Biological invasions are a population-level rather than a species-level phenomenon.

Biological invasions pose a rapidly expanding threat to the persistence, functioning and service provisioning of ecosystems globally, and to socio-economic interests. The stages of successful invasions are driven by the same mechanism that underlies adaptive changes across species in general-via natural selection on intraspecific variation in traits that influence survival and reproductive performance (i.e., fitness). Surprisingly, however, the rapid progress in the field of invasion science has resulted in a predominance of species-level approaches (such as deny lists), often irrespective of natural selection theory, local adaptation and other population-level processes that govern successful invasions. To address these issues, we analyse non-native species dynamics at the population level by employing a database of European freshwater macroinvertebrate time series, to investigate spreading speed, abundance dynamics and impact assessments among populations. Our findings reveal substantial variability in spreading speed and abundance trends within and between macroinvertebrate species across biogeographic regions, indicating that levels of invasiveness and impact differ markedly. Discrepancies and inconsistencies among species-level risk screenings and real population-level data were also identified, highlighting the inherent challenges in accurately assessing population-level effects through species-level assessments. In recognition of the importance of population-level assessments, we urge a shift in invasive species management frameworks, which should account for the dynamics of different populations and their environmental context. Adopting an adaptive, region-specific and population-focused approach is imperative, considering the diverse ecological contexts and varying degrees of susceptibility. Such an approach could improve and refine risk assessments while promoting mechanistic understandings of risks and impacts, thereby enabling the development of more effective conservation and management strategies.

Including a diverse set of voices to address biological invasions.

Inclusivity is fundamental to progress in understanding and addressing the global phenomena of biological invasions because inclusivity fosters a breadth of perspectives, knowledge, and solutions. Here, we report on how the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES) assessment on invasive alien species (IAS) prioritized inclusivity, the benefits of this approach, and the remaining challenges.

Inverse priority effects: A role for historical contingency during species losses.

Communities worldwide are losing multiple species at an unprecedented rate, but how communities reassemble after these losses is often an open question. It is well established that the order and timing of species arrival during community assembly shapes forthcoming community composition and function. Yet, whether the order and timing of species losses can lead to divergent community trajectories remains largely unexplored. Here, we propose a novel framework that sets testable hypotheses on the effects of the order and timing of species losses-inverse priority effects-and suggests its integration into the study of community assembly. We propose that the order and timing of species losses within a community can generate alternative reassembly trajectories, and suggest mechanisms that may underlie these inverse priority effects. To formalize these concepts quantitatively, we used a three-species Lotka-Volterra competition model, enabling to investigate conditions in which the order of species losses can lead to divergent reassembly trajectories. The inverse priority effects framework proposed here promotes the systematic study of the dynamics of species losses from ecological communities, ultimately aimed to better understand community reassembly and guide management decisions in light of rapid global change.

Underexplored and growing economic costs of invasive alien trees.

The high ecological impacts of many invasive alien trees have been well documented. However, to date, we lacked synthesis of their economic impacts, hampering management actions. Here, we summarize the cost records of invasive trees to (I) identify invasive trees with cost information and their geographic locations, (II) investigate the types of costs recorded and sectors impacted by invasive trees and (III) analyze the relationships between categories of uses of invasive trees and the invasion costs attributed to these uses. We found reliable cost records only for 72 invasive trees, accumulating a reported total cost of $19.2 billion between 1960 and 2020. Agriculture was the sector with the highest cost records due to invasive trees. Most costs were incurred as resource damages and losses ($3.5 billion). Close attention to the ornamental sector is important for reducing the economic impact of invasive trees, since most invasive trees with cost records were introduced for that use. Despite massive reported costs of invasive trees, there remain large knowledge gaps on most invasive trees, sectors, and geographic scales, indicating that the real cost is severely underestimated. This highlights the need for further concerted and widely-distributed research efforts regarding the economic impact of invasive trees.

Plant invasion shifts soil microbiome and physico-chemical attributes along an elevational gradient in Kashmir Himalaya.

Soil microbial communities, being situated at the interface of aboveground plant and belowground soil systems, can play a pivotal role in determining ecosystem response to the drivers of global environmental change, including invasive species. In mountains, invasive plants occurring along elevational gradients offer a unique natural experimental system to investigate the impact of invasions in determining patterns and relationships of soil microbial diversity and nutrient pools at much shorter spatial distances. Here, we studied the impact of a global plant invader, Leucanthemum vulgare, on the diversity of soil microbiome and physico-chemical attributes along an elevational gradient (1760-2880 m) in Kashmir Himalaya. We used Illumina MiSeq platform to characterize the soil microbiome in pair-wise invaded and uninvaded plots at four different sites along the gradient. We found a total of 1959 bacterial operational taxonomic units (OTUs) belonging to 152 species, and a relatively higher number of 2475 fungal OTUs belonging to 589 species. The α-diversity of soil microbiome showed a gradual increase from low to high elevation and differed significantly (p < 0.05) between the invaded and uninvaded plots. The ß-diversity revealed distinct microbiome clustering among the sampling sites. Plant invasion also altered soil physico-chemical attributes along the elevational gradient. Overall, our findings suggest that the L. vulgare-induced shifts in soil microbiome and nutrient pools may be a belowground self-reinforced mechanism to facilitate its successful invasion along the elevational gradient. Our study provides new insights into invasive plant-microbe relationships with wide implications for climate warming-driven elevational range shifts in mountains.

Predicting the impact of invasive trees from different measures of abundance.

Biological invasions produce negative impacts worldwide, causing massive economic costs and ecological impacts. Knowing the relationship between invasive species abundance and the magnitude of their impacts (abundance-impact curves) is critical to designing prevention and management strategies that effectively tackle these impacts. However, different measures of abundance may produce different abundance-impact curves. Woody plants are among the most transformative invaders, especially in grassland ecosystems because of the introduction of hitherto absent life forms. In this study, our first goal was to assess the impact of a woody invader, Pinus contorta (hereafter pine), on native grassland productivity and livestock grazing in Patagonia (Argentina), building abundance-impact curves. Our second goal, was to compare different measure of pine abundance (density, basal area and canopy cover) as predictors of pine's impact on grassland productivity. Our third goal, was to compare abundance-impact curves among the mentioned measures of pine abundance and among different measures of impact: total grassland productivity, palatable productivity and sheep stocking rate (the number of sheep that the grassland can sustainably support). Pine canopy cover, closely followed by basal area, was the measure of abundance that best explained the impact on grassland productivity, but the shape of abundance impact curves differed between measures of abundance. While increases in pine density and basal area always reduced grassland productivity, pine canopy cover below 30% slightly increased grassland productivity and higher values caused an exponential decline. This increase in grassland productivity with low levels of pine canopy cover could be explained by the amelioration of stressful abiotic conditions for grassland species. Different measures of impact, namely total productivity, palatable productivity and sheep stocking rate, drew very similar results. Our abundance-impact curves are key to guide the management of invasive pines because a proper assessment of how many invasive individuals (per surface unit) are unacceptable, according to environmental or economic impact thresholds, is fundamental to define when to start management actions.

Recommendations for making editorial boards diverse and inclusive.

Lack of diversity in editorial boards hinders multifaceted perspectives in fields such as ecology, evolution, and conservation. We outline ten key actions for editorial boards to promote equity, diversity, and inclusion, benefiting the journal in attracting a wider readership, enhancing diversity among authors, and overcoming biases in editorial decisions.

Think globally, measure locally: The MIREN standardized protocol for monitoring plant species distributions along elevation gradients.

Climate change and other global change drivers threaten plant diversity in mountains worldwide. A widely documented response to such environmental modifications is for plant species to change their elevational ranges. Range shifts are often idiosyncratic and difficult to generalize, partly due to variation in sampling methods. There is thus a need for a standardized monitoring strategy that can be applied across mountain regions to assess distribution changes and community turnover of native and non-native plant species over space and time. Here, we present a conceptually intuitive and standardized protocol developed by the Mountain Invasion Research Network (MIREN) to systematically quantify global patterns of native and non-native species distributions along elevation gradients and shifts arising from interactive effects of climate change and human disturbance. Usually repeated every five years, surveys consist of 20 sample sites located at equal elevation increments along three replicate roads per sampling region. At each site, three plots extend from the side of a mountain road into surrounding natural vegetation. The protocol has been successfully used in 18 regions worldwide from 2007 to present. Analyses of one point in time already generated some salient results, and revealed region-specific elevational patterns of native plant species richness, but a globally consistent elevational decline in non-native species richness. Non-native plants were also more abundant directly adjacent to road edges, suggesting that disturbed roadsides serve as a vector for invasions into mountains. From the upcoming analyses of time series, even more exciting results can be expected, especially about range shifts. Implementing the protocol in more mountain regions globally would help to generate a more complete picture of how global change alters species distributions. This would inform conservation policy in mountain ecosystems, where some conservation policies remain poorly implemented.

Identifying Priorities, Targets, and Actions for the Long-term Social and Ecological Management of Invasive Non-Native Species.

Formulating effective management plans for addressing the impacts of invasive non-native species (INNS) requires the definition of clear priorities and tangible targets, and the recognition of the plurality of societal values assigned to these species. These tasks require a multi-disciplinary approach and the involvement of stakeholders. Here, we describe procedures to integrate multiple sources of information to formulate management priorities, targets, and high-level actions for the management of INNS. We follow five good-practice criteria: justified, evidence-informed, actionable, quantifiable, and flexible. We used expert knowledge methods to compile 17 lists of ecological, social, and economic impacts of lodgepole pines (Pinus contorta) and American mink (Neovison vison) in Chile and Argentina, the privet (Ligustrum lucidum) in Argentina, the yellow-jacket wasp (Vespula germanica) in Chile, and grasses (Urochloa brizantha and Urochloa decumbens) in Brazil. INNS plants caused a greater number of impacts than INNS animals, although more socio-economic impacts were listed for INNS animals than for plants. These impacts were ranked according to their magnitude and level of confidence on the information used for the ranking to prioritise impacts and assign them one of four high-level actions-do nothing, monitor, research, and immediate active management. We showed that it is possible to formulate management priorities, targets, and high-level actions for a variety of INNS and with variable levels of available information. This is vital in a world where the problems caused by INNS continue to increase, and there is a parallel growth in the implementation of management plans to deal with them.

A solution for breaking the language barrier.

Global problems require global scientific solutions, but the dominance of the English language creates a large barrier for many non-English-proficient researchers to make their findings and knowledge accessible globally. Here, we propose integrating peer language proofing and translation systems in preprint platforms as a solution for promoting equity, diversity, and inclusion in science.

Making ecology really global.

Ecology must flourish globally, especially in a period of unprecedented anthropogenic global change. However, some regions dominate the ecological literature. Multiple barriers prevent global production and exchange of ecological knowledge. The first step towards solutions is acknowledging and diagnosing this inequality and embracing our geographical and cultural diversity.

Soil biotic and abiotic effects on seedling growth exhibit context-dependent interactions: evidence from a multi-country experiment on Pinus contorta invasion.

The success of invasive plants is influenced by many interacting factors, but evaluating multiple possible mechanisms of invasion success and elucidating the relative importance of abiotic and biotic drivers is challenging, and therefore rarely achieved. We used live, sterile or inoculated soil from different soil origins (native range and introduced range plantation; and invaded plots spanning three different countries) in a fully factorial design to simultaneously examine the influence of soil origin and soil abiotic and biotic factors on the growth of invasive Pinus contorta. Our results displayed significant context dependency in that certain soil abiotic conditions in the introduced ranges (soil nitrogen, phosphorus or carbon content) influenced responses to inoculation treatments. Our findings do not support the enemy release hypothesis or the enhanced mutualism hypothesis, as biota from native and plantation ranges promoted growth similarly. Instead, our results support the missed mutualism hypothesis, as biota from invasive ranges were the least beneficial for seedling growth. Our study provides a novel perspective on how variation in soil abiotic factors can influence plant-soil feedbacks for an invasive tree across broad biogeographical contexts.

Invasive trees rely more on mycorrhizas, countering the ideal-weed hypothesis.

The ideal-weed hypothesis predicts that invasive plants should be less dependent on mutualisms. However, evidence in favor of or against this hypothesis comes mainly from observational studies. Here, we experimentally tested this hypothesis using a two-factor greenhouse experiment, comparing the seedling growth response of different Pinus species (varying in invasiveness) to ectomycorrhizal fungal inoculation. Most species showed no response until they were 6 mo old, at which point inoculation increased growth between 10 and 260% among the different species. This growth response was higher for species with lower seed mass, higher dispersal ability, higher Z score (a proxy for invasiveness) and higher number of naturalized regions, all of which correspond to higher invasiveness. Our results show that timing is a crucial factor when comparing mycorrhizal dependency of different species. Dependence on mutualistic microorganisms could be part of a strategy in which invasive species produce smaller seeds, in greater number, which can disperse further, but where seedlings are more reliant on mycorrhizas to improve access to water, nutrients, and protection from pathogens. Our results suggest that reliance on mutualisms may enhance, rather than limit, nonnative species in their ability to spread, establish, and colonize.

Non-English languages enrich scientific knowledge: The example of economic costs of biological invasions.

We contend that the exclusive focus on the English language in scientific research might hinder effective communication between scientists and practitioners or policy makers whose mother tongue is non-English. This barrier in scientific knowledge and data transfer likely leads to significant knowledge gaps and may create biases when providing global patterns in many fields of science. To demonstrate this, we compiled data on the global economic costs of invasive alien species reported in 15 non-English languages. We compared it with equivalent data from English documents (i.e., the InvaCost database, the most up-to-date repository of invasion costs globally). The comparison of both databases (~7500 entries in total) revealed that non-English sources: (i) capture a greater amount of data than English sources alone (2500 vs. 2396 cost entries respectively); (ii) add 249 invasive species and 15 countries to those reported by English literature, and (iii) increase the global cost estimate of invasions by 16.6% (i.e., US$ 214 billion added to 1.288 trillion estimated from the English database). Additionally, 2712 cost entries - not directly comparable to the English database - were directly obtained from practitioners, revealing the value of communication between scientists and practitioners. Moreover, we demonstrated how gaps caused by overlooking non-English data resulted in significant biases in the distribution of costs across space, taxonomic groups, types of cost, and impacted sectors. Specifically, costs from Europe, at the local scale, and particularly pertaining to management, were largely under-represented in the English database. Thus, combining scientific data from English and non-English sources proves fundamental and enhances data completeness. Considering non-English sources helps alleviate biases in understanding invasion costs at a global scale. Finally, it also holds strong potential for improving management performance, coordination among experts (scientists and practitioners), and collaborative actions across countries. Note: non-English versions of the abstract and figures are provided in Appendix S5 in 12 languages.

The role of arbuscular mycorrhizal fungi in nonnative plant invasion along mountain roads.

Plant associated mutualists can mediate invasion success by affecting the ecological niche of nonnative plant species. Anthropogenic disturbance is also key in facilitating invasion success through changes in biotic and abiotic conditions, but the combined effect of these two factors in natural environments is understudied. To better understand this interaction, we investigated how disturbance and its interaction with mycorrhizas could impact range dynamics of nonnative plant species in the mountains of Norway. Therefore, we studied the root colonisation and community composition of arbuscular mycorrhizal (AM) fungi in disturbed vs undisturbed plots along mountain roads. We found that roadside disturbance strongly increases fungal diversity and richness while also promoting AM fungal root colonisation in an otherwise ecto-mycorrhiza and ericoid-mycorrhiza dominated environment. Surprisingly, AM fungi associating with nonnative plant species were present across the whole elevation gradient, even above the highest elevational limit of nonnative plants, indicating that mycorrhizal fungi are not currently limiting the upward movement of nonnative plants. We conclude that roadside disturbance has a positive effect on AM fungal colonisation and richness, possibly supporting the spread of nonnative plants, but that there is no absolute limitation of belowground mutualists, even at high elevation.

Sheep feeding preference as a tool to control pine invasion in Patagonia: influence of foliar toughness, terpenoids and resin content.

Herbivores modulate the structure and composition of plant communities, including plant invasions. This is conditioned by plant palatability which can be reduced by its chemical or physical traits. The effects that ungulates browsing has on pine invasions are variable and the empirical evidence on the causes of this variability is scarce. We experimentally explored how sheep browsing preference varies between seedlings of pine species with different invasiveness; Pinus contorta (high invasiveness), P. ponderosa (medium invasiveness), P. radiata (medium invasiveness) and P. jeffreyi (low invasiveness). Secondly, we quantified anti-herbivory chemical compounds and physical traits of these species and related them with sheep preference observed. The browsing incidence of P. contorta was 68%, P. ponderosa 58%, P. radiata 29%, and P. jeffreyi 84%. Among anti-herbivory traits analyzed, α-pinene concentration had a negative effect on the probability of a terminal bud being browsed and on browsing intensity. Meanwhile, foliar toughness was negatively related to browsing intensity and water concentration was positively related to browsing intensity. Also, the most invasive species, P. contorta, was highly damaged. Thus, sheep herbivory could be slowing pine invasion rate; suggesting that could be considered a tool to control early invasions, especially for this particular 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.

Invasion Science and the Global Spread of SARS-CoV-2.

Emerging infectious diseases, such as coronavirus disease 2019 (COVID-19), are driven by ecological and socioeconomic factors, and their rapid spread and devastating impacts mirror those of invasive species. Collaborations between biomedical researchers and ecologists, heretofore rare, are vital to limiting future outbreaks. Enhancing the crossdisciplinary framework offered by invasion science could achieve this goal.

Highly invasive tree species are more dependent on mutualisms.

Why some species become invasive while others do not remains an elusive question. It has been proposed that invasive species should depend less on mutualisms, because their spread would then be less constrained by the availability of mutualistic partners. We tested this idea with the genus Pinus, whose degree of invasiveness is known at the species level (being highly and negatively correlated with seed size), and which forms obligate mutualistic associations with ectomycorrhizal fungi (EMF). Mycorrhizal dependence is defined as the degree to which a plant needs the mycorrhizal fungi to show the maximum growth. In this regard, we use plant growth response to mycorrhizal fungi as a proxy for mycorrhizal dependence. We assessed the responsiveness of Pinus species to EMF using 1,206 contrasts published on 34 species, and matched these data with data on Pinus species invasiveness. Surprisingly, we found that species that are more invasive depend more on mutualisms (EMF). Seedling growth of species with smaller seeds benefited more from mutualisms, indicating a higher dependence. A higher reliance on EMF could be part of a strategy in which small-seeded species produce more seeds that can disperse further, and these species are likely to establish only if facilitated by mycorrhizal fungi. On the contrary, big-seeded species showed a lower dependence on EMF, which may be explained by their tolerance to stressful conditions during establishment. However, the limited dispersal of larger seeds may limit the spread of these species. We present strong evidence against a venerable belief in ecology that species that rely more on mutualisms are less prone to invade, and suggest that in certain circumstances greater reliance on mutualists can increase spread capacity.