A global risk assessment for the success of bird introductions

1.  Concern over the impact of invasive species has led to the development of risk assessment methodologies to identify potential invaders and prevent future ecological and economic problems. However, developing a risk assessment tool is challenging because of the difficulties of accurately predicting the outcome of species introductions.
2.  In this study, we develop a global risk assessment for birds. We integrate two approaches, generalized linear mixed models (GLMM) and hierarchical tree models, to help identify those introductions with the highest risk of establishment success.
3.  Past work has shown that the number of individuals released is the main factor influencing establishment success in animals, a conclusion that was supported in our analyses. Establishment success was also higher for species with broader ecological niches and larger brains relative to body size. These features should increase the likelihood of finding an appropriate niche in the region of introduction.
4.  The GLMM and tree model predicted the probability of establishment success of birds in Europe and Australia with high accuracy (over 80% of introductions correctly classified). This highlights that establishment risk can be reasonably assessed with information on general habitat use, brain size and the size of the founder population. When compared with an alternative risk assessment tool based on a qualitative ranking, our quantitative approaches achieved higher accuracy with less information.
5.   Synthesis and applications. Quantitative risk assessments based on traits related to establishment success are difficult but feasible, providing a useful tool for guiding preventive polices aimed at mitigating the impact of invasive species.     

Founder events predict changes in genetic diversity during human‐mediated range expansions

Intentional or accidental introduction of species to new locations is predicted to result in loss of genetic variation and increase the likelihood of inbreeding, thus reducing population viability and evolutionary potential. However, multiple introductions and large founder numbers can prevent loss of genetic diversity and may therefore facilitate establishment success and range expansion. Based on a meta‐analysis of 119 introductions of 85 species of plants and animals, we here show a quantitative effect of founding history on genetic diversity in introduced populations. Both introduction of large number of individuals and multiple introduction events significantly contribute to maintaining or even increasing genetic diversity in introduced populations. The most consistent loss of genetic diversity is seen in insects and mammals, whereas introduced plant populations tend to have higher genetic variation than native populations. However, loss or gain of genetic diversity does not explain variation in the extent to which plant or animal populations become invasive outside of their native range. These results provide strong support for predictions from population genetics theory with respect to patterns of genetic diversity in introduced populations, but suggest that invasiveness is not limited by genetic bottlenecks.     

Effects of exotic mongoose (Herpestes javanicus) on the native fauna of Amami-Oshima Island, southern Japan, estimated by distribution patterns along the historical gradient of mongoose invasion

We examined the distribution patterns of native animals on Amami-Oshima Island, southern Japan, along a historical gradient of mongoose establishment and estimated the effects of mongoose on the native fauna. To assess the relative abundance of various ground-dwelling animals, we used the following four methods; sensor cameras for exotic mammals, nighttime driving census for nocturnal native vertebrates, line census for ground-dwelling lizards, and adhesive traps for arthropods. The results indicated that seven species with larger body size, including mammals, birds, reptiles, and amphibians, were rarely observed in mongoose-infested area. By contrast, medium-sized animals showed neutral relationships with mongoose establishment. Interestingly, the densities of smaller-sized animals were higher in mongoose-infested area. It could be interpreted that smaller species have increased in abundance through top-down cascades, i.e., decreases in native predators such as frogs and lizards caused by the mongoose have resulted in increases in the abundance of smaller animals. Predation pressures by mongoose and native predators may be canceled out for medium-sized animals, causing neutral responses to mongoose by these animals. This study appears to be the first example that shows the influence of mongoose on a wide variety of native animals. In addition, our findings indicate the importance of considering the food web structure of a recipient ecosystem and contribute to the prediction and assessment of ecological risks caused not only by mongoose, but also by other invasive top predators.     

Traits across trophic levels interact to influence parasitoid establishment in biological control releases

A central goal in ecology is to predict what governs a species’ ability to establish in a new environment. One mechanism driving establishment success is individual species’ traits, but the role of trait combinations among interacting species across different trophic levels is less clear. Deliberate or accidental species additions to existing communities provide opportunities to study larger scale patterns of establishment success. Biological control introductions are especially valuable because they contain data on both the successfully established and unestablished species. Here, we used a recent dataset of importation biological control introductions to explore how life‐history traits of 132 parasitoid species and their herbivorous hosts interact to affect parasitoid establishment. We find that of five parasitoid and herbivore traits investigated, one parasitoid trait—host range—weakly predicts parasitoid establishment; parasitoids with higher levels of phylogenetic specialization have higher establishment success, though the effect is marginal. In addition, parasitoids are more likely to establish when their herbivore host has had a shorter residence time. Interestingly, we do not corroborate earlier findings that gregarious parasitoids and endoparasitoids are more likely to establish. Most importantly, we find that life‐history traits of the parasitoid species and their hosts can interact to influence establishment. Specifically, parasitoids with broader host ranges are more likely to establish when the herbivore they have been released to control is also more of a generalist. These results provide insight into how multiple species’ traits and their interactions, both within and across trophic levels, can influence establishment of species of higher trophic levels.     

Historical legacies and ecological determinants of grass naturalizations worldwide

The global distribution of exotic species is the result of abiotic, biotic and dispersal filtering processes that shape the movement and success of species outside their native range. In this study we aim to understand how these filtering processes drive the fluxes of grass species among regions, the factors that influence which species establish outside of their native range, and where they do so. We used national and subnational checklists of native and introduced grass species to determine the extent to which each region was a source or recipient of exotic grass species. We asked how species traits may distinguish those grass species that have naturalized outside their native range from those that have not, and how environmental conditions are related to the distribution of exotic grass species. We found that exotic grass establishment is shaped by an array of factors including characteristics of regions, traits of species and their interactions. Regions with a longer history of human occupation and larger numbers of native grass species were generally the most important sources of exotic species. Global flows of species were mostly driven by a climate match between the native and exotic ranges, but were also highly asymmetric, with regions with recent human arrival being the major hosts of exotic grass species. Tall, annual and C4 grass species exhibited particularly high probabilities of establishment outside their native range. Despite the idiosyncrasy and stochasticity characteristic of exotic species establishment, this biogeographical analysis revealed important generalities across this large plant group. Our results suggest that grass species that have co-occurred with humans for a longer time may be better adapted to living in anthropogenic landscapes, explaining the global asymmetry in species introductions.     

Thermal tolerance experiments help establish survival probabilities for tilapia, a group of potentially invasive aquatic species

1. Estimating the probability of establishment is an important step toward managing ecological risk posed by natural or intentional introductions. Species introductions in certain geographic areas may pose less of a threat when environmental conditions are unlikely to promote long-term establishment. Species commonly referred to as 'tilapia' have become widespread in certain areas of the southeastern United States, yet concerns remain regarding their potential spread to other areas.
2. We created a model to predict the survival of tilapia in Georgia (U.S.A.) based on individual 'thermal experience'. Laboratory experiments were conducted to measure the duration of survival of two strains of tilapia, Oreochromis nilotica and O. nilotica  ×  O. aureus hybrids, exposed to a variety of temperature regimes with three minimum temperatures and two rates of temperature decline.
3. We used Mayfield hierarchical logistic regression (MHLR) to describe the daily probability of survival as a function of the rate of temperature decline, average sustained temperature, tilapia strain and mass–length ratio.
4. Tilapia generally survived constant minimum temperatures over 12 °C but survival rate was mediated by the rate of decline, mass–length ratio and strain. For every 1 °C increase in minimum temperature, tilapia were 2.76 times more likely to survive. Fish exposed to rapid temperature decline were less likely to survive than those exposed to slowly decreasing temperature. More robust fish (higher mass–length ratio) were more likely to survive, and strain had a minimal effect despite being supported by the best MHLR model.
5. Our model can be used to estimate survival probability for tilapia under known surface water temperature regimes. Using MLHR in conjunction with experimental tolerance data may be useful for estimating the likelihood of establishment of potentially invasive species.     

Regional ecological risk assessment for the introduction of Gambusia affinis (western mosquitofish) into Montana watersheds

Qualitative risk assessment methodologies were used to assess the risk of establishment and consequent impacts on native minnows and species of concern (SOC) associated with the intentional or unintentional introduction of the mosquito biological control agent, Gambusia affinis, to various Montana watersheds. Gambusia affinis introductions for mosquito control have been made throughout the world; some introductions have resulted in deleterious effects on native species. We used average January water temperatures, the presence of dams, and the presence of native minnows and SOC to define endpoints for our assessment. Our results suggest that a section of the Missouri River running between Wolf Creek and Landusky had the highest overall risk score, which corresponds to the highest likelihood of establishment and effects on native minnows and SOC. We also demonstrate how rivers with the highest temperatures are not necessarily at the highest risk of having deleterious effects on minnows and SOC; conversely, these rivers were found to be at highest risk of a population of G. affinis establishing.     

The island biogeography of exotic bird species

Aim   A recent upsurge of interest in the island biogeography of exotic species has followed from the argument that they may provide valuable information on the natural processes structuring island biotas. Here, we use data on the occurrence of exotic bird species across oceanic islands worldwide to demonstrate an alternative and previously untested hypothesis that these distributional patterns are a simple consequence of where humans have released such species, and hence of the number of species released.
Location   Islands around the world.
Methods   Statistical analysis of published information on the numbers of exotic bird species introduced to, and established on, islands around the world.
Results   Established exotic birds showed very similar species–area relationships to native species, but different species–isolation relationships. However, in both cases the relationship for established exotics simply mimicked that for the number of exotic bird species introduced. Exotic bird introductions scaled positively with human population size and island isolation, and islands that had seen more native species extinctions had had more exotic species released.
Main conclusion   The island biogeography of exotic birds is primarily a consequence of human, rather than natural, processes.     

Assessing the potential for Grey Squirrels Sciurus carolinensis to compete with birds at supplementary feeding stations

Supplementary feeding of birds, particularly in urban areas, is often associated with increased population size and fecundity. In the UK, the non‐native Grey Squirrel Sciurus carolinensis is common in rural and urban habitats. It exploits supplementary feeders and may induce interference competition by excluding birds, but empirical evidence of this is unavailable. Using controlled model presentation experiments, we demonstrate that Grey Squirrels could reduce bird use of supplementary feeders and induce interference competition. Total bird resource use was reduced by 98% and most species exhibited similar sensitivities. The likelihood and magnitude of interference competition will depend on how rapidly displaced birds find alternative food sources; it will be greatest where there are high Grey Squirrel densities and few supplementary feeders. Other studies suggest that supplementary feeding increases Grey Squirrel numbers, and the species is also predicted to expand its non‐native range across most of Europe. Our data indicate that Grey Squirrels may eventually alter the net effect of supplementary feeding on bird populations across the European continent; increased use of squirrel‐proof feeders may help to minimize such effects.     

Across island and continents, mammals are more successful invaders than birds (Reply)

Aim  The establishment success of exotic species is calculated as the fraction of introduced species that have become established, and invasion success is estimated as the fraction of established species that have spread significantly from their points of introduction. Records on species introductions are highly incomplete, so strong conclusions about the tens rule and invasibility of island and continents cannot be drawn.
Location  Global.
Methods  Using Jeschke (2008) as an example, we explain the issue of the inadequacy of data to draw conclusions about the tens rule and invasibility of island and continents.
Results  Lack of adequate data.
Main Conclusion  Jeschke (2008) probably overestimates the establishment and invasion success rates, so his conclusions about violation of the tens rule and that islands are not more susceptible to invasion than continents are misleading.     

Widespread primary,but geographically restricted secondary,human introductions of wall lizards,Podarcis muralis

Establishing the introduction pathways of alien species is a fundamental task in invasion biology. The common wall lizard, Podarcis muralis, has been widely introduced outside of its native range in both Europe and North America, primarily through escaped pets or deliberate release of animals from captive or wild populations. Here, we use Bayesian clustering, approximate Bayesian computation (ABC) methods and network analyses to reconstruct the origin and colonization history of 23 non‐native populations of wall lizards in England. Our analyses show that established populations in southern England originate from at least nine separate sources of animals from native populations in France and Italy. Secondary introductions from previously established non‐native populations were supported for eleven (47%) populations. In contrast to the primary introductions, secondary introductions were highly restricted geographically and appear to have occurred within a limited time frame rather than being increasingly common. Together, these data suggest that extant wall lizard populations in England are the result of isolated accidental and deliberate releases of imported animals since the 1970s, with only local translocation of animals from established non‐native populations. Given that populations introduced as recently as 25 years ago show evidence of having adapted to cool climate, discouraging further translocations may be important to prevent more extensive establishment on the south coast of England.     

Establishment success of introduced amphibians increases in the presence of congeneric species

Darwin's naturalization hypothesis predicts that the success of alien invaders will decrease with increasing taxonomic similarity to the native community. Alternatively, shared traits between aliens and the native assemblage may preadapt aliens to their novel surroundings, thereby facilitating establishment (the preadaptation hypothesis). Here we examine successful and failed introductions of amphibian species across the globe and find that the probability of successful establishment is higher when congeneric species are present at introduction locations and increases with increasing congener species richness. After accounting for positive effects of congeners, residence time, and propagule pressure, we also find that invader establishment success is higher on islands than on mainland areas and is higher in areas with abiotic conditions similar to the native range. These findings represent the first example in which the preadaptation hypothesis is supported in organisms other than plants and suggest that preadaptation has played a critical role in enabling introduced species to succeed in novel environments.     

High genetic diversity is not essential for successful introduction

Some introduced populations thrive and evolve despite the presumed loss of diversity at introduction. We aimed to quantify the amount of genetic diversity retained at introduction in species that have shown evidence of adaptation to their introduced environments. Samples were taken from native and introduced ranges of Arctotheca populifolia and Petrorhagia nanteuilii. Using microsatellite data, we identified the source for each introduction, estimated genetic diversity in native and introduced populations, and calculated the amount of diversity retained in introduced populations. These values were compared to those from a literature review of diversity in native, confamilial populations and to estimates of genetic diversity retained at introduction. Gene diversity in the native range of both species was significantly lower than for confamilials. We found that, on average, introduced populations showing evidence of adaptation to their new environments retained 81% of the genetic diversity from the native range. Introduced populations of P. nanteuilii had higher genetic diversity than found in the native source populations, whereas introduced populations of A. populifolia retained only 14% of its native diversity in one introduction and 1% in another. Our literature review has shown that most introductions demonstrating adaptive ability have lost diversity upon introduction. The two species studied here had exceptionally low native range genetic diversity. Further, the two introductions of A. populifolia represent the largest percentage loss of genetic diversity in a species showing evidence of substantial morphological change in the introduced range. While high genetic diversity may increase the likelihood of invasion success, the species examined here adapted to their new environments with very little neutral genetic diversity. This finding suggests that even introductions founded by small numbers of individuals have the potential to become invasive.     

Spread of an introduced parasite across the Hawaiian archipelago independent of its introduced host

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Status of the alien pathogen Paranosema locustae (Microsporidia) in grasshoppers (Orthoptera: Acridoidea) of the Argentine Pampas

After experimental introductions from North America in 1978–1982, the biocontrol agent Paranosema locustae became established in grasshopper communities of the western Pampas region of Argentina. The use and establishment of P. locustae in Argentina constitute both a case of neoclassical or new association biological control (use of an alien species against native pests) and a case of pathogen pollution (anthropogenic introduction–establishment of an infectious disease in populations of native species). Since P. locustae is a multihost pathogen among grasshoppers, its presence in the western Pampas represents an additional factor disrupting grasshopper communities according to the differential susceptibility of each species and possibly threatening some species. Microscopic examination of 504 grasshopper samples (the mean number of individuals per sample = 185) belonging to 43 species from 93 localities throughout the Pampas revealed an establishment area of approximate 90,000 km² from about 35° North to 38° South and from 61° East to 65° West. Field infections by P. locustae have now been detected in 21 grasshopper species in the western Pampas. Susceptible species with geographic distributions mostly restricted to the establishment area and with numerically small populations, like the melanopline Scotussa daguerrei, are predicted to be the ones facing higher risks of negative impacts.     

Non-indigenous species (NIS) of polychaetes (Annelida: Polychaeta) from the Atlantic and Mediterranean coasts of the Iberian Peninsula: an annotated checklist

This study provides an updated catalogue of non-indigenous species (NIS) of polychaetes reported from the continental coasts of the Iberian Peninsula based on the available literature. A list of 23 introduced species were regarded as established and other 11 were reported as casual, with 11 established and nine casual NIS in the Atlantic coast of the studied area and 14 established species and seven casual ones in the Mediterranean side. The most frequent way of transport was shipping (ballast water or hull fouling), which according to literature likely accounted for the introductions of 14 established species and for the presence of another casual one. To a much lesser extent aquaculture (three established and two casual species) and bait importation (one established species) were also recorded, but for a large number of species the translocation pathway was unknown. About 25% of the reported NIS originated in the Warm Western Atlantic region, followed by the Tropical Indo West-Pacific region (18%) and the Warm Eastern Atlantic (12%). In the Mediterranean coast of the Iberian Peninsula, nearly all the reported NIS originated from warm or tropical regions, but less than half of the species recorded from the Atlantic side were native of these areas. The effects of these introductions in native marine fauna are largely unknown, except for one species (Ficopomatus enigmaticus) which was reported to cause serious environmental impacts. In other cases, the displacement of native species or economic damages are suspected.     

Predicting introduction, establishment and potential impacts of smallmouth bass

Aim The introduction of non‐indigenous species has resulted in wide‐ranging ecological and economic impacts. Predictive modelling of the introduction and establishment of non‐indigenous species is imperative to identify areas at high risk of invasion to effectively manage non‐indigenous species and conserve native populations. Smallmouth bass (Micropterus dolomieu), a warm water fish species native to central North America has negatively impacted native fish communities, including cyprinids and salmonid populations, as a result of intentional introductions. We predicted the introduction risk; species establishment based on habitat suitability; identified lakes at high risk of invasion; and finally assessed the consequential impacts on native salmon, trout and cyprinid populations. Location Ontario and British Columbia, Canada. Methods Classification tree and logistic regression models were developed and validated to predict the introduction and establishment of smallmouth bass for thousands of lakes. Results Densely human populated areas and larger lake surface areas successfully identify lakes associated with the introduction of smallmouth bass (introduction model) in British Columbia. Climate, lake morphology and water chemistry variables were the driving environmental parameters to define suitable smallmouth bass habitat (establishment model). A combination of the introduction and establishment model identified 138 lakes that are currently at risk in British Columbia to the introduction and establishment of smallmouth bass. Of these 138 high‐risk lakes, 95% of them contain at least one species of salmon, trout or cyprinid, thereby increasing the potential impact of an invasion by smallmouth bass. Main conclusions Our framework can be applied to other terrestrial and aquatic species to obtain a better understanding of the potential risk posed by a non‐indigenous species to an ecosystem. Furthermore, our methodology can be used to focus management efforts on areas at higher risk (e.g. number of potential releases, more favourable habitats) to control future introductions of non‐indigenous species, thereby conserving native populations.     

Can life‐history traits predict the fate of introduced species? A case study on two cyprinid fish in southern France

1. The ecological and economic costs of introduced species can be high. Ecologists try to predict the probability of success and potential risk of the establishment of recently introduced species, given their biological characteristics.
2. In 1990 gudgeon, Gobio gobio , were released in a drainage canal of the Rhône delta of southern France. The Asian topmouth gudgeon, Pseudorasbora parva, was found for the first time in the same canal in 1993. Those introductions offered a unique opportunity to compare in situ the fate of two closely related fish in the same habitat.
3. Our major aims were to assess whether G. gobio was able to establish in what seemed an unlikely environment, to compare populations trends and life-history traits of both species and to assess whether we could explain or could have predicted our results, by considering their life-history strategies.
4. Data show that both species have established in the canal and have spread. Catches of P. parva have increased strongly and are now higher than those of G. gobio .
5. The two cyprinids have the same breeding season and comparable traits (such as short generation time, small body, high reproductive effort), so both could be classified as opportunists. The observed difference in their success (in terms of population growth and colonization rate) could be explained by the wider ecological and physiological tolerance of P. parva .
6. In conclusion, our field study seems to suggest that invasive vigour also results from the ability to tolerate environmental changes through phenotypic plasticity, rather than from particular life-history features pre-adapted to invasion. It thus remains difficult to define a good invader simply on the basis of its life-history features.     

Effects of environmental variation on extinction and establishment

Theoretical models predict that increasing environmental variation increases the probability of extinction, decreases the probability of establishment, and influences the distribution of times to extinction or establishment. We conducted an experiment with 281 independent populations of Daphnia magna under controlled laboratory conditions to test these predictions. Consistent with the theory, the fraction of populations going extinct increased and the fraction of populations establishing self‐sustaining populations decreased under higher levels of environmental variation compared with controls. Time to extinction decreased under higher levels of environmental variation, but we found no effect on time to establishment. These results are consistent with theoretical predictions from models of extinction. They therefore support the use of stochastic population models to predict the fates of introductions of non‐indigenous species or native endangered species based on historic fluctuations and/or expected future conditions.     

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.