Comparison of invertebrate herbivores on native and non‐native Senecio species: Implications for the enemy release hypothesis

The enemy release hypothesis posits that non‐native plant species may gain a competitive advantage over their native counterparts because they are liberated from co‐evolved natural enemies from their native area. The phylogenetic relationship between a non‐native plant and the native community may be important for understanding the success of some non‐native plants, because host switching by insect herbivores is more likely to occur between closely related species. We tested the enemy release hypothesis by comparing leaf damage and herbivorous insect assemblages on the invasive species Senecio madagascariensis Poir. to that on nine congeneric species, of which five are native to the study area, and four are non‐native but considered non‐invasive. Non‐native species had less leaf damage than natives overall, but we found no significant differences in the abundance, richness and Shannon diversity of herbivores between native and non‐native Senecio L. species. The herbivore assemblage and percentage abundance of herbivore guilds differed among all Senecio species, but patterns were not related to whether the species was native or not. Species‐level differences indicate that S. madagascariensis may have a greater proportion of generalist insect damage (represented by phytophagous leaf chewers) than the other Senecio species. Within a plant genus, escape from natural enemies may not be a sufficient explanation for why some non‐native species become more invasive than others.     

Relationships among leaf damage, natural enemy release, and abundance in exotic and native prairie plants

The Enemy Release hypothesis holds that exotic plants may have an advantage over native plants because their specialized natural enemies are absent. We tested this hypothesis by measuring leaf damage and plant abundance for naturally-occurring plants in prairies, and by removing natural enemies in an enemy exclusion experiment. We classified plants as invasive exotic, noninvasive exotic, or native, to determine if their degree of invasiveness influenced their relationships with natural enemies. Our field surveys showed that invasive exotic plants generally had significantly lower levels of foliar damage than native species while there was no consistent pattern for noninvasive exotics compared to natives. The relationship between damage and abundance was different for exotic and native plants: foliar damage decreased with increasing abundance for exotic plants while the trend was positive for native plants. While these results from the field surveys supported the Enemy Release Hypothesis, the enemy exclusion experiment did not. There was no relationship between a species?? status as exotic or native and its degree of release from herbivory. Pastinaca sativa, the invasive exotic in this experiment, experienced gains in leaf area and vegetative biomass when treated with pesticides, indicating substantial herbivore pressure in the introduced range. These results show that foliar damage may not accurately predict the amount of herbivore pressure that plants actually experience, and that the Enemy Release hypothesis is not sufficient to explain the invasiveness of P. sativa in prairies.     

Invasive plants do not display greater phenotypic plasticity than their native or non‐invasive counterparts: a meta‐analysis

Phenotypic plasticity is commonly considered as a trait associated with invasiveness in alien plants because it may enhance the ability of plants to occupy a wide range of environments. Although the evidence of greater phenotypic plasticity in invasive plants is considerable, it is not yet conclusive. We used a meta‐analysis approach to evaluate whether invasive plant species show greater phenotypic plasticity than their native or non‐invasive counterparts. The outcome of such interspecific comparisons may be biased when phylogenetic relatedness is not taken into account. Consequently, species pairs belonged to the same genus, tribe or family. The meta‐analysis included 93 records from 35 studies reporting plastic responses to light, nutrients, water, CO₂, herbivory and support availability. Contrary to what is often assumed, overall, phenotypic plasticity was similar between invasive plants and native or non‐invasive closely related species. The same result was found when separate analyses were conducted for trait plasticity to nutrients, light and water availability. Thus, invasive plant species and their native or non‐invasive counterparts are equally capable of displaying functional responses to environmental heterogeneity. The colonization of a wide range of environments by invasive plants could be due to their capacity to undergo adaptive ecotypic differentiation rather than to their ability to display plastic responses. Alternatively, phenotypic plasticity might play a role in plant invasion, but only during the initial phases, when tolerance of the novel environment is essential for plant survival. Afterwards, once alien plants are identified as invaders, the magnitude of phenotypic plasticity might be reduced after selection of the optimum phenotypes in each habitat. The identification of plant traits that consistently predict invasiveness might be a futile task because different traits favor invasiveness in different environments. Approaches at the local scale, focusing on the ecology of specific invasive plants, could be more fruitful than global macro‐analyses.     

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.     

Post‐fire vegetation dynamics in nutrient‐enriched and non‐enriched sclerophyll woodland

Abstract Exotic plant invasions are a significant problem in urban bushland in Sydney, Australia. In low‐nutrient Hawkesbury Sandstone communities, invasive plants are often associated with urban run‐off and subsequent increases in soil nutrients, particularly phosphorus. Fire is an important aspect of community dynamics in Sydney vegetation, and is sometimes used in bush regeneration projects as a tool for weed control. This study addressed the question: ‘Are there differences in post‐fire resprouting and germination of native and exotic species in nutrient‐enriched communities, compared with communities not disturbed by nutrient enrichment?’ We found that in non‐enriched areas, few exotic species emerged, and those that did were unable to achieve the rapid growth that was seen in exotic plants in the nutrient‐enriched areas. Therefore, fire did not promote the invasion of exotic plants into areas that were not nutrient‐enriched. In nutrient‐enriched areas after fire, the diversity of native species was lower than in the non‐enriched areas. Some native species were able to survive and compete with the exotic species in terms of abundance, per cent cover and plant height. However, these successful species were a different suite of natives to those commonly found in the non‐enriched areas. We suggest that although fire can be a useful tool for short‐term removal of exotic plant biomass from nutrient‐enriched areas, it does not promote establishment of native species that were not already present.     

Plant invasion is associated with higher plant–soil nutrient concentrations in nutrient‐poor environments

Plant invasion is an emerging driver of global change worldwide. We aimed to disentangle its impacts on plant–soil nutrient concentrations. We conducted a meta‐analysis of 215 peer‐reviewed articles and 1233 observations. Invasive plant species had globally higher N and P concentrations in photosynthetic tissues but not in foliar litter, in comparison with their native competitors. Invasive plants were also associated with higher soil C and N stocks and N, P, and K availabilities. The differences in N and P concentrations in photosynthetic tissues and in soil total C and N, soil N, P, and K availabilities between invasive and native species decreased when the environment was richer in nutrient resources. The results thus suggested higher nutrient resorption efficiencies in invasive than in native species in nutrient‐poor environments. There were differences in soil total N concentrations but not in total P concentrations, indicating that the differences associated to invasive plants were related with biological processes, not with geochemical processes. The results suggest that invasiveness is not only a driver of changes in ecosystem species composition but that it is also associated with significant changes in plant–soil elemental composition and stoichiometry.     

Testing the enemy release hypothesis: a review and meta-analysis

One of the most cited hypotheses explaining the inordinate success of a small proportion of introduced plants that become pests is the ‘natural enemies hypothesis’. This states that invasive introduced plants spread rapidly because they are liberated from their co-evolved natural enemies. This hypothesis had not been properly tested until recently. Previous reviews on this topic have been narrative and vote counting in nature. In this review, we carried out quantitative synthesis and meta-analysis using existing literature on plants and their herbivores to test the different components of the enemy release hypothesis. We found supporting evidence in that (1) insect herbivore fauna richness is significantly greater in the native than introduced ranges, and the reduction is skewed disproportionally towards specialists and insects feeding on reproductive parts; and (2) herbivore damage levels are greater on native plants than on introduced invasive congeners. However, herbivore damage levels are only marginally greater for plants in native than in introduced ranges, probably due to the small numbers of this type of study. Studies quantifying herbivore impacts on plant population dynamics are too scarce to make conclusions for either comparison of plants in native vs introduced ranges or of co-occurring native and introduced congeners. For future research, we advocate that more than two-way comparisons between plants in native and introduced ranges, or native and introduced congeners are needed. In addition, the use of herbivore exclusions to quantify the impacts of herbivory on complete sets of population vital rates of native vs introduced species are highly desirable. Furthermore, three-way comparisons among congeners of native plants, introduced invasive, and introduced non-invasive plants can also shed light on the importance of enemy release. Finally, simultaneously testing the enemy release hypothesis and other competing hypotheses will provide significant insights into the mechanisms governing the undesirable success of invasive species.     

Reduced tolerance to simulated herbivory on clonal organs in alien genotypes: a multi-species experiment with native and introduced origins

Clonal growth may increase the likelihood for alien plants becoming invasive, as it is an efficient foraging and spatial exploration strategy. Here, we investigated the effect of artificial herbivory on organs of clonal growth and its potential to drive post-introduction evolutionary change. Based on the assumption that tolerance traits are costly and that clonal alien species may benefit from investing freed resources into growth, fecundity or nutrient acquisition, we tested the hypothesis of lower tolerance to herbivory on organs of clonal growth in alien plants. In a common-garden experiment we studied divergence in plants from native German and alien New Zealand populations of six species with different clonal growth forms. A nutrient treatment testing the plant’s acquisition abilities, was combined with artificial herbivory on clonal organs. We investigated origin-dependent differences in sexual reproduction, plant growth and the production of clonal organs. For aboveground and clonal organ biomass, alien plants showed lower tolerance to artificial herbivory on clonal organs than native plants. In the combined herbivory and nutrient treatment, alien plants of four species grew fewer clonal organs when compared to the nutrient treatment alone. Alien plants of the other two species produced more clonal organs, regardless of treatment. All species revealed significant differences in flower production between origins, with five of them producing more flowers on alien than on native plants. The results support the hypothesis that a release of herbivory on clonal organs has lead to subtle evolutionary changes in tolerance of alien plants and to a species-dependent increase in plant vigour, clonal growth and/or sexual reproduction that may enhance their invasive success.     

Effects of invasive alien kahili ginger (Hedychium gardnerianum) on native plant species regeneration in a Hawaiian rainforest

Questions: Does the invasive alien Hedychium gardnerianum (1) replace native understory species, (2) suppress natural regeneration of native plant species, (3) increase the invasiveness of other non‐native plants and (4) are native forests are able to recover after removal of H. gardnerianum. Location: A mature rainforest in Hawai'i Volcanoes National Park on the island of Hawai'i (about 1200 m a.s.l.; precipitation approximately 2770 mm yr^?1). Study sites included natural plots without effects of alien plants, ginger plots with a H. gardnerianum‐dominated herb layer and cleared plots treated with herbicide to remove alien plants. Methods: Counting mature trees, saplings and seedlings of native and alien plant species. Using non‐parametric H‐tests to compare impact of H. gardnerianum on the structure of different sites. Results: Results confirmed the hypothesis that H. gardnerianum has negative effects on natural forest dynamics. Lower numbers of native tree seedlings and saplings were found on ginger‐dominated plots. Furthermore, H. gardnerianum did not show negative effects on the invasive alien tree species Psidium cattleianum. Conclusions: This study reveals that where dominance of H. gardnerianum persists, regeneration of the forest by native species will be inhibited. Furthermore, these areas might experience invasion by P. cattleianum, resulting in displacement of native canopy species in the future, leading to a change in forest structure and loss of other species dependent on natural rainforest, such as endemic birds. However, if H. gardnerianum is removed the native Hawaiian forest is likely to regenerate and regain its natural structure.     

Invasive plants differentially affect soil biota through litter and rhizosphere pathways: a meta‐analysis

Invasive plants affect soil biota through litter and rhizosphere inputs, but the direction and magnitude of these effects are variable. We conducted a meta‐analysis to examine the different effects of litter and rhizosphere of invasive plants on soil communities and nutrient cycling. Our results showed that invasive plants increased bacterial biomass by 16%, detritivore abundance by 119% and microbivore abundance by 89% through litter pathway. In the rhizosphere, invasive plants reduced bacterial biomass by 12%, herbivore abundance by 55% and predator abundance by 52%, but increased AM fungal biomass by 36%. Moreover, CO₂ efflux, N mineralisation rate and enzyme activities were all higher in invasive than native rhizosphere soils. These findings indicate that invasive plants may support more decomposers that in turn stimulate nutrient release via litter effect, and enhance nutrient uptake by reducing root grazing but forming more symbioses in the rhizosphere. Thus, we hypothesise that litter‐ and root‐based loops are probably linked to generate positive feedback of invaders on soil systems through stimulating nutrient cycling, consequently facilitating plant invasion. Our findings from limited cases with diverse contexts suggest that more studies are needed to differentiate litter and rhizosphere effects within single systems to better understand invasive plant‐soil interactions.     

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.     

Diversity and abundance patterns of phytophagous insect communities on alien and native host plants in the Brassicaceae

The herbivore load (abundance and species richness of herbivores) on alien plants is supposed to be one of the keys to understand the invasiveness of species. We investigate the phytophagous insect communities on cabbage plants (Brassicaceae) in Europe. We compare the communities of endophagous and ectophagous insects as well as of Coleoptera and Lepidoptera on native and alien cabbage plant species. Contrary to many other reports, we found no differences in the herbivore load between native and alien hosts. The majority of insect species attacked alien as well as native hosts. Across insect species, there was no difference in the patterns of host range on native and on alien hosts. Likewise the similarity of insect communities across pairs of host species was not different between natives and aliens. We conclude that the general similarity in the community patterns between native and alien cabbage plant species are due to the chemical characteristics of this plant family. All cabbage plants share glucosinolates. This may facilitate host switches from natives to aliens. Hence the presence of native congeners may influence invasiveness of alien plants.     

Relatedness with plant species in native community influences ecological consequences of range expansions

Global warming is enabling many plant species to expand their range to higher latitudes and altitudes, where they may suffer less from natural aboveground and belowground enemies. Reduced control by natural enemies can enable climate warming‐induced range expanders to gain an advantage in competition with natives and become disproportionally abundant in their new range. However, so far studies have only examined individual growth of range expanders, which have common congeneric plant species in their new range. Thus it is not known how general is this reduced effect of above‐ and belowground enemies and how it operates in communities, where multiple plant species also interact with each other. Here we show that range‐expanding plant species with and without congenerics in the invaded habitats differ in their ecological interactions in the new range. In a community‐level experiment, range‐expanding plant species, both with and without congenerics, suppressed the growth of a herbivore. However, only range expanders without congenerics reduced biomass production of the native plant species. In the present study, range expanders without congenerics allocated more biomass aboveground compared to native plant species, which can explain their competitive advantage. Competitive interaction and also biomass allocation of native plants and their congeneric range expanders were similar. Our results highlight that information about species phylogenetic relatedness with native flora can be crucial for improving predictions about the consequences of climate warming‐induced range expansions.     

Virulence of oomycete pathogens from Phragmites australis‐invaded and noninvaded soils to seedlings of wetland plant species

Soil pathogens affect plant community structure and function through negative plant–soil feedbacks that may contribute to the invasiveness of non‐native plant species. Our understanding of these pathogen‐induced soil feedbacks has relied largely on observations of the collective impact of the soil biota on plant populations, with few observations of accompanying changes in populations of specific soil pathogens and their impacts on invasive and noninvasive species. As a result, the roles of specific soil pathogens in plant invasions remain unknown. In this study, we examine the diversity and virulence of soil oomycete pathogens in freshwater wetland soils invaded by non‐native Phragmites australis (European common reed) to better understand the potential for soil pathogen communities to impact a range of native and non‐native species and influence invasiveness. We isolated oomycetes from four sites over a 2‐year period, collecting nearly 500 isolates belonging to 36 different species. These sites were dominated by species of Pythium, many of which decreased seedling survival of a range of native and invasive plants. Despite any clear host specialization, many of the Pythium species were differentially virulent to the native and non‐native plant species tested. Isolates from invaded and noninvaded soils were equally virulent to given individual plant species, and no apparent differences in susceptibility were observed between the collective groups of native and non‐native plant species.     

Linking biotic homogenization to habitat type, invasiveness and growth form of naturalized alien plants in North America

Aim Biotic homogenization is a growing phenomenon and has recently attracted much attention. Here, we analyse a large dataset of native and alien plants in North America to examine whether biotic homogenization is related to several ecological and biological attributes. Location North America (north of Mexico). Methods We assembled species lists of native and alien vascular plants for each of the 64 state‐ and province‐level geographical units in North America. Each alien species was characterized with respect to habitat (wetland versus upland), invasiveness (invasive versus non‐invasive), life cycle (annual/biennial versus perennial) and habit (herbaceous versus woody). We calculated a Jaccard similarity index separately for native, for alien, and for native and alien species. We used the average of Jaccard dissimilarity index (1 ? Jaccard index) of all paired localities as a measure of the mean beta diversity of alien species for each set of localities examined in an analysis. We used a homogenization index to quantify the effect of homogenization or differentiation. Results We found that (1) wetland, invasive, annual/biennial and herbaceous alien plants markedly homogenized the state‐level floras whereas non‐invasive and woody alien plants tended to differentiate the floras; (2) beta diversity was significantly lower for wetland, invasive, annual/biennial and herbaceous alien plants than their counterparts (i.e. upland, non‐invasive, perennial and woody alien plants, respectively); and (3) upland and perennial alien plants each played an equal role in homogenizing and differentiating the state‐level floras. Main conclusions Our study shows that biotic homogenization is clearly related to habitat type (e.g. wetland versus uplands), species invasiveness and life‐history traits such as life cycle (e.g. annual/biennial and herbaceous versus woody species) at the spatial scale examined. These observations help to understand the process of biotic homogenization resulting from alien vascular plants in North America.     

Not all non‐natives are equally unequal: reductions in herbivore β‐diversity depend on phylogenetic similarity to native plant community

Effects of host plant α‐ and β‐diversity often confound studies of herbivore β‐diversity, hindering our ability to predict the full impact of non‐native plants on herbivores. Here, while controlling host plant diversity, we examined variation in herbivore communities between native and non‐native plants, focusing on how plant relatedness and spatial scale alter the result. We found lower absolute magnitudes of β‐diversity among tree species and among sites on non‐natives in all comparisons. However, lower relative β‐diversity only occurred for immature herbivores on phylogenetically distinct non‐natives vs. natives. Locally in that comparison, non‐native gardens had lower host specificity; while among sites, the herbivores supported were a redundant subset of species on natives. Therefore, when phylogenetically distinct non‐natives replace native plants, the community of immature herbivores is likely to be homogenised across landscapes. Differences in communities on closely related non‐natives were subtler, but displayed community shifts and increased generalisation on non‐natives within certain feeding guilds.     

Do alien plants escape from natural enemies of congeneric residents? Yes but not from all

As predicted by the enemy release hypothesis, plants are supposedly less attacked by herbivores in their introduced range than in their native range. However, the nature of the natural enemies, in particular their degree of specificity may also affect the level of enemy escape. It is therefore expected that ectophagous invertebrate species, being generally considered as more generalists than endophagous species, are more prompt to colonise alien plants. In Swiss, Siberian and Russian Far East arboreta, we tested whether alien woody plants are less attacked by native herbivorous insects than native congeneric woody plant species. We also tested the hypothesis that leaf miners and gall makers show stronger preference for native woody plants than external leaf chewers. In all investigated regions, leaf miners and gall makers were more abundant and showed higher species richness on native woody plants than on congeneric alien plants. In contrast, external leaf chewers did not cause more damage to native plants than to alien plants, possibly because leaf chewers are, in general, less species specific than leaf miners and gall makers. These results, obtained over a very large number of plant-enemy systems, generally support the hypothesis that alien plants partly escape from phytophagous invertebrates but also show that different feeding guilds may react differently to the introduction of alien plants.     

The invertebrate fauna on broom,Cytisus scoparius,in two native and two exotic habitats

This study quantifies the invertebrate fauna found on broom, Cytisus scoparius, L. (Link), in two countries where it grows as a native plant (France and England) and two countries where it grows as an alien plant (New Zealand and Australia). The data are used to test three hypotheses concerning the predicted differences in invertebrate community structure in native versus exotic habitats: (1) Are generalist phytophages dominant in exotic habitats and specialist phytophages dominant in native habitats? (2) Are there empty phytophage niches in exotic habitats? (3) As a plant species accumulates phytophages, do these in turn accumulate natural enemies? The broom fauna was sampled at five sites in each country by beating five broom bushes per site. The sampling efficiency of beating was quantified at one field site and it was shown to collect 87 % of invertebrate abundance, 95 % of invertebrate biomass and 100 % of phytophagous species found on the branches. Generalist phytophages were dominant on broom in exotic habitats and specialists dominant on broom in the native habitats. Thus, the two countries where broom grows as a native plant had higher numbers of total phytophage species and a higher abundance of specialist phytophages per bush. There was no significant difference in the average abundance of generalist phytophage species found per bush in native and alien habitats. Phytophages were assigned to seven feeding niches: suckers, root feeders, external chewers, flower feeders, seed feeders, miners and pollen feeders. Empty niches were found in the exotic habitats; species exploiting structurally specific parts of the host plant, such as flowers and seeds, were absent in the countries where broom grows as an alien plant. The pattern of niche occupancy was similar between native and exotic habitats when just the generalist phytophages were considered. As phytophage abundance and biomass increased, there were concomitant increases in natural enemy abundance and biomass. Thus, it appears that as plants accumulate phytophages, the phytophages in turn accumulate natural enemies and a food web develops around the plant. Moreover, in the native countries, the history of association between the natural enemies and their prey has been sufficient for specialist predators and parasitoids, feeding on the specialist phytophages, to have evolved.     

Invertebrate,not small vertebrate,herbivory interacts with nutrient availability to impact tallgrass prairie community composition and forb biomass

The effects of herbivores and their interactions with nutrient availability on primary production and plant community composition in grassland systems is expected to vary with herbivore type. We examined the effects of invertebrate and small vertebrate herbivores and their interactions with nutrient availability on grassland plant community composition and aboveground biomass in a tallgrass prairie ecosystem. The abundance of forbs relative to grasses increased with invertebrate herbivore removals. This increase in forb abundance led to a shift in community composition, where invertebrate removals resulted in greater plant species evenness as well as a divergence in composition among plots. In contrast, vertebrate herbivore removals did not affect plant community composition or aboveground biomass. Nutrient additions alone resulted in a decrease in plant species richness and an increase in the abundance of the dominant grass, but the dominant grass species did not greatly increase in abundance when nutrient additions were combined with invertebrate removals. Rather, several subdominant forbs came to dominate the plant community. Additionally, the combined nutrient addition and invertebrate herbivore removal treatment increased forb biomass, suggesting that invertebrate herbivores suppress the responses of forb species to chronic nutrient additions. Overall, the release of forbs from invertebrate herbivore pressure may result in large shifts in species composition, with consequences for aboveground biomass and forage quality due to altered grass:forb ratios in grassland systems.     

Comparison of the herbivore defense and competitive ability of ancestral and modern genotypes of an invasive plant,Lespedeza cuneata

The evolution of increased competitive ability (EICA) hypothesis provides a compelling explanation for the success of invasive species. It contends that because alien plants have escaped their coevolved natural enemies, selection pressures favor a diversion of resources from herbivore defense to traits that confer increased competitive ability. Here, we provide evidence for EICA in the noxious grassland invader Lespedeza cuneata, by comparing the ancestral genotype introduced to North America in 1930 with modern‐day invasive (North American) and native (Japanese) genotypes. We found that the invasive genotype was a better competitor than either the native or the ancestral genotype. Further, the invasive genotype exhibited greater induced resistance but lower constitutive resistance than the ancestral and native genotypes. Our results suggest that selection has played a pivotal role in shaping this invasive plant species into a more aggressive, but less constitutively defended competitor.