Comparisons of arthropod assemblages on an invasive and native trees: abundance,diversity and damage

The success of exotic plants may be due to lower herbivore loads than those on native plants (Enemies Release Hypothesis). Predictions of this hypothesis include lower herbivore abundances, diversity, and damage on introduced plant species compared to native ones. Greater density or diversity of predators and parasitoids on exotic versus native plants may also reduce regulation of exotic plants by herbivores. To test these predictions, we measured arthropod abundance, arthropod diversity, and foliar damage on invasive Chinese tallow tree (Triadica sebifera) and three native tree species: silver maple (Acer saccharinum), sycamore (Platanus occidentalis), and sweetgum (Liquidambar styraciflua). Arthropod samples were collected with canopy sweep nets from six 20 year old monoculture plots of each species at a southeast Texas site. A total of 2,700 individuals and 285 species of arthropods were caught. Overall, the species richness and abundance of arthropods on tallow tree were similar to the natives. But, ordination (NMS) showed community composition differed on tallow tree compared to all three native trees. It supported an arthropod community that had relatively lower herbivore abundance but relatively more predator species compared to the native species examined. Leaves were collected to determine damage. Tallow tree experienced less mining damage than native trees. The results of this study supported the Enemies Release Hypothesis predictions that tallow tree would have low herbivore loads which may contribute to its invasive success. Moreover, a shift in the arthropod community to fewer herbivores without a reduction in predators may further limit regulation of this exotic species by herbivores in its introduced range.     

Phylogenetic isolation increases plant success despite increasing susceptibility to generalist herbivores

Aim Theory suggests that introduced species that are phylogenetically distant from their recipient communities should be more successful than closely related introduced species because they can exploit open niches and escape enemies in their new range, i.e. Darwin’s Naturalization Hypothesis. Alternatively, it has also been hypothesized that closely related invaders might be more successful than novel invaders because they are pre‐adapted to conditions in their new range; a paradox coined Darwin’s Naturalization Conundrum. To date, these hypotheses have been tested primarily at the regional scale, not within local plant communities where introduced species colonize, compete and encounter herbivores. Location Global. Methods and Results We used community phylogenetics to analyse data from 49 published experiments to examine the importance of phylogenetic relatedness and generalist herbivory on native and exotic plant success at the community level. Plants that were categorized as ‘invasive’ were indeed less related to the recipient community than ‘non‐pest’ exotic plants. Distantly related exotic plants were also more abundant than closely related species. Phylogenetic relatedness predicted herbivore impact, but in a way that was opposite to predictions, as herbivores had stronger, not lesser, impacts on distantly related plants. Importantly, these same patterns generally held for native plants, as distantly related native plants were more abundant and more susceptible to herbivores than closely related species, ultimately resulting in herbivores suppressing community‐level phylogenetic diversity. Main conclusions Distantly related plants were more locally successful despite experiencing stronger control by generalist herbivores, a finding that was robust across native and exotic species. To our knowledge, this is the first evidence that phylogenetic matching influences the local success of both native and exotic species and that herbivores can influence community phylodiversity. Phylogenetic relatedness explained a relatively small portion of the variance in the data even after taking herbivory into account, however, suggesting that phylogenetic matching works in combination with other factors to influence community assembly.     

Proportion of exotics and relatedness of host species mediate the positive effect of plant richness on the species richness of fruit flies

1. All else being equal, the greater the local species richness of plants, the greater the number of associated herbivore species. Because most herbivore insects feed on a subset of closely related plant species, plant phylogenetic diversity is expected to play a key role in determining the number of herbivore species. What is not well known, however, is how an increase in the species richness of exotic plants affects the species richness of herbivores. 2. In this study, we used plant–fruit fly interactions to investigate the influence of the proportion and species richness of exotic host plants on the species richness of herbivorous insects. We also tested whether the phylogenetic diversity of host plants increases when the number of exotic plant species increases. 3. We found that the species richness of fruit flies is more accurately predicted by the richness of native host plants than by total plant species richness (including both native and exotic species). The proportion of exotic host species and the phylogenetic diversity of host plants had negative and positive effects, respectively, on the species richness of fruit flies. 4. Our findings suggest that a positive effect of plant richness on herbivore richness occurs only when an increase in plant diversity involves plant species with which native herbivores share some evolutionary history.     

Population,community and ecosystem effects of exotic herbivores: A growing global concern

Exotic herbivores represent a serious threat to native biodiversity, producing large scale changes in native communities and altering ecosystem processes. In this special issue, we present a series of case studies and reviews from different areas of the world that highlight (1) the consequences of herbivore introductions are a global problem; (2) they can result in wholesale shifts in the distribution of dominant plants on the landscape and; (3) the effects of herbivore introductions extend from the population to the community and ecosystem level. These studies suggest that introduced herbivores often retard ecosystem recovery after disturbance, facilitate invasion of plant species and can act as selective agents on native plant communities. These studies also suggest that several topics, including facilitation between exotic herbivores and exotic plants and animals (i.e., invasional meltdown) and the effect of exotic herbivores on ecosystem processes, require more research attention. Overall the papers in this special feature suggest that introduced herbivores are a global problem with wide-ranging ecological and evolutionary effects.     

Contrasting Effects of Land Use Intensity and Exotic Host Plants on the Specialization of Interactions in Plant-Herbivore Networks

Human land use tends to decrease the diversity of native plant species and facilitate the invasion and establishment of exotic ones. Such changes in land use and plant community composition usually have negative impacts on the assemblages of native herbivorous insects. Highly specialized herbivores are expected to be especially sensitive to land use intensification and the presence of exotic plant species because they are neither capable of consuming alternative plant species of the native flora nor exotic plant species. Therefore, higher levels of land use intensity might reduce the proportion of highly specialized herbivores, which ultimately would lead to changes in the specialization of interactions in plant-herbivore networks. This study investigates the community-wide effects of land use intensity on the degree of specialization of 72 plant-herbivore networks, including effects mediated by the increase in the proportion of exotic plant species. Contrary to our expectation, the net effect of land use intensity on network specialization was positive. However, this positive effect of land use intensity was partially canceled by an opposite effect of the proportion of exotic plant species on network specialization. When we analyzed networks composed exclusively of endophagous herbivores separately from those composed exclusively of exophagous herbivores, we found that only endophages showed a consistent change in network specialization at higher land use levels. Altogether, these results indicate that land use intensity is an important ecological driver of network specialization, by way of reducing the local host range of herbivore guilds with highly specialized feeding habits. However, because the effect of land use intensity is offset by an opposite effect owing to the proportion of exotic host species, the net effect of land use in a given herbivore assemblage will likely depend on the extent of the replacement of native host species with exotic ones.     

Plant‐mediated indirect effects and the persistence of parasitoid–herbivore communities

We have examined the effects of herbivore diversity on parasitoid community persistence and stability, mediated by nonspecific information from herbivore‐infested plants. First, we investigated host location and patch time allocation in the parasitoid Cotesia glomerata in environments where host and/or nonhost herbivores were present on Brassica oleracea leaves. Parasitoids were attracted by infochemicals from leaves containing nonhost herbivores. They spent considerable amounts of time on such leaves. Thus, when information from the plant is indistinct, herbivore diversity is likely to weaken interaction strengths between parasitoids and hosts. In four B. oleracea fields, all plants contained herbivores, often two or more species. We modelled parasitoid–herbivore communities increasing in complexity, based on our experiments and field data. Increasing herbivore diversity promoted the persistence of parasitoid communities. However, at a higher threshold of herbivore diversity, parasitoids became extinct due to insufficient parasitism rates. Thus, diversity can potentially drive both persistence and extinctions.     

Exotic weevil invasion increases floral herbivore community density,function, and impact on a native plant

Consumer communities are being re‐arranged through unprecedented rates of human‐mediated invasions and extinctions. Such changes in consumer diversity potentially alter community function and impact on resource populations. Although insect herbivore invasions are increasingly common, the influence of such species additions on native resident herbivore guilds, along with their individual and combined effects on native plant resources, are rarely investigated. Here, we used site‐to‐site and plant‐to‐plant variation in herbivore composition to examine how the addition of an invasive exotic weevil, Rhinocyllus conicus, combines with a guild of native floral herbivores (tephritid flies, pyralid moths) to influence two key components of herbivore community function – aggregate herbivore densities and cumulative levels of seed destruction – on a native thistle, Cirsium canescens. Invasion of a site by R. conicus more than doubled aggregate herbivore density, resulting in increased levels of seed destruction and a halving of seed production by the native thistle. Further, herbivore function was significantly higher on individual plants attacked by R. conicus, compared to plants attacked only by native herbivores. Insect densities and levels of seed destruction on plants attacked by multiple herbivore taxa never exceeded those observed for plants attacked by R. conicus alone, suggesting that increases in herbivore community function with invasion resulted from the inclusion of a functionally dominant insect rather than any complementarity effects. Some evidence for interference between insects emerged, with a trend towards reduced moth and weevil densities in two and three taxon mixtures compared with plants attacked by each taxon alone. However, density compensation was limited so that, overall, the addition of a novel herbivore to the floral guild was associated with a significant increase in herbivore community function and impact on seed production. The results suggest that invasion of a functionally dominant herbivore into an unsaturated recipient community can augment function within a resource guild.     

Relationship between land uses and diversity of dung beetles (Coleoptera: Scarabaeinae) in the southern Atlantic forest of Argentina: which are the key factors?

The loss of natural habitats is one of the main drivers of biodiversity decline. Anthropogenic land uses preserving biotic and abiotic conditions of the native ecosystem are more suitable to preserve the native biodiversity. In this study, we explored changes in species richness and composition in different land uses of the southern Atlantic forest, considering three independent factors: (1) canopy (presence–absence), (2) type of vegetation (native–exotic) and (3) livestock (presence–absence). We expected a gradient of response in the richness and composition of the native forest dung beetle community, from land uses preserving canopy and native vegetation to open land uses with exotic vegetation. Dung beetles were sampled in protected native forests and four land uses, using two potential food resources: human dung and carrion. The species richness and composition of each habitat, as well as differences in composition and the influence of factors over diversity, were then analyzed. As expected, our results showed that land uses preserving canopy and native vegetation maintain the dung beetle diversity of the native forest. Moreover, while the three factors analyzed influenced dung beetle diversity, canopy cover was the main driver of dung beetle diversity loss. The main conclusion of this study is that the conservation of canopy (either native or exotic) is determinant to preserve highly diverse dung beetle communities and subsequently, the ecological functions performed by this taxon. However, the ecophysiological mechanism behind the response of dung beetles to habitat disturbance is poorly understood.     

Fungal Phyllosphere Communities are Altered by Indirect Interactions Among Trophic Levels

Trophic interactions involving predators, herbivores, and plants have been described in terrestrial systems. However, there is almost no information on the effect of trophic interactions on microbial phyllosphere community abundance, diversity, or structure. In this study, the interaction between a parasitoid, an insect herbivore, and the fungal phyllosphere community is examined. Parasitoid wasps have an indirect negative impact on fungal community diversity. On the citrus phyllosphere, the exotic wasp species, Amitus hesperidum and Encarsia opulenta, may parasitize the citrus blackfly (Aleurocanthus woglumi). If parasitism levels are low, the blackfly may produce significant amounts of honeydew secretions on the surface of the leaf. Honeydew deposition provides a carbon-rich substrate for the development of fungal growth persisting as sooty mold on the leaves. Leaves from sooty mold-infested grapefruit (Citrus paradisi) trees were collected from multiple orchards in south Texas. The effect of different levels of exotic parasite activity, citrus blackfly, and sooty mold infestation on phyllosphere mycobiota community structure and diversity was examined. Our results suggest the presence of the parasitoid may lead to a top–down trophic cascade affecting phyllosphere fungal community diversity and structure. Additionally, persistent sooty mold deposits that have classically been referred to as Capnodium citri (and related asexual morphological forms) actually comprise a myriad of fungal species including many saprophytes and potential fruit and foliar pathogens of citrus.     

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

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

Evolutionary responses of native plants to novel community members

Both ecological and evolutionary processes can influence community assembly and stability, and native community members may respond both ecologically and evolutionarily as additional species enter established communities. Biological invasions provide a unique opportunity to examine these responses of native community members to novel species additions. Here, I use reciprocal transplant experiments among naturally invaded and uninvaded environments, along with experimental removals of exotic species, to determine whether exotic plant competitors and exotic insect herbivores evoke evolutionary changes in native plants. Specifically, I address whether the common native plant species Lotus wrangelianus has responded evolutionarily to a series of biological invasions by adapting to the presence of the exotic plant Medicago polymorpha and the exotic insect herbivore Hypera brunneipennis. Despite differences in selection regimes between invaded and uninvaded environments and the presence of genetic variation for traits relevant to the novel competitive and plant-herbivore interactions, these experiments failed to reveal evidence that Lotus has responded evolutionarily to the double invasion of Medicago followed by H. brunneipennis. However, when herbivory from H. brunneipennis was experimentally reduced, Lotus plants from source populations invaded by Medicago outperformed plants from uninvaded source populations when transplanted into heavily invaded destination environments. Therefore, Lotus showed evidence of adaptation to Medicago invasion but not to the newer invasion of an exotic shared herbivore. The presence of this exotic insect herbivore alters the outcome of evolutionary responses in this system and counteracts adaptation by the native Lotus to invasion by the exotic plant Medicago. This result has broad implications for the conservation of native communities. While native species may be able to adapt to the presence of one or a few exotics, a multitude of invasions may limit the ability of natives to respond evolutionarily to the novel and frequently changing selection pressures that arise with subsequent invasions.     

Coexistence between native and exotic species is facilitated by asymmetries in competitive ability and susceptibility to herbivores

Differences between native and exotic species in competitive ability and susceptibility to herbivores are hypothesized to facilitate coexistence. However, little fieldwork has been conducted to determine whether these differences are present in invaded communities. Here, we experimentally examined whether asymmetries exist between native and exotic plants in a community invaded for over 200 years and whether removing competitors or herbivores influences coexistence. We found that natives and exotics exhibit pronounced asymmetries, as exotics are competitively superior to natives, but are more significantly impacted by herbivores. We also found that herbivore removal mediated the outcome of competitive interactions and altered patterns of dominance across our field sites. Collectively, these findings suggest that asymmetric biotic interactions between native and exotic plants can help to facilitate coexistence in invaded communities.     

Fungal grass endophytes and arthropod communities: lessons from plant defence theory and multitrophic interactions

Alkaloids produced by systemic fungal endophytes of grasses are thought to act as defensive agents against herbivores. Endophytic alkaloids may reduce arthropod herbivore abundances and diversity in agronomic grasses. Yet, accumulating evidence, particularly from native grasses, shows that herbivore preference, abundances and species richness are sometimes greater on endophyte-infected plants, even those with high alkaloids, contrary to the notion of defensive mutualism. We argue that these conflicting results are entirely consistent with well-developed concepts of plant defence theory and tri-trophic interactions. Plant secondary chemicals and endophytic alkaloids often fail to protect plants because: (1) specialist herbivores evolve to detoxify and use defensive chemicals for growth and survival; and (2) natural enemies of herbivores may be more negatively affected by alkaloids than are herbivores. Endophytes and their alkaloids may have profound, but often highly variable, effects on communities, which are also consistent with existing theories of plant defence and community genetics.     

Exotic vertebrate and invertebrate herbivores differ in their impacts on native and exotic plants: a meta-analysis

Herbivores modify various ecological processes including interactions between native and exotic plants that may affect invasion success by the exotic plants. It is unknown whether different types of exotic herbivores have similar effects on native and exotic plants. Using two distinct data sets, we ran meta-analyses to compare exotic vertebrate and invertebrate herbivore preferences for, and effects on performance and population sizes of native and exotic plants. We found that exotic vertebrate herbivores have positive effects on exotic plant performance and population sizes, and no significant effects on native plants. Exotic invertebrates have significant negative effects on performance and population sizes of both exotic and native plants. Vertebrates prefer to feed on native plants relative to exotic plants, while invertebrates prefer the exotic plants to native plants. Thus the exotic vertebrate herbivores may aid invasiveness of exotic plants, in accordance with the invasional meltdown hypothesis, while exotic invertebrate herbivores probably have no net effect on invasion process of the exotic plants. Invertebrate herbivore preferences for exotic plants support the biotic resistance hypothesis, as the native plants probably resist the invertebrate herbivory. We also tested an evolutionary logic that posits that herbivores with similar evolutionary history as plants will affect the plants less negatively than plants with which they have not co-evolved. Our results indicate that there is no consistent pattern in effects of exotic vertebrate and invertebrate herbivores on exotic plants with or without which they have co-evolved.     

Long-Term Functional Dynamics of an Aphidophagous Coccinellid Community Remain Unchanged despite Repeated Invasions

Aphidophagous coccinellids (ladybeetles) are important providers of herbivore suppression ecosystem services. In the last 30 years, the invasion of exotic coccinellid species, coupled with observed declines in native species, has led to considerable interest in the community dynamics and ecosystem function of this guild. Here we examined a 24-year dataset of coccinellid communities in nine habitats in southwestern Michigan for changes in community function in response to invasion. Specifically we analyzed their temporal population dynamics and species diversity, and we modeled the community’s potential to suppress pests. Abundance of coccinellids varied widely between 1989 and 2012 and became increasingly exotic-dominated. More than 71% of 57,813 adult coccinellids captured over the 24-year study were exotic species. Shannon diversity increased slightly over time, but herbivore suppression potential of the community remained roughly constant over the course of the study. However, both Shannon diversity and herbivore suppression potential due to native species declined over time in all habitats. The relationship between Shannon diversity and herbivore suppression potential varied with habitat type: a positive relationship in forest and perennial habitats, but was uncorrelated in annual habitats. This trend may have been because annual habitats were dominated by a few, highly voracious exotic species. Our results indicated that although the composition of the coccinellid community in southwestern Michigan has changed dramatically in the past several decades, its function has remained relatively unchanged in both agricultural and natural habitats. While this is encouraging from the perspective of pest management, it should be noted that losses of one of the dominant exotic coccinellids could result in a rapid decline in pest suppression services if the remaining community is unable to respond.     

Role of dominant tree species on diversity of herbivorous insect community in temperate forests

Oak (Quercus spp.) and hornbeam (Carpinus spp.) are one of dominant tree species in East Asian temperate broad leaf deciduous forests and many insect species, including more than 65% of Lepidoptera species, feed on these trees. We sampled lepidopteran caterpillars from two 0.1 ha plots in a temperate forest to investigate the role of dominant trees (oaks and hornbeams) in herbivore community. In total, we identified 738 caterpillars from 223 Lepidopteran species on 34 tree species. Most caterpillar species were from species-rich families such as Geometridae 25% (56 spp.), Noctuidae, and Tortricidae. After excluding dominant trees, plant-herbivore network analyses showed increased network specialization and nestedness and decreased generality and vulnerability. These results suggest that oaks and hornbeams support a large Lepidopteran herbivore community, and co-occurring plant species support diverse but specific herbivores. Geographical distribution and plant community are closely related to diversity of the herbivore community. Future work is needed to investigate the likelihood that specialist herbivores become relatively more abundant in the forest as oaks are succeeded by hornbeams.     

Impacts of large herbivorous mammals on bird diversity and abundance in an African savanna

Large native mammals are declining dramatically in abundance across Africa, with strong impacts on both plant and animal community dynamics. However, the net effects of this large-scale loss in megafauna are poorly understood because responses by several ecologically important groups have not been assessed. We used a large-scale, replicated exclusion experiment in Kenya to investigate the impacts of different guilds of native and domestic large herbivores on the diversity and abundance of birds over a 2-year period. The exclusion of large herbivorous native mammals, including zebras (Equus burchelli), giraffes (Giraffa camelopardalis), elephants (Loxodonta africana), and buffalos (Syncerus caffer), increased the diversity of birds by 30%. Most of this effect was attributable to the absence of elephants and giraffes; these megaherbivores reduced both the canopy area of subdominant woody vegetation and the biomass of ground-dwelling arthropods, and both of these factors were good predictors of the diversity of birds. The canopy area of subdominant trees was positively correlated with the diversity of granivorous birds. The biomass of ground-dwelling arthropods was positively correlated with the diversity of insectivorous birds. Our results suggest that most native large herbivores are compatible with an abundant and diverse bird fauna, as are cattle if they are at a relatively low stocking rate. Future research should focus on determining the spatial arrangements and densities of megaherbivores that will optimize both megaherbivore abundance and bird diversity. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

With or without you: Effects of the concurrent range expansion of an herbivore and its natural enemy on native species interactions

Global climatic changes may lead to the arrival of multiple range‐expanding species from different trophic levels into new habitats, either simultaneously or in quick succession, potentially causing the introduction of manifold novel interactions into native food webs. Unraveling the complex biotic interactions between native and range‐expanding species is critical to understand the impact of climate change on community ecology, but experimental evidence is lacking. In a series of laboratory experiments that simulated direct and indirect species interactions, we investigated the effects of the concurrent arrival of a range‐expanding insect herbivore in Europe, Spodoptera littoralis, and its associated parasitoid Microplitis rufiventris, on the native herbivore Mamestra brassicae, and its associated parasitoid Microplitis mediator, when co‐occurring on a native plant, Brassica rapa. Overall, direct interactions between the herbivores were beneficial for the exotic herbivore (higher pupal weight than the native herbivore), and negative for the native herbivore (higher mortality than the exotic herbivore). At the third trophic level, both parasitoids were unable to parasitize the herbivore they did not coexist with, but the presence of the exotic parasitoid still negatively affected the native herbivore (increased mortality) and the native parasitoid (decreased parasitism rate), through failed parasitism attempts and interference effects. Our results suggest different interaction scenarios depending on whether S. littoralis and its parasitoid arrive to the native tritrophic system separately or concurrently, as the negative effects associated with the presence of the parasitoid were dependent on the presence of the exotic herbivore. These findings illustrate the complexity and interconnectedness of multitrophic changes resulting from concurrent species arrival to new environments, and the need for integrating the ecological effects of such arrivals into the general theoretical framework of global invasion patterns driven by climatic change.     

Trade‐off between early emergence and herbivore susceptibility mediates exotic success in an experimental California plant community

Ecological trade‐offs are fundamental to theory in community ecology; critical for understanding species coexistence in diverse plant communities, as well as the evolution of diverse life‐history strategies. Invasions by exotic species can provide insights into the importance of trade‐offs in community assembly, because the ecological strategies of invading species often differ from those present in the native species pool. Exotic annual species have invaded many Mediterranean‐climate areas around the globe, and often germinate and emerge earlier in the growing season than native species. Early‐season growth can enable exotic annual species to preempt space and resources, competitively suppressing later‐emerging native species; however, early‐emerging individuals may also be more apparent to herbivores. This suggests a potential trade‐off between seasonal phenology and susceptibility to herbivory. To evaluate this hypothesis, we monitored the emergence and growth of 12 focal species (six each native and exotic) in monoculture and polyculture, while experimentally excluding generalist herbivores both early and later in the growing season. Consistent with past studies, the exotic species emerged earlier than native species. Regardless of species origin, earlier‐emerging species achieved greater biomass by the end of the experiment, but were more negatively impacted by herbivory, particularly in the early part of the growing season. This greater impact of early‐season herbivory on early‐active species led to a reduction in the competitive advantage of exotic species growing in polyculture, and improved the performance of later‐emerging natives. Such a trade‐off between early growth and susceptibility to herbivores could be an important force in community assembly in seasonal herbaceous‐dominated ecosystems. These results also show how herbivore exclusion favors early‐active exotic species in this system, with important implications for management in many areas invaded by early‐active exotic species.     

An invasive plant provides refuge to native plant species in an intensely grazed ecosystem

Invasion by exotic plant species and herbivory can individually alter native plant species diversity, but their interactive effects in structuring native plant communities remain little studied. Many exotic plant species escape from their co-evolved specialized herbivores in their native range (in accordance with the enemy release hypothesis). When these invasive plants are relatively unpalatable, they may act as nurse plants by reducing herbivore damage on co-occurring native plants, thereby structuring native plant communities. However, the potential for unpalatable invasive plants to structure native plant communities has been little investigated. Here, we tested whether presence of an unpalatable exotic invader Opuntia ficus-indica was associated with the structure of native plant communities in an ecosystem with a long history of grazing by ungulate herbivores. Along 17 transects (each 1000 m long), we conducted a native vegetation survey in paired invaded and uninvaded plots. Plots that harboured O. ficus-indica had higher native plant species richness and Shannon–Wiener diversity H′ than uninvaded plots. However, mean species evenness J was similar between invaded and uninvaded plots. There was no significant correlation between native plant diversity and percentage plot cover by O. ficus-indica. Presence of O. ficus-indica was associated with a compositional change in native community assemblages between paired invaded and uninvaded plots. Although these results are only correlative, they suggest that unpalatable exotic plants may play an important ecological role as refugia for maintenance of native plant diversity in intensely grazed ecosystems.