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.

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.

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.

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.

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.

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.

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.

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.

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.

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.

Global Actions for Managing Cactus Invasions.

The family Cactaceae Juss. contains some of the most widespread and damaging invasive alien plant species in the world, with Australia (39 species), South Africa (35) and Spain (24) being the main hotspots of invasion. The Global Cactus Working Group (IOBC GCWG) was launched in 2015 to improve international collaboration and identify key actions that can be taken to limit the impacts caused by cactus invasions worldwide. Based on the results of an on-line survey, information collated from a review of the scientific and grey literature, expertise of the authors, and because invasiveness appears to vary predictably across the family, we (the IOBC GCWG): (1) recommend that invasive and potentially invasive cacti are regulated, and to assist with this propose five risk categories; (2) recommend that cactus invasions are treated physically or chemically before they become widespread; (3) advocate the use of biological control to manage widespread invasive species; and (4) encourage the development of public awareness and engagement initiatives to integrate all available knowledge and perspectives in the development and implementation of management actions, and address conflicts of interest, especially with the agricultural and ornamental sectors. Implementing these recommendations will require global co-operation. The IOBC GCWG aims to assist with this process though the dissemination of information and experience.

Emerging infectious diseases and biological invasions: a call for a One Health collaboration in science and management.

The study and management of emerging infectious diseases (EIDs) and of biological invasions both address the ecology of human-associated biological phenomena in a rapidly changing world. However, the two fields work mostly in parallel rather than in concert. This review explores how the general phenomenon of an organism rapidly increasing in range or abundance is caused, highlights the similarities and differences between research on EIDs and invasions, and discusses shared management insights and approaches. EIDs can arise by: (i) crossing geographical barriers due to human-mediated dispersal, (ii) crossing compatibility barriers due to evolution, and (iii) lifting of environmental barriers due to environmental change. All these processes can be implicated in biological invasions, but only the first defines them. Research on EIDs is embedded within the One Health concept-the notion that human, animal and ecosystem health are interrelated and that holistic approaches encompassing all three components are needed to respond to threats to human well-being. We argue that for sustainable development, biological invasions should be explicitly considered within One Health. Management goals for the fields are the same, and direct collaborations between invasion scientists, disease ecologists and epidemiologists on modelling, risk assessment, monitoring and management would be mutually beneficial.

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.

Prioritising surveillance for alien organisms transported as stowaways on ships travelling to South Africa.

The global shipping network facilitates the transportation and introduction of marine and terrestrial organisms to regions where they are not native, and some of these organisms become invasive. South Africa was used as a case study to evaluate the potential for shipping to contribute to the introduction and establishment of marine and terrestrial alien species (i.e. establishment debt) and to assess how this varies across shipping routes and seasons. As a proxy for the number of species introduced (i.e. 'colonisation pressure') shipping movement data were used to determine, for each season, the number of ships that visited South African ports from foreign ports and the number of days travelled between ports. Seasonal marine and terrestrial environmental similarity between South African and foreign ports was then used to estimate the likelihood that introduced species would establish. These data were used to determine the seasonal relative contribution of shipping routes to South Africa's marine and terrestrial establishment debt. Additionally, distribution data were used to identify marine and terrestrial species that are known to be invasive elsewhere and which might be introduced to each South African port through shipping routes that have a high relative contribution to establishment debt. Shipping routes from Asian ports, especially Singapore, have a particularly high relative contribution to South Africa's establishment debt, while among South African ports, Durban has the highest risk of being invaded. There was seasonal variation in the shipping routes that have a high relative contribution to the establishment debt of the South African ports. The presented method provides a simple way to prioritise surveillance effort and our results indicate that, for South Africa, port-specific prevention strategies should be developed, a large portion of the available resources should be allocated to Durban, and seasonal variations and their consequences for prevention strategies should be explored further.

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.

The global distribution of bamboos: assessing correlates of introduction and invasion.

There is a long history of species being moved around the world by humans. These introduced species can provide substantial benefits, but they can also have undesirable consequences. We explore the importance of human activities on the processes of species dissemination and potential invasions using the grass subfamily Bambusoideae ("bamboos"), a group that contains taxa that are widely utilised and that are often perceived as weedy. We (1) compiled an inventory of bamboo species and their current distributions; (2) determined which species have been introduced and become invasive outside their native ranges; and (3) explored correlates of introduction and invasion. Distribution data were collated from Kew's GrassBase, the Global Biodiversity Information Facility and other online herbarium information sources. Our list comprised 1662 species in 121 genera, of which 232 (14%) have been introduced beyond their native ranges. Twelve species were found to be invasive. A non-random selection of bamboos have been introduced and become invasive. Asiatic species in particular have been widely introduced. There was a clear over-representation of introduced species in the genera Bambusa and Phyllostachys which also contain most of the listed invasive species. The introduction of species also correlated with certain traits: taxa with larger culm dimensions were significantly more likely to have been moved to new areas; and those with many cultivars had a higher rate of dissemination and invasion. It is difficult to determine whether the patterns of introduction and invasion are due simply to differences in propagule pressure, or whether humans have deliberately selected inherently invasive taxa. In general, we suggest that human usage is a stronger driver of introductions and invasions in bamboos than in other taxa that have been well studied. It is likely that as bamboos are used more widely, the number and impact of invasions will increase unless environmental risks are carefully managed.

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.