Invasive grass litter facilitates native shrubs through abiotic effects

Questions: Plant invasions are considered one of the top threats to the biodiversity of native taxa, but clearly documenting the causal links between invasions and the decline of native species remains a major challenge of invasion biology. Most studies have focused on impacts of invaders' living biomass, rather than on mechanisms mediated by litter. However, invasive plant litter, which is often of a very different type and quantity than a system's native plant litter, can have multiple important effects on ecosystem processes – such as nitrogen cycling and soil microclimate – that may influence native plants. Location: We studied effects of litter of invasive grass species that are widespread throughout western North America on native shrubs in southern California's semi‐arid habitat of coastal sage scrub. Methods: We combined a 3‐year field manipulation of non‐native litter with structural equation modeling to understand interacting effects on non‐native grasses, native shrubs, soil nitrogen (available and total), and soil moisture. Results: Litter addition facilitated non‐native grass growth, revealing a positive feedback likely to enhance invasion success. Contrary to a major paradigm of invasion biology – that competition with invasive plant species causes declines of native plants – we found that litter also facilitated growth of the native dominant shrub, a result supported by observational trends. Structural equation models indicated that enhanced soil moisture mediated the positive effects of litter on shrub growth. Conclusions: We demonstrate that invasive plants, via their litter, can facilitate dominant native plants by altering soil moisture. Our results highlight that understanding the impacts and mechanisms of plant invasions may be enhanced by considering the role of invasive plant litter on native plants and ecosystem properties.     

Native and non‐native grasses generate common types of plant–soil feedbacks by altering soil nutrients and microbial communities

Soil conditioning occurs when plants alter features of their soil environment. When these alterations affect subsequent plant growth, it is a plant soil feedback. Plant–soil feedbacks are an important and understudied aspect of aboveground–belowground linkages in plant ecology that influence plant coexistence, invasion and restoration. Here, we examine plant–soil feedback dynamics of seven co‐occurring native and non‐native grass species to address the questions of how plants modify their soil environment, do those modifications inhibit or favor their own species relative to other species, and do non‐natives exhibit different plant–soil feedback dynamics than natives. We used a two‐phase design, wherein a first generation of plants was grown to induce species‐specific changes in the soil and a second generation of plants was used as a bioassay to determine the effects of those changes. We also used path‐analysis to examine the potential chain of effects of the first generation on soil nutrients and soil microbial composition and on bioassay plant performance. Our findings show species‐specific (rather than consistent within groups of natives and non‐natives) soil conditioning effects on both soil nutrients and the soil microbial community by plants. Additionally, native species produced plant–soil feedback types that benefit other species more than themselves and non‐native invasive species tended to produce plant–soil feedback types that benefit themselves more than other species. These results, coupled with previous field observations, support hypotheses that plant–soil feedbacks may be a mechanism by which some non‐native species increase their invasive potential and plant–soil feedbacks may influence the vulnerability of a site to invasion.     

Neighbour tolerance,not suppression,provides competitive advantage to non‐native plants

High competitive ability has often been invoked as a key determinant of invasion success and ecological impacts of non‐native plants. Yet our understanding of the strategies that non‐natives use to gain competitive dominance remains limited. Particularly, it remains unknown whether the two non‐mutually exclusive competitive strategies, neighbour suppression and neighbour tolerance, are equally important for the competitive advantage of non‐native plants. Here, we analyse data from 192 peer‐reviewed studies on pairwise plant competition within a Bayesian multilevel meta‐analytic framework and show that non‐native plants outperform their native counterparts due to high tolerance of competition, as opposed to strong suppressive ability. Competitive tolerance ability of non‐native plants was driven by neighbour's origin and was expressed in response to a heterospecific native but not heterospecific non‐native neighbour. In contrast to natives, non‐native species were not more suppressed by hetero‐ vs. conspecific neighbours, which was partially due to higher intensity of intraspecific competition among non‐natives. Heterogeneity in the data was primarily associated with methodological differences among studies and not with phylogenetic relatedness among species. Altogether, our synthesis demonstrates that non‐native plants are competitively distinct from native plants and challenges the common notion that neighbour suppression is the primary strategy for plant invasion success.     

Phenotypes of Two Generations of Sporobolus airoides Seedlings Derived from Acroptilon repens‐invaded and Non‐invaded Grass Populations

Although the ecological impacts of invasive species are well known, the evolutionary impacts on recipient native grass communities are not. We suggest that remnant native plants may provide desirable seed sources for restoration and native plant production. Native populations exposed to the selective pressures associated with exotic invasion may retain traits that increase their ability to coexist with invasive species. Two generations of Sporobolus airoides Torr. (Alkali sacaton) plants derived from lineages collected from within long‐term invaded areas of Acroptilon repens (L.) DC (Russian knapweed) and from adjacent non‐invaded areas were propagated in a greenhouse to evaluate generational changes in phenotypic traits from the production environment. Given the difference in invasion history of the two populations, we hypothesized that invaded and non‐invaded subpopulations would differ phenotypically. Phenotypic measurements revealed that invaded subpopulations had greater vegetative growth, whereas non‐invaded subpopulations had increased sexual reproduction. Phenotypic expression changed from the first to the second generation, predominantly in the invaded subpopulation. Generational phenotypic shifts are disadvantageous for native seed production which requires a standard product to sell commercially. However, phenotypic variation may improve field seed survival. This research demonstrates the potential value of targeting post‐invasion remnant grass populations for restoration.     

Bumble bees selectively use native and exotic species to maintain nutritional intake across highly variable and invaded local floral resource pools

1. Changes to plant community composition after invasion are well documented but how these shifts directly affect higher trophic levels is still poorly understood. One potentially important factor is the change in nutritional availability after an invasion. Shifts in nutrient availability could affect the nutrient intake of organisms that live in invaded habitats, causing reduced fecundity and survival. 2. The effects of the interaction among nutrient availability, selection, and diet on nutrient intake of a native bumble bee were examined. No nutritional differences were found between exotic and native pollen or collected and non‐collected pollen in protein or amino acid content, suggesting that differences in nutrient intake from random are based on selection. 3. Nutrient intake was simulated when pollen was selected randomly across all available plant species and when selection was restricted to native plants only or exotic plants only using a permutation model and compared with observed collection. The results suggest that pollen collection is non‐random and that selecting only native or exotic plants cannot provide the protein or amino acid intake observed. 4. These results may help to explain why the responses of native bees to exotic plants are so variable. If the exotic plants in a community can supply the necessary nutrients, bees may readily incorporate them into their diets, but if not, exotic plants may be avoided.     

Exotic weevil invasion increases floral herbivore community density,function, and impact on a native plant

Consumer communities are being re‐arranged through unprecedented rates of human‐mediated invasions and extinctions. Such changes in consumer diversity potentially alter community function and impact on resource populations. Although insect herbivore invasions are increasingly common, the influence of such species additions on native resident herbivore guilds, along with their individual and combined effects on native plant resources, are rarely investigated. Here, we used site‐to‐site and plant‐to‐plant variation in herbivore composition to examine how the addition of an invasive exotic weevil, Rhinocyllus conicus, combines with a guild of native floral herbivores (tephritid flies, pyralid moths) to influence two key components of herbivore community function – aggregate herbivore densities and cumulative levels of seed destruction – on a native thistle, Cirsium canescens. Invasion of a site by R. conicus more than doubled aggregate herbivore density, resulting in increased levels of seed destruction and a halving of seed production by the native thistle. Further, herbivore function was significantly higher on individual plants attacked by R. conicus, compared to plants attacked only by native herbivores. Insect densities and levels of seed destruction on plants attacked by multiple herbivore taxa never exceeded those observed for plants attacked by R. conicus alone, suggesting that increases in herbivore community function with invasion resulted from the inclusion of a functionally dominant insect rather than any complementarity effects. Some evidence for interference between insects emerged, with a trend towards reduced moth and weevil densities in two and three taxon mixtures compared with plants attacked by each taxon alone. However, density compensation was limited so that, overall, the addition of a novel herbivore to the floral guild was associated with a significant increase in herbivore community function and impact on seed production. The results suggest that invasion of a functionally dominant herbivore into an unsaturated recipient community can augment function within a resource guild.     

Experimental plant invasion reduces arthropod abundance and richness across multiple trophic levels

Plant invasions are known to have negative impacts on native plant communities, yet their influence on higher trophic levels has not been well documented. Past studies investigating the effects of invasive plants on herbivores and carnivores have been largely observational in nature and thus lack the ability to tease apart whether differences are a cause or consequence of the invasion. In addition, understanding how plant traits and plant species compositions change in invaded habitats may increase our ability to predict when and where invasive plants will have effects that cascade to animals. To assess effects on arthropods, we experimentally introduced a non‐native plant (Microstegium vimineum, Japanese stiltgrass) in a community re‐assembly experiment. We also investigated possible mechanisms through which the invader could affect associated arthropods, including changes in native plant species richness, above‐ground plant biomass, light availability and vegetation height. In experimentally invaded plots, arthropod abundance was reduced by 39%, and species richness declined by 19%. Carnivores experienced greater reductions in abundance than herbivores (61% vs 31% reduction). Arthropod composition significantly diverged between experimentally invaded and control plots, and particular species belonging to the abundant families Aphididae (aphids), Formicidae (ants) and Phalacridae (shining flower beetles) contributed the most to compositional differences. Among the mechanisms we investigated, only the reduction in native plant species richness caused by invasion was strongly correlated with total arthropod abundance and richness. In sum, our results demonstrate negative impacts of M. vimineum invasion on higher trophic levels and suggest that these effects occur, in part, indirectly through invader‐mediated reductions in the richness of the native plant community. The particularly strong response of carnivores suggests that plant invasion could reduce top–down control of herbivorous species for native plants.     

Comparisons of Mycorrhizal Responsiveness with Field Soil and Commercial Inoculum for Six Native Montane Species and Bromus tectorum

Reestablishing native perennial plants and reducing invasive species are pivotal for many ecological restoration projects. The interactions among plant species, arbuscular mycorrhizal fungi, and soil P availability may be critical determinants of the success of native and non‐native plants in restoration and species invasions. Here we assessed mycorrhizal responsiveness for three late‐successional and three early‐successional plant species native to Rocky Mountain National Park and for the non‐native Downy brome, cheatgrass (Bromus tectorum L.) using field soil and commercial inoculum. Factorial greenhouse experiments were conducted to compare biomass of plant species with and without field soil and commercial inoculum treatments along a phosphorus (P) gradient, which ranged from ambient field levels to 12% of field levels, using dilutions of native soils. The two field soil inoculum treatments resulted in significant biomass differences for all species studied. Late‐successional species responded positively to field inoculum, whereas early‐successional species responded negatively. The two commercial inocula had low colonization rates (14 of 166 inoculated plants). The commercial inocula substrates had significant treatment effects on five of seven species included in the study in the apparent absence of mycorrhizal symbiosis. Soil P levels influenced mycorrhizal responsiveness in only one species, Smooth blue aster (Aster laevis L.). Our results show that, at least for the species studied here, locally collected field inoculum is the best choice for reestablishment of late‐successional native plant species.     

Predicting novel trophic interactions in a non‐native world

Humans are altering the global distributional ranges of plants, while their co‐evolved herbivores are frequently left behind. Native herbivores often colonise non‐native plants, potentially reducing invasion success or causing economic loss to introduced agricultural crops. We developed a predictive model to forecast novel interactions and verified it with a data set containing hundreds of observed novel plant–insect interactions. Using a food network of 900 native European butterfly and moth species and 1944 native plants, we built an herbivore host‐use model. By extrapolating host use from the native herbivore–plant food network, we accurately forecasted the observed novel use of 459 non‐native plant species by native herbivores. Patterns that governed herbivore host breadth on co‐evolved native plants were equally important in determining non‐native hosts. Our results make the forecasting of novel herbivore communities feasible in order to better understand the fate and impact of introduced plants.     

The role of plant–mycorrhizal mutualisms in deterring plant invasions: Insights from an individual‐based model

Understanding the factors that determine invasion success for non‐native plants is crucial for maintaining global biodiversity and ecosystem functioning. One hypothesized mechanism by which many exotic plants can become invasive is through the disruption of key plant–mycorrhizal mutualisms, yet few studies have investigated how these disruptions can lead to invader success. We present an individual‐based model to examine how mutualism strengths between a native plant (Impatiens capensis) and mycorrhizal fungus can influence invasion success for a widespread plant invader, Alliaria petiolata (garlic mustard). Two questions were investigated as follows: (a) How does the strength of the mutualism between the native I. capensis and a mycorrhizal fungus affect resistance (i.e., native plant maintaining >60% of final equilibrium plant density) to garlic mustard invasion? (b) Is there a non‐linear relationship between initial garlic mustard density and invasiveness (i.e., garlic mustard representing >60% of final equilibrium plant density)? Our findings indicate that either low (i.e., facultative) or high (i.e., obligate) mutualism strengths between the native plant and mycorrhizal fungus were more likely to lead to garlic mustard invasiveness than intermediate levels, which resulted in higher resistance to garlic mustard invasion. Intermediate mutualism strengths allowed I. capensis to take advantage of increased fitness when the fungus was present but remained competitive enough to sustain high numbers without the fungus. Though strong mutualisms had the highest fitness without the invader, they proved most susceptible to invasion because the loss of the mycorrhizal fungus resulted in a reproductive output too low to compete with garlic mustard. Weak mutualisms were more competitive than strong mutualisms but still led to garlic mustard invasion. Furthermore, we found that under intermediate mutualism strengths, the initial density of garlic mustard (as a proxy for different levels of plant invasion) did not influence its invasion success, as high initial densities of garlic mustard did not lead to it becoming dominant. Our results indicate that plants that form weak or strong mutualisms with mycorrhizal fungi are most vulnerable to invasion, whereas intermediate mutualisms provide the highest resistance to an allelopathic invader.     

Invasive plants negatively impact native,but not exotic,animals

Despite our growing understanding of the impacts of invasive plants on ecosystem structure and function, important gaps remain, including whether native and exotic species respond differently to plant invasion. This would elucidate basic ecological interactions and inform management. We performed a meta‐analytic review of the effects of invasive plants on native and exotic resident animals. We found that invasive plants reduced the abundance of native, but not exotic, animals. This varied by animal phyla, with invasive plants reducing the abundance of native annelids and chordates, but not mollusks or arthropods. We found dissimilar impacts among “wet” and “dry” ecosystems, but not among animal trophic levels. Additionally, the impact of invasive plants increased over time, but this did not vary with animal nativity. Our review found that no studies considered resident nativity differences, and most did not identify animals to species. We call for more rigorous studies of invaded community impacts across taxa, and most importantly, explicit consideration of resident biogeographic origin. We provide an important first insight into how native and exotic species respond differently to invasion, the consequences of which may facilitate cascading trophic disruptions further exacerbating global change consequences to ecosystem structure and function.     

丛枝菌根真菌在外来植物入侵演替中的作用与机制

外来植物入侵不仅是环境、经济和社会问题,也是一个生理学和生态学问题,尤其是入侵植物与本地植物、入侵植物和本地土壤生物之间的相互作用决定外来植物入侵程度。丛枝菌根真菌（AMF）作为土壤中一类极为重要的功能生物,在外来植物入侵演替过程中发挥多种不同作用。文章系统总结了AMF对入侵植物个体和群体的影响,入侵植物与本地植物竞争中AMF发挥的促进和抑制作用;探讨了AMF与入侵植物的相互作用关系,以及环境因子对AMF一入侵植物关系的影响：对AMF在外来植物入侵演替中的作用机制进行了讨论。旨在为探索控制生物入侵的新途径、为我国开展外来植物入侵研究与防控实践提供新思路。     

Local and landscape correlates of non‐native species invasion in restored wetlands

Vulnerability of natural communities to invasion by non‐native plants has been linked to factors such as recent disturbance and high resource availability, suggesting that recently restored habitats may be especially invasible. Because non‐native plants can interfere with restoration goals, monitoring programs should anticipate which sites are most susceptible to invasion and which species are likely to become problematic at a site. Restored sites of larger area and those with high rates of propagule input should have higher species richness of both natives and non‐natives, leading to a positive correlation between the two. However, in restored wetlands, urbanization, riparian landscape settings, and nitrogen enrichment likely favor non‐native relative to native species. We sampled 28 restored wetlands in Illinois, USA, modeled the responses of native richness, non‐native richness and non‐native cover to local and landscape predictors with linear regression, and modeled the presence/absence of 21 non‐native species with logistic regressions. Unexpectedly, native and non‐native richness were uncorrelated, suggesting different responses to environmental factors. Native richness declined with increasing available soil nitrogen and urbanization in the surrounding landscape. Non‐native richness, the richness of non‐natives relative to natives, and the likelihood of invasion by several individual invasive species decreased with increasing distance from the city of Chicago, likely in response to decreasing non‐native propagule pressure. Total cover of non‐natives, however, as well as cover by non‐native Phalaris arundinacea, increased with nitrogen availability. Our results indicate that although non‐native richness was better predicted by factors related to propagule pressure, non‐native species dominance was more closely related to local abiotic factors. Non‐native richness in restoration sites may be beyond the control of restoration practitioners, and furthermore, may be of limited relevance for conservation goals. In contrast, limiting the relative dominance of non‐natives should be a restoration priority and may be achievable through management of nutrient availability.     

When do herbivores affect plant invasion? Evidence for the natural enemies and biotic resistance hypotheses

Two venerable hypotheses, widely cited as explanations for either the success or failure of introduced species in recipient communities, are the natural enemies hypothesis and the biotic resistance hypothesis. The natural enemies hypothesis posits that introduced organisms spread rapidly because they are liberated from their co‐evolved predators, pathogens and herbivores. The biotic resistance hypothesis asserts that introduced species often fail to invade communities because strong biotic interactions with native species hinder their establishment and spread. We reviewed the evidence for both of these hypotheses as they relate to the importance of non‐domesticated herbivores in affecting the success or failure of plant invasion.
To evaluate the natural enemies hypothesis, one must determine how commonly native herbivores have population‐level impacts on native plants. If native herbivores seldom limit native plant abundance, then there is little reason to think that introduced plants benefit from escape from these enemies. Studies of native herbivore‐native plant interactions reveal that plant life‐history greatly mediates the strength with which specialist herbivores suppress plant abundance. Relatively short‐lived plants that rely on current seed production for regeneration are most vulnerable to herbivory that reduces seed production. As such, these plants may gain the greatest advantage from escaping their specialist enemies in recipient communities. In contrast, native plants that are long lived or that possess long‐lived seedbanks may not be kept “in check” by native herbivores. For these species, escape from native enemies may have little to do with their success as exotics; they are abundant both where they are native and introduced.
Evidence for native herbivores providing biotic resistance to invasion by exotics is conflicting. Our review reveals that: 1) introduced plants can attract a diverse assemblage of native herbivores and that 2) native herbivores can reduce introduced plant growth, seed set and survival. However, the generality of these impacts is unclear, and evidence that herbivory actually limits or reduces introduced plant spread is scarce. The degree to which native herbivores provide biotic resistance to either exotic plant establishment or spread may be greatly determined by their functional and numerical responses to exotic plants, which we know little about. Generalist herbivores, through their direct effects on seed dispersal and their indirect effects in altering the outcome of native–non‐native plant competitive interactions, may have more of a facilitative than negative effect on exotic plant abundance.     

Impacts and interactions of multiple human perturbations in a California salt marsh

Multiple disturbances to ecosystems can influence community structure by modifying resistance to and recovery from invasion by non-native species. Predicting how invasibility responds to multiple anthropogenic impacts is particularly challenging due to the variety of potential stressors and complex responses. Using manipulative field experiments, we examined the relative impact of perturbations that primarily change abiotic or biotic factors to promote invasion in coastal salt marsh plant communities. Specifically we test the hypotheses that nitrogen enrichment and human trampling facilitate invasion of upland weeds into salt marsh, and that the ability of salt marsh communities to resist and/or recover from invasion is modified by hydrological conditions. Nitrogen enrichment affected invasion of non-native upland plants at only one of six sites, and increased aboveground native marsh biomass at only two sites. Percent cover of native marsh plants declined with trampling at all sites, but recovered earlier at tidally flushed sites than at tidally restricted sites. Synergistic interactions between trampling and restricting tidal flow resulted in significantly higher cover of non-native upland plants in trampled plots at tidally restricted sites. Percent cover of non-native plants recovered to pre-trampling levels in fully tidal sites, but remained higher in tidally restricted sites after 22 months. Thus, perturbations that reduce biotic resistance interact with perturbations that alter abiotic conditions to promote invasion. This suggests that to effectively conserve or restore native biodiversity in altered systems, one must consider impacts of multiple human disturbances, and the interactions between them. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

西双版纳外来入侵植物及其共存种叶片氮、磷化学计量特征

诊断外来植物和本地种的资源利用方式是入侵生态学研究的热点问题。叶片氮(N)、磷(P)含量和化学计量特征可以反映受侵地区植物的N、P吸收能力和限制状况, 为把握外来植物的入侵能力、为本地植物共存或消失的机制提供基础科学依据。该研究以我国西南地区典型外来入侵植物(飞机草(Chromolaena odorata)、紫茎泽兰(又叫破坏草, Ageratina adenophora))及其共存的本地植物为对象, 探讨不同入侵条件下入侵种及其共存的本地植物的N、P利用策略。在云南西双版纳孔明山研究区, 调查了无入侵条件下和飞机草与紫茎泽兰的不同入侵程度(按入侵种生物量比例划分)下的物种数量和生物量, 分析了主要植物的叶片N、P含量和N:P值。结果显示: 尽管群落地上生物量随飞机草和紫茎泽兰入侵程度增加而增加, 但本地植物种数随飞机草和紫茎泽兰入侵程度增加而显著减少。飞机草与紫茎泽兰的叶片N、P平均含量无显著差异, 但均显著高于无入侵条件下的本地植物以及与其共存的本地植物。两种入侵植物的N、P含量均随它们所占样方总生物量的比例增大而升高, 本地植物N含量也有相似的变化趋势。当对比入侵和无入侵两类样方的同种植物N、P和N:P变化时发现多数本地种叶片P含量呈降低趋势, N含量和N:P呈升高趋势。根据叶片N、P绝对含量和N:P值及其随入侵的变化规律, 推测入侵可能提高了植物的N可利用性, 但本地植物仍然受N限制; 入侵植物N:P值总体小于10与其具有相对于N吸收的较高的P吸收能力有关。该研究揭示了西南典型外来入侵植物具有较强的N、P吸收富集能力。     

Foliar nitrogen and phosphorus stoichiometry of alien invasive plants and co-occurring natives in Xishuangbanna

Aims How alien invasive plants and co-occurring native plants utilize nutrients is one of major issues in invasion ecology. Foliar nitrogen (N) and phosphorus (P) contents and stoichiometry can elucidate the uptake ability and limitation status of nutrients in plants, which provides basic knowledge for understanding the invading ability and co-occurrence or disappearance of plants.
Methods Based on typical alien invasive plants (Chromolaena odorata, Ageratina adenophora) and native plants in southwestern China, this study focused on strategies of N and P utilization among invasive plants and native plants under different invasion conditions. The species compositions, aboveground biomass, leaf N and P contents and leaf N:P were investigated for plants in plots with no invasion and with different invasion extents (estimated by the plot-based percentage of invaders’ biomass in total community) at Mt. Kongming in Xishuangbanna region, Yunnan Province, China.
Important findings The species number decreased significantly with the invasion extent of both C. odorata and A. adenophora, although the aboveground biomass was greatly enhanced. Leaf N and P contents did not differ between the two studied invaders, but they showed significantly higher N and P levels than both co-occurring and only native species (p < 0.05). Besides, leaf N and P contents of invaders increased with the invasion extent, and leaf N of native plants also showed an increasing trend with the invasion extent. When the influence of invasion was checked for the same species, leaf P contents decreased, whereas leaf N and N:P increased for most native plants under invasion. Based on the absolute foliar N and P contents, N:P values, we inferred that native plants were still limited by N, although N availability might be enhanced by invasion. Both invasive plants had leaf N:P values lower than 10, suggesting a higher P uptake relative to N uptake. All above results highlighted a higher N and P uptake of typical alien invasive plants in southwestern China.     

Impacts of alien plant invasion on native plant communities are mediated by functional identity of resident species,not resource availability

Alien plant invasion and nutrient enrichment as a result of anthropogenic landscape modification seriously threaten native plant community diversity. It is poorly understood, however, whether these two disturbances interact with the functional identity of recipient native plants to drive community change. We performed a mesocosm experiment to examine whether the interactive effects of invasion by a stoloniferous turf‐grass Stenotaphrum secundatum and nutrient enrichment vary across different plant growth forms of an endangered coastal plant community. Communities contained 18 species (drawn without replacement from a pool of 31 species) with either runner, tufted or woody growth forms. Species were well‐established and reproductively mature prior to S. secundatum introduction. Species growth (% cover), reproductive output, soil temperature and light availability were monitored for two growing seasons. Invasion and nutrient enrichment (two levels: ‘natural control’ and ‘enriched’) had no effect on species richness, community composition, reproductive output, soil temperature or light penetration. There was no interactive effect of nutrients and invasion on community productivity (i.e. final biomass), such that invasion caused a reduction in community biomass at both natural and enriched nutrient levels. This was driven only by reduced biomass of functionally‐similar native runner species, which share similar root morphologies and nutrient‐acquisition strategies with S. secundatum. Our study indicates that impacts of invasion are dependent upon the functional identity of species within recipient communities, not the availability of resources. This shows that management cannot buffer invader effects by manipulating resource availability. Revegetation strategies should target functionally‐similar natives for replacement following invader control.     

Multiple anthropogenic changes cause biodiversity loss through plant invasion

Non‐native invasive plants are a widely acknowledged threat to global biodiversity. However, our understanding of the mechanisms underlying plant invasion, and the relative importance of multiple rather than single drivers, remains poor. Here, we provide a case study using time‐series data to reconstruct patterns of change, and field experiments to test for causality. We show how, over a 50‐year period, a series of unrelated human‐induced changes created highly favorable conditions for the non‐native tree mallow (Lavatera arborea) to turn invasive, causing loss of native vegetation and seabird breeding habitat. The combination of three drivers: human‐introduced disease, climate warming and a fisheries‐mediated increase in seabird populations, removed major constraints on plant population growth, (i.e. grazer control, climatic control, germination opportunity and nutrient limitation). Collectively, these changes created optimal conditions for the rapid expansion of tree mallow. The resulting dramatic impact on both the native vegetation and breeding seabirds, notably puffins (Fratercula arctica), exemplifies how non‐native invasive plant species can transform terrestrial ecosystems. While climate change is regarded as a key factor behind plant invasion, we highlight that multiple rather than single factors may be critical to biodiversity loss.     

Phenology in a warming world: differences between native and non‐native plant species

Phenology is a harbinger of climate change, with many species advancing flowering in response to rising temperatures. However, there is tremendous variation among species in phenological response to warming, and any phenological differences between native and non‐native species may influence invasion outcomes under global warming. We simulated global warming in the field and found that non‐native species flowered earlier and were more phenologically plastic to temperature than natives, which did not accelerate flowering in response to warming. Non‐native species' flowering also became more synchronous with other community members under warming. Earlier flowering was associated with greater geographic spread of non‐native species, implicating phenology as a potential trait associated with the successful establishment of non‐native species across large geographic regions. Such phenological differences in both timing and plasticity between native and non‐natives are hypothesised to promote invasion success and population persistence, potentially benefiting non‐native over native species under climate change.