Curbing the major and growing threats from invasive alien species is urgent and achievable.

Although invasive alien species have long been recognized as a major threat to nature and people, until now there has been no comprehensive global review of the status, trends, drivers, impacts, management and governance challenges of biological invasions. The Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES) Thematic Assessment Report on Invasive Alien Species and Their Control (hereafter 'IPBES invasive alien species assessment') drew on more than 13,000 scientific publications and reports in 15 languages as well as Indigenous and local knowledge on all taxa, ecosystems and regions across the globe. Therefore, it provides unequivocal evidence of the major and growing threat of invasive alien species alongside ambitious but realistic approaches to manage biological invasions. The extent of the threat and impacts has been recognized by the 143 member states of IPBES who approved the summary for policymakers of this assessment. Here, the authors of the IPBES assessment outline the main findings of the IPBES invasive alien species assessment and highlight the urgency to act now.

Habitat modifies the relationship between grass and herbivore species richness in a South African savanna.

The savanna ecosystem is dominated by grasses, which are a key food source for many species of grazing animals. This relationship creates a diverse mosaic of habitats and contributes to the high grass species richness of savannas. However, how grazing interacts with environmental conditions in determining grass species richness and abundance in savannas is still insufficiently understood. In the Kruger National Park, South Africa, we recorded grass species and estimated their covers in 60 plots 50 × 50 m in size, accounting for varying proximity to water and different bedrocks. To achieve this, we located plots (i) near perennial rivers, near seasonal rivers, and on crests that are distant from all water sources and (ii) on nutrient-rich basaltic and nutrient-poor granitic bedrock. The presence and abundance of large herbivores were recorded by 60 camera traps located in the same plots. Grass cover was higher at crests and seasonal rivers than at perennial rivers and on basalts than on granites. The relationship between grass species richness and herbivore abundance or species richness was positive at crests, while that between grass species richness and herbivore species richness was negative at seasonal rivers. We found no support for controlling the dominance of grasses by herbivores in crests, but herbivore-induced microsite heterogeneity may account for high grass species richness there. In contrast, the decrease in grass species richness with herbivore species richness at seasonal rivers indicates that the strong grazing pressure over-rides the resistance of some species to grazing and trampling. We suggest that the relationships between grasses and herbivores may work in both directions, but the relationship is habitat-dependent, so that in less productive environments, the effect of herbivores on vegetation prevails, while in more productive environments along rivers the effect of vegetation and water supply on herbivores is more important.

Water availability, bedrock, disturbance by herbivores, and climate determine plant diversity in South-African savanna.

To identify factors that drive plant species richness in South-African savanna and explore their relative importance, we sampled plant communities across habitats differing in water availability, disturbance, and bedrock, using the Kruger National Park as a model system. We made plant inventories in 60 plots of 50 × 50 m, located in three distinct habitats: (i) at perennial rivers, (ii) at seasonal rivers with water available only during the rainy season, and (iii) on crests, at least ~ 5 km away from any water source. We predicted that large herbivores would utilise seasonal rivers' habitats less intensely than those along perennial rivers where water is available throughout the year, including dry periods. Plots on granite harboured more herbaceous and shrub species than plots on basalt. The dry crests were poorer in herb species than both seasonal and perennial rivers. Seasonal rivers harboured the highest numbers of shrub species, in accordance with the prediction of the highest species richness at relatively low levels of disturbance and low stress from the lack of water. The crests, exposed to relatively low pressure from grazing but stressed by the lack of water, are important from the conservation perspective because they harbour typical, sometimes rare savanna species, and so are seasonal rivers whose shrub richness is stimulated and maintained by the combination of moderate disturbance imposed by herbivores and position in the middle of the water availability gradient. To capture the complexity of determinants of species richness in KNP, we complemented the analysis of the above local factors by exploring large-scale factors related to climate, vegetation productivity, the character of dominant vegetation, and landscape features. The strongest factor was temperature; areas with the highest temperatures reveal lower species richness. Our results also suggest that Colophospermum mopane, a dominant woody species in the north of KNP is not the ultimate cause of the lower plant diversity in this part of the park.

Thirteen moth species (Lepidoptera, Erebidae, Noctuidae) newly recorded in South Africa, with comments on their distribution.

Background: Thanks to the high diversity of ecosystems and habitats, South Africa harbours tremendous diversity of insects. The Kruger National Park, due to its position close to the border between two biogeographic regions and high heterogeneity of environmental conditions, represents an insufficiently studied hotspot of lepidopteran diversity. During our ecological research in the Kruger National Park, we collected abundant moth material, including several interesting faunistic records reported in this study. New information: We reported 13 species of moths which had not yet been recorded in South Africa. In many cases, our records represented an important extension of the species' known distribution, including two species (Ozarbagaedei and O.persinua) whose distribution ranges extended into the Zambezian biogeographic region. Such findings confirmed the poor regional knowledge of lepidopteran diversity.

The invasive cactus Opuntia stricta creates fertility islands in African savannas and benefits from those created by native trees.

The patchy distribution of trees typical of savannas often results in a discontinuous distribution of water, nutrient resources, and microbial communities in soil, commonly referred to as "islands of fertility". We assessed how this phenomenon may affect the establishment and impact of invasive plants, using the invasion of Opuntia stricta in South Africa's Kruger National Park as case study. We established uninvaded and O. stricta-invaded plots under the most common woody tree species in the study area (Vachellia nilotica subsp. kraussiana and Spirostachys africana) and in open patches with no tree cover. We then compared soil characteristics, diversity and composition of the soil bacterial communities, and germination performance of O. stricta and native trees between soils collected in each of the established plots. We found that the presence of native trees and invasive O. stricta increases soil water content and nutrients, and the abundance and diversity of bacterial communities, and alters soil bacterial composition. Moreover, the percentage and speed of germination of O. stricta were higher in soils conditioned by native trees compared to soils collected from open patches. Finally, while S. africana and V. nilotica trees appear to germinate equally well in invaded and uninvaded soils, O. stricta had lower and slower germination in invaded soils, suggesting the potential release of phytochemicals by O. stricta to avoid intraspecific competition. These results suggest that the presence of any tree or shrub in savanna ecosystems, regardless of origin (i.e. native or alien), can create favourable conditions for the establishment and growth of other plants.

The dominating influence of efficacy above management strategy in the long-term success of alien plant clearing programmes.

Conservation managers are required to make decisions in complex and uncertain contexts. To strengthen the robustness of conservation decisions, several approaches have been proposed to facilitate stakeholder engagement in the setting of conservation objectives and priority actions. While such processes have led to the formulation of several invasive alien plant management strategies to achieve specific objectives, the long-term consequences and trade-offs inherent in these strategies have not been tested. The performance of five of these strategies over 50 years was tested in the protected area context using empirical data from Table Mountain National Park, South Africa. A simulation model based on data for invasive Acacia species in a fire-driven ecosystem, focused on the interaction between strategy performance and clearing efficacy in achieving a management goal or reducing Acacia density to below 1 plant per hectare. At near perfect levels of clearing efficacy, all strategies converged towards reaching the management goal, while at lower efficacy levels the strategies diverged in their ability to achieve desired outcomes. Despite working across the largest area, strategies that focussed on clearing low density invasions, maintained the least area in a maintenance state over time. In contrast, strategies that focussed on a mix of post-fire, low density areas and high altitude areas cleared less area annually, but maintained a much larger area in a maintenance state. At higher levels of efficacy, strategies that return to previously worked areas were more successful than a post-fire strategy. Strategies that focused solely on securing water, performed poorly in maintaining low overall density of aliens. However, the influence of efficacy was significant and substantial and a much larger difference in area reaching the management goal was achieved by varying efficacy than varying strategy. As such, improving quality of work and implementation will have a far greater effect than which areas are prioritized or how this prioritization is done. While acacias are likely to persist in the long-term, improving work quality coupled with correct strategy selection will ensure continued gains in the area under maintenance and improved return on investment over time.

Metabolomic differences between invasive alien plants from native and invaded habitats.

Globalization facilitated the spread of invasive alien species (IAS), undermining the stability of the world's ecosystems. We investigated the metabolomic profiles of three IAS species: Chromolaena odorata (Asteraceae) Datura stramonium (Solanaceae), and Xanthium strumarium (Asteraceae), comparing metabolites of individual plants in their native habitats (USA), to their invasive counterparts growing in and around Kruger National Park (South Africa, ZA). Metabolomic samples were collected using RApid Metabolome Extraction and Storage (RAMES) technology, which immobilizes phytochemicals on glass fiber disks, reducing compound degradation, allowing long-term, storage and simplifying biochemical analysis. Metabolomic differences were analyzed using ultra-performance liquid chromatography-mass spectrometry (UPLC-MS) of samples eluted from RAMES disks. Partial Least Squares-Discriminant Analysis (PLS-DA) of metabolomes of individual plants allowed statistical separation of species, native and invasive populations of each species, and some populations on the same continent. Invasive populations of all species were more phytochemically diverse than their native counterparts, and their metabolomic profiles were statistically distinguishable from their native relatives. These data may elucidate the mechanisms of successful invasion and rapid adaptive evolution of IAS. Moreover, RAMES technology combined with PLS-DA statistical analysis may allow taxonomic identification of species and, possibly, populations within each species.

Scientists' warning on invasive alien species.

Biological invasions are a global consequence of an increasingly connected world and the rise in human population size. The numbers of invasive alien species - the subset of alien species that spread widely in areas where they are not native, affecting the environment or human livelihoods - are increasing. Synergies with other global changes are exacerbating current invasions and facilitating new ones, thereby escalating the extent and impacts of invaders. Invasions have complex and often immense long-term direct and indirect impacts. In many cases, such impacts become apparent or problematic only when invaders are well established and have large ranges. Invasive alien species break down biogeographic realms, affect native species richness and abundance, increase the risk of native species extinction, affect the genetic composition of native populations, change native animal behaviour, alter phylogenetic diversity across communities, and modify trophic networks. Many invasive alien species also change ecosystem functioning and the delivery of ecosystem services by altering nutrient and contaminant cycling, hydrology, habitat structure, and disturbance regimes. These biodiversity and ecosystem impacts are accelerating and will increase further in the future. Scientific evidence has identified policy strategies to reduce future invasions, but these strategies are often insufficiently implemented. For some nations, notably Australia and New Zealand, biosecurity has become a national priority. There have been long-term successes, such as eradication of rats and cats on increasingly large islands and biological control of weeds across continental areas. However, in many countries, invasions receive little attention. Improved international cooperation is crucial to reduce the impacts of invasive alien species on biodiversity, ecosystem services, and human livelihoods. Countries can strengthen their biosecurity regulations to implement and enforce more effective management strategies that should also address other global changes that interact with invasions.

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.

Scenarios for the management of invasive Acacia species in a protected area: Implications of clearing efficacy.

In many protected areas in South Africa, invasive Australian Acacia species pose on-going management challenges, perpetuating high long-term management costs. Due to limited availability of resources, conservation actions need to be prioritised within and across Protected Areas (PA). We draw on comprehensive datasets spanning over 20 years from the Table Mountain National Park to model long-term outcomes of clearing Acacia species at different levels of management clearing efficacy. We test a 50 year outlook based on current and 38 incremental levels of management efficacy, ranging from 5 to 100%, to assess under which scenarios a management goal of reducing Acacia density to below 1 plant per hectare for the 22,671 ha protected area is achieved. With the current clearing resources and maximum clearing efficacy (100% control), it would take between 32 and 42 years to attain the management goal. The modelling revealed two main drivers of Acacia persistence. Firstly, germination of seeds added to the seedbank from standing plants made a significantly larger contribution to future clearing requirements than fire stimulated seed germination or the existing (pre-management) seedbank. Secondly the relationship between the number of hectares and management units that could be treated and the efficacy of the treatment was non-linear. When clearing efficacy was decreased from 100% to the current project minimum target of 80% efficacy, the goal was not achieved in all areas, but the area that reached a density of <1 plant per hectare was significantly reduced to 53% of the PA for the simulated 50 years. Results emphasize the need to differentiate between increasing financial resources and increasing efficacy. While increasing financial resources allows for increased effort, this is of little value for Acacia management in the absence of an increase in clearing efficacy, as low quality implementation perpetuates the need for large budgets over time. Conversely, improving efficacy allows for decreased budget requirements over time, allowing fund re-direction to additional areas of alien species management such as the early detection and rapid control of newly introduced species.

Rapid, field-deployable method for collecting and preserving plant metabolome for biochemical and functional characterization.

Study of plant metabolome is a growing field of science that catalogs vast biochemical and functional diversity of phytochemicals. However, collecting and storing samples of plant metabolome, sharing these samples across the scientific community and making them compatible with bioactivity assays presents significant challenges to the advancement of metabolome research. We have developed a RApid Metabolome Extraction and Storage (RAMES) technology that allows efficient, highly compact, field-deployable collection and storage of libraries of plant metabolome. RAMES technology combines rapid extraction with immobilization of extracts on glass microfiber filter discs. Two grams of plant tissue extracted in ethanol, using a specially adapted Dremel® rotary tool, produces 25-35 replicas of 10 mm glass fiber discs impregnated with phytochemicals. These discs can be either eluted with solvents (such as 70% ethanol) to study the metabolomic profiles or used directly in a variety of functional assays. We have developed simple, non-sterile, anti-fungal, anti-bacterial, and anti-oxidant assays formatted for 24-multiwell plates directly compatible with RAMES discs placed inside the wells. Using these methods we confirmed activity in 30 out of 32 randomly selected anti-microbial medicinal plants and spices. Seven species scored the highest activity (total kill) in the anti-bacterial (bacteria from human saliva) and two anti-fungal screens (Fusarium spp. and Saccharomyces cerevisiae), providing functional validation of RAMES technology. RAMES libraries showed limited degradation of compounds after 12 months of storage at -20°C, while others remained stable. Fifty-eight percent of structures characterized in the extracts loaded onto RAMES discs could be eluted from the discs without significant losses. Miniaturized RAMES technology, as described and validated in this manuscript offers a labor, cost, and time-effective alternative to conventional collection of phytochemicals. RAMES technology enables creation of comprehensive metabolomic libraries from various ecosystems and geographical regions in a format compatible with further biochemical and functional studies.

The role and value of conservation agency research.

Governments charge their conservation agencies to safeguard biodiversity through protected areas and threat mitigation. Increasingly, conservation management and policy need to be supported by rigorous evidence provided by science. As such, institutional arrangements should consider and enable effective scientific research and information dissemination. What role can in-house agency research play in responding to this challenge? We examined the research capabilities of three conservation agencies from Australia, South Africa, and United States. Seven indicators were used to characterize the reliability and relevance of agency research. We found similarities among agencies in their patterns of peer-reviewed publication, cultures of research collaboration, and tendencies to align research with organizational objectives. Among the many and diverse activities that constitute the role of a contemporary agency researcher, we emphasize two key research dimensions: reliability, achieved through peer-reviewed research output, and relevance, achieved through active stakeholder engagement. Amid increasingly challenging realities for conservation of ecosystems, agency science functions are vital to providing the evidence base required for effective management and policy development.

Genetic analysis shows low levels of hybridization between African wildcats (Felis silvestris lybica) and domestic cats (F. s. catus) in South Africa.

Hybridization between domestic and wild animals is a major concern for biodiversity conservation, and as habitats become increasingly fragmented, conserving biodiversity at all levels, including genetic, becomes increasingly important. Except for tropical forests and true deserts, African wildcats occur across the African continent; however, almost no work has been carried out to assess its genetic status and extent of hybridization with domestic cats. For example, in South Africa it has been argued that the long-term viability of maintaining pure wildcat populations lies in large protected areas only, isolated from human populations. Two of the largest protected areas in Africa, the Kgalagadi Transfrontier and Kruger National Parks, as well as the size of South Africa and range of landscape uses, provide a model situation to assess how habitat fragmentation and heterogeneity influences the genetic purity of African wildcats. Using population genetic and home range data, we examined the genetic purity of African wildcats and their suspected hybrids across South Africa, including areas within and outside of protected areas. Overall, we found African wildcat populations to be genetically relatively pure, but instances of hybridization and a significant relationship between the genetic distinctiveness (purity) of wildcats and human population pressure were evident. The genetically purest African wildcats were found in the Kgalagadi Transfrontier Park, while samples from around Kruger National Park showed cause for concern, especially combined with the substantial human population density along the park's boundary. While African wildcat populations in South Africa generally appear to be genetically pure, with low levels of hybridization, our genetic data do suggest that protected areas may play an important role in maintaining genetic purity by reducing the likelihood of contact with domestic cats. We suggest that approaches such as corridors between protected areas are unlikely to remain effective for wildcat conservation, as the proximity to human settlements around these areas is projected to increase the wild/domestic animal interface. Thus, large, isolated protected areas will become increasingly important for wildcat conservation and efforts need to be made to prevent introduction of domestic cats into these areas.

Predicting incursion of plant invaders into Kruger National Park, South Africa: the interplay of general drivers and species-specific factors.

BACKGROUND: Overcoming boundaries is crucial for incursion of alien plant species and their successful naturalization and invasion within protected areas. Previous work showed that in Kruger National Park, South Africa, this process can be quantified and that factors determining the incursion of invasive species can be identified and predicted confidently. Here we explore the similarity between determinants of incursions identified by the general model based on a multispecies assemblage, and those identified by species-specific models. We analyzed the presence and absence of six invasive plant species in 1.0×1.5 km segments along the border of the park as a function of environmental characteristics from outside and inside the KNP boundary, using two data-mining techniques: classification trees and random forests. PRINCIPAL FINDINGS: The occurrence of Ageratum houstonianum, Chromolaena odorata, Xanthium strumarium, Argemone ochroleuca, Opuntia stricta and Lantana camara can be reliably predicted based on landscape characteristics identified by the general multispecies model, namely water runoff from surrounding watersheds and road density in a 10 km radius. The presence of main rivers and species-specific combinations of vegetation types are reliable predictors from inside the park. CONCLUSIONS: The predictors from the outside and inside of the park are complementary, and are approximately equally reliable for explaining the presence/absence of current invaders; those from the inside are, however, more reliable for predicting future invasions. Landscape characteristics determined as crucial predictors from outside the KNP serve as guidelines for management to enact proactive interventions to manipulate landscape features near the KNP to prevent further incursions. Predictors from the inside the KNP can be used reliably to identify high-risk areas to improve the cost-effectiveness of management, to locate invasive plants and target them for eradication.

Protected-area boundaries as filters of plant invasions.

Human land uses surrounding protected areas provide propagules for colonization of these areas by non-native species, and corridors between protected-area networks and drainage systems of rivers provide pathways for long-distance dispersal of non-native species. Nevertheless, the influence of protected-area boundaries on colonization of protected areas by invasive non-native species is unknown. We drew on a spatially explicit data set of more than 27,000 non-native plant presence records for South Africa's Kruger National Park to examine the role of boundaries in preventing colonization of protected areas by non-native species. The number of records of non-native invasive plants declined rapidly beyond 1500 m inside the park; thus, we believe that the park boundary limited the spread of non-native plants. The number of non-native invasive plants inside the park was a function of the amount of water runoff, density of major roads, and the presence of natural vegetation outside the park. Of the types of human-induced disturbance, only the density of major roads outside the protected area significantly increased the number of non-native plant records. Our findings suggest that the probability of incursion of invasive plants into protected areas can be quantified reliably.

Ornamental plants as invasive aliens: problems and solutions in Kruger National Park, South Africa.

The most widespread invasive alien plant species in South Africa's Kruger National Park (KNP) were either introduced unintentionally along rivers and roads, or intentionally for use as ornamentals. We examine the spatial distribution of ornamental alien plants in KNP, look at the link between human population size, history, and species richness, and show how the distribution of particular species reflects the likely history of ornamental plantings. Results are used to assess whether past management actions have been appropriately directed. Two hundred and fifty-eight alien species have been recorded in the 36 tourist camps and staff villages. The number of staff housed in villages explains much of the diversity of cultivated alien plant species. Older camps also tend to have more ornamental alien plant species. However, the lack of a strong link between camp age and number of cultivated species suggests that ornamental plants have been widely spread around the KNP by humans. We also show that increased camp activity (either size or age) has led to more ornamental species, while, with the notable exception of Skukuza, camp activity has had a much smaller effect on the number of noncultivated species. Noncultivated species tend to be naturally dispersed, as opposed to directly spread by humans between camps. Past management prioritized certain species on the basis of their potential to invade KNP and on the prevailing national legislation. These species were removed manually and follow-up control was carried out. Once the priority species were deemed to be under control, less invasive species were targeted. All alien species were removed from vacated houses, regardless of the potential invasiveness of the species.

Risk assessment of riparian plant invasions into protected areas.

Protected areas are becoming increasingly isolated. River corridors represent crucial links to the surrounding landscape but are also major conduits for invasion of alien species. We developed a framework to assess the risk that alien plants in watersheds adjacent to a protected area will invade the protected area along rivers. The framework combines species- and landscape-level approaches and has five key components: (1) definition of the geographical area of interest, (2) delineation of the domain into ecologically meaningful zones, (3) identification of the appropriate landscape units, (4) categorization of alien species and mapping of their distribution and abundance, and (5) definition of management options. The framework guides the determination of species distribution and abundance through successive, easily followed steps, providing the means for the assessment of areas of concern. We applied the framework to Kruger National Park (KNP) in South Africa. We recorded 231 invasive alien plant species (of which 79 were major invaders) in the domain. The KNP is facing increasing pressure from alien species in the upper regions of the drainage areas of neighboring watersheds. On the basis of the climatic modeling, we showed that most major riparian invaders have the ability to spread across the KNP should they be transported down the rivers. With this information, KNP managers can identify areas for proactive intervention, monitoring, and resource allocation. Even for a very large protected area such as the KNP, sustainable management of biodiversity will depend heavily on the response of land managers upstream managing alien plants. We suggest that this framework is applicable to plants and other passively dispersed species that invade protected areas situated at the end of a drainage basin.