Biological and ecological characteristics of invasive species: a gammarid study

Knowledge of characteristics helpful in screening potential invaders and in elaborating strategies to limit their success is highly desirable. We focused on gammarid amphipods from Western Europe and North America to discover biological and/or ecological traits that may explain successful invasion by these species. Two typologies were considered: an analytical one, with groups built on the basis of biological or ecological similarities, and an empirical one, with groups constituted a priori according to a species’ invasive status and its fresh or brackish water origin. The results obtained are discussed in the light of three hypotheses that may influence invasiveness: biotic potential, species size and euryoeciousness. The analysis revealed a particular ecological profile for invaders, with a strong influence of salinity tolerance, but no typology was found based on biological characteristics. Invasiveness cannot be predicted from a limited number of criteria, and is the result of a combination of several characteristics. Invasive species therefore exhibit a particular ecological profile rather than a biological one, contrary to most classical explanations.     

The invasiveness of an introduced species does not predict its impact

Inconsistent use of terminology plagues the study and management of biological invasions. The term “invasive” has been used to describe inter alia (1) any introduced non-indigenous species; (2) introduced species that spread rapidly in a new region; and (3) introduced species that have harmful environmental impacts, particularly on native species. The second definition in various forms is more commonly used by ecologists, while the third definition is pervasive in policy papers and legislation. We tested the relationship between the invasiveness of an introduced species and its impact on native biodiversity. We quantified a species’ invasiveness by both its rate of establishment and its rate of spread, while its impact was assigned a categorical ranking based on the documented effects of the invader on native species populations. We found no correlations between these variables for introduced plants, mammals, fishes, invertebrates, amphibians and reptiles, suggesting that the mechanisms of invasion and impact are not strongly linked. Our results support the view that the term “invasive” should not be used to connote negative environmental impact.     

Secondary invasion: When invasion success is contingent on other invaders altering the properties of recipient ecosystems

Positive interactions between exotic species may increase ecosystem‐level impacts and potentially facilitate the entry and spread of other exotic species. Invader‐facilitated invasion success—”secondary invasion”—is a key conceptual aspect of the well‐known invasional meltdown hypothesis, but remains poorly defined and empirically underexplored. Drawing from heuristic models and published empirical studies, we explore this form of “secondary invasion” and discuss the phenomenon within the recognized conceptual framework of the determinants of invasion success. The term “secondary invasion” has been used haphazardly in the literature to refer to multiple invasion phenomena, most of which have other more accepted titles. Our usage of the term secondary invasion is akin to “invader‐facilitated invasion,” which we define as the phenomenon in which the invasion success of one exotic species is contingent on the presence, influence, and impacts of one or more other exotic species. We present case studies of secondary invasion whereby primary invaders facilitate the entry or establishment of exotic species into communities where they were previously excluded from becoming invasive. Our synthesis, discussion, and conceptual framework of this type of secondary invasion provides a useful reference to better explain how invasive species can alter key properties of recipient ecosystems that can ultimately determine the invasion success of other species. This study increases our appreciation for complex interactions following invasion and highlights the impacts of invasive species themselves as possible determinants of invasion success. We anticipate that highlighting “secondary invasion” in this way will enable studies reporting similar phenomena to be identified and linked through consistent terminology.     

Clonal integration facilitates the proliferation of smooth brome clones invading northern fescue prairies

Predicting exotic invaders and reducing their impacts on the biodiversity and function of native ecosystems require understanding of the mechanisms that facilitate their success during key stages of invasion. We determined whether clonal growth, characteristic of the majority of successful invaders of natural areas, facilitates the proliferation of Bromus inermis (smooth brome), an exotic grass invading prairie ecosystems across the Great Plains. By manipulating the below-ground connections of proliferating rhizomes as well as the levels of soil nitrogen along the margins of clones invading northern fescue prairies in Manitoba, Canada, we hypothesized that physiological integration would most benefit ramets invading low resource environments. Severing clonal connections reduced the mass of smooth brome shoots invading native prairies and was exacerbated by the immobilization of soil nutrients with glucose. Clonal connections were equally important in the maintenance of smooth brome density and the horizontal proliferation of ramets. Our results demonstrate the role of physiological integration in the proliferation of a clonal exotic invader and may help explain the success of clonal invaders in other regions. Although integration among invading ramets suggests several possibilities for successful management, future research must continue to elucidate differences in the invasiveness of native versus exotic species as well as the persistence of clonal connections among exotic invaders.     

Propagule Pressure: A Null Model for Biological Invasions

Invasion ecology has been criticised for its lack of general principles. To explore this criticism, we conducted a meta-analysis that examined characteristics of invasiveness (i.e. the ability of species to establish in, spread to, or become abundant in novel communities) and invasibility (i.e. the susceptibility of habitats to the establishment or proliferation of invaders). There were few consistencies among invasiveness characteristics (3 of 13): established and abundant invaders generally occupy similar habitats as native species, while abundant species tend to be less affected by enemies; germination success and reproductive output were significantly positively associated with invasiveness when results from both stages (establishment/spread and abundance/impact) were combined. Two of six invasibility characteristics were also significant: communities experiencing more disturbance and with higher resource availability sustained greater establishment and proliferation of invaders. We also found that even though ‘propagule pressure’ was considered in only ~29% of studies, it was a significant predictor of both invasiveness and invasibility (55 of 64 total cases). Given that nonindigenous species are likely introduced non-randomly, we contend that ‘propagule biases’ may confound current paradigms in invasion ecology. Examples of patterns that could be confounded by propagule biases include characteristics of good invaders and susceptible habitats, release from enemies, evolution of ‘invasiveness’, and invasional meltdown. We conclude that propagule pressure should serve as the basis of a null model for studies of biological invasions when inferring process from patterns of invasion. An erratum to this article can be found at     

The grass may not always be greener: projected reductions in climatic suitability for exotic grasses under future climates in Australia

Climate change presents a new challenge for the management of invasive exotic species that threaten both biodiversity and agricultural productivity. The invasion of exotic perennial grasses throughout the globe is particularly problematic given their impacts on a broad range of native plant communities and livelihoods. As the climate continues to change, pre-emptive long-term management strategies for exotic grasses will become increasingly important. Using species distribution modelling we investigated potential changes to the location of climatically suitable habitat for some exotic perennial grass species currently in Australia, under a range of future climate scenarios for the decade centred around 2050. We focus on eleven species shortlisted or declared as the Weeds of National Significance or Alert List species in Australia, which have also become successful invaders in other parts of the world. Our results indicate that the extent of climatically suitable habitat available for all of the exotic grasses modelled is projected to decrease under climate scenarios for 2050. This reduction is most severe for the three species of Needle Grass (genus Nassella) that currently have infestations in the south-east of the continent. Combined with information on other aspects of establishment risk (e.g. demographic rates, human-use, propagule pressure), predictions of reduced climatic suitability provide justification for re-assessing which weeds are prioritised for intensive management as the climate changes.     

Aquatic pollution increases the relative success of invasive species

Although individual ecosystems vary greatly in the degree to which they have been invaded by exotic species, it has remained difficult to isolate mechanisms influencing invader success. One largely anecdotal observation is that polluted or degraded areas will accumulate more invaders than less-impacted sites. However, the role of abiotic factors alone in influencing invisibility has been difficult to isolate, often because the supply of potential invaders is confounded with conditions thought to increase vulnerability to invasion. Here, we conducted a field experiment to test how the assemblages of exotic versus native marine invertebrates changed during community assembly under different exposure levels of a common pollutant, copper. The experiment was conducted by deploying fouling panels in a Randomized Block Design in San Francisco Bay. Panels were periodically removed, placed into buckets with differing copper concentrations, and returned to the field after 3 days. This design allowed propagule availability to the plates to be statistically independent of short-term copper exposure. The results demonstrate that copper caused significant differences in community structure. Average native species richness was significantly affected by copper exposure, but average exotic richness was not. The total native species pool within treatments exhibited a greater than 40% decline within increasing copper, while the exotic species pool did not change significantly. These results confirm that anthropogenic alteration of abiotic factors influences invader success, indicating that management strategies to reduce invader impacts should include both efforts to improve environmental conditions as well as reduce invader supply.     

Potential for successful population establishment of the nonindigenous sacred ibis in the Florida Everglades

The nonindigenous sacred ibis (Threskiornis aethiopicus) was first discovered breeding in the Florida Everglades in 2005 in the Arthur R. Marshall Loxahatchee National Wildlife Refuge. Prior to this, sacred ibises were seen periodically throughout South Florida since the mid 1990s, with occasional confirmed breeding occurrences in Miami-Dade and Palm Beach counties. We used a logistic regression model developed by Allen (Biol Invasions 8:491–500, 2006) to predict the probability of successful establishment of sacred ibis in the Florida Everglades ecosystem. Empirical data collected from several sacred ibis nests and chicks were used to validate those findings. The probability of successful establishment was estimated to be 73%. The physiological condition of nestlings suggested that this species was able to fledge chicks in good condition, thus adding to the potential to increase their breeding population. Exponential population growth rates and expanding distribution of the nonindigenous sacred ibis in France demonstrate this species’ potential for becoming invasive in Florida. We suggest that the most prudent and effective management strategy is eradication of the few pioneering individuals that are nesting in the Everglades as well as the urban source population.     

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.     

Exotic species richness and native species endemism increase the impact of exotic species on islands

Aim Exotic species pose one of the most significant threats to biodiversity, especially on islands. The impacts of exotic species vary in severity among islands, yet little is known about what makes some islands more susceptible than others. Here we determine which characteristics of an island influence how severely exotic species affect its native biota. Location We studied 65 islands and archipelagos from around the world, ranging from latitude 65° N to 54° S. Methods We compiled a global database of 10 island characteristics for 65 islands and determined the relative importance of each characteristic in predicting the impact of exotic species using multivariate modelling and hierarchical partitioning. We defined the impact of exotic species as the number of bird, amphibian and mammal (BAM) species listed by the International Union for Conservation of Nature (IUCN) as threatened by exotics, relative to the total number of BAM species on that island. Results We found that the impact of exotic species is more severe on islands with more exotic species and a greater proportion of native species that are endemic. Unexpectedly, the level of anthropogenic disturbance did not influence an island's susceptibility to the impacts of exotic species. Main conclusions By coupling our results with studies on the introduction and establishment of exotic species, we conclude that colonization pressure, or invasion opportunities, influences all stages of the invasion process. However, species endemism, the other important factor determining the impact of exotic species, is not known to contribute to introduction and establishment success on islands. This demonstrates that different factors correlate with the initial stages of the invasion process and the subsequent impacts of those invaders, highlighting the importance of studying the impacts of exotic species directly. Our study helps identify islands that are at risk of impact by exotics and where investment should focus on preventing further invasions.     

Intraspecific variation in allelochemistry determines an invasive species’ impact on soil microbial communities

Invasive species can benefit from altered species interactions in their new range, and by interfering with species interactions among native competitors. Since exotic invasions are generally studied at the species level, relatively little is known about intraspecific variation in the traits that determine an invader’s effect on native species. Alliaria petiolata is a widespread and aggressive invader of forest understories that succeeds in part by interfering with mutualistic interactions between native plants and soil fungi. Here, I show that the impact of A. petiolata on soil microbial communities varied among individuals due to variation in their allelochemical concentrations. The differential impacts translated into varied effects on native tree growth, partly because A. petiolata’s allelochemicals preferentially affected the most mutualistic fungal taxa. These results highlight the importance of considering the spatial and temporal variation in an invasive species’ impacts for understanding and managing the invasion process.     

Importation of Exotic Ticks into the United Kingdom via the International Trade in Reptiles

Due to the increased trade in exotic reptiles, many of which are infested with ticks of various genera, there is an increased risk of exotic ticks and pathogens being introduced and potentially becoming established in new geographical areas. Despite the number and variety of tick species collected from reptiles, their role in the maintenance of pathogens and transmission to humans has not been fully investigated for all species. The potential public health risks to the United Kingdom have been examined through a preliminary study involving the collection and identification of exotic tick species imported via the reptile trade. Reptiles imported into Heathrow Airport, London (July 2003–October 2004) were carefully examined for ectoparasites. Five species of tick were collected: Aponomma exornatum, Ap. latum, Amblyomma rotundatum, Am. dissimile and Am. nuttalli. This is the first report of Am. dissimile identified from an interception at Heathrow airport. The potential for establishment of exotic ticks in the UK and the public health risks are discussed.     

Woody exotic plant invasions and fire: reciprocal impacts and consequences for native ecosystems

Fire regimes influence and are influenced by the structure and composition of plant communities. This complex reciprocal relationship has implications for the success of plant invasions and the subsequent impact of invasive species on native biota. Although much attention has been given to the role of invasive grasses in transforming fire regimes and native plant communities, little is known about the relationship between woody invasive species and fire regime. Despite this, prescribed burning is frequently used for managing invasive woody species. In this study we review relationships between woody exotic plant invasions and fire in invaded ecosystems worldwide. Woody invaders may increase or decrease aspects of the fire regime, including fire frequency, intensity and extent. This is in contrast to grass invaders which almost uniformly increase fire frequency. Woody plant invasion can lead to escape from a grass-fire cycle, but the resulting reduction in fire frequency can sometimes lead to a cycle of rare but more intense fires. Prescribed fires may be a useful management tool for controlling woody exotic invaders in some systems, but they are rarely sufficient to eliminate an invasive species, and a dearth of controlled experiments hampers evaluation of their benefits. Nevertheless, because some woody invaders have fuel properties that differ substantially from native species, understanding and managing the impacts of woody invaders on fire regimes and on prescribed burns should become an important component of resource and biodiversity management.     

Co-invasion of similar invaders results in analogous ecological impact niches and no synergies

Exotic species can cause changes to their invaded ecosystems, which can be large and long lasting. Despite most landscapes being invaded by multiple exotic plant species, >90 % of impact studies only characterize the impacts of single species. Therefore, our knowledge of invasive plant impacts does not reflect the co-invaded nature of most landscapes, potentially ignoring complex interactions among exotic species. Our objective was to characterize potential invader synergies (positive interactions) on biotic and abiotic ecological parameters among the important forest invaders Japanese stiltgrass (Microstegium vimineum) and wavyleaf basketgrass (Oplismenus undulatifolius), which co-invade eastern US deciduous forests. To characterize synergies, we used a factorial selective removal study, as well as an observational study to further explore invader cover-impact relationships. Although both invaders can reduce native plant richness by 70 % individually or in combination, there were no impact synergies. Total cover of any combination of the two invaders had a negative quadratic effect on total, exotic, and native plant richness; i.e., all community metrics were greatest at intermediate levels of total invader cover and lowest at maximum invader cover. Native richness was more greatly affected than exotic richness by the co-invasion. Soil metrics had no clear trend in either study. Japanese stiltgrass and wavyleaf basketgrass appear to have overlapping impact niches—the number, magnitude, and direction of biotic and abiotic changes to the invaded ecosystem—that only vary in impact magnitude, not breadth. As a result of their overlapping impact niches and non-synergies in this co-invaded system, the addition of the recent invader wavyleaf basketgrass has not resulted in additional changes to the invaded forests. Future impact studies should focus on multiple species and identifying synergies, especially as they relate to invader cover, which informs ecological interactions and management prioritization.     

Ecological and environmental factors constrain sprouting ability in tropical trees

Most theories of forest biodiversity focus on the role of seed dispersal and seedling establishment in forest regeneration. In many ecosystems, however, sprouting by damaged stems determines which species occupies a site. Damaged trees can quickly recover from disturbance and out-compete seedlings. Links among species’ traits, environmental conditions and sprouting could offer insight into species’ resilience to changes in climate, land use, and disturbance. Using data for 25 Neotropical tree species at two sites with contrasting rainfall and soil, we tested hypotheses on how four functional traits (seed mass, leaf mass per area, wood density and nitrogen fixation) influence species’ sprouting responses to disturbance and how these relationships are mediated by a tree’s environmental context. Most species sprouted in response to cutting, and many species’ sprouting rates differed significantly between sites. Individual traits showed no direct correlation with sprouting. However, interactions among traits and site variables did affect sprouting rates. Many species showed increased sprouting in the higher-quality site. Most nitrogen-fixing species showed the opposite trend, sprouting more frequently where resources are scarce. This study highlights the use of functional traits as a proxy for life histories, and demonstrates the importance of environmental effects on demography.     

Invasive alien Crustacea: dispersal,establishment, impact and control

The subphylum Crustacea includes the most successful species among aquatic alien invaders. The impacts of invasive alien crustaceans (IAC) are often substantial, due to the complex trophic role of most of these species leading to cascading effects throughout the invaded ecosystems. IAC also have the potential to cause a shift in the ‘keystone’ ecosystem functions, changing energy flux and nutrient cycles which together affect critical ecosystem services such as biodiversity, fisheries yield and water quality. Although no individual trait appears to be a good predictor of invasion success, a combination of some characteristics such as eurytolerance, omnivory and certain r-selected life-history traits results in a high probability of alien crustacean species becoming invasive. Both environmental factors, such as habitat heterogeneity in the invaded ecosystems, and evolutionary factors, such as adaptations to new environmental conditions, also play important roles during establishment. Therefore, individual environmental niche models, including genetic algorithm, have the highest likelihood of providing useful predictive information about invasion success and spread of alien Crustacea. Attempts to control IAC through biocides or mechanical removal have had mixed success in the past but a strategic combination of different methods may lead to some success in the future.     

Patterns of non-randomness in the exotic avifauna of Florida

It is now generally recognized that human-mediated biological invasion is a multistage process, successively comprising transport, introduction, establishment, and spread, and that a complete understanding of the causes of invasion requires studies of all stages. However, while many studies address the characteristics that influence establishment, relatively few address the characteristics that influence whether or not a species transits the earlier stages of transport and introduction. Here, we use data on the rich exotic avifauna of Florida to assess non-randomness in the identities of species that have passed through the transport and introduction stages. Bird species transported and introduced to Florida are non-random with respect to their taxonomic affiliations, body mass, native geographical range size, and region of origin: introductions are more likely for widespread, large-bodied species from the Neotropics and belonging to the Anatidae, Psittacidae, Ciconiidae, and Passeridae. Data on the identities of species that have attempted to breed but failed, and on the breeding population size for most established species, also allowed us to assess the extent to which the same variables influenced various aspects of post-introduction establishment. Only native geographical range size and latitudinal range mid-point distinguish between these different classes of exotic species. Geographical range size is the most general correlate of different classes of invaders in our analyses.     

A predictive framework and review of the ecological impacts of exotic plant invasions on reptiles and amphibians

The invasive spread of exotic plants in native vegetation can pose serious threats to native faunal assemblages. This is of particular concern for reptiles and amphibians because they form a significant component of the world's vertebrate fauna, play a pivotal role in ecosystem functioning and are often neglected in biodiversity research. A framework to predict how exotic plant invasion will affect reptile and amphibian assemblages is imperative for conservation, management and the identification of research priorities. Here, we present a new predictive framework that integrates three mechanistic models. These models are based on exotic plant invasion altering: (1) habitat structure; (2) herbivory and predator‐prey interactions; (3) the reproductive success of reptile and amphibian species and assemblages. We present a series of testable predictions from these models that arise from the interplay over time among three exotic plant traits (growth form, area of coverage, taxonomic distinctiveness) and six traits of reptiles and amphibians (body size, lifespan, home range size, habitat specialisation, diet, reproductive strategy). A literature review provided robust empirical evidence of exotic plant impacts on reptiles and amphibians from each of the three model mechanisms. Evidence relating to the role of body size and diet was less clear‐cut, indicating the need for further research. The literature provided limited empirical support for many of the other model predictions. This was not, however, because findings contradicted our model predictions but because research in this area is sparse. In particular, the small number of studies specifically examining the effects of exotic plants on amphibians highlights the pressing need for quantitative research in this area. There is enormous scope for detailed empirical investigation of interactions between exotic plants and reptile and amphibian species and assemblages. The framework presented here and further testing of predictions will provide a basis for informing and prioritising environmental management and exotic plant control efforts.     

Native granivores reduce the establishment of native grasses but not invasive <Emphasis Type="Italic">Bromus tectorum</Emphasis>

Seed predation can structure plant communities by imposing strong population controls on some species but not others. In this context, studies from various ecosystems report that native granivores selectively forage for seeds from native species over seeds from exotic invaders, which could disproportionately favor the establishment of invaders and facilitate their dominance in communities. However, few studies have connected selective foraging for native seeds to differential patterns of establishment among native and invasive species. Thus, the extent to which preferential foraging for native seeds favors the establishment of invasive plants is unclear. Here, we used experimental seed additions and exclosure treatments at five field sites distributed across?≈?80,000 km² of the Great Basin Desert, USA to compare the effects of rodent foraging on the establishment of less-preferred cheatgrass (Bromus tectorum—an annual species native to Eurasia that is exotic and highly invasive across the Great Basin) and four species of more-preferred native grasses that commonly co-occur with cheatgrass. Rodent foraging reduced the establishment of each native species by at least 80% but had no effect on the establishment of cheatgrass, and this finding was consistent across study sites. Our results suggest that selective foraging for native species may favor the establishment of cheatgrass over native grasses, potentially exacerbating one of the most extensive plant invasions in North America.     

Regeneration niche differentiates functional strategies of desert woody plant species

Plant communities vary dramatically in the number and relative abundance of species that exhibit facilitative interactions, which contributes substantially to variation in community structure and dynamics. Predicting species’ responses to neighbors based on readily measurable functional traits would provide important insight into the factors that structure plant communities. We measured a suite of functional traits on seedlings of 20 species and mature plants of 54 species of shrubs from three arid biogeographic regions. We hypothesized that species with different regeneration niches—those that require nurse plants for establishment (beneficiaries) versus those that do not (colonizers)—are functionally different. Indeed, seedlings of beneficiary species had lower relative growth rates, larger seeds and final biomass, allocated biomass toward roots and height at a cost to leaf mass fraction, and constructed costly, dense leaf and root tissues relative to colonizers. Likewise at maturity, beneficiaries had larger overall size and denser leaves coupled with greater water use efficiency than colonizers. In contrast to current hypotheses that suggest beneficiaries are less “stress-tolerant” than colonizers, beneficiaries exhibited conservative functional strategies suited to persistently dry, low light conditions beneath canopies, whereas colonizers exhibited opportunistic strategies that may be advantageous in fluctuating, open microenvironments. In addition, the signature of the regeneration niche at maturity indicates that facilitation expands the range of functional diversity within plant communities at all ontogenetic stages. This study demonstrates the utility of specific functional traits for predicting species’ regeneration niches in hot deserts, and provides a framework for studying facilitation in other severe environments.