The EICAT+ framework enables classification of positive impacts of alien taxa on native biodiversity.

Species introduced through human-related activities beyond their native range, termed alien species, have various impacts worldwide. The IUCN Environmental Impact Classification for Alien Taxa (EICAT) is a global standard to assess negative impacts of alien species on native biodiversity. Alien species can also positively affect biodiversity (for instance, through food and habitat provisioning or dispersal facilitation) but there is currently no standardized and evidence-based system to classify positive impacts. We fill this gap by proposing EICAT+, which uses 5 semiquantitative scenarios to categorize the magnitude of positive impacts, and describes underlying mechanisms. EICAT+ can be applied to all alien taxa at different spatial and organizational scales. The application of EICAT+ expands our understanding of the consequences of biological invasions and can inform conservation decisions.

Considerations for developing and implementing a safe list for alien taxa.

Many species have been intentionally introduced to new regions for their benefits. Some of these alien species cause damage, others do not (or at least have not yet). There are several approaches to address this problem: prohibit taxa that will cause damage, try to limit damages while preserving benefits, or promote taxa that are safe. In the present article, we unpack the safe list approach, which we define as "a list of taxa alien to the region of interest that are considered of sufficiently low risk of invasion and impact that the taxa can be widely used without concerns of negative impacts." We discuss the potential use of safe lists in the management of biological invasions; disentangle aspects related to the purpose, development, implementation, and impact of safe lists; and provide guidance for those considering to develop and implement such lists.

Arthropods on imported plant products: Volumes predict general trends while contextual details enhance predictive power.

Agricultural biosecurity interventions are aimed at minimizing introductions of harmful non-native organisms to new areas via agricultural trade. To prioritize such interventions, historical data on interceptions have been used to elucidate which factors determine the likelihood that a particular import is carrying a harmful organism. Here we use an interception data set of arthropod contaminants recorded on plant imports arriving in South Africa from 2005 to 2019, comprising 13,566 samples inspected for arthropod contaminants, of which 4902 were positive for the presence of at least one arthropod. We tested 29 predictor variables that have previously been used to explain variation in rates of detection and three variables describing possible sources of additional variation and grouped these into six mutually exclusive "factor classes." We used boosted regression trees as a non-parametric stochastic machine-learning method to build models for each factor class and interactions between them. We explored the influence of these variables with data split either randomly or chronologically. While we identified some specific patterns that could be explained post-hoc by historical events, only inspected volumes were reliably correlated with detection of arthropod contaminants across the whole data set. However, inspected volumes could not predict future interceptions of arthropods, which instead relied on contextual factors such as country, crop or year of import. This suggests that, although certain factors may be important in certain circumstances or for particular crops or commodities, there is little general predictive power in the current data. Instead, an idiographic approach would be most beneficial in biosecurity to ascertain the details of why a particular pest arrived on a particular pathway and how it might move (and be stopped) in future.

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

BACKGROUND AND AIMS: Invasive 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. METHODS: We 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 RESULTS: Levels 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. CONCLUSION: We attribute our findings to the impressive movement, introduction effort and human usage of Australian acacias around the world.

Stronger regional biosecurity is essential to prevent hundreds of harmful biological invasions.

Biological invasions often transcend political boundaries, but the capacity of countries to prevent invasions varies. How this variation in biosecurity affects the invasion risks posed to the countries involved is unclear. We aimed to improve the understanding of how the biosecurity of a country influences that of its neighbours. We developed six scenarios that describe biological invasions in regions with contiguous countries. Using data from alien species databases, socio-economic and biodiversity data and species distribution models, we determined where 86 of 100 of the world's worst invasive species are likely to invade and have a negative impact in the future. Information on the capacity of countries to prevent invasions was used to determine whether such invasions could be avoided. For the selected species, we predicted 2,523 discrete invasions, most of which would have significant negative impacts and are unlikely to be prevented. Of these invasions, approximately a third were predicted to spread from the country in which the species first establishes to neighbouring countries where they would cause significant negative impacts. Most of these invasions are unlikely to be prevented as the country of first establishment has a low capacity to prevent invasions or has little incentive to do so as there will be no impact in that country. Regional biosecurity is therefore essential to prevent future harmful biological invasions. In consequence, we propose that the need for increased regional co-operation to combat biological invasions be incorporated in global biodiversity targets.

The first management of a marine invader in Africa: The importance of trials prior to setting long-term management goals.

Biological invasions severely impact on marine ecosystems around the world, but to date management is rare and has not previously been attempted in Africa. This study documents a trial management programme aimed at informing a national management strategy for the invasive European shore crab, Carcinus maenas, in South Africa. The approach involved testing control methods used elsewhere (baited traps, crab condos, diver collections and sediment dredging) and adapting these to the local context. Following these trials, baited traps were deployed over the course of the year, and the catch per unit effort (CPUE) tracked. A total of 36,244 crabs were collected during the management period, six times more than a pre-management population estimate. The population was not extirpated and CPUE increased once trapping ceased. The cost of attempting nationwide eradication is prohibitive, particularly given the lack of current impacts by this crab in this region and the possibility of reintroduction. We highlight key administrative challenges encountered, and the importance of such pilot trials in setting long-term goals when attempting alien species management interventions.

A framework for engaging stakeholders on the management of alien species.

Alien species can have major ecological and socioeconomic impacts in their novel ranges and so effective management actions are needed. However, management can be contentious and create conflicts, especially when stakeholders who benefit from alien species are different from those who incur costs. Such conflicts of interests mean that management strategies can often not be implemented. There is, therefore, increasing interest in engaging stakeholders affected by alien species or by their management. Through a facilitated workshop and consultation process including academics and managers working on a variety of organisms and in different areas (urban and rural) and ecosystems (terrestrial and aquatic), we developed a framework for engaging stakeholders in the management of alien species. The proposed framework for stakeholder engagement consists of 12 steps: (1) identify stakeholders; (2) select key stakeholders for engagement; (3) explore key stakeholders' perceptions and develop initial aims for management; (4) engage key stakeholders in the development of a draft management strategy; (5) re-explore key stakeholders' perceptions and revise the aims of the strategy; (6) co-design general aims, management objectives and time frames with key stakeholders; (7) co-design a management strategy; (8) facilitate stakeholders' ownership of the strategy and adapt as required; and (9) implement the strategy and monitor management actions to evaluate the need for additional or future actions. In case additional management is needed after these actions take place, some extra steps should be taken: (10) identify any new stakeholders, benefits, and costs; (11) monitor engagement; and (12) revise management strategy. Overall, we believe that our framework provides an effective approach to minimize the impact of conflicts created by alien species management.

A unified classification of alien species based on the magnitude of their environmental impacts.

Species moved by human activities beyond the limits of their native geographic ranges into areas in which they do not naturally occur (termed aliens) can cause a broad range of significant changes to recipient ecosystems; however, their impacts vary greatly across species and the ecosystems into which they are introduced. There is therefore a critical need for a standardised method to evaluate, compare, and eventually predict the magnitudes of these different impacts. Here, we propose a straightforward system for classifying alien species according to the magnitude of their environmental impacts, based on the mechanisms of impact used to code species in the International Union for Conservation of Nature (IUCN) Global Invasive Species Database, which are presented here for the first time. The classification system uses five semi-quantitative scenarios describing impacts under each mechanism to assign species to different levels of impact-ranging from Minimal to Massive-with assignment corresponding to the highest level of deleterious impact associated with any of the mechanisms. The scheme also includes categories for species that are Not Evaluated, have No Alien Population, or are Data Deficient, and a method for assigning uncertainty to all the classifications. We show how this classification system is applicable at different levels of ecological complexity and different spatial and temporal scales, and embraces existing impact metrics. In fact, the scheme is analogous to the already widely adopted and accepted Red List approach to categorising extinction risk, and so could conceivably be readily integrated with existing practices and policies in many regions.

Level of environmental threat posed by horticultural trade in Cactaceae.

Ornamental horticulture has been identified as an important threat to plant biodiversity and is a major pathway for plant invasions worldwide. In this context, the family Cactaceae is particularly challenging because it is considered the fifth most threatened large taxonomic group in the world; several species are among the most widespread and damaging invasive species; and Cactaceae is one of the most popular horticultural plant groups. Based on the Convention on International Trade in Endangered Species of Wild Flora and Fauna and the 11 largest online auction sites selling cacti, we documented the international cactus trade. To provide an in-depth look at the dynamics of the industry, we surveyed the businesses involved in the cactus trade in South Africa (a hotspot of cactus trade and invasions). We purchased seeds of every available species and used DNA barcoding to identify species to the genus level. Although <20% of this trade involved threatened species and <3% involved known invasive species, many species were identified by a common name. However, only 0.02% of the globally traded cacti were collected from wild populations. Despite a large commercial network, all South African imports (of which 15% and 1.5% were of species listed as threatened and invasive, respectively) came from the same source. With DNA barcoding, we identified 24% of the species to genus level. Based on our results, we believe that if trade restrictions are placed on the small proportion of cacti that are invasive and there is no major increase in harvesting of native populations, then the commercial trade in cactus poses a negligible environmental threat. However, there are currently no effective methods for easily identifying which cacti are traded, and both the illicit harvesting of cacti from the wild and the informal trade in invasive taxa pose on-going conservation challenges.

Border control for stowaway alien species should be prioritised based on variations in establishment debt.

Border control is one of the major approaches used by countries to limit the number of organisms introduced as stowaways. However, it is not feasible to inspect all passengers, cargo and vehicles entering a country, and so efforts need to be prioritised. Here we use South Africa as a case study to assess, based on tourism and trade data and climate matching techniques, the number of stowaway species that might be introduced ('colonisation pressure') and the likelihood that once introduced, these organisms will establish ('likelihood of establishment'). These results were used to explore how the number of species that are likely to establish ('establishment debt') varies across donor regions and seasons. A simple theoretical model was then used to compare four strategies for prioritising border control inspections: no prioritisation; based on colonisation pressure; based on likelihood of establishment; and based on both colonisation pressure and likelihood of establishment. Establishment debt was greatest in southern hemisphere spring and autumn when South Africa is climatically similar to northern hemisphere countries with which there are strong, consistent trade and tourism links (i.e. colonisation pressure varied little seasonally, but likelihood of establishment did vary across the seasons). Prioritising inspections based on both colonisation pressure and the likelihood of establishment was clearly the most effective strategy, with this strategy detecting at least 6% more potential invaders than the other strategies. While there are many practical limitations to the implementation of such prioritised inspection strategies, the results highlight the importance of national and regional studies of establishment debt.

Resolving a Prickly Situation: Involving Stakeholders in Invasive Cactus Management in South Africa.

The regulation and management of alien species can be contentious, particularly when the stakeholders who benefit from alien species are different from those who suffer the costs. We propose a consultative process involving relevant stakeholders in invasive species management decisions. The process involves (1) the identification of relevant stakeholders, (2) assessing their perceptions, (3) enhancing interaction between stakeholders, (4) assessing changes in stakeholders' perceptions following interactions with other stakeholders, and (5) developing management recommendations in collaboration with stakeholders. We demonstrate the application of the process using the family Cactaceae ('cacti') in South Africa. Many species of cacti have been introduced to the country over the past two centuries, mostly for horticulture, food and fodder, and hundreds of other species have been introduced in the past few decades (or are likely to be introduced soon) for horticulture. Using the proposed process enabled the negotiation and participation of all stakeholders in decision making and helped minimize contentious situations by clarifying stakeholder's beliefs and exploring consensus solutions. Consequently, management objectives were broadly supported by all stakeholders. These results will be included in a national cactus management strategy for South Africa.

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.

Elucidating the native sources of an invasive tree species, Acacia pycnantha, reveals unexpected native range diversity and structure.

BACKGROUND AND AIMS: Understanding the introduction history of invasive plant species is important for their management and identifying effective host-specific biological control agents. However, uncertain taxonomy, intra- and interspecific hybridization, and cryptic speciation may obscure introduction histories, making it difficult to identify native regions to explore for host-specific agents. The overall aim of this study was to identify the native source populations of Acacia pycnantha, a tree native to south-eastern Australia and invasive in South Africa, Western Australia and Portugal. Using a phylogeographical approach also allowed an exploration of the historical processes that have shaped the genetic structure of A. pycnantha in its native range. METHODS: Nuclear (nDNA) and plastid DNA sequence data were used in network and tree-building analyses to reconstruct phylogeographical relationships between native and invasive A. pycnantha populations. In addition, mismatch distributions, relative rates and Bayesian analyses were used to infer recent demographic processes and timing of events in Australia that led to population structure and diversification. KEY RESULTS: The plastid network indicated that Australian populations of A. pycnantha are geographically structured into two informally recognized lineages, the wetland and dryland forms, whereas the nuclear phylogeny showed little geographical structure between these two forms. Moreover, the dryland form of A. pycnantha showed close genetic similarity to the wetland form based on nDNA sequence data. Hybrid zones may explain these findings, supported here by incongruent phylogenetic placement of some of these taxa between nuclear and plastid genealogies. CONCLUSIONS: It is hypothesized that habitat fragmentation due to cycles of aridity inter-dispersed with periods of abundant rainfall during the Pleistocene (approx. 100 kya) probably gave rise to native dryland and wetland forms of A. pycnantha. Although the different lineages were confined to different ecological regions, we also found evidence for intraspecific hybridization in Victoria. The invasive populations in Portugal and South Africa represent wetland forms, whereas some South African populations resemble the Victorian dryland form. The success of the biological control programme for A. pycnantha in South Africa may therefore be attributed to the fact that the gall-forming wasp Trichilogaster signiventris was sourced from South Australian populations, which closely match most of the invasive populations in South Africa.

Human usage in the native range may determine future genetic structure of an invasion: insights from Acacia pycnantha.

BACKGROUND: The influence of introduction history and post-introduction dynamics on genetic diversity and structure has been a major research focus in invasion biology. However, genetic diversity and structure in the invasive range can also be affected by human-mediated processes in the native range prior to species introductions, an aspect often neglected in invasion biology. Here we aim to trace the native provenance of the invasive tree Acacia pycnantha by comparing the genetic diversity and structure between populations in the native Australian range and the invasive range in South Africa. This approach also allowed us to explore how human actions altered genetic structure before and after the introduction of A. pycnantha into South Africa. We hypothesized that extensive movement and replanting in A. pycnantha's Australian range prior to its introduction to South Africa might result in highly admixed genotypes in the introduced range, comparable genetic diversity in both ranges, and therefore preclude an accurate determination of native provenance(s) of invasive populations. RESULTS: In the native range Bayesian assignment tests identified three genetic clusters with substantial admixture and could not clearly differentiate previously identified genetic entities, corroborating admixture as a result of replantings within Australia. Assignment tests that included invasive populations from South Africa indicated similar levels of admixture compared to Australian populations and a lack of genetic structure. Invasive populations of A. pycnantha in South Africa are as genetically diverse as native populations, and could not be assigned to particular native range regions. CONCLUSIONS: Our results indicate that the genetic structure of A. pycnantha in Australia has been greatly altered through various planting initiatives. Specifically, there is little geographic structure and high levels of admixture. While numerous introduction history scenarios may explain the levels of admixture observed in South Africa, planting records of A. pycnantha in Australia suggest that populations were probably already admixed before propagules were introduced to South Africa. These findings have important implications for the management of invasive A. pycnantha populations in South Africa, especially for classical biological control, and more broadly, for studies that aim to understand the evolutionary dynamics of the invasion process.

An inventory of native-alien populations in South Africa.

Species can be both native and alien to a given administrative region. Here we present the first consolidated inventory of these 'native-alien populations' for South Africa, and provide an overview of the data it contains. To gather data, literature searches were performed and experts were consulted both directly and via an on-line survey. Putative native-alien populations were then scored based on a newly developed protocol. The final inventory contains information on 77 native species from 49 families across nine classes that have formed 132 native-alien populations across the terrestrial, freshwater, and marine environments. The phenomenon is rare when compared to the prevalence of related phenomena, such as alien species introduced from other countries (2033 alien species in South Africa), but is under-reported. However, they pose a specific problem for regulators and managers and their importance will likely increase with global change. These data will be integrated with an existing alien species list and, we hope, will provide a useful foundation to address the issue. We encourage those working on biodiversity to contribute more records.

Cultivation shapes genetic novelty in a globally important invader.

Acacia saligna is a species complex that has become invasive in a number of countries worldwide where it has caused substantial environmental and economic impacts. Understanding genetic and other factors contributing to its success may allow managers to limit future invasions of closely related species. We used three molecular markers to compare the introduced range (South Africa) to the native range (Western Australia). Nuclear markers showed that invasive populations are divergent from native populations and most closely related to a cultivated population in Western Australia. We also found incongruence between nuclear and chloroplast data that, together with the long history of cultivation of the species, suggest that introgressive hybridization (coupled with chloroplast capture) may have occurred within A. saligna. While we could not definitively prove introgression, the genetic distance between cultivated and native A. saligna populations was comparable to known interspecific divergences among other Acacia species. Therefore, cultivation, multiple large-scale introductions and possibly introgressive hybridization have rapidly given rise to the divergent genetic entity present in South Africa. This may explain the known global variation in invasiveness and inaccuracy of native bioclimatic models in predicting potential distributions.

Addressing context dependence in ecology.

Context dependence is widely invoked to explain disparate results in ecology. It arises when the magnitude or sign of a relationship varies due to the conditions under which it is observed. Such variation, especially when unexplained, can lead to spurious or seemingly contradictory conclusions, which can limit understanding and our ability to transfer findings across studies, space, and time. Using examples from biological invasions, we identify two types of context dependence resulting from four sources: mechanistic context dependence arises from interaction effects; and apparent context dependence can arise from the presence of confounding factors, problems of statistical inference, and methodological differences among studies. Addressing context dependence is a critical challenge in ecology, essential for increased understanding and prediction.

GIRAE: a generalised approach for linking the total impact of invasion to species' range, abundance and per-unit effects.

The total impact of an alien species was conceptualised as the product of its range size, local abundance and per-unit effect in a seminal paper by Parker et al. (Biol Invasions 1:3-19, 1999). However, a practical approach for estimating the three components has been lacking. Here, we generalise the impact formula and, through use of regression models, estimate the relationship between the three components of impact, an approach we term GIRAE (Generalised Impact = Range size × Abundance × per-unit Effect). We discuss how GIRAE can be applied to multiple types of impact, including environmental impacts, damage and management costs. We propose two methods for applying GIRAE. The species-specific method computes the relationship between impact, range size, abundance and per-unit effect for a given species across multiple invaded sites or regions of different sizes. The multi-species method combines data from multiple species across multiple sites or regions to calculate a per-unit effect for each species and is computed using a single regression model. The species-specific method is more accurate, but it requires a large amount of data for each species and assumes a constant per-unit effect for a species across the invaded area. The multi-species method is more easily applicable and data-parsimonious, but assumes the same relationship between impact, range size and abundance for all considered species. We illustrate these methods using data about money spent managing plant invasions in different biomes of South Africa. We found clear differences between species in terms of money spent per unit area invaded, with per-unit expenditure varying substantially between biomes for some species-insights that are useful for monitoring and evaluating management. GIRAE offers a versatile and practical method that can be applied to many different types of data to better understand and manage the impacts of biological invasions. Supplementary Information: The online version contains supplementary material available at 10.1007/s10530-022-02836-0.

Contaminant organisms recorded on plant product imports to South Africa 1994-2019.

Biosecurity interception records are crucial data underlying efforts to predict and manage pest and pathogen introductions. Here we present a dataset containing information on imported plant products inspected by the South African Department of Agriculture's laboratories between 1994 and 2019 and the contaminant organisms found on them. Samples were received from border inspectors as either propagation material (e.g. plants) or material for immediate use (e.g. fruit). Material for immediate use was further divided into two sample categories, depending on if contaminants were seen/suspected by the border official or not: intervention or audit samples. The final dataset consists of 25,279 records, of which 30% tested positive (i.e. had at least one contaminant) and 13% had multiple contaminants. Of the 13,731 recorded contaminants, fungi (41%), mites (37%) and insects (19%) were most common. This dataset provides insight into the suite of taxa transported along the plant import pathway and provides an important resource for analyses of contaminant organisms in international trade, which can inform strategies for risk assessment, pathway management and biosecurity protocols.