The Development of a Plant Risk Evaluation (PRE) Tool for Assessing the Invasive Potential of Ornamental Plants

Weed Risk Assessment (WRA) methods for evaluating invasiveness in plants have evolved rapidly in the last two decades. Many WRA tools exist, but none were specifically designed to screen ornamental plants prior to being released into the environment. To be accepted as a tool to evaluate ornamental plants for the nursery industry, it is critical that a WRA tool accurately predicts non-invasiveness without falsely categorizing them as invasive. We developed a new Plant Risk Evaluation (PRE) tool for ornamental plants. The 19 questions in the final PRE tool were narrowed down from 56 original questions from existing WRA tools. We evaluated the 56 WRA questions by screening 21 known invasive and 14 known non-invasive ornamental plants. After statistically comparing the predictability of each question and the frequency the question could be answered for both invasive and non-invasive species, we eliminated questions that provided no predictive power, were irrelevant in our current model, or could not be answered reliably at a high enough percentage. We also combined many similar questions. The final 19 remaining PRE questions were further tested for accuracy using 56 additional known invasive plants and 36 known non-invasive ornamental species. The resulting evaluation demonstrated that when “needs further evaluation” classifications were not included, the accuracy of the model was 100% for both predicting invasiveness and non-invasiveness. When “needs further evaluation” classifications were included as either false positive or false negative, the model was still 93% accurate in predicting invasiveness and 97% accurate in predicting non-invasiveness, with an overall accuracy of 95%. We conclude that the PRE tool should not only provide growers with a method to accurately screen their current stock and potential new introductions, but also increase the probability of the tool being accepted for use by the industry as the basis for a nursery certification program.     

Predicting plant invaders in the Mediterranean through a weed risk assessment system

Risk assessment schemes have been developed to identify potential invasive species, prevent their spread and reduce their damaging effects. One of the most promising tools for detecting plant invaders is the weed risk assessment (WRA) scheme developed for Australia. Our study explores whether the Australian WRA can satisfactorily predict the invasion status of alien plants in the Mediterranean Basin by screening 100 invasive and 97 casual species in Spain. Furthermore, we analysed whether the factors taken into account in the WRA are linked to invasion likelihood (i.e., invasion status) or to impacts. The outcome was that 94% of the invasive species were rejected, 50% of the casual species were rejected and 29% of them required further evaluation. The accuracy for casuals is lower than in other studies that have tested non-invasive (i.e., casuals or non-escaped) alien species. We postulate that low accuracy for casual species could result from: (1) an incorrect “a priori” expert classification of the species status, (2) a high weight of the WRA scores given to potential impacts, and (3) casual species being prone to becoming invasive when reaching a minimum residence time threshold. Therefore, the WRA could be working as a precaution early-warning system to identify casual species with potential to become invasive.     

Use of a weed risk assessment for the Mediterranean region of Central Italy to prevent loss of functionality and biodiversity in agro-ecosystems

The capacity to predict invasiveness of plant species is important for the conservation and management of natural habitats, especially within agro-ecosystems. Many factors increase the creation of newly available niches in this type of anthropogenic manipulated ecosystem. Consequently, the presence and establishment of invasive alien species with the potential to spread and cause harm, or constrain elements of semi-natural habitat or vegetation remnants, may increase. The invasiveness of weedy germplasm may be accelerated by the presence of cultivated species that are able to escape from fields also through crop movement or on livestock. The future use of agricultural land for widespread and intensive cultivation of biofuel crops for energy production increases the need for a pre-entry screening tool both for species that are new to the Italian cropping system and for the management of existing weedy species. This study aimed to assess the effectiveness of adapting the Australian and New Zealand Weed Risk Assessment (WRA) to the geographic, climatic and weed management context of Italy. We evaluated the performance of the adapted WRA on several alien plant species of known invasiveness in Mediterranean Central Italy. WRA score results were compared with a priori independent opinions of botanists with field experience in the evaluated region. The assessment procedure correctly identified 93% of invasive species and 75% of non-invasive species. Further evaluation was needed for 20% of the tested species and was conducted through a secondary screening. Throughout the whole process, only one (5%) of the investigated species could not be assessed. The results of the Receiver Operating Characteristic analysis, the consistency of the outcomes with those found in other WRA studies, the Chi Square testing categories and the high correlation between the a priori and WRA score corroborated the predictive accuracy of the WRA for determining invasive from non-invasive species. This confirmed the effectiveness of the screening process and an assessment was subsequently carried out on proposed biofuel species detecting some potential invaders. The WRA can thus be used to assess the introduction of new cropping systems and for weed management.     

Risk assessment for invasiveness differs for aquatic and terrestrial plant species

Predictive tools for preventing introduction of new species with high probability of becoming invasive in the U.S. must effectively distinguish non-invasive from invasive species. The Australian Weed Risk Assessment system (WRA) has been demonstrated to meet this requirement for terrestrial vascular plants. However, this system weights aquatic plants heavily toward the conclusion of invasiveness. We evaluated the accuracy of the WRA for 149 non-native aquatic species in the U.S., of which 33 are major invaders, 32 are minor invaders and 84 are non-invaders. The WRA predicted that all of the major invaders would be invasive, but also predicted that 83% of the non-invaders would be invasive. Only 1% of the non-invaders were correctly identified and 16% needed further evaluation. The resulting overall accuracy was 33%, dominated by scores for invaders. While the overall accuracy increased to 57% when the points assigned to aquatic life forms were removed, 57% of the non-invaders required further evaluation rather than were identified as having low probability of naturalizing. Discrimination between non-invaders and invaders would require an increase in the threshold score from the standard of 6 for this system to 19. That higher threshold resulted in accurate identification of 89% of the non-invaders and over 75% of the major invaders. Either further testing for definition of the optimal threshold or a separate screening system will be necessary for accurately predicting which freshwater aquatic plants are high risks for becoming invasive.     

Quantifying Uncertainty in Predictions of Invasiveness

Using the Australian weed risk assessment (WRA) model as an example, we applied a combination of bootstrapping and Bayesian techniques as a means for explicitly estimating the posterior probability of weediness as a function of an import risk assessment model screening score. Our approach provides estimates of uncertainty around model predictions, after correcting for verification bias arising from the original training dataset having a higher proportion of weed species than would be the norm, and incorporates uncertainty in current knowledge of the prior (base-rate) probability of weediness. The results confirm the high sensitivity of the posterior probability of weediness to the base-rate probability of weediness of plants proposed for importation, and demonstrate how uncertainty in this base-rate probability manifests itself in uncertainty surrounding predicted probabilities of weediness. This quantitative estimate of the weediness probability posed by taxa classified using the WRA model, including estimates of uncertainty around this probability for a given WRA score, would enable bio-economic modelling to contribute to the decision process, should this avenue be pursued. Regardless of whether or not this avenue is explored, the explicit estimates of uncertainty around weed classifications will enable managers to make better informed decisions regarding risk. When viewed in terms of likelihood of weed introduction, the current WRA model outcomes of ‘accept’, ‘further evaluate’, or ‘reject’, whilst not always accurate in terms of weed classification, appear consistent with a high expected cost of mistakenly introducing a weed. The methods presented have wider application to the quantitative prediction of invasive species for situations where the base-rate probability of invasiveness is subject to uncertainty, and the accuracy of the screening test imperfect     

Quantifying uncertainty in predictions of invasiveness, with emphasis on weed risk assessment

Using the Australian Weed Risk Assessment (WRA) model as an example, we applied a combination of bootstrapping and Bayesian techniques as a means of explicitly estimating the posterior probability of weediness as a function of an import risk assessment model screening score. Our approach provides estimates of uncertainty around model predictions, after correcting for verification bias arising from the original training dataset having a higher proportion of weed species than would be the norm, and incorporates uncertainty in current knowledge of the prior (base-rate) probability of weediness. The results confirm the high sensitivity of the posterior probability of weediness to the base-rate probability of weediness of plants proposed for importation, and demonstrate how uncertainty in this base-rate probability manifests itself in uncertainty surrounding predicted probabilities of weediness. This quantitative estimate of the weediness probability posed by taxa classified using the WRA model, including estimates of uncertainty around this probability for a given WRA score, would enable bio-economic modelling to contribute to the decision process, should this avenue be pursued. Regardless of whether or not this avenue is explored, the explicit estimates of uncertainty around weed classifications will enable managers to make better informed decisions regarding risk. When viewed in terms of likelihood of weed introduction, the current WRA model outcomes of ‘accept’, ‘further evaluate’ or ‘reject’, whilst not always accurate in terms of weed classification, appear consistent with a high-expected cost of mistakenly introducing a weed. The methods presented have wider application to the quantitative prediction of invasive species for situations where the base-rate probability of invasiveness is subject to uncertainty, and the accuracy of the screening test imperfect.     

Trait Values,Not Trait Plasticity,Best Explain Invasive Species' Performance in a Changing Environment

The question of why some introduced species become invasive and others do not is the central puzzle of invasion biology. Two of the principal explanations for this phenomenon concern functional traits: invasive species may have higher values of competitively advantageous traits than non-invasive species, or they may have greater phenotypic plasticity in traits that permits them to survive the colonization period and spread to a broad range of environments. Although there is a large body of evidence for superiority in particular traits among invasive plants, when compared to phylogenetically related non-invasive plants, it is less clear if invasive plants are more phenotypically plastic, and whether this plasticity confers a fitness advantage. In this study, I used a model group of 10 closely related Pinus species whose invader or non-invader status has been reliably characterized to test the relative contribution of high trait values and high trait plasticity to relative growth rate, a performance measure standing in as a proxy for fitness. When grown at higher nitrogen supply, invaders had a plastic RGR response, increasing their RGR to a much greater extent than non-invaders. However, invasive species did not exhibit significantly more phenotypic plasticity than non-invasive species for any of 17 functional traits, and trait plasticity indices were generally weakly correlated with RGR. Conversely, invasive species had higher values than non-invaders for 13 of the 17 traits, including higher leaf area ratio, photosynthetic capacity, photosynthetic nutrient-use efficiency, and nutrient uptake rates, and these traits were also strongly correlated with performance. I conclude that, in responding to higher N supply, superior trait values coupled with a moderate degree of trait variation explain invasive species'' superior performance better than plasticity per se.     

Consistent accuracy of the Australian weed risk assessment system across varied geographies

The ecological and economic advantages of preventing introduction of species likely to become invasive have increased interest in implementing effective screening tools. We compared the accuracy of the Australian Weed Risk Assessment (WRA) system with that across the six geographies in which it has been tested (New Zealand, Hawaii, Hawaii and Pacific Islands, Czech Republic, Bonin Islands and Florida). Inclusion in four of the tests of a secondary screening tool, developed to reduce the number of species requiring further evaluation, decreased the number of species with that outcome by over 60% on average. Averaging across all tests demonstrated that the WRA system accurately identified major invaders 90%, and non-invaders 70%, of the time. Examined differently, a species of unknown invasive potential is on average likely to be correctly accepted or rejected over 80% of the time for all of these geographies when minor invaders are categorized as invasive. Whereas increasing consistency in definitions and implementation would facilitate understanding of the general application of the WRA system, we believe that this tool functions similarly across islands and continents in tropical and temperate climates and has been sufficiently tested to be adopted as an initial screen for plant species proposed for introduction to a new geography.     

Big troubles are already here: risk assessment protocol shows high risk of many alien plants present in Portugal

Invasion by alien plants results in serious adverse impacts on biodiversity, ecosystem services, economy and social welfare, and is an ever-increasing challenge for nature conservation. Control of established invasive species is frequently very difficult and costly. Therefore, predicting which species have risk of becoming invasive is crucial both to prevent introduction of new invaders and to target high risk species already present in order to avoid their spread, particularly to areas with high conservation value. In Portugal more than 600 alien plant species are present as casuals or naturalized, and decision support tools are needed to discriminate which of these have higher probability of becoming invasive. We aimed to assess the effectiveness of the Australian Weed Risk Assessment, adapted to the Portuguese conditions (P-WRA), by evaluating 172 plant species: 49 considered as invasive; and, 123 as non-invasive species. The results showed that the P-WRA correctly identified all invasive species. As for non-invasive species, 17% were accepted, 78% rejected and 5% required further evaluation. Receiver Operating Characteristic (ROC) curve reflected high accuracy of predictions despite lower than that found in other regions. The best cutoff level for P-WRA score, maximizing the ability for classification of the protocol, was 13 resulting in more non-invasive species accepted (46%). In the end, this assessment informed that in addition to the 49 species already invasive, a high number (66) of alien plant species have invasive risk. Amongst these, a list of 20 species is proposed for targeted priority management aiming to prevent their spread. Comparison of the scores obtained with A-WRA for other regions with Mediterranean climate revealed that 17% of the species compared (78) reached different outcomes, signaling the need to be careful when extrapolating the use of previous scores. The P-WRA can be a promising screening system post-border for predicting invasive species already present in Portugal and contribute to the targeting of species for priority intervention, particularly in natural areas with high conservation value.     

Speciation genetics of biological invasions with hybridization

The negative effects of human‐induced habitat disturbance and modification on multiple dimensions of biological diversity are well chronicled ( Turner 1996 ; Harding et al. 1998 ; Lawton et al. 1998 ; Sakai et al. 2001 ). Among the more insidious consequences is secondary contact between formerly allopatric taxa ( Anderson & Hubricht 1938 ; Perry et al. 2002 ; Seehausen 2006 ). How the secondary contact will play out is unpredictable ( Ellstrand et al. 2010 ), but if the taxa are not fully reproductively isolated, hybridization is likely, and if the resulting progeny are fertile, the eventual outcome is often devastating from a conservation perspective ( Rhymer & Simberloff 1996 ; Wolf et al. 2001 ; McDonald et al. 2008 ). In this issue of Molecular Ecology, Steeves et al. (2010) present an analysis of hybridization between two avian species, one of which is critically endangered and the other of which is invasive. Their discovery that the endangered species has not yet been hybridized to extinction is promising and not what one would necessarily expect from theory.     

How robust is the Australian Weed Risk Assessment protocol? A test using pine invasions in the Northern and Southern hemispheres

The Australian Weed Risk Assessment protocol (WRA) is often considered the standard approach for pre-border screening of new plant introductions. Here we assess its robustness against three key criteria: ability to discriminate success or failure of species at three stages of the invasion process (introduction, naturalisation and spread); sensitivity to taxonomic range and target region; and dependence on knowledge of invasive behaviour elsewhere. We address these issues by retrospectively testing the WRA using pine (Pinus) introductions to New Zealand and Great Britain. For both regions we calculated WRA scores for 115 species, and classified all species according to whether they had been introduced, which of these had naturalised, and the extent of their naturalised distribution (spread). Using regression models, we assessed whether WRA scores could predict success at each stage. We repeated this procedure using WRA scores calculated without information on species naturalisation behaviour elsewhere. In both regions, the WRA could discriminate among species in the same genus at the introduction and naturalisation stages, but not at the spread stage. The outcome at the naturalisation stage depended on prior knowledge of naturalisation behaviour elsewhere. Without this information the WRA may be unable to distinguish among closely related species, and should be used cautiously where data on invasive behaviour elsewhere is lacking. Human selection played a strong role in the invasion process both through introducing pine species likely to naturalise in New Zealand and Great Britain in the first instance, and subsequent use of many of these species for forestry in the target regions.     

Body size and invasion success in marine bivalves

The role of body size in marine bivalve invasions has been the subject of debate. Roy et al . found that large-bodied species of marine bivalves were more likely to be successful invaders, consistent with patterns seen during Pleistocene climatic change, but Miller et al . argued that such selectivity was largely driven by the inclusion of mariculture species in the analysis and that size-selectivity was absent outside of mariculture introductions. Here we use data on non-mariculture species from the north-eastern Pacific coast and from a global species pool to test the original hypothesis of Roy et al . that range limits of larger bivalves are more fluid than those of smaller species. First, we test the hypothesis that larger bivalve species are more successful than small species in expanding their geographical ranges following introduction into new regions. Second, we compare body sizes of indigenous and non-indigenous species for 299 of the 303 known intertidal and shelf species within the marine bivalve clade that contains the greater number of non-mariculture invaders, the Mytilidae. The results from both tests provide additional support for the view that body size plays an important role in mediating invasion success in marine bivalves, in contrast to Miller et al . Thus range expansions in Recent bivalves are consistent with patterns seen in Pleistocene faunas despite the many differences in the mechanisms.     

Persistence and colonisation as measures of success in bog restoration for aquatic invertebrates: a question of detection

1. van Duinen et al. (Freshwater Biol., 2006) raise an interesting point regarding Mazerolle et al.’s (Freshwater Biol., 2006, 51 , p. 333) conclusion on the ability of invertebrates, especially sedentary species, to colonise newly created bogs pools. We wish to clarify that Mazerolle et al. (2006) targeted large arthropods and the absence of smaller sedentary species was purely a result of sampling design. 2. van Duinen et al. (2006) postulate that colonisation rates by bog specialists should be higher in Canada than in the Netherlands, given the extensive amount of intact peatlands in Canada. Here, we emphasise the importance of taking the regional context into account when assessing restoration success as our study site occurs in a landscape where most bog pools have been drained. 3. An evaluation of restoration efforts should focus on both sedentary and vagile invertebrates, to resolve the importance of persistence and colonisation. Such patterns are difficult to interpret, however, when sampling designs and analyses do not account for the probability of detection: an absence may be due to non‐detection or true absence. We strongly urge investigators to directly estimate detection probability in addition to the parameters of interest (e.g. presence, abundance) to provide the best information possible regarding restoration success.     

Ecological Invasion,Roughened Fronts,and a Competitor’s Extreme Advance: Integrating Stochastic Spatial-Growth Models

Both community ecology and conservation biology seek further understanding of factors governing the advance of an invasive species. We model biological invasion as an individual-based, stochastic process on a two-dimensional landscape. An ecologically superior invader and a resident species compete for space preemptively. Our general model includes the basic contact process and a variant of the Eden model as special cases. We employ the concept of a “roughened” front to quantify effects of discreteness and stochasticity on invasion; we emphasize the probability distribution of the front-runner’s relative position. That is, we analyze the location of the most advanced invader as the extreme deviation about the front’s mean position. We find that a class of models with different assumptions about neighborhood interactions exhibits universal characteristics. That is, key features of the invasion dynamics span a class of models, independently of locally detailed demographic rules. Our results integrate theories of invasive spatial growth and generate novel hypotheses linking habitat or landscape size (length of the invading front) to invasion velocity, and to the relative position of the most advanced invader.     

Competition between similar invasive species: modeling invasional interference across a landscape

As the number of biological invasions increases, interactions between different invasive species will become increasingly important. Several studies have examined facilitative invader–invader interactions, potentially leading to invasional meltdown. However, if invader interactions are negative, invasional interference may lead to lower invader abundance and spread. To explore this possibility, we develop models of two competing invaders. A landscape simulation model examines the patterns created by two such species invading into the same region. We then apply the model to a case study of Carduus nutans L. and C. acanthoides L., two economically important invasive weeds that exhibit a spatially segregated distribution in central Pennsylvania, USA. The results of these spatially-explicit models are generally consistent with the results of classic Lotka–Volterra competition models, with widespread coexistence predicted if interspecific effects are weaker than intraspecific effects for both species. However, spatial segregation of the two species (with lower net densities and no further spread) may arise, particularly when interspecific competition is stronger than intraspecific competition. A moving area of overlap may result when one species is a superior competitor. In the Carduus system, our model suggests that invasional interference will lead to lower levels of each species when together, but a similar net level of thistle invasion due to the similarity of intra- and interspecific competition. Thus, invasional interference may have important implications for the distribution and management of invasive species.     

Persistence of an invasive fish (Neogobius melanostomus) in a contaminated ecosystem

Post-establishment dynamics of invasive species have been under-studied. However, understanding these dynamics is particularly important for the management of invasive species known to impact native communities. Following the invasion of a highly invasive species, the round goby (Neogobius melanostomus), we document long-term population changes after establishment and address how population dynamics of a successful invader change through persistence and integration. Round goby present a threat to the areas they invade by out-competing native species for resources. Furthermore, as a pollution tolerant species, round goby present a second threat by acting as a possible vector for contaminant transfer to higher trophic levels in invaded ecosystems with areas of contamination. We sampled round goby for 11 years (2002–2012) at four low contamination sites and two high contamination sites within Hamilton Harbour ON, Canada, an International Joint Commission Area of Concern. Across sampling years, we show that round goby abundance has declined at low contamination sites, while remaining stable at high contamination sites. Moreover, we show that average body size decreased and reproductive investment increased both across sampling years and between sites of low and high contamination. Our results document population demographic shifts in a persisting invasive species, and underscore the importance of management practices for this species in contaminated environments.     

Predicted versus actual invasiveness of climbing vines in Florida

Climbing vines cause substantial ecological and economic harm, and are disproportionately represented among invasive plant species. Thus, the ability to identify likely vine invaders would enhance the effectiveness of both prevention and management. We tested whether the Weed Risk Assessment (WRA) accurately predicted the current invasion status of 84 non-native climbing vines in Florida. Seventeen percent of the species require further evaluation before risk of invasion can be determined. Of the remaining 70 species, the WRA predicted that 70% were at high risk for invasion, but only 50% of the 84 species are currently invasive in Florida. The status and risk prediction were inconsistent for 27% of the species: 15 non-invaders were predicted to be of high risk for invasion (i.e., false positive) and 4 invaders were predicted to be of low risk (i.e., false negative). Longer residence time in the flora was significantly correlated with higher invasion risk. Further investigation is necessary to identify whether residence time explains inconsistencies between risk and status conclusions, or whether the WRA over-predicts invasion risk. Nevertheless, the effects of invasive vines on native systems coupled with the influence of time on invasion status, suggest a precautionary approach is warranted when considering the introduction and management of non-native vines.     

North Atlantic marine communities through time

How and why ecological communities change their species membership over time and space is a central issue in ecology and evolution. Phylogeographic approaches based on animal mitochondrial DNA sequences have been important for revealing historical patterns of individual species and can provide qualitative comparisons among species. Exciting new methods, particularly implementing approximate Bayesian computation (ABC – Beaumont et al. 2002 ), now allow model‐based quantitative comparisons among species and permit the probabilistic exploration of alternative community‐level hypotheses (see review by Hickerson et al. 2010 ). In this issue of Molecular Ecology, Ilves et al. (2010) use an ABC approach to bring fresh insights into the well‐studied question of how North Atlantic coastal species contracted and expanded their ranges in response to late Pleistocene/Holocene climate fluctuations.