Experimental evidence that the introduced fire ant, Solenopsis invicta, does not competitively suppress co-occurring ants in a disturbed habitat

1. The fire ant, Solenopsis invicta, is a globally distributed invasive ant that is largely restricted to disturbed habitats in its introduced range. For more than half a century, biologists have believed its success results from superior competitive abilities relative to native ant species, as well as an escape from their natural enemies. 2. We used large volumes of hot water to kill fire ant colonies, and only fire ant colonies, on experimental plots in pastures, and found that populations and diversity of co-occurring ants did not subsequently increase. 3. These results are contrary to classical predictions and indicate that S. invicta is not a superior competitor that suppresses native ants, and that the low diversity and abundance of native ants in degraded ecosystems does not result from interaction with fire ants. Instead, other factors such as prior disturbance and recruitment limitation may be the primary limiting factors for native species in these habitats.     

Conspecific plant–soil feedback scales with population size in Lobelia siphilitica (Lobeliaceae)

Plant–soil interactions directly affect plant success in terms of establishment, survival, growth and reproduction. Negative plant–soil feedback on such traits may therefore reduce the density and abundance of plants of a given species at a given site. Furthermore, if conspecific feedback varies among population sites, it could help explain geographic variation in plant population size. We tested for among-site variation in conspecific plant–soil feedback in a greenhouse experiment using seeds and soils from 8 natural populations of Lobelia siphilitica hosting 30–330 plants. The first cohort of seeds was grown on soil collected from each native site, while the second cohort was grown on the soil conditioned by the first. Our goal was to distinguish site-specific effects mediated by biotic and/or abiotic soil properties from those inherent in seed sources. Cohort 1 plants grown from seeds produced in small populations performed better in terms of germination, growth, and survival compared to plants produced in large populations. Plant performance decreased substantially between cohorts, indicating strong negative feedback. Most importantly, the strength of negative feedback scaled linearly (i.e., was less negative) with increasing size of the native plant population, particularly for germination and survival, and was better explained by soil- rather than seed-source effects. Even with a small number of sites, our results suggest that the potential for negative plant–soil feedback varies among populations of L. siphilitica, and that small populations were more susceptible to negative feedback. Conspecific plant–soil feedback may contribute to plant population size variation within a species’ native range.     

Measuring the effects of invasive species on the demography of a rare endemic plant

The impacts of invasive species are among the greatest threats to the persistence of native species and communities. Yet most work on rare plants has focused on issues such as habitat fragmentation and genetic diversity, while few studies have quantified the impacts of invasive plants on native ones or investigated the underlying mechanisms of those impacts. I used removal experiments to assess the effects of invasive grasses on the seedling and adult demography of an endangered California endemic, Oenothera deltoidesssp. howellii. Invasive plant removal significantly increased O. deltoides seedling recruitment, but had no effect on adult plants. Differences in seedling recruitment were primarily driven by greatly increased seedling emergence rates in removal plots, although there was also some evidence of higher seedling survivorship with invasives removal. Differences in habitat type strongly influenced both the effects of removal treatments and O. deltoides demography, with areas that support natural recruitment showing weaker treatment effects and higher overall recruitment, but lower adult survivorship, compared to those under restoration through planting. These results indicate that inhibition of germination due to reduced soil disturbance, rather than resource competition, appears to be the strongest impact of invasive plants on this rare endemic. Although previous work has documented the importance of changed disturbance regimes in generating invasion impacts, invader effects on rare plants are generally presumed to result from resource competition. Studies like this one highlight the need for a greater emphasis on understanding the mechanisms by which invasive plants impact native ones, and the importance of such information in designing conservation and management strategies.     

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.     

Release from natural enemies mitigates inbreeding depression in native and invasive Silene latifolia populations

Inbreeding and enemy infestation are common in plants and can synergistically reduce their performance. This inbreeding ×environment (I × E) interaction may be of particular importance for the success of plant invasions if introduced populations experience a release from attack by natural enemies relative to their native conspecifics. Here, we investigate whether inbreeding affects plant infestation damage, whether inbreeding depression in growth and reproduction is mitigated by enemy release, and whether this effect is more pronounced in invasive than native plant populations. We used the invader Silene latifolia and its natural enemies as a study system. We performed two generations of experimental out‐ and inbreeding within eight native (European) and eight invasive (North American) populations under controlled conditions using field‐collected seeds. Subsequently, we exposed the offspring to an enemy exclusion and inclusion treatment in a common garden in the species’ native range to assess the interactive effects of population origin (range), breeding treatment, and enemy treatment on infestation damage, growth, and reproduction. Inbreeding increased flower and leaf infestation damage in plants from both ranges, but had opposing effects on fruit damage in native versus invasive plants. Inbreeding significantly reduced plant fitness; whereby, inbreeding depression in fruit number was higher in enemy inclusions than exclusions. This effect was equally pronounced in populations from both distribution ranges. Moreover, the magnitude of inbreeding depression in fruit number was lower in invasive than native populations. These results support that inbreeding has the potential to reduce plant defenses in S. latifolia, which magnifies inbreeding depression in the presence of enemies. However, future studies are necessary to further explore whether enemy release in the invaded habitat has actually decreased inbreeding depression and thus facilitated the persistence of inbred founder populations and invasion success.     

Disturbance facilitates invasion: the effects are stronger abroad than at home

Disturbance is one of the most important factors promoting exotic invasion. However, if disturbance per se is sufficient to explain exotic success, then "invasion" abroad should not differ from "colonization" at home. Comparisons of the effects of disturbance on organisms in their native and introduced ranges are crucial to elucidate whether this is the case; however, such comparisons have not been conducted. We investigated the effects of disturbance on the success of Eurasian native Centaurea solstitialis in two invaded regions, California and Argentina, and one native region, Turkey, by conducting field experiments consisting of simulating different disturbances and adding locally collected C. solstitialis seeds. We also tested differences among C. solstitialis genotypes in these three regions and the effects of local soil microbes on C. solstitialis performance in greenhouse experiments. Disturbance increased C. solstitialis abundance and performance far more in nonnative ranges than in the native range, but C. solstitialis biomass and fecundity were similar among populations from all regions grown under common conditions. Eurasian soil microbes suppressed growth of C. solstitialis plants, while Californian and Argentinean soil biota did not. We suggest that escape from soil pathogens may contribute to the disproportionately powerful effect of disturbance in introduced regions.     

Biotic interactions and plant invasions

Introduced plant populations lose interactions with enemies, mutualists and competitors from their native ranges, and gain interactions with new species, under new abiotic conditions. From a biogeographical perspective, differences in the assemblage of interacting species, as well as in abiotic conditions, may explain the demographic success of the introduced plant populations relative to conspecifics in their native range. Within invaded communities, the new interactions and conditions experienced by the invader may influence both its demographic success and its effects on native biodiversity. Here, we examine indirect effects involving enemies, mutualists and competitors of introduced plants, and effects of abiotic conditions on biotic interactions. We then synthesize ideas building on Darwin's idea that the kinds of new interactions gained by an introduced population will depend on its relatedness to native populations. This yields a heuristic framework to explain how biotic interactions and abiotic conditions influence invader success. We conclude that species introductions generally alter plants' interactions with enemies, mutualists and competitors, and that there is increasing evidence that these altered interactions jointly influence the success of introduced populations.     

Shading by invasive shrub reduces seed production and pollinator services in a native herb

Plant invasions disrupt native plant reproduction directly via competition for light and other resources and indirectly via competition for pollination. Furthermore, shading by an invasive plant may reduce pollinator visitation and therefore reproduction in native plants. Our study quantifies and identifies mechanisms of these direct and indirect effects of an invasive shrub on pollination and reproductive success of a native herb. We measured pollinator visitation rate, pollen deposition, and female reproductive success in potted arrays of native Geranium maculatum in deciduous forest plots invaded by the non-native shrub Lonicera maackii and in two removal treatments: removal of aboveground L. maackii biomass and removal of flowers. We compared fruit and seed production between open-pollinated and pollen-supplemented plants to test for pollen and light limitation of reproduction. Plots with L. maackii had significantly lower light, pollinator visitation rate, and conspecific pollen deposition to G. maculatum than biomass removal plots. Lonicera maackii flower removal did not increase pollinator visitation or pollen deposition compared to unmanipulated invaded plots, refuting the hypothesis of competition for pollinators. Thus, pollinator-mediated impacts of invasive plants are not limited to periods of co-flowering or pollinator sharing between potential competitors. Geranium maculatum plants produced significantly fewer seeds in plots containing L. maackii than in plant removal plots. Seed set was similar between pollen-supplemented and open-pollinated plants, but pollen-supplemented plants exhibited higher seed set in plant removal plots compared to invaded plots. Therefore, we conclude that the mechanism of impact of L. maackii on G. maculatum reproduction was increased understory shade.     

Exotic cheatgrass and loss of soil biota decrease the performance of a native grass

Soil disturbances can alter microbial communities including arbuscular mycorrhizal (AM) fungi, which may in turn, affect plant community structure and the abundance of exotic species. We hypothesized that altered soil microbial populations owing to disturbance would contribute to invasion by cheatgrass (Bromus tectorum), an exotic annual grass, at the expense of the native perennial grass, squirreltail (Elymus elymoides). Using a greenhouse experiment, we compared the responses of conspecific and heterospecific pairs of cheatgrass and squirreltail inoculated with soil (including live AM spores and other organisms) collected from fuel treatments with high, intermediate and no disturbance (pile burns, mastication, and intact woodlands) and a sterile control. Cheatgrass growth was unaffected by type of soil inoculum, whereas squirreltail growth, reproduction and nutrient uptake were higher in plants inoculated with soil from mastication and undisturbed treatments compared to pile burns and sterile controls. Squirreltail shoot biomass was positively correlated with AM colonization when inoculated with mastication and undisturbed soils, but not when inoculated with pile burn soils. In contrast, cheatgrass shoot biomass was negatively correlated with AM colonization, but this effect was less pronounced with pile burn inoculum. Cheatgrass had higher foliar N and P when grown with squirreltail compared to a conspecific, while squirreltail had lower foliar P, AM colonization and flower production when grown with cheatgrass. These results indicate that changes in AM communities resulting from high disturbance may favor exotic plant species that do not depend on mycorrhizal fungi, over native species that depend on particular taxa of AM fungi for growth and reproduction.     

Plant–soil biota interactions and spatial distribution of black cherry in its native and invasive ranges

One explanation for the higher abundance of invasive species in their non‐native than native ranges is the escape from natural enemies. But there are few experimental studies comparing the parallel impact of enemies (or competitors and mutualists) on a plant species in its native and invaded ranges, and release from soil pathogens has been rarely investigated. Here we present evidence showing that the invasion of black cherry (Prunus serotina) into north‐western Europe is facilitated by the soil community. In the native range in the USA, the soil community that develops near black cherry inhibits the establishment of neighbouring conspecifics and reduces seedling performance in the greenhouse. In contrast, in the non‐native range, black cherry readily establishes in close proximity to conspecifics, and the soil community enhances the growth of its seedlings. Understanding the effects of soil organisms on plant abundance will improve our ability to predict and counteract plant invasions.     

Prey naïveté rather than enemy release dominates the relation of an invasive spider toward a native predator

Ecosystems may suffer from the impact of invasive species. Thus, understanding the mechanisms contributing to successful invasions is fundamental for limiting the effects of invasive species. Most intuitive, the enemy release hypothesis predicts that invasive species might be more successful in the exotic range than resident sympatric species owing to the absence of coevolution with native enemies. Here, we test the enemy release hypothesis for the invasion of Europe by the North American spider Mermessus trilobatus. We compare the susceptibility of invasive Mermessus trilobatus and a native species with similar life history to a shared predator with which both species commonly co‐occur in Europe. Contrary to our expectations, invasive Mermessus trilobatus were consumed three times more frequently by native predators than their native counterparts. Our study shows that invasive Mermessus trilobatus is more sensitive to a dominant native predator than local sympatric species. This suggests that the relation between the invasive spider and its native predator is dominated by prey naïveté rather than enemy release. Further studies investigating evolutionary and ecological processes behind the invasion success of Mermessus trilobatus, including testing natural parasites and rapid reproduction, are needed to explain its invasion success in Europe.     

永州广聚萤叶甲和豚草卷蛾的种群动态及对豚草的控制效果

【目的】豚草是一种重要的入侵杂草,严重危害农业生产和人类健康。广聚萤叶甲和豚草卷蛾是豚草的专一性天敌。研究这2种天敌在永州的种群动态及其对豚草的控制效果,可以为永州豚草的防控及这2种天敌的有效利用提供依据。【方法】在湖南省永州市江永县豚草大面积发生区域,人工释放广聚萤叶甲和豚草卷蛾,调查这2种天敌在释放区和扩散区的种群动态和对豚草的防治效果,以及这2种天敌在扩散区的越冬情况。【结果】广聚萤叶甲和豚草卷蛾的扩散能力强。释放1个月后,在释放区,广聚萤叶甲各虫态及豚草卷蛾虫瘿均被发现。整体上,随时间延长,广聚萤叶甲各虫态虫口密度先增后减,而豚草卷蛾虫瘿密度呈逐渐降低趋势。释放2个月后,在距释放区边缘10 km的豚草发生区,发现了广聚萤叶甲和豚草卷蛾,且成功建立了种群并顺利越冬。释放区豚草株高几乎没有增加,且叶片最终被取食精光,几乎全部死亡;扩散区豚草株高略有增加,最终近75%叶片被取食。【结论】广聚萤叶甲和豚草卷蛾可在永州成功建立种群并安全越冬,还能自行扩散,可持续控制野外豚草的发生。     

Developmental plasticity and reduced susceptibility to natural enemies following host plant defoliation in a specialized herbivore

Host-specific phytophagous insects that are short lived and reliant on ephemeral plant tissues provide an excellent system in which to investigate the consequences of disruption in the timing of resource availability on consumer populations and their subsequent interactions with higher tropic levels. The specialist herbivore, Belonocnema treatae (Hymenoptera: Cynipidae) induces galls on only newly flushed leaves of live oak, Quercus fusiformis. In central Texas (USA) episodic defoliation of the host creates variation in the timing of resource availability and results in heterogeneous populations of B. treatae that initiate development at different times. We manipulated the timing of leaf flush in live oak via artificial defoliation to test the hypothesis that a 6- to 8-week delay in the availability of resources alters the timing of this gall former’s life cycle events, performance and survivorship on its host, and susceptibility to natural enemies. B. treatae exhibits plasticity in development time, as the interval from egg to emergence was significantly reduced when gallers oviposited into the delayed leaf flush. As a consequence, the phenologies of gall maturation and adult emergence remain synchronized in spite of variation in the timing of resource availability. Per capita gall production and gall-former performance are not significantly affected by the timing of resource availability. The timing of resource availability and natural enemies interact, however, to produce strong effects on survivorship: when exposed to natural enemies, B. treatae developing in galls initiated by delayed oviposition exhibited an order-of-magnitude increase in survivorship. Developmental plasticity allows this gall former to circumvent disruptions in resource availability, maintain synchrony of life cycle events, and results in reduced vulnerability to natural enemies following defoliation of the host plant.     

Evolutionary history and distance dependence control survival of dipterocarp seedlings

One important hypothesis to explain tree-species coexistence in tropical forests suggests that increased attack by natural enemies near conspecific trees gives locally rare species a competitive advantage. Host ranges of natural enemies generally encompass several closely related plant taxa suggesting that seedlings should also do poorly around adults of closely related species. We investigated the effects of adult Parashorea malaanonan on seedling survival in a Bornean rain forest. Survival of P. malaanonan seedlings was highest at intermediate distances from parent trees while heterospecific seedlings were unaffected by distance. Leaf herbivores did not drive this relationship. Survival of seedlings was lowest for P. malaanonan , and increased with phylogenetic dissimilarity from this species, suggesting that survival of close relatives of common species is reduced. This study suggests that distance dependence contributes to species coexistence and highlights the need for further investigation into the role of shared plant enemies in community dynamics.
Ecology Letters (2010) 13: 51–59     

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.     

Soil biota and invasive plants

Interactions between plants and soil biota resist invasion by some nonnative plants and facilitate others. In this review, we organize research and ideas about the role of soil biota as drivers of invasion by nonnative plants and how soil biota may fit into hypotheses proposed for invasive success. For example, some invasive species benefit from being introduced into regions of the world where they encounter fewer soil-borne enemies than in their native ranges. Other invasives encounter novel but strong soil mutualists which enhance their invasive success. Leaving below-ground natural enemies behind or encountering strong mutualists can enhance invasions, but indigenous enemies in soils or the absence of key soil mutualists can help native communities resist invasions. Furthermore, inhibitory and beneficial effects of soil biota on plants can accelerate or decelerate over time depending on the net effect of accumulating pathogenic and mutualistic soil organisms. These 'feedback' relationships may alter plant-soil biota interactions in ways that may facilitate invasion and inhibit re-establishment by native species. Although soil biota affect nonnative plant invasions in many different ways, research on the topic is broadening our understanding of why invasive plants can be so astoundingly successful and expanding our perspectives on the drivers of natural community organization.     

Herbarium specimens reveal a historical shift in phylogeographic structure of common ragweed during native range disturbance

Invasive plants provide ample opportunity to study evolutionary shifts that occur after introduction to novel environments. However, although genetic characters pre‐dating introduction can be important determinants of later success, large‐scale investigations of historical genetic structure have not been feasible. Common ragweed (Ambrosia artemisiifolia L.) is an invasive weed native to North America that is known for its allergenic pollen. Palynological records from sediment cores indicate that this species was uncommon before European colonization of North America, and ragweed populations expanded rapidly as settlers deforested the landscape on a massive scale, later becoming an aggressive invasive with populations established globally. Towards a direct comparison of genetic structure now and during intense anthropogenic disturbance of the late 19th century, we sampled 45 natural populations of common ragweed across its native range as well as historical herbarium specimens collected up to 140 years ago. Bayesian clustering analyses of 453 modern and 473 historical samples genotyped at three chloroplast spacer regions and six nuclear microsatellite loci reveal that historical ragweed's spatial genetic structure mirrors both the palaeo‐record of Ambrosia pollen deposition and the historical pattern of agricultural density across the landscape. Furthermore, for unknown reasons, this spatial genetic pattern has changed substantially in the intervening years. Following on previous work relating morphology and genetic expression between plants collected from eastern North America and Western Europe, we speculate that the cluster associated with humans’ rapid transformation of the landscape is a likely source of these aggressive invasive populations.     

Association with mature plants protects seedlings from predation in an arid grassland shrub,Gutierrezia microcephala

Summary Survivorship of Gutierrezia microcephala (Compositae) seedlings was studied in an undisturbed arid grassland and in experimental plots where various components of the natural vegetation were removed following seed germination. The major causes of seedling mortality were herbivore damage from the specialist grasshopper, Hesperotettix viridis, and drought stress associated with competition from established plants. The relative intensity of these mortality factors varied strongly with seedling size. Large seedlings had higher overall survivorship but were most likely to be killed by defoliation; most small seedlings died of drought stress.In plots where all perennial grasses were removed (leaving established G. microcephala plants), seedling survivorship was 5 times greater than in undisturbed vegetation. Surviorship in plots where both grasses and mature G. microcephala were removed was slightly poorer than in undisturbed vegetation due to a large increase in mortality from defoliation. From May–August, when the herbivore H. viridis was abundant, seedling survivorship was better in the immediate vicinity of mature conspecific plants than in plots lacking mature G. microcephala, both in the presence and absence of perennial grasses. These results provide a counter example to theories predicting that the impact of specialist herbivores on seedling recruitment is greatest in the vicinity of parent plants.     

Emulating Natural Disturbances for Declining Late-Successional Species: A Case Study of the Consequences for Cerulean Warblers (Setophaga cerulea)

Forest cover in the eastern United States has increased over the past century and while some late-successional species have benefited from this process as expected, others have experienced population declines. These declines may be in part related to contemporary reductions in small-scale forest interior disturbances such as fire, windthrow, and treefalls. To mitigate the negative impacts of disturbance alteration and suppression on some late-successional species, strategies that emulate natural disturbance regimes are often advocated, but large-scale evaluations of these practices are rare. Here, we assessed the consequences of experimental disturbance (using partial timber harvest) on a severely declining late-successional species, the cerulean warbler (Setophaga cerulea), across the core of its breeding range in the Appalachian Mountains. We measured numerical (density), physiological (body condition), and demographic (age structure and reproduction) responses to three levels of disturbance and explored the potential impacts of disturbance on source-sink dynamics. Breeding densities of warblers increased one to four years after all canopy disturbances (vs. controls) and males occupying territories on treatment plots were in better condition than those on control plots. However, these beneficial effects of disturbance did not correspond to improvements in reproduction; nest success was lower on all treatment plots than on control plots in the southern region and marginally lower on light disturbance plots in the northern region. Our data suggest that only habitats in the southern region acted as sources, and interior disturbances in this region have the potential to create ecological traps at a local scale, but sources when viewed at broader scales. Thus, cerulean warblers would likely benefit from management that strikes a landscape-level balance between emulating natural disturbances in order to attract individuals into areas where current structure is inappropriate, and limiting anthropogenic disturbance in forests that already possess appropriate structural attributes in order to maintain maximum productivity.     

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

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