Alien Plant Species: Environmental Risks in Agricultural and Agro-Forest Landscapes Under Climate Change

Insights into ecological drivers of alien plant invasions can be gained through comparative studies of growth and fecundity of invasive alien plants versus those of co‐occurring non‐invasive alien plants and native plants across environmental conditions in common garden settings. Habitats that harbour alien plant species in many ecosystems globally are presently experiencing light pollution resulting from artificial light at night (ALAN) and increased rates of nutrient enrichment of the soil. However, the potential interactive effects of ALAN and nutrient enrichment on invasiveness of alien plant species remain unknown. Here, we performed a common‐garden experiment to test the interactive effects of ALAN and soil nutrient enrichment on the growth of a random set of ten alien (five invasive and five non‐invasive naturalized) and seven co‐occurring native ornamental plant species that are commonly cultivated within urban and peri‐urban areas of Nairobi city in Kenya. We predicted that a simultaneous increase in photoperiod via ALAN and nutrient enrichment will favour growth of invasive alien plant species over that of non‐invasive alien and native plant species. We grew the 17 plant species under natural daylight (ALAN−) versus natural daylight followed by ALAN (ALAN+) and fully crossed with two levels of nutrient enrichment (low vs. high) and competition (competition vs. no‐competition against a native plant Ocimum gratissimum ) treatments. Under simultaneous high‐nutrient and no‐competition treatments, ALAN enhanced mean total biomass of invasive and non‐invasive naturalized alien species by 61.1% and 131.4%, respectively but decreased that of native plant species by 34%. In contrast, under simultaneous high‐nutrient and competition treatments, ALAN enhanced mean total biomass of invasive alien plant species by 68.6% and that of non‐invasive naturalized alien species by 51.9% and native species by 35.4%. High‐nutrient treatment enhanced flower formation more strongly in invasive and non‐invasive naturalized alien plants than in native plants. The invasive and non‐invasive naturalized alien species grew taller than native species across the light, nutrient, and competition treatments. Synthesis : The present findings suggest that light pollution and nutrient enrichment may jointly confer growth advantage to invasive alien plant species over that of co‐occurring native plant species and enhance invasiveness of alien plant species.

Invasive alien plant species: From the molecular to the economic approach Invasive alien species (IAS) are plants, animals, pathogens and other organisms that are not natural in a certain ecosystem and that can cause economic or environmental damage or adversely affect human health (Ljubojevićet al., 2022) . Plant invasiveness is a globally recognized environmental and economic problem (Bang et al., 2022) . Today, the loss of biodiversity occurs due to intensive anthropogenic activities, like over-exploitation of species, urbanization and industrialization, environmental pollution, land-use shift, the introduction of IAS as well as climate change (Ren and Duan, 2017) . IAS introduction has many advantages and disadvantages, leading to numerous ecosystem services and disservices. Fostered by climate changes (Kariyawasam et al., 2019) and inherited (inner) ability to adapt to various conditions, IAS treat to suppress natural vegetation. Theoharides and Dukes (2007) defined four main stages of invasion: introduction, naturalization, colonization, and dispersal. Milanovićet al. (2020) stated that alien (especially invasive) plant species differ from native species in different morphological characteristics such as specific leaf area, height, seed size or flowering duration, where invasive species showed significant dominance in the investigated characteristics. A list of invasion-promoting traits is being amended as novel research is being conducted, from seed characteristics (Ljubojevićet al., 2021) to the whole-plant level (Bajwa et al., 2016) . With such high adaptive potential alien species develop traits that allow them to successfully cope with the changes in climate or habitats (Dukes and Mooney, 1999) . However, being very adaptive, fast-growing, and not infrequently highly ornamental, those species provide numerous ecosystem services. If viewed exclusively as 'weeds' then many resources would be invested in the eradication measures of invasive alien species, with little certainty about the desired outcome. Thus, this Research Topic gathered recent findings from around the world, providing insight into differences in gene expression, morphology, physiology and resource utilization by invasive alien species, that provided them with successful habitat invasion. As shown by (Nunes et al.) weedy invaders spread rapidly due to successful adaptation and naturalization, affecting both natural and agricultural ecosystems. One of the most invasive weed species in the sub-tropical and continental climate is Arundo donax L. from Poaceae family. Owing to the chemical and histological modifications, this species successfully inhabit the Mediterranean basin. Another invasion pathway includes soil Frontiers in Plant Science frontiersin.org 01

AimTo examine how native plant native communities, environment and geography are associated with alien plant species invasion in tropical island forests.LocationFour US national parks in Hawai'i and American Samoa.MethodsWe analysed the richness, coverage and composition of alien plant species communities located across 204 plots, five islands and two archipelagos. We used generalized linear mixed‐effects models and generalized dissimilarity models to determine the relative importance of native plant species communities (richness, structure and composition), environment (temperatures, precipitation, and soil age) and geography (elevation, geographical distance between plots, identity of islands and archipelagos) on alien plant species communities.ResultsAlien plant species were found in >90% of plots with on average four species covering ~15% of the understorey. Variation in alien species richness and coverage was primarily explained by archipelago and island identity and elevation. Plots located on younger islands tended to exhibit higher richness and coverage of alien plants. Native tree density and canopy height were negatively correlated with alien species coverage but not with alien species richness. Elevation was negatively correlated with both the richness and coverage of alien species. Differences in elevation, mean annual precipitation and geographical distances explained about 40% of the observed spatial turnover in alien species composition with differences in elevation being the main driver. Turnover in alien species composition was weakly correlated with turnover in native species.Main conclusionsIn the studied islands, higher elevations seem to be relatively spared from alien species invasion so far. However, our results suggest that the negative effect of elevation on alien species coverage may be larger than on alien species richness. Maintenance of tall and dense canopy may limit the coverage of alien species but not the number of alien species. Our results also support that alien species invasion may be particularly high on isolated archipelagos and younger islands.

The expansion of alien plant species is of global concern, yet our understanding of their dispersal mechanisms is limited. Here we address the potential of alien plant seeds to disperse via ingestion, transport and egestion in waterfowl (endozoochory). Based on their general rapid expansions, we expected alien plant species to have several advantages for endozoochory compared to native plant species. We hypothesised that seeds of alien species would have higher passage rate, longer gut retention times, higher germinability after gut passage and shorter time‐to‐germination after egestion by waterfowl. In order to test our hypotheses, we compared the endozoochorous dispersal ability of six pairs of congeneric alien and native wetland plant species in a feeding experiment with mallards (Anas platyrhynchos). We focused on differences in seed survival, gut retention time, germinability and time‐to‐germination. In the analyses we corrected for seed shape and volume as these seed traits are known to have important effects. With gut passage, alien species had higher passage rates and germinated slower, whereas native species had shorter retention times and greater germinability. Controlling for seed traits did not alter these conclusions, but seed traits affected all aspects of the endozoochory process. This suggests that alien species may have particular traits correlated with a higher endozoochory potential. Among control seeds, alien seeds germinated faster and their germinability was higher than natives. Seed traits explained differences in germinability and time‐to‐germination in control seeds. Seeds of alien plant species have traits that correlate with successful endozoochory. This may provide alien species with a competitive advantage over native plant species by ensuring higher endozoochory rates in new environments, potentially enabling their rapid expansions. Our study underlines the important role of seed traits in the endozoochory potential of alien and native plant species, notably through their influence on retention time and germination.

Rising human activity in the Arctic, combined with a warming climate, increases the probability of introduction and establishment of alien plant species. While settlements are known hotspots for persistent populations, little is known about colonization of particularly susceptible natural habitats. Systematic monitoring is lacking and available survey methods vary greatly. Here, we present the most comprehensive survey of alien vascular plant species in the high‐Arctic archipelago of Svalbard to date, aimed at (i) providing a status within settlements; (ii) surveying high‐risk habitats such as those with high visitor numbers and nutrient enrichment from sea bird colonies; (iii) presenting a systematic monitoring method that can be implemented in future work on alien plant species in Arctic environments; and (iv) discuss possibilities for mapping alien plant habitats using unmanned aerial vehicles. The systematic grid survey, covering 1.7 km2 over three settlements and six bird cliffs, detected 36 alien plant species. Alien plant species were exclusively found in areas of human activity, particularly areas associated with current or historic animal husbandry. The survey identified the successful eradication of Anthriscus sylvestris in Barentsburg, as well as the rapid expansion of Taraxacum sect. Ruderalia over the last few decades. As there is currently no consistent method for monitoring alien plant species tailored to polar environments, we propose a systematic methodology that could be implemented within a structured monitoring regime as part of an adaptive monitoring strategy towards alien species in the Arctic.

:This study explores the correlates of alien plant species richness in South Africa at the scale of quarter-degree squares (QDS; ª 25 ¥ 27 km; 675 km2). We considered all alien plant species for which we had records and a subset of these – those that invade natural and semi-natural vegetation. The main source of data for species richness of indigenous and alien plant species was a national database based on herbarium specimens. For invasive alien species, data were from a national atlassing project. First, we explored the importance of energy availability and habitat heterogeneity as correlates of indigenous, alien, and invasive alien plant species richness. Linear regression models showed that species richness in the three groups of plants was explained by the same variables: a principal component of climatic factors and topographic roughness were the top-ranking variables for all groups. Next, we examined the role of indigenous species richness together with a range of environmental and human-activity variables in explaining species richness of alien and invasive alien plants. Results reveal an interplay of natural features and variables that quantify the dimension of human activities. If indigenous species richness is ignored, human-activity variables are more strongly correlated with alien species richness than with invasive alien species richness. Numbers of alien and invasive species in QDSs are significantly correlated with indigenous plant species richness in the 1,597 QDSs selected for analysis, a pattern consistent with findings from other parts of the world. Analysis of residuals between observed and predicted values showed that patterns differed between biomes. The results are useful for planning long-term intervention policy at the national scale; they suggest that areas with rich native biodiversity will face a sustained onslaught from invasive alien species and that ongoing management actions will be required to reduce and mitigate impacts from biological invasions in these areas.

Invasive process of alien species is affected by not only the invaded habitats, but also the surrounding landscape matrix. Understanding the effects of landscape matrix on alien species is of great significance for controlling invasive alien species. We surveyed plant communities along the water-level fluctuating zone (WLFZ) of the Three Gorges Reservoir. Invasive status of alien plant species was evaluated. Totally 10 spatial scales of the surrounding landscape matrix in the scope of 2000 m (including WLFZ) were classified, and 14 landscape indices were applied to analyze the landscape matrix composition and configuration. Using the principal component analysis and correlation analysis, the effects of landscape matrix on the alien invasive plant species and associated scale effect were tested. Results showed that a total of 42 alien invasive plant species were found in the WLFZ, belonging to 17 families and 36 genera. Fuling was a dividing place to differentiate invasive species distribution. The number of the alien invasive species between Fuling and the Three Gorges Dam was found more than that between Fuling and Jiangjin. For the all scales (within 2000 m). The higher the landscape matrix fragmentation was, the more difficult the alien species invading. The higher landscape connectivity was, the easier the alien species invading. The effects of landscape matrix composition and configuration on the invasive plant diversity at large scales (1200-2000 m) was more significant than those at small scales (200-1000 m), in which landscape matrix composition and configuration at 1200-1400 m showed the strongest effect, demonstrating a significant spatial scale effect. Different invasive plant species showed the scale effects of landscape matrix composition and configuration. At all scales, Xanthium strumarium and Bidens frondosa showed weak correlations with landscape indices, but Bidens tripartita and Erigeron canadensis showed strong correlations. Landscape matrix was closely related to invasive plant species, and demonstrated a significant scale effect. The alien invasive plant species could be traced to the landscape matrix at large scales. Grassland and forest patches at the small scales could be used as the 'stepping stone' for the alien species transiting before they arrived at the WLFZ. In order to control alien plants in the WLFZ, land-use management and optimization should be strengthened at different scales of landscape matrix on the basis of enhancement of habitat management. A diversified comprehensive control for alien species should thus be taken into account.

Understanding the factors affecting the species richness of alien and native plant is a key issue for predicting the spread of alien species and protecting rare and endangered native species in nature reserves. To investigate the factors affecting alien and native species richness in temperate nature reserves of China, we used a database of 25 nature reserves in Shandong Province of northern China, and studied the relationship of alien and native plant species richness with climatic and anthropogenic factors, as well as area and elevation range. We found that most of the nature reserves in Shandong Province have been invaded by alien plant species. The distribution of alien and native species responds to the same climatic factors, and temperature and precipitation exert strong effects on both groups. Alien and native plant species richness are positively correlated. Human activity is more effective for explaining richness of alien than for native species. Simultaneously, human activity has stronger effects on alien herbaceous than on alien woody plants. Our results suggest that native species richness is mainly explained by climatic factors, whereas alien species richness is mainly explained by climatic factors and human activity together.

Biological invasions pose an increasing risk to nature, social security and the economy, being ranked amongst the top five threats to biodiversity. Managing alien and invasive species is a priority for the European Union, as outlined in the EU Biodiversity Strategy for 2030 and the Kunming-Montreal Global Biodiversity Framework. Alien plant species are acknowledged to impact the economy and biodiversity; thus, analysing the distribution of such species provides valuable inputs for the management and decision-making processes. The database presented in the current study is the first consolidated checklist of alien plant species that are present in Romania, both of European Union concern and of national interest. This database complements a prior published distribution, based only on records from literature, bringing new information regarding the occurrence of alien plants in Romania, as revealed by a nationwide field survey. We consider this database a valuable instrument for managing biological invasions at both national and regional levels, as it can be utilised in further research studies and in drafting management and action plans, assisting stakeholders in making informed decisions and implementing management actions. We present the results of the first nationwide survey of alien plant species in Romania, conducted between 2019 and 2022, in the framework of a national project coordinated by the Ministry of Environment, Waters and Forests and the University of Bucharest. The present database complements and updates the database published by Sirbu et. al (2022), which included occurrence records published until 2019. The new database includes 98323 occurrence records for 396 alien plant species in 77 families, with most species belonging to the Asteraceae family. One alien plant species in our database, the black locust Robiniapseudoacacia L., had more than 10,000 occurrence records. The distribution database also includes information on newly-reported invasive alien plant species of European Union concern in Romania (i.e. the floating primrose-willow Ludwigiapeploides (Kunth) P.H.Raven) and documents the presence of plants in 44 additional families compared to Sirbu et al. (2022). Each entry includes information on species taxonomy, location, year, person who recorded and identified the alien plant, geographical coordinates and taxon rank.

Functional trait variation predicts distribution of alien plant species across the light gradient in a temperate rainforest

The flora of the steppe Dnieper region is characterized by an abundance of naturalized alien species, some of which colonised over the last decade. Climate change, associated primarily with increasing temperature, became clearly manifested in this period. We tested the hypothesis that there is an association between climate change and the initiation of invasiveness of some alien plant species in the steppe Dnieper region. For this purpose, comparative studies of the distribution boundaries of naturalized alien trees, shrubs and herbaceous plants were conducted. Along the research route numerous 5–10-year-old broadleaf linden trees ( Tilia platyphyllos Scop.) were found in the man-made plantation communities of Dnipro city in areas with moist soil; seeded undergrowth was located at a significant distance from the adult linden plants. Numerous groups of young 7–10-year-old plants of the smoke trees ( Cotinus coggygria Scop.), which had a seed origin, were found in the shelterbelt and urban recreational plantations. Young 10–12-year-old virginal and generative plants of the black cherry ( Padus serotina Ehrh.) were found in large numbers in both the semi-natural and artificial plant communities at great distances from the adult trees. The alien plant species common hackberry ( Celtis occidentalis L.) showed the ability to form fairly sparse seminal seedlings, which was presented by the plants at the age of 4–7 years in both the natural and urban plant communities. The perennial herbaceous plant common milkweed ( Asclepias syriaca L.) was found in the course of the research in ruderal habitats, urban plant communities, and also in the agrocoenoses. The common milkweed reached its greatest abundance in fields of winter crops, where the spread of this species was accompanied by a sharp decrease in the number of other species of segetal plants. Our study results confirm that the extension of the distribution boundaries of alien species over the last decade was not related to the ground conditions of the steppe Dnieper region. At the same time, changes in climatic conditions were favourable for some naturalized alien species because they have created the opportunity for seed reproduction of species away from the maternal plants. Alien species C. coggigria , P. serotina and A. syriaca were also the most sensitive to the influence of the climate changes. Consequently, these species have the greatest potential for increasing their level of invasiveness and endangering the biodiversity in the steppe Dnieper region under conditions of climate change. We suggest that a simultaneous initiation of invasiveness of these several alien species leads to an increase in the degree of threat to the diversity of natural plants in the region. The study results confirm the urgent need for analysis and forecasting of the consequences of introduction of alien species, in order to prevent the undesirable effects that this would bring for the region’s native vegetation.

Invasion of alien plant species is one of the major drivers that alter ecosystems, and threaten native plant diversity. In this study, we compared the diversity of native and alien plant species (naturalized and invasive) in three forest types in the mid-hills of central Nepal. We selected eight community-managed forests in three districts with similar management practices. Altogether, 24 plots (50 m × 20 m each) were sampled to collect vegetation data. We recorded 274 (88%) native and 24 (8%) alien plant species. The species richness did not vary significantly across the forest types. Among the alien plants, 12 species were naturalized, 11 were invasive including the globally worst species Chromolaena odorata, and the remaining one was casual. Native plant diversity was high in the Pinus roxburghii forest while alien plant diversity was high in the Schima-Castanopsis forest. The Shorea robusta forest with relatively mature trees, low anthropogenic disturbances, and dense canopy cover had the lowest diversity of invasive alien plants followed by Pinus roxburghii and Schima-Castanopsis forests. Contrary to the expectation, there was no significant relationship between canopy cover and species richness of native and alien plant species. Although the three studied forest types did not significantly vary in plant species richness the recorded number of alien species indicates that the community forests of the mid-hills are being invaded rapidly by alien plants. Implementation of appropriate control measures is recommended to reduce the abundance of invasive alien plants in forests and avert likely negative impacts on native species and ecosystems.

The majority of plant species rely, at least partly, on animals for pollination. Our knowledge on whether pollinator visitation differs between native and alien plant species, and between invasive and non-invasive alien species is still limited. Additionally, because numerous invasive plant species are escapees from horticulture, the transition from human-assisted occurrence in urbanized habitats to unassisted persistence and spread in (semi-)natural habitats requires study. To address whether pollinator visitation differs between native, invasive alien and non-invasive alien species, we did pollinator observations for a total of 17 plant species representing five plant families. To test whether pollinator visitation to the three groups of species during the initial stage of invasion depends on habitat type, we did the study in three urbanized habitats and three semi-natural grasslands, using single potted plants. Native plants had more but smaller flower units than alien plants, and invasive alien plants had more but smaller flowers than non-invasive alien plants. After accounting for these differences in floral display, pollinator visitation was higher for native than for alien plant species, but did not differ between invasive and non-invasive alien plant species. Pollinator visitation was on average higher in semi-natural than in urbanized habitats, irrespective of origin or status of the plant species. This might suggest that once an alien species has managed to escape from urbanized into more natural habitats, pollinator limitation will not be a major barrier to establishment and invasion.

Summary Plants interact with many organisms, such as microbes and herbivores, and these interactions are likely to affect the establishment and spread of plants. In the context of plant invasions, mycorrhizal fungi and constitutive and induced resistance of plants against herbivores have received attention independently of each other. However, plants are frequently involved in complex multi‐trophic interactions, which might differ between invasive and non‐invasive alien plants. In a multi‐species comparative experiment, we aimed to improve our understanding of plant traits associated with invasiveness. We tested whether eight invasive alien plant species use the mycorrhizal symbiosis in a more beneficial way, and have higher levels of constitutive or induced resistance against two generalist bioassay herbivores, than nine non‐invasive alien species. We further assessed whether the presence of mycorrhizal fungi altered the resistance of the plant species, and whether this differed between invasive and non‐invasive alien species. While invasive species produced more biomass, they did not differ in their biomass response to mycorrhizal fungi from non‐invasive alien species. Invasive species also did not have higher levels of constitutive or induced resistance against the two generalist herbivores. Mycorrhizal fungi greatly affected the resistance of our plant species, however, this was also unrelated to whether the alien species were invasive or not. Our study confirms the previous findings that invasive species generally grow faster and produce more biomass than non‐invasive alien species. We further show that alien plant species used a variety of defence strategies, and also varied in their interactions with mycorrhizal fungi. These multi‐trophic interactions were not consistently related to invasiveness of the alien plant species. We suggest that awareness of the fact that alien plant species are involved in multi‐trophic interactions might lead to a more complete understanding of the factors contributing to a plant's success.

Alien species can drastically disrupt ecological processes such as those involving plant–pollinator interactions, performing central roles that may affect the structure of native pollination networks. However, most studies to date have focused on a single trophic level of alien species, evaluating either the impacts of an alien pollinator or an alien plant species, and have neglected their joint effects on the seasonal dynamics of mutualistic networks. Here, we aim to fill this gap by investigating how alien plant and flower visitor (here used as a proxy for pollinator) species structure temporal networks, and how these species affect the beta‐diversity of interactions across a flowering season. Our study system is located in the South Island of New Zealand, where 68% of the sampled plant species and 14% of the flower visitor species that interact with them are alien. Alien flower visitor species exhibited higher interaction degree, specialization and strength than their native counterparts, while alien plant species showed the opposite pattern. We found that invader complexes (in which alien species interact significantly more with each other than with native species) were established across the season, and interactions involving alien plant species were the main connectors of the temporal networks. Both alien plant and flower visitor species increased total interaction turnover through the flowering season by promoting interaction rewiring in the case of alien plants and by increasing species turnover in the case of alien flower visitors. Synthesis. This study provides one of the first empirical reports of alien species shaping the seasonal dynamics of plant–flower visitor networks. We demonstrate that the presence of alien species may simultaneously lead to a homogenization of plant species composition and increase the diversity of plant–flower visitor interactions through a flowering season. Additionally, we highlight the importance of considering the role of different trophic levels when analysing the impact of alien species in plant–flower visitor communities. Overall, our findings suggest that management strategies should pay particular attention to the timescale at which interactions with alien species dissolve or form, and to the consequences and drivers of such seasonal dynamics.