Simulating plant invasion dynamics in mountain ecosystems under global change scenarios

Across the globe, invasive alien species cause severe environmental changes, altering species composition and ecosystem functions. So far, mountain areas have mostly been spared from large‐scale invasions. However, climate change, land‐use abandonment, the development of tourism and the increasing ornamental trade will weaken the barriers to invasions in these systems. Understanding how alien species will react and how native communities will influence their success is thus of prime importance in a management perspective. Here, we used a spatially and temporally explicit simulation model to forecast invasion risks in a protected mountain area in the French Alps under future conditions. We combined scenarios of climate change, land‐use abandonment and tourism‐linked increases in propagule pressure to test if the spread of alien species in the region will increase in the future. We modelled already naturalized alien species and new ornamental plants, accounting for interactions among global change components, and also competition with the native vegetation. Our results show that propagule pressure and climate change will interact to increase overall species richness of both naturalized aliens and new ornamentals, as well as their upper elevational limits and regional range‐sizes. Under climate change, woody aliens are predicted to more than double in range‐size and herbaceous species to occupy up to 20% of the park area. In contrast, land‐use abandonment will open new invasion opportunities for woody aliens, but decrease invasion probability for naturalized and ornamental alien herbs as a consequence of colonization by native trees. This emphasizes the importance of interactions with the native vegetation either for facilitating or potentially for curbing invasions. Overall, our work highlights an additional and previously underestimated threat for the fragile mountain flora of the Alps already facing climate changes, land‐use transformations and overexploitation by tourism.     

Congener diversity,topographic heterogeneity and human‐assisted dispersal predict spread rates of alien herpetofauna at a global scale

Understanding the factors that determine rates of range expansion is not only crucial for developing risk assessment schemes and management strategies for invasive species, but also provides important insight into the ability of species to disperse in response to climate change. However, there is little knowledge on why some invasions spread faster than others at large spatiotemporal scales. Here, we examine the effects of human activities, species traits and characteristics of the invaded range on spread rates using a global sample of alien reptile and amphibian introductions. We show that spread rates vary remarkably among invaded locations within a species, and differ across biogeographical realms. Spread rates are positively related to the richness of native congeneric species and human‐assisted dispersal in the invaded range but are negatively correlated with topographic heterogeneity. Our findings highlight the importance of environmental characteristics and human‐assisted dispersal in developing robust frameworks for predicting species' range shifts.     

Birds as key vectors for the dispersal of some alien species: Further thoughts

W. Solarz, K. Najberek, A. Pociecha & E. Wilk‐Wo?niak ( 2017 , Diversity and Distributions, 23 , 113–117) published a letter in Diversity and Distributions debating our view that waterbirds are important vectors of alien species (C. Reynolds, N. A. F. Miranda & G. S. Cumming, 2015 Diversity and Distributions, 21 , 744–754; A. J. Green, 2016 Diversity and Distributions, 22 , 239–247) and question whether future research into the mechanisms under‐pinning this phenomenon can be advantageous for the practical management of alien species. Additionally, Solarz et al. suggest that human activities are the primary source of all alien species introductions and that waterbirds may only act as vectors of secondary dispersal. In this letter, we respond to several arguments raised by the authors surrounding the relevance of waterbird‐mediated dispersal in the introduction and spread of alien species. We emphasize the partly deterministic nature of waterbird dispersal and the significance of long‐distance dispersal events (and hence the potential for primary introductions of new alien species across political boundaries). Finally, we reaffirm the importance of further research into dispersal by birds to improve our capacity to foresee and manage invasions of those alien species with strong capacity to spread via avian vectors.     

The abiotic and biotic factors limiting establishment of predatory fishes at their expanding northern range boundaries in Ontario,Canada

There is a poor understanding of the importance of biotic interactions in determining species distributions with climate change. Theory from invasion biology suggests that the success of species introductions outside of their historical ranges may be either positively (biotic acceptance) or negatively (biotic resistance) related to native biodiversity. Using data on fish community composition from two survey periods separated by approximately 28 years during which climate was warming, we examined the factors influencing the establishment of three predatory centrarchids: Smallmouth Bass (Micropterus dolomieu), Largemouth Bass (M. salmoides), and Rock Bass (Ambloplites rupestris) in lakes at their expanding northern range boundaries in Ontario. Variance partitioning demonstrated that, at a regional scale, abiotic factors play a stronger role in determining the establishment of these species than biotic factors. Pairing lakes within watersheds where each species had established with lakes sharing similar abiotic conditions where the species had not established revealed both positive and negative relationships between the establishment of centrarchids and the historical presence of other predatory species. The establishment of these species near their northern range boundaries is primarily determined by abiotic factors at a regional scale; however, biotic factors become important at the lake‐to‐lake scale. Studies of exotic species invasions have previously highlighted how spatial scale mediates the importance of abiotic vs. biotic factors on species establishment. Our study demonstrates how concepts from invasion biology can inform our understanding of the factors controlling species distributions with changing climate.     

Recent non‐native invertebrate plant pest establishments in Great Britain: origins,pathways, and trends

1 An appraisal of non‐native invertebrate plant pest establishments in Great Britain, between 1970 and 2004, was carried out to improve our understanding of current invasion processes by non‐native plant pests, and to assist national strategies in managing the risks they pose. 2 A total of 164 establishments, comprising 50 natural colonists and 114 human‐assisted introductions, were recorded across 13 major taxonomic groups. 3 The mean rate of establishment was 22.1 species per 5‐year period: 19.1 and 3.0 species outside and inside protected cultivation, respectively. Despite the continuing rapid growth in international trade and a general perception that rates of pest invasions are accelerating, no significant temporal trends in the rate of establishments in Great Britain were detected, either for natural colonists or human‐assisted introductions, or for pests of plants grown indoors or outside. 4 The plant trade, particularly in ornamental plants, accounted for nearly 90% of human‐assisted introductions; apiculture, biological control, timber imports, transport stowaways and intentional releases each contributed less than 5%. Only eight (4.9%) of the establishments could be considered as having no direct potential economic impact because all other species have been recorded as feeding on cultivated plants. A greater proportion of establishments by both natural colonists and human‐assisted introductions occurred on non‐native, woody plants. 5 The present study confirms previous work in other European countries that highlight the predominant role of the ornamental plant trade in introducing new plant pests to the European continent, mainly from Asia and North America.     

Establishment success of invasive ring‐necked and monk parakeets in Europe

Aim Invasive alien species are a growing threat to biodiversity, and identifying the mechanisms that enable these species to establish viable populations in their new environment is paramount for management of the problems they pose. Using an unusually large number of both failed and successful documented introductions of parakeets (Aves: Psittacidae) in Europe, we test two of the major hypotheses on the establishment success of invading species, namely the climate‐matching and the human‐activity hypothesis. Location European human population centres where ring‐necked parakeet (Psittacula krameri) and/or monk parakeet (Myiopsitta monachus) introductions have occurred. Methods Data on ring‐necked and monk parakeet introductions in Europe were gathered from various sources, including published books and articles, but also from unpublished reports and local grey literature. Information was verified with experts from the region under consideration. In order to test the climate‐matching hypothesis, we verified whether the climatic factors that determine the parakeets’ native ranges also explain establishment success in Europe. Parakeet occurrence data from the native ranges were analysed using the presence‐only modelling method Maxent , and correlations between parakeet establishment and climatic and anthropogenic variables in Europe were assessed using both stepwise logistic regression and the information‐theoretic model selection approach. Results The establishment success of ring‐necked and monk parakeets was found to be positively associated with human population density, and, both in the native and in the introduced regions, parakeet occurrence was negatively correlated with the number of frost days. Thus, parakeets are more likely to establish in warmer and human‐dominated areas. Main conclusions The large number of independent parakeet introductions in Europe allows us to test the often‐used climate‐matching and human‐activity hypotheses at the species level. We show that both hypotheses offer insight into the invasion process of monk and ring‐necked parakeets. Our results suggest that, in the future, parakeet establishment probability may increase even further because global warming is likely to cause a decrease in the number of frost days and because urbanization and human populations are still increasing.     

Anthropogenic fires increase alien and native annual species in the Chilean coastal matorral

Aim We tested the hypothesis that anthropogenic fires favour the successful establishment of alien annual species to the detriment of natives in the Chilean coastal matorral. Location Valparaíso Region, central Chile. Methods We sampled seed rain, seedbank emergence and establishment of species in four paired burned and unburned areas and compared (using GLMM) fire resistance and propagule arrival of alien and native species. To assess the relative importance of seed dispersal and seedbank survival in explaining plant establishment after fire, we compared seed rain and seedbank structure with post‐fire vegetation using ordination analyses. Results Fire did not change the proportion of alien species in the coastal matorral. However, fire increased the number of annual species (natives and aliens) of which 87% were aliens. Fire reduced the alien seedbank and not the native seedbank, but alien species remained dominant in burned soil samples (66% of the total species richness). Seed rain was higher for alien annuals than for native annuals or perennials, thus contributing to their establishment after fire. Nevertheless, seed rain was less important than seedbank survival in explaining plant establishment in burned areas. Main conclusions Anthropogenic fires favoured alien and native annuals. Thus, fire did not increase the alien/native ratio but increased the richness of alien species. The successful establishment of alien annuals was attributable to their ability to maintain rich seedbanks in burned areas and to the greater propagule arrival compared to native species. The native seedbank also survived fire, indicating that the herbaceous community has become highly resilient after centuries of human disturbances. Our results demonstrate that fire is a relevant factor for the maintenance of alien‐dominated grasslands in the matorral and highlight the importance of considering the interactive effect of seed rain and seedbank survival to understand plant invasion patterns in fire‐prone ecosystems.     

Propagule pressure and an invasion syndrome determine invasion success in a plant community model

The success of species invasions depends on multiple factors, including propagule pressure, disturbance, productivity, and the traits of native and non‐native species. While the importance of many of these determinants has already been investigated in relative isolation, they are rarely studied in combination. Here, we address this shortcoming by exploring the effect of the above‐listed factors on the success of invasions using an individual‐based mechanistic model. This approach enables us to explicitly control environmental factors (temperature as surrogate for productivity, disturbance, and propagule pressure) as well as to monitor whole‐community trait distributions of environmental adaptation, mass, and dispersal abilities. We simulated introductions of plant individuals to an oceanic island to assess which factors and species traits contribute to invasion success. We found that the most influential factors were higher propagule pressure and a particular set of traits. This invasion trait syndrome was characterized by a relative similarity in functional traits of invasive to native species, while invasive species had on average higher environmental adaptation, higher body mass, and increased dispersal distances, that is, had greater competitive and dispersive abilities. Our results highlight the importance in management practice of reducing the import of alien species, especially those that display this trait syndrome and come from similar habitats as those being managed.     

Effects of climate change and horticultural use on the spread of naturalized alien garden plants in Europe

Climate warming is supposed to enlarge the area climatically suitable to the naturalization of alien garden plants in temperate regions. However, the effects of a changing climate on the spread of naturalized ornamentals have not been evaluated by spatially and temporarily explicit range modelling at larger scales so far. Here, we assess how climate change and the frequency of cultivation interactively determine the spread of 15 ornamental plants over the 21st century in Europe. We coupled species distribution modelling with simulations of demography and dispersal to predict range dynamics of these species in annual steps across a 250 × 250 m raster of the study area. Models were run under four scenarios of climate warming and six levels of cultivation intensity. Cultivation frequency was implemented as size of the area used for planting a species. Although the area climatically suitable to the 15 species increases, on average, the area predicted to be occupied by them in 2090 shrinks under two of the three climate change scenarios. This contradiction obviously arises from dispersal limitations that were pronounced although we assumed that cultivation is spatially adapting to the changing climate. Cultivation frequency had a much stronger effect on species spread than climate change, and this effect was non‐linear. The area occupied increased sharply from low to moderate levels of cultivation intensity, but levelled off afterwards. Our simulations suggest that climate warming will not necessarily foster the spread of alien garden plants in Europe over the next decades. However, climatically suitable areas do increase and hence an invasion debt is likely accumulating. Restricting cultivation of species can be effective in preventing species spread, irrespective of how the climate develops. However, for being successful, they depend on high levels of compliance to keep propagule pressure at a low level.     

Non‐native fishes and climate change: predicting species responses to warming temperatures in a temperate region

1. Temperate regions with fish communities dominated by cold‐water species (physiological optima <20 °C) are vulnerable to the effects of warming temperatures caused by climate change, including displacement by non‐native cool‐water (physiological optima 20–28 °C) and warm‐water fishes (physiological optima >28 °C) that are able to establish and invade as the thermal constraints on the expression of their life history traits diminish. 2. England and Wales is a temperate region into which at least 38 freshwater fishes have been introduced, although 14 of these are no longer present. Of the remaining 24 species, some have persisted but failed to establish, some have established populations without becoming invasive and some have become invasive. The aim of the study was to predict the responses of these 24 non‐native fishes to the warming temperatures of England and Wales predicted under climate change in 2050. 3. The predictive use of climate‐matching models and an air and water temperature regression model suggested that there are six non‐native fishes currently persistent but not established in England and Wales whose establishment and subsequent invasion would benefit substantially from the predicted warming temperatures. These included the common carp Cyprinus carpio and European catfish Silurus glanis, fishes that also exert a relatively high propagule pressure through stocking to support angling and whose spatial distribution is currently increasing significantly, including in open systems. 4. The potential ecological impacts of the combined effects of warming temperatures, current spatial distribution and propagule pressure on the establishment and invasion of C. carpio and S. glanis were assessed. The ecological consequences of C. carpio invasion were assessed as potentially severe in England and Wales, with impacts likely to relate to habitat destruction, macrophyte loss and increased water turbidity. However, evidence of ecological impacts of S. glanis elsewhere in their introduced range was less clear and so their potential impacts in England and Wales remain uncertain.     

What affects the probability of biological invasions in Antarctica? Using an expanded conceptual framework to anticipate the risk of alien species expansion

Successful alien species invasion depends on many factors studied mostly in post invasion habitats, and subsequently summarized in frameworks tailored to describe the studied invasion. We used an existing expanded framework with three groups of contributing factors: habitat invisibility, system context and species invasiveness, to analyze the probability of alien species invasions in terrestrial communities of Maritime Antarctic in the future. We focused on the first two factor groups. We tested if the expanded framework could be used under a different scenario. We chose Point Thomas Oasis on King George Island to perform our analysis. Strong geographical barrier, low potential bioclimatic suitability and resource availability associated with habitat invasibility significantly reduce the likelihood of biological invasion in Antarctica. An almost full enemy release (low pressure of consumers), the high patchiness of the habitat, and the prevalence of open gaps also associated with habitat invasibility increase the possibility of invasion. The dynamics of functional connectivity, propagule pressure and spatio-temporal patterns of propagule arrival associated with human activity and climate change belonging to the system context contribute to an increase in the threat of invasions. Due to the still low land transport activity migration pathways are limited and will reduce the spread of alien terrestrial organisms by land. An effective way of preventing invasions in Antarctica seems to lie in reducing propagule pressure and eliminating alien populations as early as possible. The expanded conceptual framework opens up wider possibilities in analyzing invasions taking place in different systems and with multiple taxa.     

Predicting establishment success for alien reptiles and amphibians: a role for climate matching

We examined data comprising 1,028 successful and 967 failed introduction records for 596 species of alien reptiles and amphibians around the world to test for factors influencing establishment success. We found significant variations between families and between genera. The number of jurisdictions where a species was introduced was a significant predictor of the probability the species had established in at least one jurisdiction. All species that had been introduced to more than 10 jurisdictions (34 species) had established at least one alien population. We also conducted more detailed quantitative comparisons for successful (69 species) and failed (116 species) introductions to three jurisdictions (Great Britain, California and Florida) to test for associations with climate match, geographic range size, and history of establishment success elsewhere. Relative to failed species, successful species had better climate matches between the jurisdiction where they were introduced and their geographic range elsewhere in the world. Successful species were also more likely to have high establishment success rates elsewhere in the world. Cross-validations indicated our full model correctly categorized establishment success with 78–80% accuracy. Our findings may guide risk assessments for the import of live alien reptiles and amphibians to reduce the rate new species establish in the wild.     

Marine range shifts and species introductions: comparative spread rates and community impacts

Aim Shifts in species ranges are a predicted and realized effect of global climate change; however, few studies have addressed the rates and consequence of such shifts, particularly in marine systems. Given ecological similarities between shifting and introduced species, we examined how our understanding of range shifts may be informed by the more established study of non‐native species introductions. Location Marine systems world‐wide. Methods Database and citation searches were used to identify 129 marine species experiencing range shifts and to determine spread rates and impacts on recipient communities. Analyses of spread rates were based on studies for which post‐establishment spread was reported in linear distance. The sizes of the effects of community impacts of shifting species were compared with those of functionally similar introduced species having ecologically similar impacts. Results Our review and meta‐analyses revealed that: (1) 75% of the range shifts found through the database search were in the poleward direction, consistent with climate change scenarios, (2) spread rates of range shifts were lower than those of introductions, (3) shifting species spread over an order of magnitude faster in marine than in terrestrial systems, and (4) directions of community effects were largely negative and magnitudes were often similar for shifters and introduced species; however, this comparison was limited by few data for range‐shifting species. Main conclusions Although marine range shifts are likely to proceed more slowly than marine introductions, the community‐level effects could be as great, and in the same direction, as those of introduced species. Because it is well‐established that introduced species are a primary threat to global biodiversity, it follows that, just like introductions, range shifts have the potential to seriously affect biological systems. In addition, given that ranges shift faster in marine than terrestrial environments, marine communities might be affected faster than terrestrial ones as species shift with climate change. Regardless of habitat, consideration of range shifts in the context of invasion biology can improve our understanding of what to expect from climate change‐driven shifts as well as provide tools for formal assessment of risks to community structure and function.     

Drivers of future alien species impacts: An expert‐based assessment

Understanding the likely future impacts of biological invasions is crucial yet highly challenging given the multiple relevant environmental, socio‐economic and societal contexts and drivers. In the absence of quantitative models, methods based on expert knowledge are the best option for assessing future invasion trajectories. Here, we present an expert assessment of the drivers of potential alien species impacts under contrasting scenarios and socioecological contexts through the mid‐21st century. Based on responses from 36 experts in biological invasions, moderate (20%–30%) increases in invasions, compared to the current conditions, are expected to cause major impacts on biodiversity in most socioecological contexts. Three main drivers of biological invasions—transport, climate change and socio‐economic change—were predicted to significantly affect future impacts of alien species on biodiversity even under a best‐case scenario. Other drivers (e.g. human demography and migration in tropical and subtropical regions) were also of high importance in specific global contexts (e.g. for individual taxonomic groups or biomes). We show that some best‐case scenarios can substantially reduce potential future impacts of biological invasions. However, rapid and comprehensive actions are necessary to use this potential and achieve the goals of the Post‐2020 Framework of the Convention on Biological Diversity.     

Hurricanes overcome migration lag and shape intraspecific genetic variation beyond a poleward mangrove range limit

Expansion of many tree species lags behind climate change projections. Extreme storms can rapidly overcome this lag, especially for coastal species, but how will storm‐driven expansion shape intraspecific genetic variation? Do storms provide recruits only from the nearest sources, or from more distant sources? Answers to these questions have ecological and evolutionary implications, but empirical evidence is absent from the literature. In 2017, Hurricane Irma provided an opportunity to address this knowledge gap at the northern range limit of the neotropical black mangrove (Avicennia germinans) on the Atlantic coast of Florida, USA. We observed massive post‐hurricane increases in beach‐stranded A. germinans propagules at, and past, this species’ present day range margin when compared to a previously surveyed nonhurricane year. Yet, propagule dispersal does not guarantee subsequent establishment and reproductive success (i.e., effective dispersal). We also evaluated prior effective dispersal along this coastline with isolated A. germinans trees identified beyond the most northern established population. We used 12 nuclear microsatellite loci to genotype 896 hurricane‐driven drift propagules from nine sites and 10 isolated trees from four sites, determined their sources of origin, and estimated dispersal distances. Almost all drift propagules and all isolated trees came from the nearest sources. This research suggests that hurricanes are a prerequisite for poleward range expansion of a coastal tree species and that storms can shape the expanding gene pool by providing almost exclusively range‐margin genotypes. These insights and empirical estimates of hurricane‐driven dispersal distances should improve our ability to forecast distributional shifts of coastal species.     

Insights from modeling studies on how climate change affects invasive alien species geography

Climate change and biological invasions are threatening biodiversity and ecosystem services worldwide. It has now been widely acknowledged that climate change will affect biological invasions. A large number of studies have investigated predicted shifts and other changes in the geographic ranges of invasive alien species related to climate change using modeling approaches. Yet these studies have provided contradictory evidence, and no consensus has been reached. We conducted a systematic review of 423 modeling case studies included in 71 publications that have examined the predicted effects of climate change on those species. We differentiate the approaches used in these studies and synthesize their main results. Our results reaffirm the major role of climate change as a driver of invasive alien species distribution in the future. We found biases in the literature both regarding the taxa, toward plants and invertebrates, and the areas of the planet investigated. Despite these biases, we found for the plants and vertebrates studied that climate change will more frequently contribute to a decrease in species range size than an increase in the overall area occupied. This is largely due to oceans preventing terrestrial invaders from spreading poleward. In contrast, we found that the ranges of invertebrates and pathogens studied are more likely to increase following climate change. An important caveat to these findings is that researchers have rarely considered the effects of climate change on transport, introduction success, or the resulting impacts. We recommend closing these research gaps, and propose additional avenues for future investigations, as well as opportunities and challenges for managing invasions under climate change.     

Simulating the effects of climate change on the distribution of an invasive plant,using a high resolution,local scale,mechanistic approach: challenges and insights

The growing economic and ecological damage associated with biological invasions, which will likely be exacerbated by climate change, necessitates improved projections of invasive spread. Generally, potential changes in species distribution are investigated using climate envelope models; however, the reliability of such models has been questioned and they are not suitable for use at local scales. At this scale, mechanistic models are more appropriate. This paper discusses some key requirements for mechanistic models and utilises a newly developed model (PSS[gt]) that incorporates the influence of habitat type and related features (e.g., roads and rivers), as well as demographic processes and propagule dispersal dynamics, to model climate induced changes in the distribution of an invasive plant (Gunnera tinctoria) at a local scale. A new methodology is introduced, dynamic baseline benchmarking, which distinguishes climate‐induced alterations in species distributions from other potential drivers of change. Using this approach, it was concluded that climate change, based on IPCC and C4i projections, has the potential to increase the spread‐rate and intensity of G. tinctoria invasions. Increases in the number of individuals were primarily due to intensification of invasion in areas already invaded or in areas projected to be invaded in the dynamic baseline scenario. Temperature had the largest influence on changes in plant distributions. Water availability also had a large influence and introduced the most uncertainty in the projections. Additionally, due to the difficulties of parameterising models such as this, the process has been streamlined by utilising methods for estimating unknown variables and selecting only essential parameters.     

A conceptual framework for understanding arthropod predator and parasitoid invasions

Arthropods make up the largest group of invasive alien species (IAS) worldwide. Although invasion research has been biased towards alien plants and vertebrates, it has suggested potential mechanisms for the success of IAS and provided a theoretical framework for further investigation. Here we address key concepts from invasion biology that are essential to our understanding of the success of invasive alien arthropod predators and parasitoids including human intervention, environmental characteristics, propagule pressure, biological traits, and biological interactions. To gain a greater understanding of the factors most likely to influence the different stages of invasion (arrival, establishment, and spread) for alien arthropod predators and parasitoids, we use a comparative approach to compare and contrast the differential success of invasions by alien phytophagous and carnivorous arthropods. Insights gained from this comparison suggest that future research will require a multitrophic approach in order to enhance our understanding of invasions at higher trophic levels.     

A framework integrating physiology,dispersal and land‐use to project species ranges under climate change

To study the potential effects of climate change on species, one of the most popular approaches are species distribution models (SDMs). However, they usually fail to consider important species‐specific biological traits, such as species’ physiological capacities or dispersal ability. Furthermore, there is consensus that climate change does not influence species distributions in isolation, but together with other anthropogenic impacts such as land‐use change, even though studies investigating the relative impacts of different threats on species and their geographic ranges are still rare. Here we propose a novel integrative approach which produces refined future range projections by combining SDMs based on distribution, climate, and physiological tolerance data with empirical data on dispersal ability as well as current and future land‐use. Range projections based on different combinations of these factors show strong variation in projected range size for our study species Emberiza hortulana. Using climate and physiological data alone, strong range gains are projected. However, when we account for land‐use change and dispersal ability, future range‐gain may even turn into a future range loss. Our study highlights the importance of accounting for biological traits and processes in species distribution models and of considering the additive effects of climate and land‐use change to achieve more reliable range projections. Furthermore, with our approach we present a new tool to assess species’ vulnerability to climate change which can be easily applied to multiple species.     

Fishing pressure and species traits affect stream fish invasions both directly and indirectly

Aim

We sought to identify direct and indirect effects of factors contributing to establishment and spread of 272 stream fish species.

Location

Two hundred and ninety‐seven watersheds in the eastern United States.

Methods

We modelled two variables: (1) whether a species had become established outside its native range (establishment) and (2) the number of watersheds in which species established outside their native range (spread). We estimated these variables by comparing historical distributions to a rich data set of contemporary sampling. We calculated metrics of human use (indexing propagule pressure), and gathered species trait data from an open‐access database. We then used piecewise path analysis to estimate direct and indirect effects of human use, native range size and species traits on the two metrics of species introductions.

Results

We identified a hierarchical causal structure in which native range size and fishing pressure were important direct determinants of introductions. Species traits had some direct effects, but played a more indirect role. Native range size was significantly affected by thermal tolerance and diet breadth. Likewise, fishing pressure was significantly affected by life history strategy: larger‐bodied, longer‐living and more fecund species were positively associated with fishing pressure.

Main conclusions

Functional traits can confer an advantage to some species during the establishment phase, but human use is important for subsequent dispersal throughout the non‐native range. However, human use is non‐random, and is largely a function of species traits. Considering both direct and indirect effects of traits across stages of the invasion process can help to elucidate the full role of traits in species invasions.