Using species distribution models for IUCN Red Lists of threatened species

Red Lists have been used for years globally and regionally in many countries to highlight species that need special attention because of the rarity or rapid decline of their populations. To ensure homogenized classification at the global and regional level, the International Union for Conservation of Nature (IUCN) defined categories of threat, and criteria to attribute the taxa to these categories. Nevertheless, the strict application of the criteria is not always straightforward, especially for invertebrates, because of the difficulties associated with precise estimates of the size and viability of their populations. This paper presents a method for the estimation of extent of occurrence (EOO) and area of occupancy (AOO) based on species distribution models using multivariate adaptive regression splines. To achieve this, presence data have been modeled against topographical and climatic explanatory variables. Predictions from the statistical distribution models have then been cut using the minimal convex hull around (EOO) or the watersheds in which (AOO) the species have really been observed in recent years. This allows us to delimit the EOO and AOO according to the IUCN criteria, and better take into account the ecological requirements of the species. Furthermore, the method allows for the use of historical data (e.g. from museum’s collections) and the direct comparison of historical and recent distributions of species. The method has been tested on six species of butterflies. The results show the possibility of using species distribution models to define the Red Lists status according to the IUCN guidelines, and shows that the results are consistent with previous Red Lists assessments.     

Risk of extinction of plant communities: Risk and assessment categories

There are many proposals for the assessment of plant communities, based on different criteria, but very few proposals for categorization of the risk of extinction of plant communities (syntaxa). In this paper, concepts related to extent of occurence (EOO), area of occupancy (AOO), extinction and regeneration of plant communities are defined. Also, we propose and define several ranks of extinction risk based on quantitative criteria of the EOO, AOO and processes of decline. The proposals are global, for application to any type of cormophytic vegetation, without geographical restrictions. To check the adequacy of the proposed ranks and its thresholds, the method has been applied to the vegetation of the Iberian Peninsula based on the EOO (122 plant communities) and the AOO (2224 plant communities).     

Least concern to endangered: Applying climate change projections profoundly influences the extinction risk assessment for wild Arabica coffee

Arabica coffee (Coffea arabica) is a key crop in many tropical countries and globally provides an export value of over US$13 billion per year. Wild Arabica coffee is of fundamental importance for the global coffee sector and of direct importance within Ethiopia, as a source of harvestable income and planting stock. Published studies show that climate change is projected to have a substantial negative influence on the current suitable growing areas for indigenous Arabica in Ethiopia and South Sudan. Here we use all available future projections for the species based on multiple general circulation models (GCMs), emission scenarios, and migration scenarios, to predict changes in Extent of Occurrence (EOO), Area of Occupancy (AOO), and population numbers for wild Arabica coffee. Under climate change our results show that population numbers could reduce by 50% or more (with a few models showing over 80%) by 2088. EOO and AOO are projected to decline by around 30% in many cases. Furthermore, present‐day models compared to the near future (2038), show a reduction for EOO of over 40% (with a few cases over 50%), although EOO should be treated with caution due to its sensitivity to outlying occurrences. When applying these metrics to extinction risk, we show that the determination of generation length is critical. When applying the International Union for Conservation of Nature's Red list of Threatened Species (IUCN Red List) criteria, even with a very conservative generation length of 21 years, wild Arabica coffee is assessed as Threatened with extinction (placed in the Endangered category) under a broad range of climate change projections, if no interventions are made. Importantly, if we do not include climate change in our assessment, Arabica coffee is assessed as Least Concern (not threatened) when applying the IUCN Red List criteria.     

The use of spatial ecological modelling as a tool for improving the assessment of geographic range size of threatened species

We analysed endemic threatened tree and reptile species of Socotra Island (Yemen), characterised by different ecological requirements and spatial distribution, in order to evaluate the usefulness of spatial ecological modelling in the estimation of species extent of occurrence (EOO) and area of occupancy (AOO). Point occurrences for the entire species range were used to model their spatial distribution by Random Forest (RF) and Generalised Linear Model (GLM). For each species the suitability area (SA) was obtained by applying the 0% omission error criterion on the probability map, and compared or integrated with EOO and AOO area obtained by topological methods such as the minimum convex polygon (MCP), α-hull and 2 km × 2 km grid.RF showed a lower prediction error than GLM. Higher accuracy was achieved for species with higher number of occurrences and narrower ecological niche. SA was always greater than AOO measured with the 2 km × 2 km grid method. SA was greater than EOO, measured by both MCP and α-hull methods, for species with localised distribution, while it was smaller for widely distributed species. EOO-α-hull area was equal or smaller than that calculated by MCP depending on the spatial distribution of species. AOO measured considering the SA within the EOO-MCP was greater than that measured using the standard 2 km × 2 km grid. Conversely, AOO calculated considering the suitable area within the EOO-α-hull showed variable results, being smaller or greater than the 2 km × 2 km grid AOO depending on the ecological niche and spatial distribution of species. According to our results, SEM does not provide an effective alternative to topological methods for the estimate of EOO and AOO. However, it may be considered a useful tool to estimate AOO within the boundaries of EOO measured by the α-hull method, because it reduces some potential sources of inconsistency and bias.     

Supporting Red List threat assessments with GeoCAT: geospatial conservation assessment tool

GeoCAT is an open source, browser based tool that performs rapid geospatial analysis to ease the process of Red Listing taxa. Developed to utilise spatially referenced primary occurrence data, the analysis focuses on two aspects of the geographic range of a taxon: the extent of occurrence (EOO) and the area of occupancy (AOO). These metrics form part of the IUCN Red List categories and criteria and have often proved challenging to obtain in an accurate, consistent and repeatable way. Within a familiar Google Maps environment, GeoCAT users can quickly and easily combine data from multiple sources such as GBIF, Flickr and Scratchpads as well as user generated occurrence data. Analysis is done with the click of a button and is visualised instantly, providing an indication of the Red List threat rating, subject to meeting the full requirements of the criteria. Outputs including the results, data and parameters used for analysis are stored in a GeoCAT file that can be easily reloaded or shared with collaborators. GeoCAT is a first step toward automating the data handling process of Red List assessing and provides a valuable hub from which further developments and enhancements can be spawned.     

Spatial ecology and conservation of the microendemic ovenbird Cipo Cinclodes (Cinclodes espinhacensis) from the Brazilian highlands

Birds in the genus Cinclodes are habitat specialists, with most restricted to the highlands of South America. The recently described Cipo Cinclodes (C. espinhacensis) is isolated in the southern Espinhaço Range of Brazil and is considered Endangered in Brazil and Near Threatened by the IUCN, but as a subspecies of Long‐tailed Cinclodes (C. pabsti). We examined the population and spatial ecology of Cipo Cinclodes at two geographic scales to improve our understanding of their basic biology and conservation status. We monitored 30 birds at Serra do Breu and found relatively large home ranges (mean = 9.3 ha), a density of paired adults of 0.09/ha, a male‐skewed adult sex ratio (males/total adults = 0.57) due to territories occupied by unpaired males, and long‐term site fidelity. Cipo Cinclodes used all habitat types available in our study area, including rocky outcrops, grasslands, and riparian areas, but habitat selection analyses revealed the importance of riparian areas for foraging and rocky outcrops for nesting. At the species distribution scale, we compiled known and novel recorded occurrence points and used them to calculate the extent of occurrence (EOO) and the area of occupancy (AOO). We used a Maxent species distribution model to generate a binary map to estimate upper limits for EOO (EOO around the model predicted area) and AOO (comprised by the model predicted area within the EOO). We obtained 41 locations, resulting in an EOO of 890.7 km² (up to 1748.7 km²) and an AOO of 100 km² (up to 327.5 km²). The global population is estimated to be between 880 and 2882 birds, which is concerning because small populations are at risk of extinction due to demographic stochasticity, genetic drift, and the interaction of these factors. As such, our results support the designation of Cipo Cinclodes as Endangered on the Brazilian red list.     

Cross-scale management strategies for optimal control of trees invading from source plantations

Biological invasion by non-native tree species can transform landscapes, and as a consequence, has received growing attention from researchers and managers alike. This problem is driven primarily by the naturalisation and invasion of tree species escaping from cultivation or forestry plantations. Furthermore, these invasions can be strongly influenced by the land-use matrix of the surrounding region, specific management of the source populations, and environmental conditions that influence seed dispersal or habitat quality for the invader. A major unresolved challenge for managing tree invasions in landscapes is how management should be deployed to contain or slow the spread of invading populations from one or more sources (e.g. plantations). We develop a spatial simulation model to test: (1) how to best prioritise the control of invasive tree populations spatially to slow or contain the biological invader when habitat quality varies in the landscape, and (2) how to allocate control effort among different management units when trees spread from many source populations. We first show that to slow down spread effectively, management strategy is less important than management effort. We then identify the conditions affecting the relative performance of different management strategies. At the landscape scale, targeting peripheral stands consistently yielded the best results whereas at the regional scale, management strategies needed to account for both habitat quality and tree life-history. Overall, our findings demonstrate that knowledge of how habitat affects tree life-history stages can improve management to contain or slow tree invasions by improving the spatial match between management effort and efficacy.     

Global networks for invasion science: benefits,challenges and guidelines

Much has been done to address the challenges of biological invasions, but fundamental questions (e.g., which species invade? Which habitats are invaded? How can invasions be effectively managed?) still need to be answered before the spread and impact of alien taxa can be effectively managed. Questions on the role of biogeography (e.g., how does biogeography influence ecosystem susceptibility, resistance and resilience against invasion?) have the greatest potential to address this goal by increasing our capacity to understand and accurately predict invasions at local, continental and global scales. This paper proposes a framework for the development of ‘Global Networks for Invasion Science’ to help generate approaches to address these critical and fundamentally biogeographic questions. We define global networks on the basis of their focus on research questions at the global scale, collection of primary data, use of standardized protocols and metrics, and commitment to long-term global data. Global networks are critical for the future of invasion science because of their potential to extend beyond the capacity of individual partners to identify global priorities for research agendas and coordinate data collection over space and time, assess risks and emerging trends, understand the complex influences of biogeography on mechanisms of invasion, predict the future of invasion dynamics, and use these new insights to improve the efficiency and effectiveness of evidence-based management techniques. While the pace and scale of global change continues to escalate, strategic and collaborative global networks offer a powerful approach to inform responses to the threats posed by biological invasions.     

Elusive species distribution modelling: The case of Natrix natrix cypriaca

Improving knowledge on species distribution is a fundamental part of ecological research and imperative when it comes to species conservation and management. For rare, endemic, and elusive species the effort to obtain reliable ecological data is extremely challenging. Data sparsity, or the use of outdated data, for species that are in the greatest need of protection, hampers the ability to develop efficient conservation and management schemes.In the present research, we use SDMs to examine the predictive distribution of elusive species exemplified by the Cyprus grass snake (Natrix natrix cypriaca). We use two different methods (i.e. Maxent and Ensemble) and three different sets of environmental envelopes (i.e. bioclimatic, biophysical, combined) for predicting species distribution. We compare the results of each model with the Extent of Occurrence (EOO) and Area of Occupancy (AOO) of the species and map high potential occurrence areas for future survey efforts indicating areas of interest outside the species previously known distribution.This research is the first effort to predict and map the potential distribution of this elusive species. The results have significantly improved past estimations clearly indicating a geographic range larger and wider that what was previously thought, providing new perspectives on the species ecology and conservation. This study demonstrates that SDMs can be employed successfully for elusive species with limited current localities providing insights on their distribution guiding monitoring schemes and conservation actions.     

ConR: An R package to assist large‐scale multispecies preliminary conservation assessments using distribution data

The Red List Categories and the accompanying five criteria developed by the International Union for Conservation of Nature (IUCN) provide an authoritative and comprehensive methodology to assess the conservation status of organisms. Red List criterion B, which principally uses distribution data, is the most widely used to assess conservation status, particularly of plant species. No software package has previously been available to perform large‐scale multispecies calculations of the three main criterion B parameters [extent of occurrence (EOO), area of occupancy (AOO) and an estimate of the number of locations] and provide preliminary conservation assessments using an automated batch process. We developed ConR, a dedicated R package, as a rapid and efficient tool to conduct large numbers of preliminary assessments, thereby facilitating complete Red List assessment. ConR (1) calculates key geographic range parameters (AOO and EOO) and estimates the number of locations sensu IUCN needed for an assessment under criterion B; (2) uses this information in a batch process to generate preliminary assessments of multiple species; (3) summarize the parameters and preliminary assessments in a spreadsheet; and (4) provides a visualization of the results by generating maps suitable for the submission of full assessments to the IUCN Red List. ConR can be used for any living organism for which reliable georeferenced distribution data are available. As distributional data for taxa become increasingly available via large open access datasets, ConR provides a novel, timely tool to guide and accelerate the work of the conservation and taxonomic communities by enabling practitioners to conduct preliminary assessments simultaneously for hundreds or even thousands of species in an efficient and time‐saving way.     

Mapping of the Asian longhorned beetle’s time to maturity and risk to invasion at contiguous United States extent

Anoplophora glabripennis, the Asian Longhorned Beetle (ALB), is an invasive species of high economic and ecological relevance given the potential it has to cause tree damage, and sometimes mortality, in the United States. Because this pest is introduced by transport in wood-packing products from Asia, ongoing trade activities pose continuous risk of transport and opportunities for introduction. Therefore, a geographic understanding of the spatial distribution of risk factors associated with ALB invasions is needed. Chief among the multiple risk factors are (a) the potential for infestation based on host tree species presence/absence, and (b) the temperature regime as a determinant of ALB’s growth time to maturity. This study uses an empirical model of ALB’s time to maturity as a function of temperature, along with a model of heat transfer in the wood of the host and spatial data describing host species presence/absence data, to produce a map of risk factors across the conterminous United States to define potential for ALB infestation and relative threat of impact. Results show that the region with greatest risk of ALB infestation is the eastern half of the country, with lower risk across most of the western half due to low abundance of host species, less urban area, and prevalence of cold, high elevations. Risk is high in southeastern states primarily because of temperature, while risk is high in northeastern and northern central states because of high abundance of host species.     

Incentivizing the Public to Support Invasive Species Management: Eurasian Milfoil Reduces Lakefront Property Values

Economic evaluations of invasive species are essential for providing comprehensive assessments of the benefits and costs of publicly-funded management activities, yet many previous investigations have focused narrowly on expenditures to control spread and infestation. We use hedonic modeling to evaluate the economic effects of Eurasian milfoil (Myriophyllum spicatum) invasions on lakefront property values of single-family homes in an urban-suburban landscape. Milfoil often forms dense canopies at the water surface, diminishing the value of ecosystem services (e.g., recreation, fishing) and necessitating expensive control and management efforts. We compare 1,258 lakeshore property sale transactions (1995–2006) in 17 lakes with milfoil and 24 un-invaded lakes in King County, Washington (USA). After accounting for structural (e.g., house size), locational (e.g., boat launch), and environmental characteristics (e.g., water clarity) of lakes, we found that milfoil has a significant negative effect on property sales price ($94,385 USD lower price), corresponding to a 19% decline in mean property values. The aggregate cost of milfoil invading one additional lake in the study area is, on average, $377,542 USD per year. Our study illustrates that invasive aquatic plants can significantly impact property values (and associated losses in property taxes that reduce local government revenue), justifying the need for management strategies that prevent and control invasions. We recommend coordinated efforts across Lake Management Districts to focus institutional support, funding, and outreach to prevent the introduction and spread of milfoil. This effort will limit opportunities for re-introduction from neighboring lakes and incentivize private landowners and natural resource agencies to commit time and funding to invasive species management.     

Effects of a Major Tree Invader on Urban Woodland Arthropods

Biological invasions are a major threat to biodiversity; however, the degree of impact can vary depending on the ecosystem and taxa. Here, we test whether a top invader at a global scale, the tree Robinia pseudoacacia (black locust or false acacia), which is known to profoundly change site conditions, significantly affects urban animal diversity. As a first multi-taxon study of this kind, we analyzed the effects of Robinia dominance on 18 arthropod taxa by pairwise comparisons of woodlands in Berlin, Germany, that were dominated by R. pseudoacacia or the native pioneer tree Betula pendula. As a negative effect, abundances of five arthropod taxa decreased (Chilopoda, Formicidae, Diptera, Heteroptera, Hymenoptera); 13 others were not affected. Woodland type affected species composition of carabids and functional groups in spiders, but surprisingly did not decrease alpha and beta diversity of carabid and spider assemblages or the number of endangered species. Tree invasion thus did not induce biotic homogenization at the habitat scale. We detected no positive effects of alien dominance. Our results illustrate that invasions by a major tree invader can induce species turnover in ground-dwelling arthropods, but do not necessarily reduce arthropod species abundances or diversity and might thus contribute to the conservation of epigeal invertebrates in urban settings. Considering the context of invasion impacts thus helps to set priorities in managing biological invasions and can illustrate the potential of novel ecosystems to maintain urban biodiversity.     

The risk of invasion by angiosperms peaks at intermediate levels of human influence

Biological invasions are a growing threat to biodiversity. The control and eradication of exotic species established in earnest are of limited success despite high financial investments. Anticipating biological invasions based on species’ suitabilities is a cost-effective strategy given it helps identifying areas where exotic species can prosper, which can then translate in improving management and conservation efforts. Based on information from 191 invasive angiosperm species worldwide, we used ecological niche models to identify areas at high risk of invasion (cumulative predicted distribution of invasive species) in Mexico. Further, we explored the importance of bioclimatic and human influence variables as drivers of the distribution of invasive species and analyzed the status of the currently recognized priority conservation sites in Mexico. We found that areas with intermediate human activity scores had a high risk of invasion. Additionally, we found that many of the current priority conservation sites in Mexico had a high risk of invasion. Our findings contribute to disentangling the factors that drive environment susceptibility to invasions and urge management strategies to minimize the impacts of biological invasions in priority conservation sites.     

Current distribution and potential extent of the most invasive alien plant species on La Réunion (Indian Ocean,Mascarene islands)

Abstract La Réunion Island has the largest area of intact vegetation of the islands in the Mascarene archipelago. Biological invasions are the primary threat to biodiversity in the intact habitats of the island (those not already transformed by agriculture and urbanization). Our study aimed to identify areas to prioritize in managing invasive alien plants for biodiversity conservation. We used extensive surveys of 238 distinct untransformed areas on La Réunion to define the current distribution patterns of all invasive species. Using expert knowledge, we compiled maps of the current distribution of the 46 most widespread/important invasive plants at the habitat scale (identified according to vegetation structure). Data from 440 botanical relevés for the 20 most threatening invasive alien plant species across the island and climatic envelope models were used to derive climatic suitability surfaces; these were used to map potential distributions for these species. More than 10 species invade 16.7% of the remaining habitat. Five habitat types are invaded by 25 or more species, and eight have fewer than 10 invasive alien plant species. Cluster analysis based on presence/absence of species in the 18 habitat types produced eight groups of species that invade particular habitats. Potential distribution models show that some species have invaded large parts of their potential range (e.g. Fuchsia magellanica, Furcraea foetida, Hiptage benghalensis), whereas others have the potential to increase their range substantially (e.g. Clidemia hirta, Strobilanthes hamiltonianus, Ulex europaeus). Management implications are identified for both groups. Three broad groups of habitats were identified: (i) intact habitats with a low level of invasion (e.g. subalpine shrubland); (ii) moderately invaded habitats with varying levels of intactness (ranging from windward submountain rainforest to the Acacia heterophylla forest); and (iii) habitats with little remaining intact area and high levels of invasion (e.g. lowland rainforest). Different management interventions are appropriate for these three groups.     

Conservation study of endemic plants in serpentine landscapes: Antirrhinum rothmaleri (Plantaginaceae), a serpentinophyte with a restricted geographic distribution

Abstract

Assessing the conservation status of endemic species by monitoring population trends is one of the main aims of plant conservation studies. This article reports a detailed study of the distribution of Antirrhinum rothmaleri, a species endemic to the serpentine systems of Trás-os-Montes in NE Portugal (Lusitan-Duriensean biogeographical sector). The species' current distribution status, the size of its populations, and the threats it faces, measured according to International Union for Conservation of Nature (IUCN) 2008 Guidelines, were all assessed. The size estimates provided for the four confirmed populations were based on census data taken at the different sites. All censuses were performed by direct counting of all potentially reproductive individuals. Based on the present data, A. rothmaleri should be assigned a conservation status of “Critically Endangered” (CR). Its conservation status, according to the IUCN criteria, is mainly based on the narrow distribution, and on the area of occupancy (AOO) and extent of occurrence (EOO). The risk of extinction faced by the local populations is high due to their typically very small size. Suitable conservation strategies should be developed in order to preserve the species. The present study allowed us to characterize its distribution and population size, to collect and store ex situ the germplasm in the UIRGEMP/Banco Português de Germoplasma Vegetal, and to investigate its ecology. The conservation status of the species requires continued monitoring of the demography of its populations. Studies on the biology of the species are also needed in order to improve its management. The present results illustrate the need to protect serpentine landscapes, which contain a range of microhabitats inhabited byendemic plants.     

Developing and testing alien species indicators for Europe

Alien species indicators provide vital information to the biodiversity policy sector on the status-quo and trends of biological invasions and on the efficacy of response measures. Applicable at different geographical scales and organizational levels, alien species indicators struggle with data availability and quality. Based on policy needs and previous work on the global scale, we here present a set of six alien species indicators for Europe, which capture complementary facets of biological invasions in Europe: (a) an combined index of invasion trends, (b) an indicator on pathways of invasions, (c) the Red List Index of Invasive Alien Species (IAS), (d) an indicator of IAS impacts on ecosystem services, (e) trends in incidence of livestock diseases and (f) an indicator on costs for alien species management and research. Each of these indicators has its particular strengths and shortcomings, but combined they allow for a nuanced understanding of the status and trends of biological invasions in Europe. We found that the scale and impact of biological invasions are steadily increasing across all impact indicators, although societal response in recent years has increased. The Red List Index is fit-for-purpose and demonstrates that overall extinction risks (here shown for amphibians in Europe) are increasing. Introduction pathway dynamics have changed, with some pathways decreasing in relevance (e.g., biological control agents) and others increasing (e.g., horticultural trade) providing a leverage for targeted policy and stakeholder response. The IAS indicators presented here for the first time on a continental basis serve as a starting point for future improvements, and as a basis for monitoring the efficacy of the recent EU legislation of IAS. This will need a better workflow for data collection and management. To achieve this, all main actors must work toward improving the interoperability among existing databases and between data holders.     

Alien plant invasions in tropical and sub-tropical savannas: patterns,processes and prospects

Biological invasions affect virtually all ecosystems on earth, but the degree to which different regions and biomes are invaded, and the quality of information from different regions, varies greatly. A large body of literature exists on the invasion of savannas in the Neotropics and northern Australia where invasive plants, especially African grasses, have had major impacts. Less has been published on plant invasions in African savannas, except for those in South Africa. Negative impacts due to plant invasions in African savannas appear to be less severe than in other regions at present. As savannas cover about 60% of the continent, with tens of millions of people relying on the services they provide, it is timely to assess the current status of invasions as a threat to these ecosystems. We reviewed the literature, contrasting the African situation with that of Neotropical and Australian savannas. A number of drivers and explanatory factors of plant invasions in savannas have been described, mostly from the Neotropics and Australia. These include herbivore presence, residence time, intentional introductions for pasture improvements, fire regimes, the physiology of the introduced species, and anthropogenic disturbance. After comparing these drivers across the three regions, we suggest that the lower extent of alien plant invasions in African savannas is largely attributable to: (1) significantly lower rates of intentional plant introductions and widespread plantings (until recently); (2) the role of large mammalian herbivores in these ecosystems; (3) historical and biogeographical issues relating to the regions of origin of introduced species; and (4) the adaptation of African systems to fire. We discuss how changing conditions in the three regions are likely to affect plant invasions in the future.     

Rapid evolution in introduced species, 'invasive traits' and recipient communities: challenges for predicting invasive potential

The damaging effects of invasive organisms have triggered the development of Invasive Species Predictive Schemes (ISPS). These schemes evaluate biological and historical characteristics of species and prioritize those that should be the focus of exclusion, quarantine, and/or control. However, it is not clear how commonly these schemes take microevolutionary considerations into account. We review the recent literature and find that rapid evolutionary changes are common during invasions. These evolutionary changes include rapid adaptation of invaders to new environments, effects of hybridization, and evolution in recipient communities. Strikingly, we document 38 species in which the specific traits commonly associated with invasive potential (e.g. growth rate, dispersal ability, generation time) have themselves undergone evolutionary change following introduction, in some cases over very short (≤ 10 year) timescales. In contrast, our review of 29 ISPS spanning plant, animal, and microbial taxa shows that the majority (76%) envision invading species and recipient communities as static entities. Those that incorporate evolutionary considerations do so in a limited way. Evolutionary change not only affects the predictive power of these schemes, but also complicates their evaluation. We argue that including the evolutionary potential of species and communities in ISPS is overdue, present several metrics related to evolutionary potential that could be incorporated in ISPS, and provide suggestions for further research on these metrics and their performance. Finally, we argue that the fact of evolutionary change during invasions begs for added caution during risk assessment.     

Tree invasions: a comparative test of the dominant hypotheses and functional traits

Trees act as ecosystem engineers and invasions by exotic tree species profoundly impact recipient communities. Recently, research on invasive trees has dramatically increased, enabling the assessment of general trends in tree invasion. Analysing 90 studies dealing with 45 invasive tree species, we conducted a quantitative review and a meta-analysis to estimate the relevance of eight leading hypotheses for explaining tree invasions. We also tested whether species functional traits (growth rate, density/cover, germination, biomass and survival) equally promote tree invasiveness. Overall, our results suggest that several hypotheses, linked to invasibility or invasiveness, are pertinent to explain tree invasions. Furthermore, more than one hypothesis has been supported for a given species, which indicates that multiple factors lead to the success of invasive tree species. In addition, growth rate appears to be the most efficient predictor of invasiveness for invasive trees and could thus be used as a means to identify potential alien tree invasions. We conclude that further investigations are needed to test the consistency of some hypotheses across a broader pool of invasive tree species, whilst experimental studies with the same tree species across a larger range of sites would help to reveal the full suite of factors that affect tree invasions.