Traits linked with species invasiveness and community invasibility vary with time,stage and indicator of invasion in a long‐term grassland experiment

Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.     

A functional trait perspective on plant invasion

Background and Aims

Global environmental change will affect non-native plant invasions, with profound potential impacts on native plant populations, communities and ecosystems. In this context, we review plant functional traits, particularly those that drive invader abundance (invasiveness) and impacts, as well as the integration of these traits across multiple ecological scales, and as a basis for restoration and management.

Scope

We review the concepts and terminology surrounding functional traits and how functional traits influence processes at the individual level. We explore how phenotypic plasticity may lead to rapid evolution of novel traits facilitating invasiveness in changing environments and then ‘scale up’ to evaluate the relative importance of demographic traits and their links to invasion rates. We then suggest a functional trait framework for assessing per capita effects and, ultimately, impacts of invasive plants on plant communities and ecosystems. Lastly, we focus on the role of functional trait-based approaches in invasive species management and restoration in the context of rapid, global environmental change.

Conclusions

To understand how the abundance and impacts of invasive plants will respond to rapid environmental changes it is essential to link trait-based responses of invaders to changes in community and ecosystem properties. To do so requires a comprehensive effort that considers dynamic environmental controls and a targeted approach to understand key functional traits driving both invader abundance and impacts. If we are to predict future invasions, manage those at hand and use restoration technology to mitigate invasive species impacts, future research must focus on functional traits that promote invasiveness and invader impacts under changing conditions, and integrate major factors driving invasions from individual to ecosystem levels.     

Differences in seed properties and germination between native and introduced populations of Triadica sebifera

很多入侵植物的种子比原产地植物的种子萌发的更快,这可能有利于它们成功入侵。为了理解生物入侵过程,对入侵植物的入侵种群 和原产地种群的种子特性和萌发特征同时进行评估极为必要。本研究中,我们以乌桕(Triadica sebifera)为研究对象探究种子萌发和种子 特征的种内差异。 我们利用采于2017年的乌桕种子,来自中国的12个原产地种群和美国的12个引入地种群,测定种子的物理性状(体积、重量、种皮硬度和种皮厚度)和化学特性[种仁的粗脂肪、可溶性蛋白质、糖、赤霉素(GA)和脱落酸(ABA)],并在中国的温室中测定种子的发芽率和萌发时间。此外,我们在美国用采自2016年和2017年的种子开展了研究地点(中国和美国)和种子采集年份(2016年和2017年)对种子萌发影响的比较实验。 来自引入地的种子要比来自原产地的种子萌发更快。种子的物理和化学特性测定结果表明：与原产地种子相比,引入地种子更大、种仁内含有的赤霉素浓度以及GA:ABA比更高,但粗脂肪含量较低。种子重量、种皮硬度、种皮厚度、种仁脱落酸、可溶性蛋白质和糖含量在引入地和原产地之间没有显著差异。在中国和美国温室间的种子萌发率具有相关性,但种群的萌发率具有年际变化。我们的研究结果表明,体积较大和赤霉素含量较高的种子可能有助于快速萌发,这可能也有利于乌桕在引入地的入侵。     

Release from belowground enemies and shifts in root traits as interrelated drivers of alien plant invasion success: a hypothesis

Our understanding of the interrelated mechanisms driving plant invasions, such as the interplay between enemy release and resource‐acquisition traits, is biased by an aboveground perspective. To address this bias, I hypothesize that plant release from belowground enemies (especially fungal pathogens) will give invasive plant species a fitness advantage in the alien range, via shifts in root traits (e.g., increased specific root length and branching intensity) that increase resource uptake and competitive ability compared to native species in the alien range, and compared to plants of the invader in its native range. Such root‐trait changes could be ecological or evolutionary in nature. I explain how shifts in root traits could occur as a consequence of enemy release and contribute to invasion success of alien plants, and how they could be interrelated with other potential belowground drivers of invasion success (allelopathy, mutualist enhancement). Finally, I outline the approaches that could be taken to test whether belowground enemy release results in increased competitive ability and nutrient uptake by invasive alien plants, via changes in root traits in the alien range.     

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.     

Soil conditioning effects of Phragmites australis on native wetland plant seedling survival

Interactions between introduced plants and soils they colonize are central to invasive species success in many systems. Belowground biotic and abiotic changes can influence the success of introduced species as well as their native competitors. All plants alter soil properties after colonization but, in the case of many invasive plant species, it is unclear whether the strength and direction of these soil conditioning effects are due to plant traits, plant origin, or local population characteristics and site conditions in the invaded range. Phragmites australis in North America exists as a mix of populations of different evolutionary origin. Populations of endemic native Phragmites australis americanus are declining, while introduced European populations are important wetland invaders. We assessed soil conditioning effects of native and non‐native P. australis populations on early and late seedling survival of native and introduced wetland plants. We further used a soil biocide treatment to assess the role of soil fungi on seedling survival. Survival of seedlings in soils colonized by P. australis was either unaffected or negatively affected; no species showed improved survival in P. australis‐conditioned soils. Population of P. australis was a significant factor explaining the response of seedlings, but origin (native or non‐native) was not a significant factor. Synthesis: Our results highlight the importance of phylogenetic control when assessing impacts of invasive species to avoid conflating general plant traits with mechanisms of invasive success. Both native (noninvasive) and non‐native (invasive) P. australis populations reduced seedling survival of competing plant species. Because soil legacy effects of native and non‐native P. australis are similar, this study suggests that the close phylogenetic relationship between the two populations, and not the invasive status of introduced P. australis, is more relevant to their soil‐mediated impact on other plant species.     

Biotic resistance buffers the effects of nutrient enrichment on the success of a highly invasive aquatic plant

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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.     

Life-history correlates of plant invasiveness at regional and continental scales

We implemented cross‐species and independent‐contrasts multiple regression models to compare life‐history correlates of invasion success between regional and continental spatial scales among non‐native plants of eastern Australia. We focussed on three life‐history traits that represent major axes of variation in plant life history: specific leaf area (SLA), plant height and seed mass. After controlling for residence time and cross‐correlation with other life‐history traits, small seed mass was significantly and uniquely correlated with invasion success at continental and regional scales. High SLA was significantly and uniquely correlated with invasion success at the continental scale only. Plant height could not explain unique variation in invasion success at either spatial scale. Variation among spatial scales in the significance and strength of life‐history relationships with invasion success suggests that the search for predictive tools of invasion need not be fruitless, as long as predictive investigations are targeted at appropriate spatial scales.     

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.     

Soil pathogen communities associated with native and non‐native Phragmites australis populations in freshwater wetlands

Soil pathogens are believed to be major contributors to negative plant–soil feedbacks that regulate plant community dynamics and plant invasions. While the theoretical basis for pathogen regulation of plant communities is well established within the plant–soil feedback framework, direct experimental evidence for pathogen community responses to plants has been limited, often relying largely on indirect evidence based on above‐ground plant responses. As a result, specific soil pathogen responses accompanying above‐ground plant community dynamics are largely unknown. Here, we examine the oomycete pathogens in soils conditioned by established populations of native noninvasive and non‐native invasive haplotypes of Phragmites australis (European common reed). Our aim was to assess whether populations of invasive plants harbor unique communities of pathogens that differ from those associated with noninvasive populations and whether the distribution of taxa within these communities may help to explain invasive success. We compared the composition and abundance of pathogenic and saprobic oomycete species over a 2‐year period. Despite a diversity of oomycete taxa detected in soils from both native and non‐native populations, pathogen communities from both invaded and noninvaded soils were dominated by species of Pythium. Pathogen species that contributed the most to the differences observed between invaded and noninvaded soils were distributed between invaded and noninvaded soils. However, the specific taxa in invaded soils responsible for community differences were distinct from those in noninvaded soils that contributed to community differences. Our results indicate that, despite the phylogenetic relatedness of native and non‐native P. australis haplotypes, pathogen communities associated with the dominant non‐native haplotype are distinct from those of the rare native haplotype. Pathogen taxa that dominate either noninvaded or invaded soils suggest different potential mechanisms of invasion facilitation. These findings are consistent with the hypothesis that non‐native plant species that dominate landscapes may “cultivate” a different soil pathogen community to their rhizosphere than those of rarer native species.     

Species composition,functional and phylogenetic distances correlate with success of invasive Chromolaena odorata in an experimental test

Biotic resistance may influence invasion success; however, the relative roles of species richness, functional or phylogenetic distance in predicting invasion success are not fully understood. We used biomass fraction of Chromolaena odorata, an invasive species in tropical and subtropical areas, as a measure of ‘invasion success’ in a series of artificial communities varying in species richness. Communities were constructed using species from Mexico (native range) or China (non‐native range). We found strong evidence of biotic resistance: species richness and community biomass were negatively related with invasion success; invader biomass was greater in plant communities from China than from Mexico. Harvesting time had a greater effect on invasion success in plant communities from China than on those from Mexico. Functional and phylogenetic distances both correlated with invasion success and more functionally distant communities were more easily invaded. The effects of plant‐soil fungi and plant allelochemical interactions on invasion success were species‐specific.     

Preadapted for invasiveness: do species traits or their plastic response to shading differ between invasive and non‐invasive plant species in their native range?

Aim Species capable of vigorous growth under a wide range of environmental conditions should have a higher chance of becoming invasive after introduction into new regions. High performance across environments can be achieved either by constitutively expressed traits that allow for high resource uptake under different environmental conditions or by adaptive plasticity of traits. Here we test whether invasive and non‐invasive species differ in presumably adaptive plasticity. Location Europe (for native species); the rest of the world and North America in particular (for alien species). Methods We selected 14 congeneric pairs of European herbaceous species that have all been introduced elsewhere. One species of each pair is highly invasive elsewhere in the world, particularly so in North America, whereas the other species has not become invasive or has spread only to a limited degree. We grew native plant material of the 28 species under shaded and non‐shaded conditions in a common garden experiment, and measured biomass production and morphological traits that are frequently related to shade tolerance and avoidance. Results Invasive species had higher shoot–root ratios, tended to have longer leaf‐blades, and produced more biomass than congeneric non‐invasive species both under shaded and non‐shaded conditions. Plants responded to shading by increasing shoot–root ratios and specific leaf area. Surprisingly, these shade‐induced responses, which are widely considered to be adaptive, did not differ between invasive and non‐invasive species. Main conclusions We conclude that high biomass production across different light environments pre‐adapts species to become invasive, and that this is not mediated by plasticities of the morphological traits that we measured.     

Invasiveness of plants is predicted by size and fecundity in the native range

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Differences in life‐cycle stage components between native and introduced ranges of five woody Fabaceae species

Understanding differences in the components of life‐cycle stages of species between their native and introduced ranges can provide insights into the process of species transitioning from introduction to naturalization and invasion. We examined reproductive variables of the germination (seed predation, seed viability, time to germination), seed output (crown projection, seed production, seed weight) and dispersal (seed weight, dispersal investment) stages of five woody Fabaceae species, comparing native and introduced ranges. We predicted that each species would differ in reproductive variables of at least one life‐cycle stage between their native and introduced ranges, thus allowing us to determine the life‐cycle stage most associated with invasion success in the introduced range. Acacia melanoxylon and Paraserianthes lophantha had reduced seed predation in their introduced ranges while P. lophantha also had higher seed viability indicating that the germination life‐cycle stage is most strongly associated with their invasion success in the introduced range. Only Acacia longifolia varied between ranges for the seed output stage due to larger plant size, greater seed production and smaller seed size in its introduced range. Similar to A. longifolia, Acacia cyclops had smaller seed size in its introduced range but did not have any other variable differences between ranges suggesting that the dispersal stage is best associated with its invasion success in the introduced range. Surprisingly, Acacia saligna was the only species without a clear life‐cycle stage difference between ranges despite it being one of the more invasive acacia species in Australia. Although we found clear differences in reproductive variables associated with life‐cycle stages between native and introduced ranges of these five species, these differences were largely species‐specific. This suggests that a species invasion strategy into a novel environment is complex and varies among species depending on the environmental context, phenotypic plasticity and genotypic variation in particular traits.     

Soil pH influences patterns of plant community composition after restoration with native‐based seed mixes

Reclamation of highly disturbed lands typically includes establishing fast‐growing, non‐native plants to achieve rapid ground cover for erosion control. Establishing native plant communities could achieve ecosystem functions beyond soil erosion, such as providing wildlife habitat. Pipelines, or other disturbed corridors through a landscape, present unique challenges for establishing native plant communities given the heterogeneity of soil environments and invasive plant propagule pressure. We created two structural equation models to address multiple related hypotheses about the influence of soil pH on plant community composition (current diversity and vegetative cover of the original restoration seed mix and background flora, and invasive plant density during mix establishment and current density) of a highly disturbed landscape corridor restored with native species. To test our hypotheses we conducted a plant survey on a gas pipeline crossing two state forests in the north‐central Appalachians that had been seeded with a native‐based mixture 8 years prior. Low soil pH was a strong predictor of density of the invasive annual plant, Microstegium vimineum, and had resulted in lower species diversity and cover of the seeded mix. Overall, our data provide evidence that native‐based grass and forb mixtures can establish and persist on a wide range of soil environments and thrive in competition with invasive plants in moderately acidic to neutral soils. Advancing knowledge on restoration methods using native species is essential to improving restoration practice norms to incorporate multifunctional ecological goals.     

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.     

Differential responses to fertilization and competition among invasive,noninvasive alien,and native Bidens species

Comparative studies of invasive, noninvasive alien, and native congenic plant species can identify plant traits that drive invasiveness. In particular, functional traits associated with rapid growth rate and high fecundity likely facilitate invasive success. As such traits often exhibit high phenotypic plasticity, characterizing plastic responses to anthropogenic environmental changes such as eutrophication and disturbance is important for predicting the invasive success of alien plant species in the future. Here, we compared trait expression and phenotypic plasticity at the species level among invasive, noninvasive alien, and native Bidens species. Plants were grown under nutrient addition and competition treatments, and their functional, morphological, and seed traits were examined. Invasive B. frondosa exhibited higher phenotypic plasticity in most measured traits than did the alien noninvasive B. pilosa or native B. bipinnata. However, differential plastic responses to environmental treatments rarely altered the rank of trait values among the three Bidens species, except for the number of inflorescences. The achene size of B. frondosa was larger, but its pappus length was shorter than that of B. pilosa. Two species demonstrated opposite plastic responses of pappus length to fertilization. These results suggest that the plasticity of functional traits does not significantly contribute to the invasive success of B. frondosa. The dispersal efficiency of B. frondosa is expected to be lower than that of B. pilosa, suggesting that long‐distance dispersal is likely not a critical factor in determining invasive success.     

Interactions between alien plant species traits and habitat characteristics in agricultural landscapes in Finland

The survival and success of alien plant species is determined by species traits (i.e., invasiveness) and the characteristics of the habitats in the region of introduction (i.e., invasibility). However, little is known about species traits as related to habitat characteristics. We assessed the characteristics of successful invaders and the interaction of environmental factors and life-history traits for alien plant species. The vascular plants were recorded from 52 agricultural landscapes in Finland. We compared the traits of native and alien plant species with Fisher’s exact test and used a three table ordination analysis, RLQ analysis, to relate species traits to environmental conditions. Species were clustered according to their position on the RLQ axes, and the clusters were tested for phylogenetic independence. The successful alien plant species were associated with life form and preferences for moisture and nitrogen, but the trait composition varied according to the habitat type. Two RLQ axes explained 80.5% of the variation, and the species traits were significantly associated with environmental variables. The clustering showed that the occurrence of alien plant species in agricultural habitats was driven by invasion history, traits related to dispersal (dispersal type, seed mass) and habitat preferences, as well as environmental features, such as geographical location, temperature and the quality and disturbance regime of the habitats. All clusters were phylogenetically non-independent. Thus, the clusters of alien species comprised species of diverse taxonomic affinities, although, they shared the traits explaining their occurrence in particular habitats. This information is useful for understanding the link between species traits and the environmental conditions of the habitats, and complexity of the invasion process.     

Incomplete recovery of plant diversity in restored prairie wetlands on agricultural landscapes

Restoration efforts are being implemented globally to mitigate the degradation and loss of wetland habitat; however, the rate and success of wetland vegetation recovery post‐restoration is highly variable across wetland classes and geographies. Here, we measured the recovery of plant diversity along a chronosequence of restored temporary and seasonal prairie wetlands ranging from 0 to 23 years since restoration, including drained and natural wetlands embedded in agricultural and natural reserve landscapes in central Alberta, Canada. We assessed plant diversity using the following structural indicators: percent cover of hydrophytes, native and non‐native species, species richness, and community composition. Our findings indicate that plant diversity recovered to resemble reference wetlands in agricultural landscapes within 3–5 years of restoration; however, restored wetlands maintained significantly lower species richness and a distinct community composition compared to reference wetlands located within natural reserves. Early establishment of non‐native species during recovery, dispersal limitation, and depauperated native seed bank were probable barriers to complete recovery. Determining the success of vegetation recovery provides important knowledge that can be used to improve restoration strategies, especially considering projected future changes in land use and climate.