Deciphering the worldwide invasion of the Asian long‐horned beetle: A recurrent invasion process from the native area together with a bridgehead effect

Retracing introduction routes is crucial for understanding the evolutionary processes involved in an invasion, as well as for highlighting the invasion history of a species at the global scale. The Asian long‐horned beetle (ALB) Anoplophora glabripennis is a xylophagous pest native to Asia and invasive in North America and Europe. It is responsible for severe losses of urban trees, in both its native and invaded ranges. Based on historical and genetic data, several hypotheses have been formulated concerning its invasion history, including the possibility of multiple introductions from the native zone and secondary dispersal within the invaded areas, but none have been formally tested. In this study, we characterized the genetic structure of ALB in both its native and invaded ranges using microsatellites. In order to test different invasion scenarios, we used an approximate Bayesian “random forest” algorithm together with traditional population genetics approaches. The strong population differentiation observed in the native area was not geographically structured, suggesting complex migration events that were probably human‐mediated. Both native and invasive populations had low genetic diversity, but this characteristic did not prevent the success of the ALB invasions. Our results highlight the complexity of invasion pathways for insect pests. Specifically, our findings indicate that invasive species might be repeatedly introduced from their native range, and they emphasize the importance of multiple, human‐mediated introductions in successful invasions. Finally, our results demonstrate that invasive species can spread across continents following a bridgehead path, in which an invasive population may have acted as a source for another invasion.     

Pine invasions: climate predicts invasion success; something else predicts failure

Aim Explaining why some invasions fail while others succeed is a prevailing question in invasion biology. Different factors have been proposed to explain the success or failure of exotics. Evidence suggests that climate similarities may be crucial. We tested this using 12 species of the genus Pinus that have been widely planted and shown to be highly invasive. Pinus is among the best‐studied group of exotic species and one that has been widely introduced world‐wide, so we were able to obtain data on invasive and non‐invasive introductions (i.e. unsuccessful invasions; areas where after many decades of self‐sowing seeds there is no invasion). Location World‐wide. Methods We developed species distribution models for native ranges using a maximum entropy algorithm and projected them across the globe. We tested whether climate‐based models were able to predict both invasive and non‐invasive introductions. Results Appropriate climatic conditions seem to be required for these long‐lived species to invade because climates accurately predicted invasions. However, climate matching is necessary, but not sufficient to predict the fate of an introduction because most non‐invasive introductions were predicted to have triggered an invasion. Main conclusions Other factors, possibly including biotic components, may be the key to explaining why some introductions do not become invasions, because many areas where Pinus is not invading were predicted to be suitable for invasion based solely on climate.     

Niche dynamics of two cryptic <Emphasis Type="Italic">Prosopis</Emphasis> invading South American drylands

Whether or not species track native climatic conditions during invasions (i.e., climate match hypothesis) is fundamental to understand and prevent potential impacts of invasive species. Recent empirical work suggests that climatic mismatches between native and invasive ranges are pervasive. Whether these differences are due to adaptation to new climatic spaces in the invasive range or due to partial filling of the potential climatic space are still subject to debate. Here, we analyze climatic niche dynamics associated with the invasion of the two most common invasive plants in Brazilian semi-arid areas, Prosopis juliflora and Prosopis pallida. These species have been simultaneously introduced in the region, which creates a unique opportunity to compare their niche dynamics during invasion. Given that P. juliflora have a much wider native range size, we expect these species would present different dispersal potentials, which might translate into different unfilling levels. Using an ordination method with kernel smoother and null models, we contrasted climate spaces occupied by each species in both native and invasive ranges. We further used ecological niche models (ENMs) to compare reciprocal predictions of potentially suitable areas. Against our expectation based on differences in native range sizes, climatic niches of P. juliflora and P. pallida overlapped greatly, both in their native and invasive ranges. Our results support niche conservatism during the invasion process. Climatic mismatches among native and invaded ranges were exclusively attributed to unfilling of native climates in the invasive range. Both species showed similar unfilling levels. Likewise, ENMs predicted regions not yet occupied in the invasive range, revealing a potential for further expansion. We discuss colonization time lag and founder effect as potential mechanisms that may have prevented these species to fully occupy their native niches in the invasive range.     

Severity of impacts of an introduced species corresponds with regional eco‐evolutionary experience

Invasive plant impacts vary widely across introduced ranges. We tested the hypothesis that differences in the eco‐evolutionary experience of native communities with the invader correspond with the impacts of invasive species on native vegetation, with impacts increasing with ecological novelty. We compared plant species richness and composition beneath Pinus contorta to that in adjacent vegetation and other P. contorta stands across a network of sites in its native (Canada and USA) and non‐native (Argentina, Chile, Finland, New Zealand, Scotland, Sweden) ranges. At sites in North America and Europe, within the natural distribution of the genus Pinus, P. contorta was not associated with decreases in diversity. In the Southern Hemisphere, where there are no native Pinaceae, plant communities beneath P. contorta were less diverse than in other regions and compared to uninvaded native vegetation. Effects on native vegetation were particularly pronounced where P. contorta was a more novel life form and exhibited higher growth rates. Our results support the hypothesis that the eco‐evolutionary experience of the native vegetation, and thus the novelty of the invader, determines the magnitude of invader impacts on native communities. Understanding the eco‐evolutionary context of invasions will help to better understand and predict where invasion impacts will be greatest and to prioritize invasive species management.     

Macroecology meets invasion ecology: performance of Australian acacias and eucalypts around the world revealed by features of their native ranges

Native geographical range extent has frequently emerged as a correlate of invasiveness, especially for plant species. We tested whether dimensions of the native range (measured by the area-of-occupancy and its scaling patterns) of 720 Australian eucalypts (genera Angophora, Eucalyptus and Corymbia) could explain introduction and invasion success. We also compared our results with a previous study on 979 Australian acacias (previously grouped in Acacia subgenus Phyllodineae) to investigate whether features of their native ranges explained the much higher invasion success in this group compared to eucalypts. From nine databases we found records that 373 eucalypts have been introduced to areas outside their native ranges; 82 of these have become naturalised, and eight are invasive. A similar proportion of Australian acacias have been introduced, but almost three times as many are invasive (384 species introduced, 71 of which are naturalized and 23 invasive). Eucalypts with large native ranges are more likely to have been introduced and subsequently naturalise, as is the case with acacias. Unlike acacias, however, the native range size of invasive eucalypts was not significantly greater than naturalised (but not invasive) species. Intriguingly, the human preference for introducing species with larger ranges was much greater for acacias than for eucalypts as the geometric mean range sizes of introduced, naturalised and invasive acacias are 2.04, 1.88 and 3.59 times those of eucalypts at the same stage. Moreover, the percolation exponent (i.e. the slope of occupancy scaling) becomes more extreme towards the end of the introduction–naturalization–invasion continuum, decreasing for acacias but increasing for eucalypts, with acacias, except for non-introduced species, having a lower exponent than eucalypts. The selection preference of acacias during introduction is thus for species that can rapidly expand their range; in contrast, slow-spreading eucalypts have been selected for dissemination. In other words, humans appear to have selected for highly invasive acacias but against introducing highly invasive eucalypts.     

Thermogeography predicts the potential global range of the invasive European green crab (Carcinus maenas)

Aim The highly adaptable estuarine crab (Carcinus maenas) has successfully invaded five temperate geographic regions outside of its native Europe. Here, we determine which environmental factors predict the current distribution of C. maenas and what the potential geographic range of this species might be. We also investigated whether the invasion potential of C. maenas differs with respect to the origin of a native subpopulation. Location Models were developed using global observation records of C. maenas. Methods Boosted regression trees were used to model observations from the (1) native, (2) invasive, (3) southern European, (4) northern European and (5) the combined native and invasive geographic ranges of C. maenas. Results Most established invasions were predicted mainly based on temperature. Interestingly, the environment encountered by established invasions failed to predict the majority of northern European populations; suggesting that invasion potential may differ between distinct native populations. Supporting this suggestion, a model of northern European populations, distinguished from southern European populations based on genetic structure, only predicted established invasions south of Nova Scotia. By contrast, a model of southern European populations predicted most established invasions. Main conclusions These results suggest that invasion potential depends on the European origin of an invasive population and that most invasions have arisen from southern Europe. Finally, a model based on combined native and invasive ranges of C. maenas identified potential geographic range extension along many currently invaded coastlines and the potential invasion of countries like Chile, China, Russia, Namibia and New Zealand.     

Invasion trajectory of alien trees: the role of introduction pathway and planting history

Global change is driving a massive rearrangement of the world's biota. Trajectories of distributional shifts are shaped by species traits, the recipient environment and driving forces with many of the driving forces directly due to human activities. The relative importance of each in determining the distributions of introduced species is poorly understood. We consider 11 Australian Acacia species introduced to South Africa for different reasons (commercial forestry, dune stabilization and ornamentation) to determine how features of the introduction pathway have shaped their invasion history. Projections from species distribution models (SDMs) were developed to assess how the reason for introduction influences the similarity between climatic envelopes in native and alien ranges. A lattice model for an idealized invasion was developed to assess the relative contribution of intrinsic traits and introduction dynamics on the abundance and extent over the course of simulated invasions. SDMs show that alien populations of ornamental species in South Africa occupy substantially different climate space from their native ranges, whereas species introduced for forestry occupy a similar climate space in native and introduced ranges. This may partly explain the slow spread rates observed for some alien ornamental plants. Such mismatches are likely to become less pronounced with the current drive towards ‘eco gardens’ resulting in more introductions of ornamental species with a close climate match between native and newly introduced regions. The results from the lattice model showed that the conditions associated with the introduction pathway (especially introduction pressure) dominate early invasion dynamics. The placement of introduction foci in urban areas limited the extent and abundance of invasive populations. Features of introduction events appear to initially mask the influence of intrinsic species traits on invasions and help to explain the relative success of species introduced for different purposes. Introduction dynamics therefore can have long‐lasting influences on the outcomes of species redistributions, and must be explicitly considered in management plans.     

No evidence for evolutionarily decreased tolerance and increased fitness in invasive Chromolaena odorata: implications for invasiveness and biological control

Tolerance, the degree to which plant fitness is affected by herbivory, is associated with invasiveness and biological control of introduced plant species. It is important to know the evolutionary changes in tolerance of invasive species after introduction in order to understand the mechanisms of biological invasions and assess the feasibility of biological control. While many studies have explored the evolutionary changes in resistance of invasive species, little has been done to address tolerance. We hypothesized that compared with plants from native populations, plants from invasive populations may increase growth and decrease tolerance to herbivory in response to enemy release in introduced ranges. To test this hypothesis, we compared the differences in growth and tolerance to simulated herbivory between plants from invasive and native populations of Chromolaena odorata, a noxious invader of the tropics and subtropics, at two nutrient levels. Surprisingly, flower number, total biomass (except at high nutrient), and relative increase in height were not significantly different between ranges. Also, plants from invasive populations did not decrease tolerance to herbivory at both nutrient levels. The invader from both ranges compensated fully in reproduction after 50?% of total leaf area had been damaged, and achieved substantial regrowth after complete shoot damage. This strong tolerance to damage was associated with increased resource allocation to reproductive structures and with mobilization of storage reserves in roots. The innately strong tolerance may facilitate invasion success of C. odorata and decrease the efficacy of leaf-feeding biocontrol agents. Our study highlights the need for further research on biogeographical differences in tolerance and their role in the invasiveness of exotic plants and biological control.     

Origins and distribution of invasive Rubus fruticosus L. agg. (Rosaceae) clones in the Western United States

In facultatively asexual invasive species, an understanding of the origins and diversity of clones can reveal introduction and invasion pathways and inform management efforts. In this study, we use microsatellite and chloroplast DNA markers to infer clonal diversity of the Rubus fruticosus agg. invasion in the Western United States, determine the relationship of these clones to clones found in other exotic ranges, and determine the geographic and genetic origins of the invasive clones. We found two invasive clones in the Western United States, where the invasion had previously been thought to consist of a single asexual lineage. The most common clone was genetically identical to the microspecies R. armeniacus from the native range of Germany, while the second clone was identical to the microspecies R. anglocandicans in the invaded range of Australia and closely related to samples from the native ranges of England and Serbia. A third distinct clone was identified in a collection from the exotic range of Chile. Our results demonstrate that cryptic genetic diversity may be present in asexual invasions that are thought to be homogeneous. However, the asexual relationships between R. fruticosus agg. clones in the native and multiple exotic ranges indicate that preadaptation has played an important role in invasion success in this species aggregate.     

Highly diverse and highly successful: invasive Australian acacias have not experienced genetic bottlenecks globally

Background and AimsInvasive species may undergo rapid evolution despite very limited standing genetic diversity. This so-called genetic paradox of biological invasions assumes that an invasive species has experienced (and survived) a genetic bottleneck and then underwent local adaptation in the new range. In this study, we test how often Australian acacias (genus Acacia), one of the world’s most problematic invasive tree groups, have experienced genetic bottlenecks and inbreeding.MethodsWe collated genetic data from 51 different genetic studies on Acacia species to compare genetic diversity between native and invasive populations. These studies analysed 37 different Acacia species, with genetic data from the invasive ranges of 11 species, and data from the native range for 36 species (14 of these 36 species are known to be invasive somewhere in the world, and the other 22 are not known to be invasive).Key ResultsLevels of genetic diversity are similar in native and invasive populations, and there is little evidence of invasive populations being extensively inbred. Levels of genetic diversity in native range populations also did not differ significantly between species that have and that do not have invasive populations.ConclusionWe attribute our findings to the impressive movement, introduction effort and human usage of Australian acacias around the world.     

Molecular genetic approaches to elucidate the ecological and evolutionary issues associated with biological invasions

Biological invasions may cause serious damage to the native environments and threaten the native biodiversity. Molecular genetic approaches have been found to be powerful tools for investigating the ecological and evolutionary aspects of biological invasions because the genetic structure and level of genetic variation of an invasive species are changed following its invasion. The present article reviews the use of molecular markers in addressing various aspects of invasive species. The application of these techniques has shown that many invasive species are actually "cryptic" species – species whose uniqueness is only recognizable at the genetic level. An estimation of the actual number of invasive species is essential when evaluating its ecological and economic impacts. Molecular genetic approaches have also enabled the source populations of invasive species to be identified. Reconstructions of invasion histories are crucial to preventing future invasions and conserving the native biodiversity, while comparisons of genetic variations between the native and introduced populations provide valuable opportunities to elucidate the mechanisms of rapid adaptation demonstrated by many invasive species.     

Rapid evolution and range expansion of an invasive plant are driven by provenance–environment interactions

To improve our ability to prevent and manage biological invasions, we must understand their ecological and evolutionary drivers. We are often able to explain invasions after they happen, but our predictive ability is limited. Here, we show that range expansions of introduced Pinus taeda result from an interaction between genetic provenance and climate and that temperature and precipitation clines predict the invasive performance of particular provenances. Furthermore, we show that genotypes can occupy climate niche spaces different from those observed in their native ranges and, at least in our case, that admixture is not a main driver of invasion. Genotypes respond to climate in distinct ways, and these interactions affect the ability of populations to expand their ranges. While rapid evolution in introduced ranges is a mechanism at later stages of the invasion process, the introduction of adapted genotypes is a key driver of naturalisation of populations of introduced species.     

Effects of admixture in native and invasive populations of <Emphasis Type="Italic">Lythrum salicaria</Emphasis>

Intraspecific hybridization between diverged populations can enhance fitness via various genetic mechanisms. The benefits of such admixture have been proposed to be particularly relevant in biological invasions, when invasive populations originating from different source populations are found sympatrically. However, it remains poorly understood if admixture is an important contributor to plant invasive success and how admixture effects compare between invasive and native ranges. Here, we used experimental crosses in Lythrum salicaria, a species with well-established history of multiple introductions to Eastern North America, to quantify and compare admixture effects in native European and invasive North American populations. We observed heterosis in between-population crosses both in native and invasive ranges. However, invasive-range heterosis was restricted to crosses between two different Eastern and Western invasion fronts, whereas heterosis was absent in geographically distant crosses within a single large invasion front. Our results suggest that multiple introductions have led to already-admixed invasion fronts, such that experimental crosses do not further increase performance, but that contact between different invasion fronts further enhances fitness after admixture. Thus, intra-continental movement of invasive plants in their introduced range has the potential to boost invasiveness even in well-established and successfully spreading invasive species.     

Habitat, predation, and coexistence between invasive and native crayfishes: prioritizing lakes for invasion prevention

Conditions fostering coexistence of native species with invasive species have received little attention in invasion biology, especially for closely related invasive and native species. We used long-term datasets on multiple replicate invasions to define conditions under which native virile crayfish (Orconectes virilis) can coexist with invasive rusty crayfish (O. rusticus). We examined multiple drivers of coexistence involving habitat use and predation at between-lake and within-lake scales to derive predictions that could guide prioritization efforts to prevent future introductions of rusty crayfish and mitigate impacts of existing invasions. Lakes in which native species persisted for many years had significantly less cobble and sand habitats, and significantly more vegetated habitats compared to lakes from which native crayfish have been displaced. In the presence of rusty crayfish, virile crayfish alter their habitat use to vegetated habitats relative to habitat use in the absence of rusty crayfish. Such vegetated habitats had greater plant standing crop, plant species richness, and sediment percent organic matter compared to vegetated sites occupied by rusty crayfish. Our results suggest that low abundance of cobble habitat and altered habitat use allows native crayfish to coexist with the rusty crayfish invader. At the within-lake scale, virile crayfish persist by escaping predation in the vegetated habitats, despite suboptimal abiotic conditions. By understanding these abiotic and biotic conditions that promote coexistence, managers could enhance native crayfish persistence by targeting high cobble lakes for efforts to prevent the introduction of invasive crayfish, and targeting vegetated habitats for protection in already invaded lakes.     

Phylogeographic analysis of invasive Asian earthworms (Amynthas) in the northeast United States

Phylogeographic studies are useful in reconstructing the history of species invasions, and in some instances can elucidate cryptic diversity of invading taxa. This can help in predicting or managing the spread of invasive species. Among terrestrial invasive species in North America, earthworms can have profound ecological effects. We are familiar with the centuries‐old invasions of European earthworms (Lumbricidae) and their impacts on nutrient cycling in soils. More recent invasions by Asian earthworms of the family Megascolecidae are less fully understood. We used data for two mitochondrial gene fragments, cytochrome oxidase I (COI) and 16S rRNA, to examine the relationships among populations of Asian earthworms in the megascolecid genus Amynthas in the northeast United States. Recent reports have indicated that one species in particular, Amynthas agrestis, is having detrimental effects in mixed forest ecosystems, and we were interested in understanding the invasion history for this species. We were surprised to discover three divergent mitochondrial lineages of Amynthas occurring sympatrically in upstate New York. Given the gap between intra‐ and inter‐lineage sequence divergences, we propose that these three lineages represent cryptic species of Amynthas, one of which is A. agrestis. For each of the three lineages of Amynthas, we observed shared haplotypes across broad geographic distances. This may reflect common origins for populations in each lineage, either by direct routes from native ranges or through post‐introduction spread by natural dispersal or human‐mediated transport within North America. Management efforts focused on horticultural imports from Asia, commercial nurseries within the USA, and on prohibition of bait disposal may help to reduce the further invasion success of Amynthas.     

Invaders versus endemics: alien gastropod species in ancient Lake Ohrid

Species invasions into ancient lakes are an important but little understood phenomenon. At ancient Lake Ohrid, a systematic assessment of invasive mollusc species using morphological and genetic data was conducted from 2003 to 2012. Two globally invasive gastropod species, Physa acuta and Ferrissia fragilis, have recently been discovered at 4 out of 386 sites. These sites are anthropogenically impacted. The invasive species co-occur with endemics. Phylogenetic analyses of populations from native and invaded ranges of both species confirmed their identities and provided insights into their invasion histories. Accordingly, P. acuta is genetically more diverse than F. fragilis. Both species are currently present in a considerable number of lakes on the Balkan Peninsula. Possible future trends in Lake Ohrid and the Balkans are discussed and further spread of both species is likely. Given the ongoing environmental change in Lake Ohrid, the number of observations of non-indigenous or other widespread species will probably rise in the coming years and such species and their impact on native species should be carefully monitored. Moreover, ancient lakes with recurrent invasions of alien species might serve as interesting model systems for the study of important topics of invasion biology.     

Biogeographic differences between native and non-native populations of crayfish alter species coexistence and trophic interactions in mesocosms

Biogeographical comparisons of native and non-native populations allow researchers to understand the degree to which traits contributing to invasion success are intrinsic or change during the invasion process. Here, we investigate whether traits underlying interspecific competition change following invasion and whether these alter the impacts of two crayfish congeners that have invaded into each other’s native ranges. Specifically, we compared native and non-native populations of rusty (Faxonius rusticus) and virile crayfish (F. virilis). We compared native and non-native populations of each species using laboratory assays to examine aggression and large mesocosms with the congeners in sympatry to examine growth and survival as well as impacts on lower trophic levels. We found that non-native virile crayfish were more aggressive in response to a threat than native virile crayfish and exhibited greater growth and survival in sympatry with rusty crayfish. These intraspecific differences were large enough to alter coexistence between species in the mesocosm experiment, which is consistent with patterns of coexistence between these species in the field. We did not observe differences in traits between native and non-native rusty crayfish, but rusty crayfish were consistently competitively dominant over virile crayfish in paired laboratory assays. Non-native populations of both species had greater impacts on lower trophic levels than native populations. Taken together, these findings provide new evidence that trait changes during invasions may enhance ecological impacts of invasive animals and their ability to compete with closely related native species.     

Genetic analysis reveals multiple cryptic invasive species of the hydrozoan genus Cordylophora

Understanding the patterns and dynamics of biological invasions is a crucial prerequisite to predicting and mitigating their potential ecological and economic impacts. Unfortunately, in many cases such understanding is limited not only by ignorance of invasion history, but also by uncertainty surrounding the ecology, physiology, and even systematics of the invasive taxa themselves. The invasive, colonial euryhaline hydroid Cordylophora has invaded multiple regions outside of its native Ponto-Caspian range. However, extensive morphological and ecological plasticity has prevented consensus on both species-level classification within the genus and the environmental conditions conducive to establishment. The goal of this research was to explore the invasive history and species composition of the genus Cordylophora through molecular analyses. We addressed both issues using DNA sequence data from two mitochondrial loci [the small subunit 16S rRNA and cytochrome c oxidase subunit I (COI)] and one nuclear locus (28S large nuclear rRNA), generated from 27 invasive Cordylophora populations collected throughout the global range of the taxon. Phylogenetic analysis and comparisons of genetic distances between populations suggest the presence of multiple cryptic species within the genus. This conclusion is further supported by the observation of significantly different habitat preferences between invasive lineages. Geographic distribution of lineages is consistent with the introduction of multiple lineages to some non-native regions, indicating that repeated introductions may contribute to the current global distribution of Cordylophora. Applying molecular and morphological analyses to additional populations of Cordylophora is likely to assist in clarifying the taxonomy of this genus and in providing a better understanding of the invasive history of this hydroid.     

Invasion Is a Community Affair: Clandestine Followers in the Bacterial Community Associated to Green Algae,Caulerpa racemosa,Track the Invasion Source

Biological invasions rank amongst the most deleterious components of global change inducing alterations from genes to ecosystems. The genetic characteristics of introduced pools of individuals greatly influence the capacity of introduced species to establish and expand. The recently demonstrated heritability of microbial communities associated to individual genotypes of primary producers makes them a potentially essential element of the evolution and adaptability of their hosts. Here, we characterized the bacterial communities associated to native and non-native populations of the marine green macroalga Caulerpa racemosa through pyrosequencing, and explored their potential role on the strikingly invasive trajectory of their host in the Mediterranean. The similarity of endophytic bacterial communities from the native Australian range and several Mediterranean locations confirmed the origin of invasion and revealed distinct communities associated to a second Mediterranean variety of C . racemosa long reported in the Mediterranean. Comparative analysis of these two groups demonstrated the stability of the composition of bacterial communities through the successive steps of introduction and invasion and suggested the vertical transmission of some major bacterial OTUs. Indirect inferences on the taxonomic identity and associated metabolism of bacterial lineages showed a striking consistency with sediment upheaval conditions associated to the expansion of their invasive host and to the decline of native species. These results demonstrate that bacterial communities can be an effective tracer of the origin of invasion and support their potential role in their eukaryotic host’s adaptation to new environments. They put forward the critical need to consider the ''meta-organism'' encompassing both the host and associated micro-organisms, to unravel the origins, causes and mechanisms underlying biological invasions.     

Can Invasive Species Facilitate Native Species? Evidence of How, When, and Why These Impacts Occur

Although the predatory and competitive impacts of biological invasions are well documented, facilitation of native species by non-indigenous species is frequently overlooked. A search through recent ecological literature found that facilitative interactions between invasive and native species occur in a wide range of habitats, can have cascading effects across trophic levels, can re-structure communities, and can lead to evolutionary changes. These are critical findings that, until now, have been mostly absent from analyses of ecological impacts of biological invasions. Here I present evidence for several mechanisms that exemplify how exotic species can facilitate native species. These mechanisms include habitat modification, trophic subsidy, pollination, competitive release, and predatory release. Habitat modification is the most frequently documented mechanism, reflecting the drastic changes generated by the invasion of functionally novel habitat engineers. Further, I predict that facilitative impacts on native species will be most likely when invasive species provide a limiting resource, increase habitat complexity, functionally replace a native species, or ameliorate predation or competition. Finally, three types of facilitation (novel, substitutive, and indirect) define why exotic species can lead to facilitation of native species. It is evident that understanding biological invasions at the community and ecosystem levels will be more accurate if we integrate facilitative interactions into future ecological research. Since facilitative impacts of biological invasions can occur with native endangered species, and can have wide-ranging impacts, these results also have important implications for management, eradication, and restoration.Contribution Number 2293, Bodega Marine Laboratory, University of California at Davis.