High-performing plastic clones best explain the spread of yellow monkeyflower from lowland to higher elevation areas in New Zealand.

The relative contribution of adaptation and phenotypic plasticity can vary between core and edge populations, with implications for invasive success. We investigated the spread of the invasive yellow monkeyflower, Erythranthe gutatta in New Zealand, where it is spreading from lowland agricultural land into high-elevation conservation areas. We investigated the extent of phenotypic variation among clones from across the South Island, looked for adaptation and compared degrees of plasticity among lowland core versus montane range-edge populations. We grew 34 clones and measured their vegetative and floral traits in two common gardens, one in the core range at 9 m a.s.l. and one near the range-edge at 560 m a.s.l. Observed trait variation was explained by a combination of genotypic diversity (as identified through common gardens) and high phenotypic plasticity. We found a subtle signature of local adaptation to lowland habitats but all clones were plastic and able to survive and reproduce in both gardens. In the range-edge garden, above-ground biomass was on average almost double and stolon length almost half that of the same clone in the core garden. Clones from low-elevation sites showed higher plasticity on average than those from higher elevation sites. The highest performing clones in the core garden were also top performers in the range-edge garden. These results suggest some highly fit general-purpose genotypes, possibly pre-adapted to New Zealand montane conditions, best explains the spread of E. gutatta from lowland to higher elevation areas.

Comparing the Above and Below-Ground Chemical Defences of Three Rumex Species Between Their Native and Introduced Provenances.

Compared to their native range, non-native plants often experience reduced levels of herbivory in the introduced range. This may result in reduced pressure to produce chemical defences that act against herbivores. We measured the most abundant secondary metabolites found in Rumex spp., namely oxalates, phenols and tannins. To test this hypothesis, we compared native (UK) and introduced (NZ) provenances of three different Rumex species (R. obtusifolius, R. crispus and R. conglomeratus, Polygonaceae) to assess whether any significant differences existed in their levels of chemical defences in either leaves and roots. All three species have previously been shown to support a lower diversity of insect herbivores and experience less herbivory in the introduced range. We further examined leaf herbivory on plants from both provenances when grown together in a common garden experiment in New Zealand to test whether any differences in damage might be consistent with variation in the quantity of chemical defences. We found that two Rumex species (R. obtusifolius and R. crispus) showed no evidence for a reduction in chemical defences, while a third (R. conglomeratus) showed only limited evidence. The common garden experiment revealed that the leaves analysed had low levels of herbivory (~ 0.5%) with no differences in damage between provenances for any of the three study species. Roots tended to have a higher concentration of tannins than shoots, but again showed no difference between the provenances. As such, the findings of this study provide no evidence for lower plant investments in chemical defences, suggesting that other factors explain the success of Rumex spp. in New Zealand.

Importance of greater interdisciplinarity and geographic scope when tackling the driving forces behind biological invasions.

Invasive non-native species are important drivers of ecosystem change, yet the driving forces of biological invasions themselves are poorly understood. Such information is essential to ensure policies focus on the most relevant drivers, and that future scenarios capture the full range of potential outcomes for invasive non-native species. I carried out a bibliometric analysis of articles published from 2000 to 2020 that address either invasive non-native species or biodiversity and ecosystem services and that also mention 1 or more drivers of ecosystem change. I examined 5 indirect drivers (demographic, economic, governance, sociocultural, and technological) and 6 direct drivers (climate change, invasive non-native species, land-use or sea-use change, natural hazards, pollution, and resource extraction). Using the Web of Science core collection of citation indexes, I undertook searches of article titles and keywords and retrieved 27,462 articles addressing invasive non-native species and 110,087 articles dealing with biodiversity or ecosystem services. Most research to date on biological invasions as well as on biodiversity and ecosystem services has focused on anthropogenic direct drivers of ecosystem change rather than indirect drivers. Yet currently, less than 18% of articles addressing biological invasions examined drivers of ecosystem change, a similar level to that found over 20 years ago for biodiversity or ecosystem services. Knowledge of the drivers of biological invasions is limited, emphasizes tractable drivers over those that require an interdisciplinary approach, and is biased toward developed economies. Drivers generally deemed important for biological invasions, such as governance and resource extraction, accounted for less than 2% of research effort. The absence of a systematic understanding of the forces that drive invasive non-native species and how they interact means that attempts to mitigate or forecast biological invasions are likely to fail. To address biological invasions requires a much better orientation of national and international research on drivers in relation to both their actual importance as well as their policy relevance.

Importancia de un Mayor Enfoque Geográfico e Interdisciplinario al Abordar las Causas detrás de las Invasiones Biológicas Resumen Las especies invasoras no nativas son causas importantes de los cambios en el ecosistema; sin embargo, las mismas causas de las invasiones biológicas no se entienden claramente. Dicha información es esencial para asegurar que las políticas se enfoquen en las causas más relevantes, y que los escenarios futuros capturen la gama completa de resultados potenciales para las especies invasoras no nativas. Realicé un análisis biométrico de artículos publicados desde el año 2000 y hasta el 2020 que tratan el tema de las especies invasoras no nativas o el de los servicios ambientales y la biodiversidad y que también mencionan uno o más causas de los cambios en el ecosistema. Examiné cinco causas indirectas (demográficas, económicas, de gestión, socioculturales y tecnológicas) y seis causas directas (cambio climático, especie invasora no nativa, cambio en el uso de suelo o del mar, peligros naturales, contaminación y extracción de recursos). Mediante la colección central de índices de referencia de la plataforma Web of Science, realicé la búsqueda de títulos de artículos y palabras clave y recopilé 27,462 artículos sobre especies invasoras no nativas y 110,087 sobre servicios ambientales y biodiversidad. Las especies invasoras no nativas estuvieron menos estudiadas que otras causas antropogénicas directas del cambio en los ecosistemas (p. ej.: cambio climático, cambio en el uso de suelo, contaminación y extracción de recursos). Actualmente, <18% de los artículos que abordan las invasiones biológicas examinaron las causas del cambio ambiental, un nivel similar al hallado hace más de 20 años para la biodiversidad o los servicios ambientales. El conocimiento sobre las causas de las invasiones biológicas está limitado, enfatiza a las causas rastreables por encima de aquellas que requieren de un enfoque interdisciplinario y está sesgado hacia las economías desarrolladas. Las causas que generalmente se consideran importantes para las invasiones biológicas, como la gestión y la extracción de recursos, representaron <2% del esfuerzo de investigación. La ausencia de un entendimiento sistemático de las fuerzas que impulsan a las especies invasoras no nativas y cómo interactúan significa que probablemente fallen los intentos por mitigar o pronosticar las invasiones biológicas. Para abordar las invasiones biológicas se requiere una mucho mejor orientación de la investigación nacional e internacional sobre las causas en relación con su importancia actual y su relevancia política.

Import volumes and biosecurity interventions shape the arrival rate of fungal pathogens.

Global trade and the movement of people accelerate biological invasions by spreading species worldwide. Biosecurity measures seek to allow trade and passenger movements while preventing incursions that could lead to the establishment of unwanted pests, pathogens, and weeds. However, few data exist to evaluate whether changes in trade volumes, passenger arrivals, and biosecurity measures have altered rates of establishment of nonnative species over time. This is particularly true for pathogens, which pose significant risks to animal and plant health and are consequently a major focus of biosecurity efforts but are difficult to detect. Here, we use a database of all known plant pathogen associations recorded in New Zealand to estimate the rate at which new fungal pathogens arrived and established on 131 economically important plant species over the last 133 years. We show that the annual arrival rate of new fungal pathogens increased from 1880 to about 1980 in parallel with increasing import trade volume but subsequently stabilised despite continued rapid growth in import trade and recent rapid increases in international passenger arrivals. Nevertheless, while pathogen arrival rates for crop and pasture species have declined in recent decades, arrival rates have increased for forestry and fruit tree species. These contrasting trends between production sectors reflect differences in biosecurity effort and suggest that targeted biosecurity can slow pathogen arrival and establishment despite increasing trade and international movement of people.

Global rise in emerging alien species results from increased accessibility of new source pools.

Our ability to predict the identity of future invasive alien species is largely based upon knowledge of prior invasion history. Emerging alien species-those never encountered as aliens before-therefore pose a significant challenge to biosecurity interventions worldwide. Understanding their temporal trends, origins, and the drivers of their spread is pivotal to improving prevention and risk assessment tools. Here, we use a database of 45,984 first records of 16,019 established alien species to investigate the temporal dynamics of occurrences of emerging alien species worldwide. Even after many centuries of invasions the rate of emergence of new alien species is still high: One-quarter of first records during 2000-2005 were of species that had not been previously recorded anywhere as alien, though with large variation across taxa. Model results show that the high proportion of emerging alien species cannot be solely explained by increases in well-known drivers such as the amount of imported commodities from historically important source regions. Instead, these dynamics reflect the incorporation of new regions into the pool of potential alien species, likely as a consequence of expanding trade networks and environmental change. This process compensates for the depletion of the historically important source species pool through successive invasions. We estimate that 1-16% of all species on Earth, depending on the taxonomic group, qualify as potential alien species. These results suggest that there remains a high proportion of emerging alien species we have yet to encounter, with future impacts that are difficult to predict.

Projecting the continental accumulation of alien species through to 2050.

Biological invasions have steadily increased over recent centuries. However, we still lack a clear expectation about future trends in alien species numbers. In particular, we do not know whether alien species will continue to accumulate in regional floras and faunas, or whether the pace of accumulation will decrease due to the depletion of native source pools. Here, we apply a new model to simulate future numbers of alien species based on estimated sizes of source pools and dynamics of historical invasions, assuming a continuation of processes in the future as observed in the past (a business-as-usual scenario). We first validated performance of different model versions by conducting a back-casting approach, therefore fitting the model to alien species numbers until 1950 and validating predictions on trends from 1950 to 2005. In a second step, we selected the best performing model that provided the most robust predictions to project trajectories of alien species numbers until 2050. Altogether, this resulted in 3,790 stochastic simulation runs for 38 taxon-continent combinations. We provide the first quantitative projections of future trajectories of alien species numbers for seven major taxonomic groups in eight continents, accounting for variation in sampling intensity and uncertainty in projections. Overall, established alien species numbers per continent were predicted to increase from 2005 to 2050 by 36%. Particularly, strong increases were projected for Europe in absolute (+2,543 ± 237 alien species) and relative terms, followed by Temperate Asia (+1,597 ± 197), Northern America (1,484 ± 74) and Southern America (1,391 ± 258). Among individual taxonomic groups, especially strong increases were projected for invertebrates globally. Declining (but still positive) rates were projected only for Australasia. Our projections provide a first baseline for the assessment of future developments of biological invasions, which will help to inform policies to contain the spread of alien species.

Drivers of future alien species impacts: An expert-based assessment.

Understanding the likely future impacts of biological invasions is crucial yet highly challenging given the multiple relevant environmental, socio-economic and societal contexts and drivers. In the absence of quantitative models, methods based on expert knowledge are the best option for assessing future invasion trajectories. Here, we present an expert assessment of the drivers of potential alien species impacts under contrasting scenarios and socioecological contexts through the mid-21st century. Based on responses from 36 experts in biological invasions, moderate (20%-30%) increases in invasions, compared to the current conditions, are expected to cause major impacts on biodiversity in most socioecological contexts. Three main drivers of biological invasions-transport, climate change and socio-economic change-were predicted to significantly affect future impacts of alien species on biodiversity even under a best-case scenario. Other drivers (e.g. human demography and migration in tropical and subtropical regions) were also of high importance in specific global contexts (e.g. for individual taxonomic groups or biomes). We show that some best-case scenarios can substantially reduce potential future impacts of biological invasions. However, rapid and comprehensive actions are necessary to use this potential and achieve the goals of the Post-2020 Framework of the Convention on Biological Diversity.

A conceptual map of invasion biology: Integrating hypotheses into a consensus network.

BACKGROUND AND AIMS: Since its emergence in the mid-20th century, invasion biology has matured into a productive research field addressing questions of fundamental and applied importance. Not only has the number of empirical studies increased through time, but also has the number of competing, overlapping and, in some cases, contradictory hypotheses about biological invasions. To make these contradictions and redundancies explicit, and to gain insight into the field's current theoretical structure, we developed and applied a Delphi approach to create a consensus network of 39 existing invasion hypotheses. RESULTS: The resulting network was analysed with a link-clustering algorithm that revealed five concept clusters (resource availability, biotic interaction, propagule, trait and Darwin's clusters) representing complementary areas in the theory of invasion biology. The network also displays hypotheses that link two or more clusters, called connecting hypotheses, which are important in determining network structure. The network indicates hypotheses that are logically linked either positively (77 connections of support) or negatively (that is, they contradict each other; 6 connections). SIGNIFICANCE: The network visually synthesizes how invasion biology's predominant hypotheses are conceptually related to each other, and thus, reveals an emergent structure - a conceptual map - that can serve as a navigation tool for scholars, practitioners and students, both inside and outside of the field of invasion biology, and guide the development of a more coherent foundation of theory. Additionally, the outlined approach can be more widely applied to create a conceptual map for the larger fields of ecology and biogeography.

Resolving the invasion paradox: pervasive scale and study dependence in the native-alien species richness relationship.

The degree to which plant communities are vulnerable to invasion by alien species has often been assessed using the relationship between native and alien plant species richness (NAR). Variation in the direction and strength of the NAR tends to be negative for small plot sizes and study extents, but positive for large plots and extents. This invasion paradox has been attributed to different processes driving species richness at different spatial scales. However, the focus on plot size has drawn attention away from other factors influencing the NAR, in part because the influence of other factors may be obscured by or interact with plot size. Here, we test whether variation in the NAR can be explained by covariates linked to community susceptibility to invasion and whether these interact with plot size using a quantitative meta-analysis drawn from 87 field studies that examined 161 NARs. While plot size explained most variation, the NAR was less positive in grassland habitats and in the Australasian region. Other covariates did not show strong relationships with the NAR even after accounting for interactions with plot size. Instead, much of the unexplained variation is associated with article or author specific differences, suggesting the NAR depends strongly on how different authors choose their study system or study design.

Developing a list of invasive alien species likely to threaten biodiversity and ecosystems in the European Union.

The European Union (EU) has recently published its first list of invasive alien species (IAS) of EU concern to which current legislation must apply. The list comprises species known to pose great threats to biodiversity and needs to be maintained and updated. Horizon scanning is seen as critical to identify the most threatening potential IAS that do not yet occur in Europe to be subsequently risk assessed for future listing. Accordingly, we present a systematic consensus horizon scanning procedure to derive a ranked list of potential IAS likely to arrive, establish, spread and have an impact on biodiversity in the region over the next decade. The approach is unique in the continental scale examined, the breadth of taxonomic groups and environments considered, and the methods and data sources used. International experts were brought together to address five broad thematic groups of potential IAS. For each thematic group the experts first independently assembled lists of potential IAS not yet established in the EU but potentially threatening biodiversity if introduced. Experts were asked to score the species within their thematic group for their separate likelihoods of i) arrival, ii) establishment, iii) spread, and iv) magnitude of the potential negative impact on biodiversity within the EU. Experts then convened for a 2-day workshop applying consensus methods to compile a ranked list of potential IAS. From an initial working list of 329 species, a list of 66 species not yet established in the EU that were considered to be very high (8 species), high (40 species) or medium (18 species) risk species was derived. Here, we present these species highlighting the potential negative impacts and the most likely biogeographic regions to be affected by these potential IAS.

The emerging science of linked plant-fungal invasions.

Contents 1314 I. 1315 II. 1316 III. 1322 IV. 1323 V. 1325 VI. 1326 VII. 1326 VIII. 1327 1328 References 1328 SUMMARY: Invasions of alien plants are typically studied as invasions of individual species, yet interactions between plants and symbiotic fungi (mutualists and potential pathogens) affect plant survival, physiological traits, and reproduction and hence invasion success. Studies show that plant-fungal associations are frequently key drivers of plant invasion success and impact, but clear conceptual frameworks and integration across studies are needed to move beyond a series of case studies towards a more predictive understanding. Here, we consider linked plant-fungal invasions from the perspective of plant and fungal origin, simplified to the least complex representations or 'motifs'. By characterizing these interaction motifs, parallels in invasion processes between pathogen and mutualist fungi become clear, although the outcomes are often opposite in effect. These interaction motifs provide hypotheses for fungal-driven dynamics behind observed plant invasion trajectories. In some situations, the effects of plant-fungal interactions are inconsistent or negligible. Variability in when and where different interaction motifs matter may be driven by specificity in the plant-fungal interaction, the size of the effect of the symbiosis (negative to positive) on plants and the dependence (obligate to facultative) of the plant-fungal interaction. Linked plant-fungal invasions can transform communities and ecosystem function, with potential for persistent legacies preventing ecosystem restoration.

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.

New pasture plants intensify invasive species risk.

Agricultural intensification is critical to meet global food demand, but intensification threatens native species and degrades ecosystems. Sustainable intensification (SI) is heralded as a new approach for enabling growth in agriculture while minimizing environmental impacts. However, the SI literature has overlooked a major environmental risk. Using data from eight countries on six continents, we show that few governments regulate conventionally bred pasture taxa to limit threats to natural areas, even though most agribusinesses promote taxa with substantial weed risk. New pasture taxa (including species, subspecies, varieties, cultivars, and plant-endophyte combinations) are bred with characteristics typical of invasive species and environmental weeds. By introducing novel genetic and endophyte variation, pasture taxa are imbued with additional capacity for invasion and environmental impact. New strategies to prevent future problems are urgently needed. We highlight opportunities for researchers, agribusiness, and consumers to reduce environmental risks associated with new pasture taxa. We also emphasize four main approaches that governments could consider as they build new policies to limit weed risks, including (i) national lists of taxa that are prohibited based on environmental risk; (ii) a weed risk assessment for all new taxa; (iii) a program to rapidly detect and control new taxa that invade natural areas; and (iv) the polluter-pays principle, so that if a taxon becomes an environmental weed, industry pays for its management. There is mounting pressure to increase livestock production. With foresight and planning, growth in agriculture can be achieved sustainably provided that the scope of SI expands to encompass environmental weed risks.

Taxonomic similarity, more than contact opportunity, explains novel plant-pathogen associations between native and alien taxa.

Novel associations between plants and pathogens can have serious impacts on managed and natural ecosystems world-wide. The introduction of alien plants increases the potential for biogeographically novel plant-pathogen associations to arise when pathogens are transmitted from native to alien plant species and vice versa. We quantified biogeographically novel associations recorded in New Zealand over the last 150 yr between plant pathogens (fungi, oomycetes and plasmodiophorids) and vascular plants. We examined the extent to which taxonomic similarity, pathogen traits, contact opportunity and sampling effort could explain the number of novel associates for host and pathogen species. Novel associations were common; approximately one-third of surveyed plants and pathogens were recorded with at least one biogeographically novel associate. Native plants had more alien pathogens than vice versa. Taxonomic similarity between the native and alien flora and the total number of recorded associations (a measure of sampling effort) best explained the number of novel associates among species. The frequency of novel associations and the importance of sampling effort as an explanatory variable emphasize the need for effective monitoring and risk assessment tools to mitigate the potential environmental and economic impact of novel pathogen associations.

Europe's other debt crisis caused by the long legacy of future extinctions.

Rapid economic development in the past century has translated into severe pressures on species survival as a result of increasing land-use change, environmental pollution, and the spread of invasive alien species. However, though the impact of these pressures on biodiversity is substantial, it could be seriously underestimated if population declines of plants and animals lag behind contemporary environmental degradation. Here, we test for such a delay in impact by relating numbers of threatened species appearing on national red lists to historical and contemporary levels of socioeconomic pressures. Across 22 European countries, the proportions of vascular plants, bryophytes, mammals, reptiles, dragonflies, and grasshoppers facing medium-to-high extinction risks are more closely matched to indicators of socioeconomic pressures (i.e., human population density, per capita gross domestic product, and a measure of land use intensity) from the early or mid-, rather than the late, 20th century. We conclude that, irrespective of recent conservation actions, large-scale risks to biodiversity lag considerably behind contemporary levels of socioeconomic pressures. The negative impact of human activities on current biodiversity will not become fully realized until several decades into the future. Mitigating extinction risks might be an even greater challenge if temporal delays mean many threatened species might already be destined toward extinction.

Functional equivalence, competitive hierarchy and facilitation determine species coexistence in highly invaded grasslands.

Alien and native plant species often differ in functional traits. Trait differences could lead to niche differences that minimize competitive interactions and stabilize coexistence. However, trait differences could also translate into average fitness differences, leading to a competitive hierarchy that prevents coexistence. We tested whether trait differences between alien and native species translated into average fitness or stabilizing niche differences, and whether competition could explain observed coexistence within invaded grassland communities (New Zealand). Trait differences reflected marked competitive hierarchy, suggesting average fitness differences. Species coexistence was determined by a trade-off between species susceptibility to herbivory vs competitive hierarchy and facilitation. Importantly, although aliens and natives differed in their trait values, they did not differ in their competitive response, highlighting the importance of equalizing mechanisms in structuring invaded communities. Only a few alien species with a particular set of traits were able to jeopardize species coexistence when grazing was ceased. Our study explains why some alien species coexist with natives, whereas others have strong impacts on native communities. It highlights that trait differences can underlie several coexistence processes and that the demonstration of trait differences between aliens and natives is only a first step to understanding the role of biotic interactions in structuring invaded communities.

Resolving whether botanic gardens are on the road to conservation or a pathway for plant invasions.

A global conservation goal is to understand the pathways through which invasive species are introduced into new regions. Botanic gardens are a pathway for the introduction of invasive non-native plants, but a quantitative assessment of the risks they pose has not been performed. I analyzed data on the living collections of over 3000 botanic gardens worldwide to quantify the temporal trend in the representation of non-native species; the relative composition of threatened, ornamental, or invasive non-native plant species; and the frequency with which botanic gardens implement procedures to address invasive species. While almost all of the world's worst invasive non-native plants occurred in one or more living collections (99%), less than one-quarter of red-listed threatened species were cultivated (23%). Even when cultivated, individual threatened species occurred in few living collections (7.3), while non-native species were on average grown in 6 times as many botanic gardens (44.3). As a result, a botanic garden could, on average, cultivate four times as many invasive non-native species (20) as red-listed threatened species (5). Although the risk posed by a single living collection is small, the probability of invasion increases with the number of botanic gardens within a region. Thus, while both the size of living collections and the proportion of non-native species cultivated have declined during the 20th century, this reduction in risk is offset by the 10-fold increase in the number of botanic gardens established worldwide. Unfortunately, botanic gardens rarely implement regional codes of conduct to prevent plant invasions, few have an invasive species policy, and there is limited monitoring of garden escapes. This lack of preparedness is of particular concern given the rapid increase in living collections worldwide since 1950, particularly in South America and Asia, and highlights past patterns of introduction will be a poor guide to determining future invasion risks.