Risks posed by invasive species to the provision of ecosystem services in Europe.

Invasive species significantly impact biodiversity and ecosystem services, yet understanding these effects at large spatial scales remains a challenge. Our study addresses this gap by assessing the current and potential future risks posed by 94 invasive species to seven key ecosystem services in Europe. We demonstrate widespread potential impacts, particularly on outdoor recreation, habitat maintenance, crop provisioning, and soil and nitrogen retention. Exposure to invasive species was higher in areas with lower provision of ecosystem services, particularly for regulating and cultural services. Exposure was also high in areas where ecosystem contributions to crop provision and nitrogen retention were at their highest. Notably, regions vital for ecosystem services currently have low invasion suitability, but face an average 77% increase in potential invasion area. Here we show that, while high-value ecosystem service areas at the highest risk represent a small fraction of Europe (0-13%), they are disproportionally important for service conservation. Our study underscores the importance of monitoring and protecting these hotspots to align management strategies with international biodiversity targets, considering both invasion vulnerability and ecosystem service sustainability.

The importance of migratory drop-off for island colonization in birds.

Seasonal migration is an underappreciated driver of animal diversification. Changes in migratory behaviour may favour the establishment of sedentary founder populations and promote speciation if there is sufficient reproductive isolation between sedentary and migratory populations. From a systematic literature review, we here quantify the role of migratory drop-off-the loss of migratory behaviour-in promoting speciation in birds on islands. We identify at least 157 independent colonization events likely initiated by migratory species that led to speciation, including 44 cases among recently extinct species. By comparing, for all islands, the proportion of island endemic species that derived from migratory drop-off with the proportion of migratory species among potential colonizers, we showed that seasonal migration has a larger effect on island endemic richness than direct dispersal. We also found that the role of migration in island colonization increases with the geographic isolation of islands. Furthermore, the success of speciation events depends in part on species biogeographic and ecological factors, here positively associated with greater range size and larger flock sizes. These results highlight the importance of shifts in migratory behaviour in the speciation process and calls for greater consideration of migratory drop-off in the biogeographic distribution of birds.

Body size and life history shape the historical biogeography of tetrapods.

Dispersal across biogeographic barriers is a key process determining global patterns of biodiversity as it allows lineages to colonize and diversify in new realms. Here we demonstrate that past biogeographic dispersal events often depended on species' traits, by analysing 7,009 tetrapod species in 56 clades. Biogeographic models incorporating body size or life history accrued more statistical support than trait-independent models in 91% of clades. In these clades, dispersal rates increased by 28-32% for lineages with traits favouring successful biogeographic dispersal. Differences between clades in the effect magnitude of life history on dispersal rates are linked to the strength and type of biogeographic barriers and intra-clade trait variability. In many cases, large body sizes and fast life histories facilitate dispersal success. However, species with small bodies and/or slow life histories, or those with average traits, have an advantage in a minority of clades. Body size-dispersal relationships were related to a clade's average body size and life history strategy. These results provide important new insight into how traits have shaped the historical biogeography of tetrapod lineages and may impact present-day and future biogeographic dispersal.

The EICAT+ framework enables classification of positive impacts of alien taxa on native biodiversity.

Species introduced through human-related activities beyond their native range, termed alien species, have various impacts worldwide. The IUCN Environmental Impact Classification for Alien Taxa (EICAT) is a global standard to assess negative impacts of alien species on native biodiversity. Alien species can also positively affect biodiversity (for instance, through food and habitat provisioning or dispersal facilitation) but there is currently no standardized and evidence-based system to classify positive impacts. We fill this gap by proposing EICAT+, which uses 5 semiquantitative scenarios to categorize the magnitude of positive impacts, and describes underlying mechanisms. EICAT+ can be applied to all alien taxa at different spatial and organizational scales. The application of EICAT+ expands our understanding of the consequences of biological invasions and can inform conservation decisions.

Global predictors of alien plant establishment success: combining niche and trait proxies.

Biological invasions are on the rise globally. To reduce future invasions, it is imperative to determine the naturalization potential of species. Until now, screening approaches have relied largely on species-specific functional feature data. Such information is, however, time-consuming and expensive to collect, thwarting the screening of large numbers of potential invaders. We propose to resolve such data limitations by developing indicators of establishment success of alien species that can be readily derived from open-access databases. These indicators describe key features of successfully established aliens, including estimates of potential range size, niche overlap with human-disturbed environments, and proxies of species traits related to their palaeoinvasions and local dominance capacities. We demonstrate the utility of this new approach by applying it to two large and highly invasive plant groups: Australian acacias and eucalypts. Our results show that these indicators robustly predict establishment successes and failures in each clade independently, and that they can cross-predict establishment in these two clades. Interestingly, the indicator identified as most important was species potential range size on Earth, a variable too rarely considered as a predictor. By successfully identifying key features that predispose Australian plants to naturalize, we provide an objective and cost-effective protocol for flagging high-risk introductions.

Long-distance migratory birds threatened by multiple independent risks from global change.

Many species migrate long distances annually between their breeding and wintering areas1. While global change affects both ranges, impact assessments have generally focused on breeding ranges and ignore how environmental changes influence migrants across geographic regions and the annual cycle2,3. Using range maps and species distribution models, we quantified the risk of summer and winter range loss and migration distance increase from future climate and land cover changes on long-distance migratory birds of the Holarctic (n=715). Risk estimates are largely independent of each other and magnitudes vary geographically. If seasonal range losses and increased migration distances are not considered, we strongly underestimate the number of threatened species by 18-49% and the overall magnitude of risk for 17-50% species. Many of the analysed species facing multiple global change risks are not listed by IUCN as threatened or near threatened. Neglecting seasonal migration in impact assessments could thus seriously misguide species' conservation.

Frequency and intensity of facilitation reveal opposing patterns along a stress gradient.

Disentangling the different processes structuring ecological communities is a long-standing challenge. In species-rich ecosystems, most emphasis has so far been given to environmental filtering and competition processes, while facilitative interactions between species remain insufficiently studied. Here, we propose an analysis framework that not only allows for identifying pairs of facilitating and facilitated species, but also estimates the strength of facilitation and its variation along environmental gradients. Our framework combines the analysis of both co-occurrence and co-abundance patterns using a moving window approach along environmental gradients to control for potentially confounding effects of environmental filtering in the co-abundance analysis. We first validate our new approach against community assembly simulations, and exemplify its potential on a large 1,134 plant community plots dataset. Our results generally show that facilitation intensity was strongest under cold stress, whereas the proportion of facilitating and facilitated species was higher under drought stress. Moreover, the functional distance between individual facilitated species and their facilitating species significantly changed along the temperature-moisture gradient, and seemed to influence facilitation intensity, although no general positive or general negative trend was discernible among species. The main advantages of our robust framework are as follows: It enables detecting facilitating and facilitated species in species-rich systems, and it allows identifying the directionality and intensity of facilitation in species pairs as well as its variation across long environmental gradients. It thus opens numerous opportunities for incorporating functional (and phylogenetic) information in the analysis of facilitation patterns. Our case study indicated high complexity in facilitative interactions across the stress gradient and revealed new evidence that facilitation, similarly to competition, can operate between functionally similar and dissimilar species. Extending the analyses to other taxa and ecosystems will foster our understanding how complex interspecific interactions promote biodiversity.

The effects of intransitive competition on coexistence.

Coexistence theory has been developed with an almost exclusive focus on interactions between two species, often ignoring more complex and indirect interactions, such as intransitive loops, that can emerge in competition networks. In fact, intransitive competition has typically been studied in isolation from other pairwise stabilising processes, and thus little is known about how intransitivity interacts with more traditional drivers of species coexistence such as niche partitioning. To integrate intransitivity into traditional coexistence theory, we developed a metric of growth rate when rare, Δri¯, to identify and quantify the impact of intransitive competition against a backdrop of pairwise stabilising niche differences. Using this index with simulations of community dynamics, we demonstrate that intransitive loops can both stabilise or destabilise species coexistence, but the strength and importance of intransitive interactions are significantly affected by the length and the topology of these loops. We conclude by showing how Δri¯ can be used to evaluate effects of intransitivity in empirical studies. Our results emphasise the need to integrate complex mechanisms emerging from diverse interactions into our understanding of species coexistence.

Where less may be more: how the rare biosphere pulls ecosystems strings.

Rare species are increasingly recognized as crucial, yet vulnerable components of Earth's ecosystems. This is also true for microbial communities, which are typically composed of a high number of relatively rare species. Recent studies have demonstrated that rare species can have an over-proportional role in biogeochemical cycles and may be a hidden driver of microbiome function. In this review, we provide an ecological overview of the rare microbial biosphere, including causes of rarity and the impacts of rare species on ecosystem functioning. We discuss how rare species can have a preponderant role for local biodiversity and species turnover with rarity potentially bound to phylogenetically conserved features. Rare microbes may therefore be overlooked keystone species regulating the functioning of host-associated, terrestrial and aquatic environments. We conclude this review with recommendations to guide scientists interested in investigating this rapidly emerging research area.

Is There Any Evidence for Rapid, Genetically-Based, Climatic Niche Expansion in the Invasive Common Ragweed?

Climatic niche shifts have been documented in a number of invasive species by comparing the native and adventive climatic ranges in which they occur. However, these shifts likely represent changes in the realized climatic niches of invasive species, and may not necessarily be driven by genetic changes in climatic affinities. Until now the role of rapid niche evolution in the spread of invasive species remains a challenging issue with conflicting results. Here, we document a likely genetically-based climatic niche expansion of an annual plant invader, the common ragweed (Ambrosia artemisiifolia L.), a highly allergenic invasive species causing substantial public health issues. To do so, we looked for recent evolutionary change at the upward migration front of its adventive range in the French Alps. Based on species climatic niche models estimated at both global and regional scales we stratified our sampling design to adequately capture the species niche, and localized populations suspected of niche expansion. Using a combination of species niche modeling, landscape genetics models and common garden measurements, we then related the species genetic structure and its phenotypic architecture across the climatic niche. Our results strongly suggest that the common ragweed is rapidly adapting to local climatic conditions at its invasion front and that it currently expands its niche toward colder and formerly unsuitable climates in the French Alps (i.e. in sites where niche models would not predict its occurrence). Such results, showing that species climatic niches can evolve on very short time scales, have important implications for predictive models of biological invasions that do not account for evolutionary processes.

Is invasion success of Australian trees mediated by their native biogeography, phylogenetic history, or both?

For a plant species to become invasive it has to progress along the introduction-naturalization-invasion (INI) continuum which reflects the joint direction of niche breadth. Identification of traits that correlate with and drive species invasiveness along the continuum is a major focus of invasion biology. If invasiveness is underlain by heritable traits, and if such traits are phylogenetically conserved, then we would expect non-native species with different introduction status (i.e. position along the INI continuum) to show phylogenetic signal. This study uses two clades that contain a large number of invasive tree species from the genera Acacia and Eucalyptus to test whether geographic distribution and a novel phylogenetic conservation method can predict which species have been introduced, became naturalized, and invasive. Our results suggest that no underlying phylogenetic signal underlie the introduction status for both groups of trees, except for introduced acacias. The more invasive acacia clade contains invasive species that have smoother geographic distributions and are more marginal in the phylogenetic network. The less invasive eucalyptus group contains invasive species that are more clustered geographically, more centrally located in the phylogenetic network and have phylogenetic distances between invasive and non-invasive species that are trending toward the mean pairwise distance. This suggests that highly invasive groups may be identified because they have invasive species with smoother and faster expanding native distributions and are located more to the edges of phylogenetic networks than less invasive groups.

Does the legacy of historical biogeography shape current invasiveness in pines?

Why are some introduced species more successful at establishing and spreading than others? Until now, characteristics of extant species have been intensively investigated to answer this question. We propose to gain new insights on species invasiveness by exploring the long-term biogeographic and evolutionary history of lineages. We exemplify our approach using one of the best-studied invasive plant genera, Pinus. We notably estimated the historical biogeography of pines and the rates of trait evolution in pines. These estimates were analysed with regard to species invasiveness status. The results revealed that currently invasive species belong to lineages that were particularly successful at colonizing new regions in the past. We also showed that highly mobile lineages had faster rates of niche evolution, but that these rates are poor proxies for species adaptive potential in invaded regions (estimated by niche shift among native and invaded regions). In summary, working at the interface of ecology, historical biogeography and evolutionary history offers stimulating perspectives to improve our understanding of the drivers of invasion success.

Interaction between two invasive organisms on the European chestnut: does the chestnut blight fungus benefit from the presence of the gall wasp?

The impact of invasive fungal pathogens and pests on trees is often studied individually, thereby omitting possible interactions. In this study the ecological interaction between the chestnut blight fungus Cryphonectria parasitica and the chestnut gall wasp Dryocosmus kuriphilus was investigated. We determined if abandoned galls could be colonized by C. parasitica and thereby act as an entry point and a source of pathogen inoculum. Moreover we assessed the identity and diversity of other gall-colonizing fungal species. A total of 1973 galls were randomly sampled from 200 chestnut trees in eight Swiss stands. In a stand C. parasitica was isolated from 0.4-19.2% of the galls. The incidence of C. parasitica on the galls and the fungal diversity significantly increased with the residence time of D. kuriphilus in a stand. All but one C. parasitica cultures were virulent. The predominant fungus isolated from galls was Gnomoniopsis castanea whose abundance influenced negatively that of C. parasitica. This study shows that D. kuriphilus galls can be colonized by virulent strains of the chestnut blight fungus C. parasitica. This can have effects on the chestnut blight incidence even in chestnut stands where the disease is successfully controlled by hypovirulence. The gall wasp presence influences also the fungal species composition on chestnut trees.

Contrasting the effects of environment, dispersal and biotic interactions to explain the distribution of invasive plants in alpine communities.

Despite considerable efforts devoted to investigate the community assembly processes driving plant invasions, few general conclusions have been drawn so far. Three main processes, generally acting as successive filters, are thought to be of prime importance. The invader has to disperse (1st filter) into a suitable environment (2nd filter) and succeed in establishing in recipient communities through competitive interactions (3rd filter) using two strategies: competition avoidance by the use of different resources (resource opportunity), or competitive exclusion of native species. Surprisingly, despite the general consensus on the importance of investigating these three processes and their interplay, they are usually studied independently. Here we aim to analyse these three filters together, by including them all: abiotic environment, dispersal and biotic interactions, into models of invasive species distributions. We first propose a suite of indices (based on species functional dissimilarities) supposed to reflect the two competitive strategies (resource opportunity and competition exclusion). Then, we use a set of generalised linear models to explain the distribution of seven herbaceous invaders in natural communities (using a large vegetation database for the French Alps containing 5,000 community-plots). Finally, we measure the relative importance of competitive interaction indices, identify the type of coexistence mechanism involved and how this varies along environmental gradients. Adding competition indices significantly improved model's performance, but neither resource opportunity nor competitive exclusion were common strategies among the seven species. Overall, we show that combining environmental, dispersal and biotic information to model invasions has excellent potential for improving our understanding of invader success.

Scale decisions can reverse conclusions on community assembly processes.

AIM: Phylogenetic diversity patterns are increasingly being used to better understand the role of ecological and evolutionary processes in community assembly. Here, we quantify how these patterns are influenced by scale choices in terms of spatial and environmental extent and organismic scales. LOCATION: European Alps. METHODS: We applied 42 sampling strategies differing in their combination of focal scales. For each resulting sub-dataset, we estimated the phylogenetic diversity of the species pools, phylogenetic α-diversities of local communities, and statistics commonly used together with null models in order to infer non-random diversity patterns (i.e. phylogenetic clustering versus over-dispersion). Finally, we studied the effects of scale choices on these measures using regression analyses. RESULTS: Scale choices were decisive for revealing signals in diversity patterns. Notably, changes in focal scales sometimes reversed a pattern of over-dispersion into clustering. Organismic scale had a stronger effect than spatial and environmental extent. However, we did not find general rules for the direction of change from over-dispersion to clustering with changing scales. Importantly, these scale issues had only a weak influence when focusing on regional diversity patterns that change along abiotic gradients. MAIN CONCLUSIONS: Our results call for caution when combining phylogenetic data with distributional data to study how and why communities differ from random expectations of phylogenetic relatedness. These analyses seem to be robust when the focus is on relating community diversity patterns to variation in habitat conditions, such as abiotic gradients. However, if the focus is on identifying relevant assembly rules for local communities, the uncertainty arising from a certain scale choice can be immense. In the latter case, it becomes necessary to test whether emerging patterns are robust to alternative scale choices.

Darwin's naturalization hypothesis: scale matters in coastal plant communities.

Darwin proposed two seemingly contradictory hypotheses for a better understanding of biological invasions. Strong relatedness of invaders to native communities as an indication of niche overlap could promote naturalization because of appropriate niche adaptation, but could also hamper naturalization because of negative interactions with native species ('Darwin's naturalization hypothesis'). Although these hypotheses provide clear and opposing predictions for expected patterns of species relatedness in invaded communities, so far no study has been able to clearly disentangle the underlying mechanisms. We hypothesize that conflicting past results are mainly due to the neglected role of spatial resolution of the community sampling. In this study, we corroborate both of Darwin's expectations by using phylogenetic relatedness as a measure of niche overlap and by testing the effects of sampling resolution in highly invaded coastal plant communities. At spatial resolutions fine enough to detect signatures of biotic interactions, we find that most invaders are less related to their nearest relative in invaded plant communities than expected by chance (phylogenetic overdispersion). Yet at coarser spatial resolutions, native assemblages become more invasible for closely-related species as a consequence of habitat filtering (phylogenetic clustering). Recognition of the importance of the spatial resolution at which communities are studied allows apparently contrasting theoretical and empirical results to be reconciled. Our study opens new perspectives on how to better detect, differentiate and understand the impact of negative biotic interactions and habitat filtering on the ability of invaders to establish in native communities.