seasonal network). We additionally calculated functional flexibility, i.e. the ability of bird species to change their fruit preference across seasons in response to variation in plant resources. Functional specialisation of bird species varied more among species than across seasons, and phylogenetically similar bird species showed similar degrees of functional niche breadth (phylogenetic signal λ = 0·81) and functional originality (λ = 0·89). Additionally, we found that birds with high functional flexibility across seasons had narrow functional niche breadth and high functional originality per season, suggesting that birds that are seasonally specialised on particular resources are most flexible in switching to other fruit resources across seasons. The high flexibility of functionally specialised bird species to switch seasonally to other resources challenges the view that consumer species rely on functionally similar resources throughout the year. This flexibility of consumer species may be an important, but widely neglected mechanism that could potentially stabilise consumer-resource networks in response to human disturbance and environmental change.

Morphology predicts species' functional roles and their degree of specialization in plant-frugivore interactions.

Species' functional roles in key ecosystem processes such as predation, pollination or seed dispersal are determined by the resource use of consumer species. An interaction between resource and consumer species usually requires trait matching (e.g. a congruence in the morphologies of interaction partners). Species' morphology should therefore determine species' functional roles in ecological processes mediated by mutualistic or antagonistic interactions. We tested this assumption for Neotropical plant-bird mutualisms. We used a new analytical framework that assesses a species's functional role based on the analysis of the traits of its interaction partners in a multidimensional trait space. We employed this framework to test (i) whether there is correspondence between the morphology of bird species and their functional roles and (ii) whether morphologically specialized birds fulfil specialized functional roles. We found that morphological differences between bird species reflected their functional differences: (i) bird species with different morphologies foraged on distinct sets of plant species and (ii) morphologically distinct bird species fulfilled specialized functional roles. These findings encourage further assessments of species' functional roles through the analysis of their interaction partners, and the proposed analytical framework facilitates a wide range of novel analyses for network and community ecology.

Continent-scale global change attribution in European birds - combining annual and decadal time scales.

Species attributes are commonly used to infer impacts of environmental change on multiyear species trends, e.g. decadal changes in population size. However, by themselves attributes are of limited value in global change attribution since they do not measure the changing environment. A broader foundation for attributing species responses to global change may be achieved by complementing an attributes-based approach by one estimating the relationship between repeated measures of organismal and environmental changes over short time scales. To assess the benefit of this multiscale perspective, we investigate the recent impact of multiple environmental changes on European farmland birds, here focusing on climate change and land use change. We analyze more than 800 time series from 18 countries spanning the past two decades. Analysis of long-term population growth rates documents simultaneous responses that can be attributed to both climate change and land-use change, including long-term increases in populations of hot-dwelling species and declines in long-distance migrants and farmland specialists. In contrast, analysis of annual growth rates yield novel insights into the potential mechanisms driving long-term climate induced change. In particular, we find that birds are affected by winter, spring, and summer conditions depending on the distinct breeding phenology that corresponds to their migratory strategy. Birds in general benefit from higher temperatures or higher primary productivity early on or in the peak of the breeding season with the largest effect sizes observed in cooler parts of species' climatic ranges. Our results document the potential of combining time scales and integrating both species attributes and environmental variables for global change attribution. We suggest such an approach will be of general use when high-resolution time series are available in large-scale biodiversity surveys.

Contrasting changes in the abundance and diversity of North American bird assemblages from 1971 to 2010.

Although it is generally recognized that global biodiversity is declining, few studies have examined long-term changes in multiple biodiversity dimensions simultaneously. In this study, we quantified and compared temporal changes in the abundance, taxonomic diversity, functional diversity, and phylogenetic diversity of bird assemblages, using roadside monitoring data of the North American Breeding Bird Survey from 1971 to 2010. We calculated 12 abundance and diversity metrics based on 5-year average abundances of 519 species for each of 768 monitoring routes. We did this for all bird species together as well as for four subgroups based on breeding habitat affinity (grassland, woodland, wetland, and shrubland breeders). The majority of the biodiversity metrics increased or remained constant over the study period, whereas the overall abundance of birds showed a pronounced decrease, primarily driven by declines of the most abundant species. These results highlight how stable or even increasing metrics of taxonomic, functional, or phylogenetic diversity may occur in parallel with substantial losses of individuals. We further found that patterns of change differed among the species subgroups, with both abundance and diversity increasing for woodland birds and decreasing for grassland breeders. The contrasting changes between abundance and diversity and among the breeding habitat groups underscore the relevance of a multifaceted approach to measuring biodiversity change. Our findings further stress the importance of monitoring the overall abundance of individuals in addition to metrics of taxonomic, functional, or phylogenetic diversity, thus confirming the importance of population abundance as an essential biodiversity variable.

Different foraging preferences of hummingbirds on artificial and natural flowers reveal mechanisms structuring plant-pollinator interactions.

In plant-pollinator networks, the floral morphology of food plants is an important determinant of the interaction niche of pollinators. Studies on foraging preferences of pollinators combining experimental and observational approaches may help to understand the mechanisms behind patterns of interactions and niche partitioning within pollinator communities. In this study, we tested whether morphological floral traits were associated with foraging preferences of hummingbirds for artificial and natural flower types in Costa Rica. We performed field experiments with artificial feeders, differing in length and curvature of flower types, to quantify the hummingbirds' interaction niche under unlimited nectar resources. To quantify the interaction niche under real-world conditions of limited nectar resources, we measured foraging preferences of hummingbirds for a total of 34 plant species. Artificial feeders were visited by Eupherusa nigriventris and Phaethornis guy in the pre-montane forest, and Lampornis calolaemus in the lower montane forest. Under experimental conditions, all three hummingbird species overlapped their interaction niches and showed a preference for the short artificial flower type over the long-straight and the long-curved flower types. Under natural conditions, the two co-occurring hummingbird species preferred to feed on plant species with floral traits corresponding to their bill morphology. The short-billed hummingbird E. nigriventris preferred to feed on short and straight flowers, whereas the long- and curved-billed P. guy preferred long and curved natural flowers. The medium-size billed species L. calolaemus preferred to feed on flowers of medium length and did not show preferences for plant species with specific corolla curvature. Our results show that floral morphological traits constrain access by short-billed hummingbird species to nectar resources. Morphological constraints, therefore, represent one important mechanism structuring trophic networks. In addition, other factors, such as competition and differences in resource quantity or quality, define the interaction niches of consumer species in real-world communities, enforcing patterns of niche segregation between co-occurring consumer species. This suggests that experimental studies are needed to disentangle effects of morphological constraints from those of competition for resources in plant-pollinator interactions and other types of trophic interactions.

Seed perishability determines the caching behaviour of a food-hoarding bird.

Many animals hoard seeds for later consumption and establish seed caches that are often located at sites with specific environmental characteristics. One explanation for the selection of non-random caching locations is the avoidance of pilferage by other animals. Another possible hypothesis is that animals choose locations that hamper the perishability of stored food, allowing the consumption of unspoiled food items over long time periods. We examined seed perishability and pilferage avoidance as potential drivers for caching behaviour of spotted nutcrackers (Nucifraga caryocatactes) in the Swiss Alps where the birds are specialized on caching seeds of Swiss stone pine (Pinus cembra). We used seedling establishment as an inverse measure of seed perishability, as established seedlings cannot longer be consumed by nutcrackers. We recorded the environmental conditions (i.e. canopy openness and soil moisture) of seed caching, seedling establishment and pilferage sites. Our results show that sites of seed caching and seedling establishment had opposed microenvironmental conditions. Canopy openness and soil moisture were negatively related to seed caching but positively related to seedling establishment, i.e. nutcrackers cached seeds preferentially at sites where seed perishability was low. We found no effects of environmental factors on cache pilferage, i.e. neither canopy openness nor soil moisture had significant effects on pilferage rates. We thus could not relate caching behaviour to pilferage avoidance. Our study highlights the importance of seed perishability as a mechanism for seed-caching behaviour, which should be considered in future studies. Our findings could have important implications for the regeneration of plants whose seeds are dispersed by seed-caching animals, as the potential of seedlings to establish may strongly decrease if animals cache seeds at sites that favour seed perishability rather than seedling establishment.

Ecological, historical and evolutionary determinants of modularity in weighted seed-dispersal networks.

Modularity is a recurrent and important property of bipartite ecological networks. Although well-resolved ecological networks describe interaction frequencies between species pairs, modularity of bipartite networks has been analysed only on the basis of binary presence-absence data. We employ a new algorithm to detect modularity in weighted bipartite networks in a global analysis of avian seed-dispersal networks. We define roles of species, such as connector values, for weighted and binary networks and associate them with avian species traits and phylogeny. The weighted, but not binary, analysis identified a positive relationship between climatic seasonality and modularity, whereas past climate stability and phylogenetic signal were only weakly related to modularity. Connector values were associated with foraging behaviour and were phylogenetically conserved. The weighted modularity analysis demonstrates the dominating impact of ecological factors on the structure of seed-dispersal networks, but also underscores the relevance of evolutionary history in shaping species roles in ecological communities.

A comparative analysis of dispersal syndromes in terrestrial and semi-terrestrial animals.

Dispersal, the behaviour ensuring gene flow, tends to covary with a number of morphological, ecological and behavioural traits. While species-specific dispersal behaviours are the product of each species' unique evolutionary history, there may be distinct interspecific patterns of covariation between dispersal and other traits ('dispersal syndromes') due to their shared evolutionary history or shared environments. Using dispersal, phylogeny and trait data for 15 terrestrial and semi-terrestrial animal Orders (> 700 species), we tested for the existence and consistency of dispersal syndromes across species. At this taxonomic scale, dispersal increased linearly with body size in omnivores, but decreased above a critical length in herbivores and carnivores. Species life history and ecology significantly influenced patterns of covariation, with higher phylogenetic signal of dispersal in aerial dispersers compared with ground dwellers and stronger evidence for dispersal syndromes in aerial dispersers and ectotherms, compared with ground dwellers and endotherms. Our results highlight the complex role of dispersal in the evolution of species life-history strategies: good dispersal ability was consistently associated with high fecundity and survival, and in aerial dispersers it was associated with early maturation. We discuss the consequences of these findings for species evolution and range shifts in response to future climate change.

Global variation in thermal tolerances and vulnerability of endotherms to climate change.

The relationships among species' physiological capacities and the geographical variation of ambient climate are of key importance to understanding the distribution of life on the Earth. Furthermore, predictions of how species will respond to climate change will profit from the explicit consideration of their physiological tolerances. The climatic variability hypothesis, which predicts that climatic tolerances are broader in more variable climates, provides an analytical framework for studying these relationships between physiology and biogeography. However, direct empirical support for the hypothesis is mostly lacking for endotherms, and few studies have tried to integrate physiological data into assessments of species' climatic vulnerability at the global scale. Here, we test the climatic variability hypothesis for endotherms, with a comprehensive dataset on thermal tolerances derived from physiological experiments, and use these data to assess the vulnerability of species to projected climate change. We find the expected relationship between thermal tolerance and ambient climatic variability in birds, but not in mammals-a contrast possibly resulting from different adaptation strategies to ambient climate via behaviour, morphology or physiology. We show that currently most of the species are experiencing ambient temperatures well within their tolerance limits and that in the future many species may be able to tolerate projected temperature increases across significant proportions of their distributions. However, our findings also underline the high vulnerability of tropical regions to changes in temperature and other threats of anthropogenic global changes. Our study demonstrates that a better understanding of the interplay among species' physiology and the geography of climate change will advance assessments of species' vulnerability to climate change.