Differential induced response to generalist and specialist herbivores by Lindera benzoin (Lauraceae) in sun and shade

Theoretically, induced defenses should be prevalent within low resource environments like the forest understory where constitutive defenses would be costly. Also, the induced response should be stronger when the herbivore is a generalist rather than a specialist, which often have mechanisms to avoid or overcome plant defenses. These ideas have been previously tested for herbaceous species, and we examined these predictions in Lindera benzoin (spicebush), a common woody shrub of the eastern deciduous forest. Lindera benzoin plants in contrasting light environments served as control plants or were subjected to one of four treatments: application of jasmonic acid, clipping, herbivory by the specialist Epimecis hortaria (tulip tree beauty) and herbivory by the generalist Spodoptera exigua (beet armyworm). Following treatment, we assessed induced responses by measuring leaf chemistry (C/N ratio, protein content, and peroxidase activity), and by using insect bioassays with E. hortaria larvae. We found no difference in peroxidase activity between light environments in controls, plants treated with clipping or jasmonic acid. In plants subject to insect herbivory, peroxidase activity was greater in shade plants than in sun plants. The magnitude of this increase in the shade varied between the herbivores, with a 32 fold increase in plants exposed to the generalist S. exigua and a 9 fold increase in plants exposed to the specialist E. hortaria . Leaves from shade plants had more protein and lower C/N ratios than leaves from sun plants, regardless of induction treatment. In control plants, E. hortaria larvae consumed more leaf biomass and achieved greater final weights in the sun than in the shade, but these differences disappeared with induction treatments were applied. These results are among the first to show rapid induction in a woody plant, and different levels of induction with light environments and with specialist versus generalist herbivores.     

Influence of Drought and Shade on Seedling Growth of Native and Invasive Trees in the Seychelles

We studied the growth of seedlings of native and invasive tree species from secondary tropical forests on Mahé (Seychelles). We were interested in whether native or invasive species are more drought-tolerant, and therefore conducted a garden (pot) experiment comparing the growth of seedlings of five native and five invasive tree species under different light (7% and 60% transmittance) and water (natural and repeated drought stress) conditions. Differences in the responses of native and invasive species to these treatments were small. In both groups, mean relative growth rates were reduced only slightly by intermittent drought that caused wilting of leaves. However, invasive species produced clearly thinner leaves (high specific leaf area, SLA) and more root biomass than native species under high light, while these differences were small under low light. Native species performed better than invasive species under low light with low water availability. It appears that high phenotypic plasticity allows some fast-growing invasive species to cope with water stress by adjusting the relative allocation of resources to aboveground and belowground structures under high light, while this strategy is not effective when both light and water resources are limiting. We conclude that water stress may reduce the invasibility of shaded habitats by fast-growing invasive species, while water stress in unshaded habitats may have less effect on invasive species than previously recognized.     

Seedlings of the introduced invasive shrub Eugenia uniflora (Myrtaceae) outperform those of its native and introduced non-invasive congeners in Florida

One particularly compelling explanation for the success of invasive species is the ability to outperform other species in characteristics affecting fitness. Past studies have compared native or introduced non-invasive species to their invasive counterparts, while a system incorporating both native and introduced non-invasive congeners provides an opportunity for multiple controls. We used such a system of Eugenia congeners in Florida to compare seedling performance. In order to determine if invasive Eugenia uniflora seedlings outperform those of its congeners, we sowed seeds in the field and a common garden and quantified seedling emergence, growth, and survival, as well as foliar damage by insect herbivores. We obtained similar results in the field and garden experiments. Although there were no differences in seedling emergence for E. uniflora seedlings when compared to some of its introduced congeners in certain years, emergence of E. uniflora seedlings was consistently high across years. However, emergence, growth, and survival rates of native species were consistently low. In addition, E. uniflora outperformed its introduced and native congeners in growth and survival traits in most comparisons, even when sustaining higher levels of herbivore damage by an introduced weevil, Myllocerus undatus Marshall. Our results support our predictions, indicating that invasive E. uniflora may possess a competitive advantage because its seedlings perform better than or equivalently to its congeners in all of the attributes quantified. Our study further suggests that measurements of such traits may be useful in determining the likelihood of invasion by newly introduced woody plant species.     

Specific leaf area relates to the differences in leaf construction cost,photosynthesis, nitrogen allocation,and use efficiencies between invasive and noninvasive alien congeners

Comparisons between invasive and native species may not characterize the traits of invasive species, as native species might be invasive elsewhere if they were introduced. In this study, invasive Oxalis corymbosa and Peperomia pellucida were compared with their respective noninvasive alien congeners. We hypothesized that the invasive species have higher specific leaf (SLA) than their respective noninvasive alien congeners, and analyzed the physiological and ecological consequences of the higher SLA. Higher SLA was indeed the most important trait for the two invaders, which was associated with their lower leaf construction cost, higher nitrogen (N) allocation to photosynthesis and photosynthetic N use efficiency (PNUE). The higher N allocation to photosynthesis of the invaders in turn increased their PNUE, N content in photosynthesis, biochemical capacity for photosynthesis, and therefore light-saturated photosynthetic rate. The above resource capture-, use- and growth-related traits may facilitate the two invaders' invasion, while further comparative studies on a wider range of invasive and noninvasive congeners are needed to understand the generality of this pattern and to fully assess the competitive advantages afforded by these traits.     

Herbivory is related to taxonomic isolation, but not to invasiveness of tropical alien plants

Aim  The enemy release hypothesis is often invoked to explain why some alien plant species become invasive. Here, we investigated relationships between invasiveness, taxonomic isolation and leaf herbivory for tropical alien plant species introduced to a botanical garden in East Africa.
Location  Amani Botanical Garden, East Usambara mountains, northeast Tanzania.
Methods  We measured the proportion of leaves damaged, and the percentage leaf area damaged on individuals of 28 alien plant species. We extracted data on the presence/absence of native congeners and the number of native confamilial species from an inventory of the East Usambara flora. We also obtained data on planting effort for 26 species, from historical records. Linear and generalized linear models were used to analyse the relationships between invasiveness, herbivory and taxonomic isolation.
Results  Mean proportion of leaves damaged per species was significantly explained by taxonomic isolation; proportion of leaves damaged increased with the number of native confamilial species and was greater, on average, for species with native congeners than those without native congeners. The mean percentage of leaf area damaged per species could not be explained by any variables considered in this study. There was no relationship between the degree of herbivory or taxonomic isolation and alien plant species invasiveness, but more-invasive species did have a significantly greater planting effort than less-invasive species.
Conclusions  The role herbivores play in controlling alien plant invasions has been investigated relatively little in the tropics. In this study, although the amount of herbivory suffered by alien plants was related to taxonomic isolation, we found no evidence for leaf-feeding invertebrates having a significant role in invasion, suggesting that other factors may be responsible for differences in species success.     

How to induce defense responses in wild plant populations? Using bilberry (Vaccinium myrtillus) as example

Inducible plant defense is a beneficial strategy for plants, which imply that plants should allocate resources from growth and reproduction to defense when herbivores attack. Plant ecologist has often studied defense responses in wild populations by biomass clipping experiments, whereas laboratory and greenhouse experiments in addition apply chemical elicitors to induce defense responses. To investigate whether field ecologists could benefit from methods used in laboratory and greenhouse studies, we established a randomized block‐design in a pine‐bilberry forest in Western Norway. We tested whether we could activate defense responses in bilberry (Vaccinium myrtillus) by nine different treatments using clipping (leaf tissue or branch removal) with or without chemical treatment by methyljasmonate (MeJA). We subsequently measured consequences of induced defenses through vegetative growth and insect herbivory during one growing season. Our results showed that only MeJA‐treated plants showed consistent defense responses through suppressed vegetative growth and reduced herbivory by leaf‐chewing insects, suggesting an allocation of resources from growth to defense. Leaf tissue removal reduced insect herbivory equal to the effect of the MeJa treatments, but had no negative impact on growth. Branch removal did not reduce insect herbivory or vegetative growth. MeJa treatment and clipping combined did not give an additional defense response. In this study, we investigated how to induce defense responses in wild plant populations under natural field conditions. Our results show that using the chemical elicitor MeJA, with or without biomass clipping, may be a better method to induce defense response in field experiments than clipping of leaves or branches that often has been used in ecological field studies.     

Influence of an invasive ant on grazing and detrital communities and nutrient fluxes in a tropical forest

Aim  Pathways linking grazing and detrital subsystems of terrestrial ecosystems are important for ecosystem processes and function, but remain poorly understood. The invasion of a generalist predator creates a unique opportunity to study the effects of predation across these subsystems. We examine here, the effects of a non-native generalist predator, the little red fire ant ( Wasmannia auropunctata, Roger) on both grazing and detrital invertebrate communities and ecosystem processes in a rain forest understorey.
Location  Gamba Protected Area Complex, south-western Gabon, Africa.
Methods  We measured abundances and diversities of understorey grazing and detrital invertebrate communities, soil nutrients, herbivory, litter fragmentation rates and leaf chemistry of a dominant understorey shrub inside and outside of 19 separate invasion fronts. We then explored possible trophic cascades and pathways of interaction using path analysis.
Results  Results suggest that invasive ants may alter herbivory regimes, grazing and detrital communities, and may indirectly alter litter decomposition and nutrient cycling in the soil by suppressing important microbivore and detritivore populations with consequences for leaf chemistry.
Main conclusions  These results demonstrate that generalist predators may be major drivers of both grazing and detrital subsystems by inducing strong shifts in adjacent communities that ultimately affect ecosystem processes.     

Effects of generalist herbivory on resistance and resource allocation by the invasive plant,Phytolacca americana

Successful invasions by exotic plants are often attributed to a loss of co‐evolved specialists and a re‐allocation of resources from defense to growth and reproduction. However, invasive plants are rarely completely released from insect herbivory because they are frequently attacked by generalists in their introduced ranges. The novel generalist community may also affect the invasive plant's defensive strategies and resource allocation. Here, we tested this hypothesis using American pokeweed (Phytolacca americana L.), a species that has become invasive in China, which is native to North America. We examined resistance, tolerance, growth and reproduction of plant populations from both China and the USA when plants were exposed to natural generalist herbivores in China. We found that leaf damage was greater for invasive populations than for native populations, indicating that plants from invasive ranges had lower resistance to herbivory than those from native ranges. A regression of the percentage of leaf damage against mass showed that there was no significant difference in tolerance between invasive and native populations, even though the shoot, root, fruit and total mass were larger for invasive populations than for native populations. These results suggest that generalist herbivores are important drivers mediating the defensive strategies and resource allocation of the invasive American pokeweed.     

Seasonality of herbivory and communication between individuals of sagebrush

Seasonal changes in herbivore numbers and in plant defenses are well known to influence plant–herbivore interactions. Some plant defenses are induced in response to herbivore attack or cues correlated with risk of attack although seasonal variation in these defenses is relatively poorly known. We previously reported that sagebrush becomes more resistant to its herbivores when neighboring plants have been experimentally clipped with scissors. In this study we asked whether herbivory to leaves of sagebrush varied seasonally and whether there was seasonal variation in natural levels of damage when neighbors were clipped. We found that sagebrush accumulated most chewing damage early in the season, soon after the spring flush of new leaves. This damage was caused by generalist grasshoppers, deer, specialist caterpillars, beetles, gall makers, and other less common herbivores. Sagebrush showed no evidence of preferentially abscising leaves that had been experimentally clipped. Experimental clipping by Trirhabda pilosa beetle larvae caused neighbors to accumulate less herbivore damage later that season, similar to results in which clipping was done with scissors. Induced resistance caused by experimentally clipping a neighbor was affected by season; plants with neighbors clipped in May accumulated less damage throughout the season relative to plants with unclipped neighbors or neighbors clipped later in the summer. We found a correlation between seasonal herbivore pressure, damage accumulated by plants, and induced responses to experimentally clipping neighbors. The causal mechanisms responsible for this correlation are unknown although a strong seasonal effect was clear.     

Traits correlate with invasive success more than plasticity: A comparison of three Centaurea congeners

The importance of phenotypic plasticity for successful invasion by exotic plant species has been well studied, but with contradictory and inconclusive results. However, many previous studies focused on comparisons of native and invasive species that co‐occur in a single invaded region, and thus on species with potentially very different evolutionary histories. We took a different approach by comparing three closely related Centaurea species: the highly invasive C. solstitialis, and the noninvasive but exotic C. calcitrapa and C. sulphurea. These species have overlapping distributions both in their native range of Spain and in their non‐native range of California. We collected seeds from 3 to 10 populations from each region and species and grew them in common garden greenhouse conditions to obtain an F1 generation in order to reduce maternal effects. Then, F1 seeds were grown subjected to simulated herbivory, variation in nutrient availability, and competition, to explore plasticity in the responses to these conditions. We found little variation in phenotypic plasticity among species and regions, but C. solstitialis plants from California produced more biomass in competition than their Spanish conspecifics. This species also had the highest relative growth rates when in competition and when grown under low nutrient availability. Noninvasive congeners produced intermediate or opposite patterns.     

Compensation and resistance to herbivory in seagrasses: induced responses to simulated consumption by fish

Herbivory can induce changes in plant traits that may involve both tolerance mechanisms that compensate for biomass loss and resistance traits that reduce herbivore preference. Seagrasses are marine vascular plants that possess many attributes that may favour tolerance and compensatory growth, and they are also defended with mechanisms of resistance such as toughness and secondary metabolites. We quantified phenotypic changes induced by herbivore damage on the temperate seagrass Posidonia oceanica in order to identify specific compensatory and resistance mechanisms in this plant, and to assess any potential trade-offs between these two strategies of defence. We simulated three natural levels of fish herbivory by repeatedly clipping seagrass leaves during the summer period of maximum herbivory. Compensatory responses were determined by measuring shoot-specific growth, photosynthetic rate, and the concentration of nitrogen and carbon resources in leaves and rhizomes. Induced resistance was determined by measuring the concentration of phenolic secondary metabolites and by assessing the long-term effects of continued clipping on herbivore feeding preferences using bioassays. Plants showed a significant ability to compensate for low and moderate losses of leaf biomass by increasing aboveground growth of damaged shoots, but this was not supported by an increase in photosynthetic capacity. Low levels of herbivory induced compensatory growth without any measurable effects on stored resources. In contrast, nitrogen reserves in the rhizomes played a crucial role in the plant’s ability to compensate and survive herbivore damage under moderate and high levels of herbivory, respectively. We found no evidence of inducibility of long-term resistance traits in response to herbivory. The concentration of phenolics decreased with increasing compensatory growth despite all treatments having similar carbon leaf content, suggesting reallocation of these compounds towards primary functions such as cell-wall construction.     

Induction of extrafloral nectar depends on herbivore type in invasive and native Chinese tallow seedlings

Although induced defenses are widespread in nature, and a potentially important strategy used by invasive plants, it is unclear how induced defenses vary among populations and whether the intensity and duration of induced defenses depends on herbivore type. For invasive plants, low herbivore loads in their introduced ranges can lead to differences in herbivore defense compared to their native ranges, but we currently know little about how induced defenses vary among native and invasive populations. We conducted a greenhouse experiment to examine variation in one type of induced defense, extrafloral nectar (EFN) production, among native and invasive populations of Chinese tallow tree, Triadica sebifera. We experimentally manipulated herbivory from an exotic generalist scale insect, a native generalist caterpillar, both herbivores, or neither and then examined EFN production by Triadica. Damage from leaf-chewing caterpillars resulted in strongly induced EFN in both native and invasive populations while damage from phloem-feeding scales did not. Extrafloral nectar production and dissolved solute content peaked 4 days after caterpillar herbivory for both native and invasive populations. Number and proportion of leaves producing EFN, EFN volume and concentration of dissolved solutes were similar among native and invasive populations. These results suggest that selection for indirect defenses may be different than selection for other defenses in the introduced ranges of invasive plants, as constitutive and induced EFN production is retained in invasive populations.     

Long-term effects of feral goats (<Emphasis Type="Italic">Capra hircus</Emphasis>) on Mediterranean island communities: results from whole island manipulations

Islands exhibit disproportionally high biodiversity, however high levels of endemism and simplified food webs make their communities susceptible to invasive species. Introduced goats (Capra hircus), a generalist herbivore, are among the most harmful invasive species on islands. Concern about goat impacts on island communities have resulted in eradication programs, which have been generally implemented without comprehensive evaluation and monitoring. Unintended consequences may follow eradication, as grazing can have complex effects on island food webs. Using whole island manipulations, we evaluate the long-term, community-wide effects of goat herbivory, as well as their subsequent removal, in a system of 16 islands in the Aegean Sea (Greece) located within the Mediterranean biodiversity hotspot. Goat grazing on these islands is a major conservation concern, as these support endemic plant communities that have evolved in low herbivory conditions and lack appropriate defenses. We show that goat introductions lead to significant decreases in vegetation height, percent cover, and biomass but not to immediate plant species loss, as native island endemics are replaced with widespread generalist taxa carried in by the livestock. Additionally, goats contribute to the desertification of islands by initiating a long-term soil loss cycle that continues even after goats are removed; however, remaining soil structure and chemistry are not affected. Island arthropod populations do not appear to be significantly impacted by goat introduction or removal, except for a distinct increase in the order Diptera with goat presence. This study also reaffirms the role of seabirds in providing important marine subsidies, rich in nitrogen and phosphorus, to island food webs. Plant species diversity declines following goat removal, and vegetation cover returns only partially, as further recovery is being prevented by the long-term loss of soil. This suggests that following goat removal, island communities may require additional restoration efforts, including seabird reestablishment and reintroduction of extirpated plant populations, to promote island recovery.     

The relationship between leaf nitrogen,nitrogen metabolites and herbivory in two species of Nyctaginaceae from the Brazilian Cerrado

Guapira graciliflora and Neea theifera are taxonomically related species of the tribe Pisoneae. Both species are found in the same environment, the Brazilian Cerrado, and therefore, are subjected to similar selective pressures. These species occur in oligotrophic environments, yet contain high concentrations of nitrogen in their leaves. The present study was carried out to investigate the ecological role of nitrogen in herbivory on these species. The differences in the N content, compositions of secondary N-metabolites, mechanical resistance, and water content between their leaves indicate that these species have different adaptations as defense mechanisms. In both species, their high nitrogen content seems to promote herbivory. The presence of secondary nitrogen metabolites does not prevent the species from suffering intense damage by herbivores on their early leaves. The herbivory rates observed were lower for mature leaves of both species than for young leaves. In G. graciliflora, nutritional content and leaf hardness are the most important variables correlated with reduction of herbivory rates, whereas in N. theifera, N compounds are also correlated with herbivory rates. Despite the differences in the strategies of these two species, they exhibit a similar efficiency of protection against natural enemies because their total herbivory rates are similar. The difference in their N defense allocation may imply benefits for survival under Cerrado conditions. We briefly discuss the oligotrophic habitat conditions of the studied plants and possible advantages of their strategies of N accumulation and metabolic uses.     

Specificity of extrafloral nectar induction by herbivores differs among native and invasive populations of tallow tree

Background and Aims

Invasive plants can be released from specialist herbivores and encounter novel generalists in their introduced ranges, leading to variation in defence among native and invasive populations. However, few studies have examined how constitutive and induced indirect defences change during plant invasion, especially during the juvenile stage.

Methods

Constitutive extrafloral nectar (EFN) production of native and invasive populations of juvenile tallow tree (Triadica sebifera) were compared, and leaf clipping, and damage by a native specialist (Noctuid) and two native generalist caterpillars (Noctuid and Limacodid) were used to examine inducible EFN production.

Key results

Plants from introduced populations had more leaves producing constitutive EFN than did native populations, but the content of soluble solids of EFN did not differ. Herbivores induced EFN production more than simulated herbivory. The specialist (Noctuid) induced more EFN than either generalist for native populations. The content of soluble solids in EFN was higher (2·1 times), with the specialist vs. the generalists causing the stronger response for native populations, but the specialist response was always comparable with the generalist responses for invasive populations.

Conclusions

These results suggest that constitutive and induced indirect defences are retained in juvenile plants of invasive populations even during plant establishment, perhaps due to generalist herbivory in the introduced range. However, responses specific to a specialist herbivore may be reduced in the introduced range where specialists are absent. This decreased defence may benefit specialist insects that are introduced for classical biological control of invasive plants.     

Competitive ability,not tolerance,may explain success of invasive plants over natives

When entering a new community, introduced species leave behind members of their native community while simultaneously forming novel biotic interactions. Escape from enemies during the process of introduction has long been hypothesized to drive the increased performance of invasive species. However, recent studies and quantitative syntheses find that invaders often receive similar, or even more, damage from enemies than do native species. Therefore, invasives may be those more tolerant to enemy damage, or those able to maintain competitive ability in light of enemy damage. Here, we investigate whether tolerance and competitive ability could contribute to invasive plant success. We determined whether invasive plants were more competitive than native or noninvasive exotic species in both the presence and absence of simulated herbivory. We found competition and herbivory additively reduced individual performance, and affected the performance of native, invasive, and noninvasive exotic species’ to the same degree. However, invasives exerted stronger competitive effects on an abundant native species (Elymus canadensis) in both the presence and absence of herbivory. Therefore, while invasive species responded similarly to competition and simulated herbivory, their competitive effects on natives may contribute to their success in their introduced range.     

Invading from the garden? A comparison of leaf herbivory for exotic and native plants in natural and ornamental settings

Abstract The enemies release hypothesis proposes that exotic species can become invasive by escaping from predators and parasites in their novel environment. Agrawal et al. (Enemy release? An experiment with congeneric plant pairs and diverse above‐ and below‐ground enemies. Ecology, 86, 2979–2989) proposed that areas or times in which damage to introduced species is low provide opportunities for the invasion of native habitat. We tested whether ornamental settings may provide areas with low levels of herbivory for trees and shrubs, potentially facilitating invasion success. First, we compared levels of leaf herbivory among native and exotic species in ornamental and natural settings in Cincinnati, Ohio, United States. In the second study, we compared levels of herbivory for invasive and noninvasive exotic species between natural and ornamental settings. We found lower levels of leaf damage for exotic species than for native species; however, we found no differences in the amount of leaf damage suffered in ornamental or natural settings. Our results do not provide any evidence that ornamental settings afford additional release from herbivory for exotic plant species.     

Enemy release and plant invasion: patterns of defensive traits and leaf damage in Hawaii

Invasive species may be released from consumption by their native herbivores in novel habitats and thereby experience higher fitness relative to native species. However, few studies have examined release from herbivory as a mechanism of invasion in oceanic island systems, which have experienced particularly high loss of native species due to the invasion of non-native animal and plant species. We surveyed putative defensive traits and leaf damage rates in 19 pairs of taxonomically related invasive and native species in Hawaii, representing a broad taxonomic diversity. Leaf damage by insects and pathogens was monitored in both wet and dry seasons. We found that native species had higher leaf damage rates than invasive species, but only during the dry season. However, damage rates across native and invasive species averaged only 2% of leaf area. Native species generally displayed high levels of structural defense (leaf toughness and leaf thickness, but not leaf trichome density) while native and invasive species displayed similar levels of chemical defenses (total phenolics). A defense index, which integrated all putative defense traits, was significantly higher for native species, suggesting that native species may allocate fewer resources to growth and reproduction than do invasive species. Thus, our data support the idea that invasive species allocate fewer resources to defense traits, allowing them to outperform native species through increased growth and reproduction. While strong impacts of herbivores on invasion are not supported by the low damage rates we observed on mature plants, population-level studies that monitor how herbivores influence recruitment, mortality, and competitive outcomes are needed to accurately address how herbivores influence invasion in Hawaii.     

Interactive effects of herbivory and competition intensity determine invasive plant performance

Herbivory can reduce plant fitness, and its effects can be increased by competition. Though numerous studies have examined the joint effects of herbivores and competitors on plant performance, these interactive effects are seldom considered in the context of plant invasions. Here, we examined variation in plant performance within a competitive environment in response to both specialist and generalist herbivores using Chinese tallow as a model species. We combined tallow plants from native and invasive populations to form all possible pairwise combinations, and designated invasive populations as stronger neighbours and native populations as weaker neighbours. We found that when no herbivory was imposed, invasive populations always had higher total biomass than natives, regardless of their neighbours, which is consistent with our assumption of increased competitive ability. Defoliation by either generalist or specialist herbivores suppressed plant growth but the effects of specialists were generally stronger for invasive populations. Invasive populations had their lowest biomass when fed upon by specialists while simultaneously competing with stronger neighbours. The root/shoot ratios of invasive populations were lower than those of native populations under almost all conditions, and invasive plants were taller than native plants overall, especially when herbivores were present, suggesting that invasive populations may adopt an "aboveground first" strategy to cope with herbivory and competition. These results suggest that release from herbivores, especially specialists, improves an invader's performance and helps to increase its competitive ability. Therefore, increasing interspecific competition intensity by planting a stronger neighbour while simultaneously releasing a specialist herbivore may be an especially effective method of managing invasive plants.