Invasive slugs as under-appreciated obstacles to rare plant restoration: evidence from the Hawaiian Islands

Introduced slugs have invaded many parts of the world where they were recognized as important pests of gardens and agriculture, but we know little about the effects of introduced slugs on rare plants in natural areas. The Hawaiian Islands have no native slugs, but over a dozen introduced slug species are now established. We reviewed Rare Plant Recovery Plans produced by the U.S. Fish and Wildlife Service for Hawaii and found that introduced slugs were specifically mentioned as threats or potential threats to 59 rare plant species (22% of all endangered and threatened plants), based mainly on anecdotal observations by field biologists. We then initiated an experimental field study to assess the impact of slug herbivory on the growth and survival of two endangered plant species (Cyanea superba, and Schidea obovata), one non-endangered native species (Nestegis sandwicensis) and two co-occurring invasive plant species (Psidium cattleianum and Clidemia hirta). In mesic forest on the Island of Oahu, we tracked the fate of outplanted seedlings in replicated 1 m² plots, with and without slug control. Slugs decreased seedling survival of the endangered species by 51%, on average. Slugs did not significantly affect survival of the non-endangered or invasive plant species. Introduced slugs seem to be under-appreciated as a direct cause of plant endangerment. Invasive slugs may also facilitate the success of some invasive plant species by reducing competition with more palatable, native plant competitors. Slug control measures are relatively inexpensive and could facilitate rare plant establishment and population recovery.     

Restoration of mountain beech (

Fire occurs relatively frequently in beech (Nothofagus) forest in drought prone eastern areas of the South Island, New Zealand. Because beech is poorly adapted to fire, and is slow to regenerate, forest is normally replaced by scrub or grassland. Seeding was investigated as a means of restoring mountain beech (N. solandri var. cliffortioides) forest after fire destroyed 300 ha of forest at Mt. Thomas, Canterbury, in 1980. A mixture of mountain beech, Leptospermum scoparium and other small tree and shrub species was sown within a year of the fire in the presence and absence of pasture species as a cover crop, and fertiliser. Seeding of mountain beech and L. scoparium was successful, but other species were of limited success. Competition from pasture species inhibited establishment of all native species. Fertiliser increased L. scoparium plant numbers in the first year but had no other beneficial effect on establishment of native species. Leptospermum scoparium provided a dense shrub cover in plots where the native species were sown in the absence of pasture species, but mountain beech had begun to overtop the shrub canopy by 20 years after seeding. Browsing by insects or small animals in the first 2 years is suggested as the main cause of mortality in mountain beech. Mountain beech seeded at 1.4 kg/ha resulted in about 1800 saplings/ha at age 20. It is suggested that seeding the wider burn area more than 2 years after the fire would have been unsuccessful because of competition from herbaceous species, especially Agrostis capillaris, which rapidly invaded the burnt area. A strategy is outlined for establishing mountain beech over large areas when limited quantities of seed are available.     

Soil water exploitation after fire: competition between Bromus tectorum (cheatgrass) and two native species

Summary Causes for the widespread abundance of the alien grass Bromus tectorum (cheatgrass) after fire in semiarid areas of western North America may include: (1) utilization of resources freed by the removal of fireintolerant plants; and (2) successful competition between B. tectorum and individual plants that survive fire. On a site in northwestern Nevada (USA), measurements of soil water content, plant water potential, aboveground biomass production, water use efficiency, and B. tectorum tiller density were used to determine if B. tectorum competes with either of two native species (Stipa comata and Chrysothamnus viscidiflorus) or simply uses unclaimed resources. Soil water content around native species occurring with B. tectorum was significantly lower (P<0.05) than around individuals without B. tectorum nearby. Native species had significantly more negative plant water potential when they occurred with B. tectorum. Aboveground biomass was significantly higher for native species without B. tectorum. However, the carbon isotope ratio of leaves for native species with B. tectorum was not significantly different from individuals without B. tectorum. Thus, B. tectorum competes with native species for soil water and negatively affects their wate status and productivity, but the competition for water does not affect water use efficiency of the native species. These adverse effects of B. tectorum competition on the productivity and water status of native species are also evident at 12 years after a fire. This competitive ability of B. tectorum greatly enhances its capability to exploit soil resources after fire and to enhance its status in the community.     

Takahe Valley Hut: a focal point for weed invasion in an isolated area of Fiordland National Park, New Zealand

The role of backcountry huts as focal points for weed establishment and spread into New Zealand’s national parks has received little attention. In this study we describe the pattern of weed spread around Takahe Valley Hut, Murchison Mountains, Fiordland National Park. Established in 1948, the hut is located at 900 m a.s.l. at the ecotone between Nothofagus forest and valley floor shrubland/grassland. We recorded the distribution of vascular plants in quadrats (110) placed by restricted randomisation around the hut, and measured relative irradiance and distance from the hut. Nine exotic species, mostly grasses, were recorded, the most frequent being Agrostis capillaris (34%). The majority of occurrences of exotic plant species were located in the immediate vicinity (less than 5 m) of the hut but two exotic species (Agrostis capillaris and Dactylis glomerata) ranged more widely. Exotic species were present in well-drained shrubland and grassland but did not extend far into Nothofagus forest or onto infertile wetlands. The percentage of exotic species in quadrats declined significantly with distance from the hut. There was no linear relationship between the percentage of exotic species and relative irradiance. When forest quadrats were excluded, the number of native species in quadrats was negatively correlated with the number of exotic species, suggesting competitive displacement of native species by exotics in non-forest habitats. Long-term persistence of most exotic species at this site depends on physical disturbance and nutrient enrichment associated with human activities at the hut site. However the maintenance of this species pool has provided sufficient propagule pressure for some exotic species to disperse into the wider area. Weed accumulation around huts can be reduced by locating huts in vegetation types that are more resistant to invasion, and maintaining facilities to eliminate local weed infestations.     

Effects of competition and litter on a carnivorous plant, Drosera capillaris (Droseraceae)

Although carnivorous plants are widely recognized as being generally restricted to open habitats, tests of the effects of competition on individual performance are extremely rare. In this study, I examined the effects of the removal of herbaceous and shrub canopies on seedling density and growth, survival, and reproduction of phytometers of a small insectivorous plant, Drosera capillaris (pink sundew). I also examined the distribution of this species in relation to the occurrence of woody species in a frequently burned wet savanna in southeastern Mississippi. Killing plants and removing dead biomass increased seedling density in both open areas and shrub thickets. The removal of dead biomass following herbicide application was critical to increasing densities of seedlings. Killing plants with herbicide without also clearing residual litter and standing dead was not sufficient to increase seedling densities in shrub thickets. Although the removal of the groundcover canopy strongly influenced the density of seedlings, it had very little effect on survival, growth, and reproduction of small phytometers during a single growing season. Survival of phytometers was greater in open areas than in shrub thickets, regardless of whether the groundcover canopy was removed. Densities of both seedlings and adults were greater in open areas away from shrub thickets than beneath the woody canopies of thickets and were negatively correlated with the leaf area index of groundcover vegetation. Results of this study show that the establishment of this carnivorous plant species is limited in part by the effects of litter on seedling density in both open areas and shrub thickets.     

Exotic Slugs Pose a Previously Unrecognized Threat to the Herbaceous Layer in a Midwestern Woodland

Developing effective restoration strategies requires first identifying the underlying factors limiting native plant recovery. The slug Deroceras reticulatum is an important herbivore in Europe, a global agricultural pest, and is introduced and abundant throughout eastern North America, but little information is available on the effect of this exotic herbivore on the forest herbaceous layer. Here, we test the palatability of 12 forest herbs to the introduced slug D. reticulatum and use field surveys to determine the degree to which slugs are damaging plants in the field. In laboratory feeding trials, slugs readily consumed most plants, but avoided the grass Elymus virginicus, the invasive forb Alliaria petiolata (garlic mustard), and thicker leaved plants. In the field, we documented significant slug damage, with close to 50% or more of plant leaves damaged by slugs on five of the six native species tested. Slug damage in the field was predicted by laboratory‐determined acceptability, but was significantly greater on short‐statured rosette species than on erect plants for a given acceptability value. Our results identify introduced slugs as an important, but overlooked obstacle to forest herb restoration and potential drivers of larger scale understory compositional change. The relaxed herbivore pressure on A. petiolata, relative to native competitors, suggests that invasive plant removal alone may not result in the recovery of native flora. Rather, restoration of unpalatable native species should accompany invasive plant control in slug invaded areas. Erect forbs, thick‐leaved plants, and graminoids should have the greatest success where introduced slugs are abundant.     

No interaction between competition and herbivory in limiting introduced Cirsium vulgare rosette growth and reproduction

Both competition and herbivory have been shown to reduce plant survival, growth, and reproduction. Much less is known about whether competition and herbivory interact in determining plant performance, especially for introduced, weedy plant species in the invaded habitat. We simultaneously evaluated both the main and interactive effects of plant neighbors and insect herbivory on rosette growth and seed reproduction in the year of flowering for Cirsium vulgare (bull thistle, spear thistle), an introduced Eurasian species, in tallgrass prairie in 2 years. Effects of insect herbivory were strong and consistent in both years, causing reduced plant growth and seed production, whereas the effects of competition with established vegetation were weak. The amount of herbivore damage inflicted on rosettes did not depend on the presence of neighbor plants. We also found no interaction between competition and herbivory on key parameters of plant growth and fitness. The results of this study contradict the hypothesis that competitive context interacts with insect herbivory in limiting the invasiveness of this introduced thistle. Further, the results provide additional, experimental evidence that high levels of herbivory on established rosettes by native insects exert significant biotic resistance to the invasiveness of C. vulgare in western tallgrass prairie.     

Colonization by alpine native plants of a stabilized road verge on the Bogong High Plains, Victoria

Summary Although many native species are now used in the revegetation of soil disturbances in Australian alpine areas, exotic species were, until recently, the only components of seed mixes. The use of exotic species and fertilizer was justified by their availability and low cost, and the prediction that native species would replace the exotic sward, presumably once soil nutrient levels dropped to those found in native vegetation. There was no apparent regard for the invasive capacities of the exotic species used. An investigation of a road verge on the Bogong High Plains, revegetated with a mixture of exotic species in the late 1950s, has shown that colonization by native plants can be very slow. Although many native species had colonized by 1993, exotic species still provided 50% of the cover. Agrostis capillaris accounted for most of this. The major native colonizers were Carex spp., Colobanthus affinis , Euchiton spp., Leptinella filicula , Poa hiemata , Ranunculus victoriensis and Scleranthus biflorus . Poa hiemata provided by far the most cover of the native colonizers. There have been considerable increases in the cover and number of native species on the verge since 1989. Cattle grazing and trampling are likely to have limited colonization of native plants prior to the removal of livestock in 1991. Recommendations are made on approaches to future revegetation in the alpine area.     

Abundance,species richness and feeding preferences of introduced molluscs in native grasslands of Victoria,Australia

Abstract Introduced molluscs have invaded endangered, remnant native grasslands of south‐eastern Australia, but few studies have investigated their distribution, abundance or potential impact. Molluscs were surveyed in grassland sites across an urban to rural transect west of Melbourne, Australia. It was confirmed that several introduced mollusc species have invaded these areas. Three snail and five slug species were identified, none of which was native to Australia. The most common species was the brown field slug (Deroceras panormitanum). Mollusc capture and species richness were positively related to the degree of urbanization in the surrounding landscape. There was also a negative relationship with fire frequency. Feeding trials revealed selective herbivory by the black‐keeled slug (Milax gagates) among native plants. Anecdotal evidence that variable glycine (Glycine tabacina) and button wrinklewort (Rutidosis leptorrhynchoides) are palatable to molluscs was supported. Mollusc herbivory may potentially lead to reduced fitness of palatable species, and changes in community composition and structure.     

Exotic consumers interact with exotic plants to mediate native plant survival in a Midwestern forest herb layer

Consumer-facilitated invasions have been proposed as an alternative mechanism to direct competitive exclusion to explain the replacement of native plants by exotics. In a factorial field experiment manipulating competition from the exotic plant Alliaria petiolata and herbivory by exotic mollusks, we documented that mollusk herbivory significantly reduced the survival of two species of native palatable plants, but found minimal direct herbivore effects on less palatable species, including the invasive A. petiolata. These effects were evident after one growing season on younger juvenile plants of Aster cordifolius, but only after two growing seasons on older transplants of the same species, suggesting a greater vulnerability of young plants. In contrast to our expectations, A. petiolata competition alone had no effect on any of the six native species we tested. However, competition from A. petiolata did affect the survival of the most palatable native plant when mollusks were also present. While not significant for any other single species, this same pattern was observed for three of the five remaining native species tested. The selective grazing on palatable plants that we document provides novel evidence contributing to our understanding of observed shifts in the forest herbaceous layer towards the dominance of exotic plants and unpalatable species. More broadly, our results highlight the importance of the interactive effect of consumers and inter-specific competition in forest understories via its contribution to differential survival among regenerating species.     

The invasive shrub Buddleja davidii performs better in its introduced range

It is commonly assumed that invasive plants grow more vigorously in their introduced than in their native range, which is then attributed to release from natural enemies or to microevolutionary changes, or both. However, few studies have tested this assumption by comparing the performance of invasive species in their native vs. introduced ranges. Here, we studied abundance, growth, reproduction, and herbivory in 10 native Chinese and 10 invasive German populations of the invasive shrub Buddleja davidii (Scrophulariaceae; butterfly bush). We found strong evidence for increased plant vigour in the introduced range: plants in invasive populations were significantly taller and had thicker stems, larger inflorescences, and heavier seeds than plants in native populations. These differences in plant performance could not be explained by a more benign climate in the introduced range. Since leaf herbivory was substantially reduced in invasive populations, our data rather suggest that escape from natural enemies, associated with increased plant growth and reproduction, contributes to the invasion success of B. davidii in Central Europe.     

入侵植物的繁殖策略以及对本土植物繁殖的影响

理解入侵生物的繁殖策略是阐明生物入侵机制的一个重要方面。入侵植物常表现出一些共同的繁殖特征, 如以两性花为主的性系统、自动自交为主的繁育系统或不依赖传粉媒介的无融合生殖和无性繁殖以及高生殖投资的资源配置策略等。成功入侵的外来植物通过影响本土的传粉者, 在种群和群落水平上影响本土植物的有性繁殖, 甚至促使某些本土植物在繁殖对策和表型性状上发生快速转变。目前, 入侵植物繁殖策略及其生态效应的研究多侧重于入侵种的快速演化, 而有关外来植物与本土植物间的相互影响及其可能存在的协同适应研究还较为缺乏。探讨本土植物在外来种入侵压力下的繁殖对策和响应机制, 将丰富人们对物种间竞争、共存及群落构建等机制的深入了解。从繁殖和适应的角度探求入侵植物与本土植物之间的复杂关系, 将有助于解析生物入侵的机制及人类干扰下的物种演化规律, 也为预测和防控入侵植物提供科学依据。     

Adaptive introductions: How multiple experiments and comparisons to wild populations provide insights into requirements for long-term introduction success of an endangered shrub

Recovery of an imperiled plant species may require augmentation of existing populations or creation of new ones. Hundreds of such projects have been conducted over the last few decades, but there is a bias in the literature favoring successes over failures. In this paper, we evaluate a series of introductions that experimentally manipulated microhabitat and fire in an adaptive introduction framework. Between 2002 and 2012, we (and our collaborators) carried out ten introductions and augmentations of Florida ziziphus Pseudoziziphus (Condalia, Ziziphus) celata, a clonal shrub limited to very small populations and narrowly endemic to pyrogenic central Florida sandhills. Six of the introductions were designed as experiments to test hypotheses about how demographic performance was affected by microhabitat, fire, and propagule type. Introduced transplants had high survival (<90% annually), inconsistent growth, and little transition to reproduction, while introduced seeds had low germination and survival. Transplants were more efficient than seeds as translocation propagules. Shaded (vs. open) sites supported generally higher transplant and seedling survival and seed germination percentages, but growth responses varied among experiments. Supplemental irrigation increased transplant survival and seed germination, but otherwise seedling and plant survival and growth were not significantly affected. Contrary to expectations based on wild populations, introduced propagules have not been more successful in unshaded sites, suggesting that Florida ziziphus has broader microhabitat preferences than hypothesized. Compared to wild plants, introduced plants had similar survival and responses to fire, slower growth, and more delayed flowering. Introduced plants had no clonal spread. While no introduced population has demonstrated a capacity for long-term viability, one augmented population has flowered and produced viable fruits. Given that Florida ziziphus genets are long-lived, low levels of sexual reproduction may be adequate for the establishment of viable populations. Thus, after many translocations over more than a decade, it is premature to characterize any single translocation as a success or a failure, underscoring the need for a long view of translocation success.     

EFFECTS OF COMPETITION AND DISTURBANCE ON THE RESPROUTING PERFORMANCE OF THE MEDITERRANEAN SHRUB ERICA MULTIFLORA L. (ERICACEAE)

Two field experiments were designed to evaluate the importance of competition, fire, repeated disturbance, and their interactions on the vegetative and reproductive performance of the Mediterranean shrub Erica multiflora over a 2.5-yr period. In a burn experiment, fire was applied to the ground-level stumps of previously clipped 13-yr-old plants with a propane torch and competition was diminished by removal of neighboring plants. Fire resulted in a reduction of sprout vigor and biomass of flowers; mature neighbors also reduced E. multiflora sprout vigor and flowering. The interaction between fire and competition was nonsignificant. In a stand burned by a wildfire we studied the effects of regenerating neighbors on target plants by removing all neighbors or only Quercus coccifera, the most dominant species in the burned stand. In this stand we also simulated herbivory by repeatedly clipping the sprouts of E. multiflora. Regenerating neighbors did not affect target plant sprout vigor after the wildfire, but did cause a decrease in the biomass of flowers per plant. Survival decreased after repeated clipping but was not affected by neighborhood treatment. The results suggest that the importance of competition on resprouting vigor was temporally variable. Variables related to plant size rather than species determined competitive superiority: resprouting neighbors did not affect resprouting performance of target plants, but mature neighbors did. In nature, fire may directly reduce vegetative and reproductive biomass by the heating effect. But it may have an indirect positive effect on biomass, by reducing competition among plants. Frequent disturbances that removed aboveground biomass of E. multiflora had a detrimental effect on target plant survival independent of neighborhood effect.     

A phylogenetically controlled analysis of the roles of reproductive traits in plant invasions

Reproductive traits are tightly linked to plant fitness and may therefore be mechanisms driving biological invasions, including the greater success of more phylogenetically novel introduced species in some systems. We present a phylogenetic comparative analysis of “Baker’s law’’, that introduced plants with the ability to reproduce autogamous or asexually may be better able to establish on introduction. We gathered data from both published and unpublished sources on pollen limitation of 141 species, including 26 introduced species and 115 native species. Our analysis compared differences in the proportion of autonomous autogamy, asexual reproduction, and pollen limitation among native, introduced noninvasive, and introduced invasive plant species, and included the phylogenetic novelty of the introduced species to the native species in that community. Introduced species were more likely to be autogamous than native species, consistent with Baker’s law. On the other hand, introduced species were less likely to have the ability to reproduce asexually. Further, among species with no autonomous autogamy, pollen limitation was greater for introduced compared to native species. Such a result is consistent with the idea that plants entering a new continent receive lower quality or quantity of services from resident pollinators than species native to that continent. Finally, more phylogenetically novel invasive species had lower pollen limitation than less novel invasive species, potentially because they experience less competition for pollinators. This is the first evidence that enhanced pollination may be one mechanism driving the greater invasiveness of phylogenetically novel introduced species observed in some systems.     

Evolutionary increase in sexual and clonal reproductive capacity during biological invasion in an aquatic plant Butomus umbellatus (Butomaceae)

To test the hypothesis that increased allocation to reproduction is selected during biological invasion, we compared germination, survival, growth, and reproduction of native vs. introduced populations of the invasive aquatic plant Butomus umbellatus in a common greenhouse environment. Although seedling emergence and establishment did not differ consistently, survival thereafter was twice as high for eight introduced North American than eight native European populations. As predicted, introduced plants were more likely to produce sexual inflorescences and clonal asexual vegetative bulbils, and they invested much more biomass in both reproductive modes. Higher reproductive investment was due to higher proportional allocation of biomass rather than larger plant size. These results are consistent with selection for increased reproduction during range expansion. However, population genetic surveys indicate that recruitment from seed rarely occurs in introduced populations. Hence increased sexual allocation is not an adaptive response to invasion. Although increased clonal reproduction may be advantageous in expanding populations, genetic evidence from introduced populations of B. umbellatus suggests that increased clonal allocation may have arisen via stochastic processes during long-distance transport or a selective filter right at introduction, rather than incremental natural selection during range expansion.     

Native perennial grasses show evolutionary response to Bromus tectorum (cheatgrass) invasion

Invasive species can change selective pressures on native plants by altering biotic and abiotic conditions in invaded habitats. Although invasions can lead to native species extirpation, they may also induce rapid evolutionary changes in remnant native plants. We investigated whether adult plants of five native perennial grasses exhibited trait shifts consistent with evolution in response to invasion by the introduced annual grass Bromus tectorum L. (cheatgrass), and asked how much variation there was among species and populations in the ability to grow successfully with the invader. Three hundred and twenty adult plants were collected from invaded and uninvaded communities from four locations near Reno, Nevada, USA. Each plant was divided in two and transplanted into the greenhouse. One clone was grown with B. tectorum while the other was grown alone, and we measured tolerance (ability to maintain size) and the ability to reduce size of B. tectorum for each plant. Plants from invaded populations consistently had earlier phenology than those from uninvaded populations, and in two out of four sites, invaded populations were more tolerant of B. tectorum competition than uninvaded populations. Poa secunda and one population of E. multisetus had the strongest suppressive effect on B. tectorum, and these two species were the only ones that flowered in competition with B. tectorum. Our study indicates that response to B. tectorum is a function of both location and species identity, with some, but not all, populations of native grasses showing trait shifts consistent with evolution in response to B. tectorum invasion within the Great Basin.     

Are invaders moving targets? The generality and persistence of advantages in size, reproduction, and enemy release in invasive plant species with time since introduction

Successful plant invasions are often attributed to increased plant size, reproduction, or release from natural enemies, but the generality and persistence of these patterns remains widely debated. Meta-analysis was used to quantitatively assess invasive plant performance and release from enemy damage and how these change with residence time and geographic distribution. Invasive plants were compared either in their introduced and home ranges or with native congeners in the introduced range. Invasive plants in the introduced range were generally larger, allocated more to reproduction, and had lower levels of herbivore damage compared with conspecifics in the home range; pathogen attack, however, varied widely. In congener comparisons, invasive and native plants did not differ in size or herbivory, but invaders did allocate less to reproduction and had lower levels of pathogen damage. Time since introduction was a significant nonlinear predictor of enemy release for both herbivores and pathogens, with initial release in recently arrived species and little to no release after 50 to 200 years. Geographic distribution was also a significant nonlinear predictor of enemy release. The observed nonlinear relationships are consistent with dynamic invasions and may define targets for eradication efforts if these patterns hold up for individual species.     

Revealing constitutively expressed resistance genes in Agrostis species using PCR-based motif-directed RNA fingerprinting

Agrostis species are mainly used in athletic fields and golf courses. Their integrity is maintained by fungicides, which makes the development of disease-resistance varieties a high priority. However, there is a lack of knowledge about resistance (R) genes and their use for genetic improvement in Agrostis species. The objective of this study was to identify and clone constitutively expressed cDNAs encoding R gene-like (RGL) sequences from three Agrostis species (colonial bentgrass (A. capillaris L.), creeping bentgrass (A. stolonifera L.) and velvet bentgrass (A. canina L.)) by PCR-based motif-directed RNA fingerprinting towards relatively conserved nucleotide binding site (NBS) domains. Sixty-one constitutively expressed cDNA sequences were identified and characterized. Sequence analysis of ESTs and probable translation products revealed that RGLs are highly conserved among these three Agrostis species. Fifteen of them were shown to share conserved motifs found in other plant disease resistance genes such as MLA13, Xa1, YR6, YR23 and RPP5. The molecular evolutionary forces, analysed using the Ka/Ks ratio, reflected purifying selection both on NBS and leucine-rich repeat (LRR) intervening regions of discovered RGL sequences in these species. This study presents, for the first time, isolation and characterization of constitutively expressed RGL sequences from Agrostis species revealing the presence of TNL (TIR-NBS-LRR) type R genes in monocot plants. The characterized RGLs will further enhance knowledge on the molecular evolution of the R gene family in grasses.     

Allelopathy of a native grassland community as a potential mechanism of resistance against invasion by introduced plants

Successful plant invasions depend, at least partly, on interactions between introduced plants and native plant communities. While allelopathic effects of introduced invaders on native resident species have received much attention, the reverse, i.e. allelopathic effects of native residents on introduced plants, have been largely neglected. Therefore, we tested whether allelopathy of native plant communities decreases their invasibility to introduced plant species. In addition, we tested among the introduced species whether the invasive ones are more tolerant to allelopathy of native plant communities than the non-invasive ones. To test these hypotheses, we grew nine pairs of related (congeneric or confamilial) invasive and non-invasive introduced plant species (i.e. 18 species) in the presence or absence of a native grassland community, which consisted of three common forbs and three common grasses, with or without activated carbon in the soil. Activated carbon reduced the survival percentage and growth of introduced plants in the absence of the native plant community. However, its net effect on the introduced plants was neutral or even slightly positive in the presence of the native community. This might suggest that the native plant community imposed allelopathic effects on the introduced plants, and that these effects were neutralized or reduced by activated carbon. The invasive and non-invasive introduced plants, however, did not differ in their tolerance to such allelopathic effects of the native plant community. Thus, although allelopathy of native plant communities might increase their resistance against introduced plants, there was no evidence that tolerance to allelopathy of native plant communities contributes to the degree of invasiveness of introduced plants.