Determinants of vertebrate invasion success in Europe and North America

Species that are frequently introduced to an exotic range have a high potential of becoming invasive. Besides propagule pressure, however, no other generally strong determinant of invasion success is known. Although evidence has accumulated that human affiliates (domesticates, pets, human commensals) also have high invasion success, existing studies do not distinguish whether this success can be completely explained by or is partly independent of propagule pressure. Here, we analyze both factors independently, propagule pressure and human affiliation. We also consider a third factor directly related to humans, hunting, and 17 traits on each species' population size and extent, diet, body size, and life history. Our dataset includes all 2362 freshwater fish, mammals, and birds native to Europe or North America. In contrast to most previous studies, we look at the complete invasion process consisting of (1) introduction, (2) establishment, and (3) spread. In this way, we not only consider which of the introduced species became invasive but also which species were introduced. Of the 20 factors tested, propagule pressure and human affiliation were the two strongest determinants of invasion success across all taxa and steps. This was true for multivariate analyses that account for intercorrelations among variables as well as univariate analyses, suggesting that human affiliation influenced invasion success independently of propagule pressure. Some factors affected the different steps of the invasion process antagonistically. For example, game species were much more likely to be introduced to an exotic continent than nonhunted species but tended to be less likely to establish themselves and spread. Such antagonistic effects show the importance of considering the complete invasion process.     

Residence time,native range size,and genome size predict naturalization among angiosperms introduced to Australia

Although critical to progress in understanding (i) if, and (ii) at what rate, introduced plants will naturalize and potentially become invasive, establishing causal links between traits and invasion success is complicated by data gaps, phylogenetic nonindependence of species, the inability to control for differences between species in residence time and propagule pressure, and covariance among traits. Here, we focus on statistical relationships between genomic factors, life history traits, native range size, and naturalization status of angiosperms introduced to Australia. In a series of analyses, we alternately investigate the role of phylogeny, incorporate introduction history, and use graphical models to explore the network of conditional probabilities linking traits and introduction history to naturalization status. Applying this ensemble of methods to the largest publicly available data set on plant introductions and their fates, we found that, overall, residence time and native range size best predicted probability of naturalization. Yet, importantly, probability of naturalization consistently increased as genome size decreased, even when the effects of shared ancestry and residence time in Australia were accounted for, and that this pattern was stronger in species without a history of cultivation, but present across annual–biennials, and herbaceous and woody perennials. Thus, despite introduction biases and indirect effects of traits via introduction history, across analyses, reduced genome size was nevertheless consistently associated with a tendency to naturalize.     

Tracing the introduction history of a potentially invasive ornamental shrub: variation in frost hardiness and climate change

Nursery catalogues have recently been used as a method for tracing the introduction history of invasive plants. Information on species, cultivars, plant size and price can help quantify historical changes in propagule pressure. Propagule pressure of invasive ornamentals has to be combined with climatic data to understand shifting distribution patterns of (potentially) invasive plants. In this study the spread of Ilex aquifolium L. cultivars was investigated at the edge of its distribution margin in Denmark. Danish nursery catalogues from 1841–2007 were studied to 1) reconstruct the introduction history of potentially invasive ornamental genotypes of I. aquifolium in Denmark; 2) explore potential factors that could explain naturalization of the species, i.e. nursery location, plant size, hardiness, price and year; 3) investigate whether or not marketing has been more common inside the historical range of the species in western Denmark; and 4) test the hypothesis that marketing of this frost‐sensitive species has increased parallel with higher temperatures in the past 100 years. Price, plant size, hardiness, location, and year were recorded for a total of 3213 entries. Through time, increasing prices and the availability of frost‐hardy cultivars were the most significant factors contributing to the sale of I. aquifolium in Denmark. The number of nurseries per area, and hence propagule pressure, was higher in eastern Denmark, outside the natural range of the species. The proportion of very frost‐hardy cultivars increased with milder winter temperatures. The main conclusion from this combined nursery catalogue and climate study are that nurseries in an indirect way play an important role in increasing the propagule pressure of potentially invasive ornamentals and thus can affect distribution of native species.     

Spread of exotic plants in the landscape: the role of time,growth habit,and history of invasiveness

We investigated the relative contribution of minimum residence time, growth habit, and history of invasiveness to the spread of exotic plants in Michigan and California. Our data include minimum residence time as estimated by earliest herbarium collection records, growth habit, and history of invasiveness for over 2000 records from two herbaria (MI = 943, CA = 1131). Our data support the hypothesis that minimum residence time is highly associated with landscape spread, explaining 39–44% of variation in the number of counties invaded. In contrast, growth habit and history of invasiveness explained a small fraction of variation in spread in California but not Michigan. Over the past 30 years exotic plant species frequently became established in Michigan and California (≥50 species per decade), suggesting that many more species will become invasive over time. There is an urgent need to develop effective policies for exotic plant management. In both states we found significant positive correlations between minimum residence time and species occurrence on state invasive plant lists. Further, we found historical information on the pest status of a plant species introduced into a similar environment to be relevant in determining landscape spread of exotic plants. We conclude that efforts to predict exotic species spread based on biological characteristics may have limited success, and instead endorse pest risk analysis for proposed new imports coupled with rapid detection and early response for unintended and unwanted introductions.     

Niche based distribution modelling of an invasive alien plant: effects of population status, propagule pressure and invasion history

Forecasting the spatial spread of invasive species is important to inform management planning. Niche-based species distribution models offer a well-developed framework for assessing the potential range of species. However, these models assume equilibrium between the species’ distribution and its ecological requirements. During range expansion, invasive species are not in such equilibrium due to both dispersal limitation and frequent casual occurrence in sites unsuitable to persistent populations. In this article we use the example of the invasive annual plant Ambrosia artemisiifolia in Austria to evaluate if model accuracy can be enhanced in such non-equilibrium situations by taking account of propagule pressure and by restricting model calibration to naturalized populations. Moreover, we test if model accuracy increases during invasion history using distribution data from 1984 to 2005. The results suggest that models calibrated with naturalized populations are much more accurate than those based on the total set of records. Proxies of propagule pressure slightly but significantly improve goodness of fit, accuracy, and Type I and II error rates of models calibrated with all available records but have less consistent effects on models of naturalized populations. Model accuracy did not increase during the recent invasion history, probably because the species is still far from an equilibrium distribution. We conclude that even a coarse assessment of population status with records of invasive species delivers important information for predictive modelling and that proxies of propagule pressure should be included into such models at least during early to intermediate stages of the invasion history.     

Invasive invertebrates associated with highly duplicated gene content

Invasion of alien species has led to serious problems, including the destruction of native ecosystems. In general, invasive species adapt to new environments rapidly, suggesting that they have high genetic diversity that can directly influence environmental adaptability. However, it is not known how genomic architecture causes genetic diversity that leads to invasiveness. Recent studies have showed that the proportion of duplicated genes (P_D) in whole animal genomes correlate with environmental variability within a habitat. Here, we show that P_D and propagule size significantly explain the differences in species categories (invasive species, noninvasive species, and parasites). P_D correlated negatively with the propagule size. The residual values of regression of P_D on propagule size revealed that the invasive species had higher P_D values and larger propagule size than those of the noninvasive species, whereas the parasites had lower P_D values and smaller propagule size than those of others. There were no correlations between the invasive species and other genomic factors including the genome size, number of genes, and certain gene families. Our results suggest that the P_D values of a genome might be a potential genomic source causing genetic variations for adaptation to diverse environments. The results also showed that the invasiveness status of a species would be predicted by the residual values of regression of P_D on propagule size. Our innovative approach provides a measure to estimate the environmental adaptability of organisms based on genomic data.     

Getting the Right Traits: Reproductive and Dispersal Characteristics Predict the Invasiveness of Herbaceous Plant Species

To better understand the effect of species traits on plant invasion, we collected comparative data on 20 reproductive and dispersal traits of 93 herbaceous alien species in the Czech Republic, central Europe, introduced after 1500 A. D. We explain plant invasion success, expressed by two measures: invasiveness, i.e. whether the species is naturalized but non-invasive, or invasive; and dominance in plant communities expressed as the mean cover in vegetation plots. We also tested how important reproductive and dispersal traits are in models including other characteristics generally known to predict invasion outcome, such as plant height, life history and residence time. By using regression/classification trees we show that the biological traits affect invasion success at all life stages, from reproduction (seed production) to dispersal (propagule properties), and the ability to compete with resident species (height). By including species traits information not usually available in multispecies analyses, we provide evidence that traits do play important role in determining the outcome of invasion and can be used to distinguish between alien species that reach the final stage of the invasion process and dominate the local communities from those that do not. No effect of taxonomy ascertained in regression and classification trees indicates that the role of traits in invasiveness should be assessed primarily at the species level.     

Biological attributes discriminating invasive from native European stream macroinvertebrates

Rising economic and ecological costs caused by invasive organisms revived research on biological attributes associated with invasiveness, focussing on the question: do invasive taxa have biological attributes favouring (1) propagule pressure; (2) dispersal; and (3) establishment and population growth? Using a literature-derived database on 312 stream macroinvertebrate genera occurring at 527 least human-impacted European sites, we quantitatively examined this question. Compared with native genera, genera with invasive invertebrate species (1) tended to reproduce more frequently and to have higher abundances (i.e. higher propagule pressure); (2) had no particular resistance stages to survive during dispersal; and (3) had significantly more ovoviviparity (enabling colonization by a single individual that typically releases viable offspring), larger size and longer life (providing resistance against mortality), food and feeding habits exploiting food resources in streams more effectively, and tended to be more dominant in their communities (all favouring establishment and population growth). Repeating these analyses excluding “native” flying insects (i.e. genera that presumably invaded from glacial refuges unnoticed by biologists), fewer biological attributes were significantly associated with invasiveness. For both data sets (all genera or insects excluded), their affinity to few biological traits (e.g. ovoviviparity, gill respiration) assigned the same 13 of the 19 invasive genera to the top 19 ranks on a gradient of potential invasiveness, together with native genera that risk to become invasive (e.g. Pisidium, Unio), but also with one endangered native genus (Margaritifera). Overall, our data support the idea that invasiveness can be predicted using biological attributes.     

Genetic evidence for high propagule pressure and long‐distance dispersal in monk parakeet (Myiopsitta monachus) invasive populations

The monk parakeet (Myiopsitta monachus) is a successful invasive species that does not exhibit life history traits typically associated with colonizing species (e.g., high reproductive rate or long‐distance dispersal capacity). To investigate this apparent paradox, we examined individual and population genetic patterns of microsatellite loci at one native and two invasive sites. More specifically, we aimed at evaluating the role of propagule pressure, sexual monogamy and long‐distance dispersal in monk parakeet invasion success. Our results indicate little loss of genetic variation at invasive sites relative to the native site. We also found strong evidence for sexual monogamy from patterns of relatedness within sites, and no definite cases of extra‐pair paternity in either the native site sample or the examined invasive site. Taken together, these patterns directly and indirectly suggest that high propagule pressure has contributed to monk parakeet invasion success. In addition, we found evidence for frequent long‐distance dispersal at an invasive site (～100 km) that sharply contrasted with previous estimates of smaller dispersal distance made in the native range (～2 km), suggesting long‐range dispersal also contributes to the species’ spread within the United States. Overall, these results add to a growing body of literature pointing to the important role of propagule pressure in determining, and thus predicting, invasion success, especially for species whose life history traits are not typically associated with invasiveness.     

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