Dominant species identity, not community evenness, regulates invasion in experimental grassland plant communities

While there has been extensive interest in understanding the relationship between diversity and invasibility of communities, most studies have only focused on one component of diversity: species richness. Although the number of species can affect community invasibility, other aspects of diversity, including species identity and community evenness, may be equally important. While several field studies have examined how invasibility varies with diversity by manipulating species identity or evenness, the results are often confounded by resource heterogeneity, site history, or disturbance. We designed a mesocosm experiment to examine explicitly the role of dominant species identity and evenness on the invasibility of grassland plant communities. We found that the identity of the dominant plant species, but not community evenness, significantly impacted invasibility. Using path analysis, we found that community composition (dominant species identity) reduced invasion by reducing early-season light availability and increasing late-season plant community biomass. Nitrogen availability was an important factor for the survival of invaders in the second year of the experiment. We also found significant direct effects of certain dominant species on invasion, although the mechanisms driving these effects remain unclear. The magnitude of dominant species effects on invasibility we observed are comparable to species richness effects observed in other studies, showing that species composition and dominant species can have strong effects on the invasibility of a community.     

Population-level compensation impedes biological control of an invasive forb and indirect release of a native grass

The intentional introduction of specialist insect herbivores for biological control of exotic weeds provides ideal but understudied systems for evaluating important ecological concepts related to top-down control, plant compensatory responses, indirect effects, and the influence of environmental context on these processes. Centaurea stoebe (spotted knapweed) is a notorious rangeland weed that exhibited regional declines in the early 2000s, attributed to drought by some and to successful biocontrol by others. We initiated an experiment to quantify the effects of the biocontrol agent, Cyphocleonus achates, on Ce. stoebe and its interaction with a dominant native grass competitor, Pseudoroegneria spicata, under contrasting precipitation conditions. Plots containing monocultures of each plant species or equal mixtures of the two received factorial combinations of Cy. achates herbivory (exclusion or addition) and precipitation (May-June drought or "normal," defined by the 50-year average) for three years. Cy. achates herbivory reduced survival of adult Ce. stoebe plants by 9% overall, but this effect was stronger under normal precipitation compared to drought conditions, and stronger in mixed-species plots compared to monocultures. Herbivory had no effect on Ce. stoebe per capita seed production or on recruitment of seedlings or juveniles. In normal-precipitation plots of mixed composition, greater adult mortality due to Cy. achates herbivory resulted in increased recruitment of new adult Ce. stoebe. Due to this compensatory response to adult mortality, final Ce. stoebe densities did not differ between herbivory treatments regardless of context. Experimental drought reduced adult Ce. stoebe survival in mixed-species plots but did not impede recruitment of new adults or reduce final Ce. stoebe densities, perhaps due to the limited duration of the treatment. Ce. stoebe strongly depressed P. spicata reproduction and recruitment, but these impacts were not substantively alleviated by herbivory on Ce. stoebe. Population-level compensation by dominant plants may be an important factor inhibiting top-down effects in herbivore-driven and predator-driven cascades.     

Habitat-specific impacts of multiple consumers on plant population dynamics

Multiple consumers often attack seeds, seedlings, and adult plants, but their population-level consequences remain uncertain. We examined how insect and small mammal consumers influenced the demography and abundance of the perennial shrub, bush lupine (Lupinus arboreus). In grassland and dune habitats we established replicate experimental lupine populations in 81-m2 plots that were either protected from, or exposed to, herbivorous voles and granivorous mice (via fencing) and/or root feeding insects (via insecticide treatment). Populations were initiated with transplanted seedlings in 1999 and 2000. We followed the demography of these cohorts, subsequent generations, and the seed bank for 5.5 years. Voles and insects killed many seedlings in dune (1999 only) and grassland (1999 and 2000) habitats. After 2000, insects and voles had minimal effects on seedling or adult survival. Seed predation by granivorous mice, however, greatly depressed seedling recruitment, resulting in lower adult lupine abundance in control plots vs. those protected from rodents. In grasslands, initial effects of voles and insects on seedling survival produced large differences among treatments in adult plant density and the cumulative number of seeds produced throughout the experiment. Differences among grassland populations in seed rain, however, had little influence on the magnitude of seedling recruitment into this habitat. Instead, recruitment out of a preexisting seed bank compensated for the lack of seed production in populations exposed to consumers. Shading by dense adults in plots protected from consumers limited seedling establishment within these populations. Although differences among populations in cumulative seed rain did not influence adult establishment, populations protected from consumers accumulated substantially larger seed banks than controls. These results illustrate how density dependence, habitat-specific seed dynamics, and particular demographic impacts of consumers interact to shape plant population responses to consumers.     

Disturbance, Diversity, and Invasion: Implications for Conservation

Disturbance is an important component of many ecosystems, and variations in disturbance regime can affect ecosystem and community structure and functioning. The "intermediate disturbance hypothesis" suggests that species diversity should be highest at moderate levels of disturbance. However, disturbance is also known to increase the invasibility of communities. Disturbance therefore poses an important problem for conservation management, Here, we review the effects of disturbances such as fire grazing, soil disturbance and nutrient addition on plant species diversity and invasion with particular emphasis on grassland vegetation. Individual components of the disturbance regime can have marked effects on species diversity, but it is often modifications of the existing regime that have the largest influence. Similarly, disturbance can enhance invasion of natural communities, but frequently it is the interaction between different disturbances that has the largest effect. The natural disturbance regime is now unlikely to persist within conservation areas since fragmentation and human intervention have usually modified physical and biotic conditions. Active management decisions must now be made on what disturbance regime is required and this requires decisions on what species are to be encouraged or discouraged.     

Spatial and temporal patterns of seed dispersal: an important determinant of grassland invasion.

We measured spatial and temporal patterns of seed dispersal and seedling recruitment for 58 species in a grassland community to test whether seed dispersal could predict patterns of invasion after disturbance. For the 12 most abundant grasses, recruitment of native species was dependent on the propagule supply of both native and exotic species. Variability in seed rain on small spatial (1-10 m) and temporal (within season) scales led to qualitative differences in the outcome of disturbance colonization such that native species dominated disturbances when exotic seed supply was low but failed to establish when exotic seed supply was high. Local dispersal and spatial heterogeneity in species composition promoted coexistence of native and exotic species by creating refuges from high exotic seed supply within native dominated patches. Despite this, copious exotic seed production strongly limited recruitment of native species in exotic dominated patches. Most grasslands in California are presently dominated by exotic species, suggesting that competition at the seedling stage is a major barrier to native species restoration.     

Abundance declines of a native forb have nonlinear impacts on grassland invasion resistance

The effects of declining plant biodiversity on ecosystem processes are well studied, with most investigations examining the role of species richness declines rather than declines of species abundance. Using grassland mesocosms, we examined how the abundance of a native, resident species, Hemizonia congesta (hayfield tarweed), affected exotic Centaurea solstitialis (yellow starthistle) invasion. We found that progressive H. congesta abundance declines had threshold effects on invasion resistance, with initial declines resulting in minor increases in invasion and subsequent declines leading to accelerating increases in invader performance. Reduced invasion resistance was explained by increased resource availability as H. congesta declined. We also found evidence that resident abundance might indirectly affect invasion by mediating invader impact on resident competitors; C. solstitialis disproportionately reduced H. congesta biomass in low-abundance rather than high-abundance populations. H. congesta's direct and indirect effects on invasion resistance illustrate that an individual species' declining abundance can have accelerating, deleterious effects on ecosystem functions of conservation value.     

Invasibility and Effects of Amur Honeysuckle in Southwestern Ohio Forests

The Asian exotic Amur honeysuckle ( Lonicera maackii [Rupr.] Herder) has become the dominant shrub in many forests in southwestern Ohio and in some other locations in the eastern United States. Our research focused on the invasibility of forest communities and relationships of L. maackii to the abundance of tree seedlings and herbs. We surveyed 93 forest stands near Oxford, Ohio (USA) to determine L. maackii cover, time since invasion, tree canopy cover, tree basal area, and a shade tolerance index. Stepwise multiple regression indicated that greater than one-half of the variation (r ² = 0.550) in Lonicera cover was correlated to five variables (in descending order of importance): tree canopy cover, distance from Oxford, shade tolerance index, tree basal area, and time since invasion. The results suggest that high light levels and proximity to an abundant seed source increase forest invasibility. Tree seedling density, species richness of seedlings, and herb cover were all inversely related to L. maackii cover. When Lonicera becomes abundant, future structure and composition of forests could be affected and local populations of herbs threatened.     

The importance of quantifying propagule pressure to understand invasion: an examination of riparian forest invasibility

The widely held belief that riparian communities are highly invasible to exotic plants is based primarily on comparisons of the extent of invasion in riparian and upland communities. However, because differences in the extent of invasion may simply result from variation in propagule supply among recipient environments, true comparisons of invasibility require that both invasion success and propagule pressure are quantified. In this study, we quantified propagule pressure in order to compare the invasibility of riparian and upland forests and assess the accuracy of using a community's level of invasion as a surrogate for its invasibility. We found the extent of invasion to be a poor proxy for invasibility. The higher level of invasion in the studied riparian forests resulted from greater propagule availability rather than higher invasibility. Furthermore, failure to account for propagule pressure may confound our understanding of general invasion theories. Ecological theory suggests that species-rich communities should be less invasible. However, we found significant relationships between species diversity and invasion extent, but no diversity-invasibility relationship was detected for any species. Our results demonstrate that using a community's level of invasion as a surrogate for its invasibility can confound our understanding of invasibility and its determinants.     

An Integrated Approach to the Ecology and Management of Plant Invasions

Plant invasions are a serious threat to natural and managed ecosystems worldwide. The number of species involved and the extent of existing invasions renders the problem virtually intractable, and it is likely to worsen as more species are introduced to new habitats and more existing invaders move into a phase of rapid spread. We contend that current research and management approaches are inadequate to tackle the problem. The current focus is mostly on the characteristics and control of individual invading species. Much can be gained, however, by considering other important components of the invasion problem. Patterns of weed spread indicate that many species have a long lag phase following introduction before they spread explosively. Early detection and treatment of invasions before explosive spread occurs will prevent many future problems. Similarly, a focus on the invaded ecosystem and its management, rather than on the invader, is likely to be more effective. Identification of the causal factors enhancing ecosystem invasibility should lead to more-effective integrated control programs. An assessment of the value of particular sites and their degree of disturbance would allow the setting of management priorities for protection and control. Socioeconomic factors frequently play a larger part than ecological factors in plant invasions. Changes in human activities in terms of plant introduction and use, land use, and timing of control measures are all required before the plant invasion problem can be tackled adequately. Dealing with plant invasions is an urgent task that will require difficult decisions about land use and management priorities. These decisions have to be made if we want to conserve biodiversity worldwide.     

Seaweed richness and herbivory increase rate of community recovery from disturbance

The importance of herbivores and of plant diversity for community succession and recovery from disturbance is well documented. However, few studies have assessed the relative magnitude of, or potential interactions between, these factors. To determine the combined effect of herbivory and surrounding algal species richness on the recovery of a rocky intertidal community, we conducted a 27-month field experiment assessing algal recruitment and succession in cleared patches that mimic naturally forming gaps in the ambient community. We crossed two herbivore treatments, ambient and reduced abundance, with monocultures and polycultures of the four most common algal species in a mid-high rocky intertidal zone of northern California. We found that both the presence of herbivores and high surrounding algal richness increased recovery rates, and the effect of algal richness was twice the magnitude of that of herbivores. The increased recovery rate of patches containing herbivores was due to the consumption of fast-growing, early colonist species that preempt space from perennial, late-successional species. Mechanisms linking algal richness and recovery are more numerous. In polycultures, herbivore abundance and species composition is altered, desiccation rates are lower, and propagule recruitment, survival, and growth are higher compared to monocultures, all of which could contribute the observed effect of surrounding species richness. Herbivory and species richness should jointly accelerate recovery wherever palatable species inhibit late-successional, herbivore-resistant species and recruitment and survival of new colonists is promoted by local species richness. These appear to be common features of rocky-shore seaweed, and perhaps other, communities.     

Terrestrial vertebrates alter seedling composition and richness but not diversity in an Australian tropical rain forest

Although birds and mammals play important roles in several mechanisms hypothesized to maintain plant diversity in species-rich habitats, there have been few long-term, community-level tests of their importance. We excluded terrestrial birds and mammals from fourteen 6 x 7.5 m plots in Australian primary tropical rain forest and compared recruitment and survival of tree seedlings annually over the subsequent seven years to that on nearby open plots. We re-censused a subset of the plots after 13 years of vertebrate exclusion to test for longer-term effects. After two years of exclusion, seedling abundance was significantly higher (74%) on exclosure plots and remained so at each subsequent census. Richness was significantly higher on exclosure plots from 1998 to 2003, but in 2009 richness no longer differed, and rarefied species richness was higher in the presence of vertebrates. Shannon's diversity and Pielou's evenness did not differ in any year. Vertebrates marginally increased density-dependent mortality and recruitment limitation, but neither effect was great enough to increase richness or diversity on open plots relative to exclosure plots. Terrestrial vertebrates significantly altered seedling community composition, having particularly strong impacts on members of the Lauraceae. Overall, our results highlight that interactions between terrestrial vertebrates and tropical tree recruitment may not translate into strong community-level effects on diversity, especially over the short-term, despite significant impacts on individual species that result in altered species composition.     

Inversion of plant dominance-diversity relationships along a latitudinal stress gradient

Species interactions affect plant diversity through the net effects of competition and facilitation, with the latter more prevalent in physically stressful environments when plant cover ameliorates abiotic stress. One explanation for species loss in invader-dominated systems is a shift in the competition-facilitation balance, with competition intensifying in areas formerly structured by facilitation. We test this possibility with a 10-site prairie meta-experiment along a 500-km latitudinal stress gradient, quantifying the relationships among abiotic stress, exotic dominance, and native plant recruitment over five years. The latitudinal gradient is inversely correlated with abiotic stress, with lower latitudes more moisture- and nutrient-limited. We observed strong negative effects by invasive dominant grasses on plant establishment, but only in northern sites with lower-stress environments. At these locations, disturbance was critical for recruitment by reducing the suppressive dominant (invasive) canopy. In more stressful environments to the south, the impacts of the dominant invaders on plant establishment became facilitative, and diversity was more limited by seed availability. Disturbance prevented recruitment because seedling survival depended on a protective plant canopy, presumably because the canopy reduced temperature or moisture stress. Seed limitation was similarly prevalent in all sites. Our work confirms the importance of facilitation as an organizing process for plants in higher-stress environments, even with transformations of species composition and dominance. It also demonstrates that the mechanisms regulating diversity, including invader impacts, can vary within the same plant community depending on environmental context. Because limits on native plant recruitment are environmentally contingent, management strategies that seek to increase diversity, including invader eradication, must account for site-level variations in the balance between biotic and abiotic constraints.     

Herbivory limits recruitment in an old-field seed addition experiment

Environmental variability can promote coexistence by creating establishment sites for rare plants, but low diversity in anthropogenic grasslands suggests that this variability may be eliminated (homogenization hypothesis) or inaccessible (barrier hypothesis). We explore these alternatives on the northern Great Plains, where 11 million hectares have been transformed by multiple environmental changes, but the causes of species loss are unclear. In a degraded grassland, we increased environmental variability by manipulating competition and herbivory along gradients of fertility and disturbance, and we circumvented dispersal barriers by adding 1.2 million seeds of five functionally distinct species at varying densities. The experiment ended after 12 weeks due to the direct and indirect effects of unapparent small native herbivores, which were barriers to population establishment by the added species. The direct cause of recruitment failure was browsing. The indirect cause was associated with competition from invasive plants that appeared to be more tolerant or resistant to herbivory. Variability in fertility, disturbance, propagule pressure, and competition had relatively minor impacts on colonization by the added species because herbivores controlled recruitment in most environments. Recruitment outside the herbivore exclosures was mostly by unpalatable exotics, suggesting a possible link between invasion success and herbivore resistance for some introduced plants.     

Community assembly in experimental grasslands: suitable environment or timely arrival?

It is hard to defend the view that biotic communities represent a simple and predictable response to the abiotic environment. Biota and the abiotic environment interact, and the environment of an individual certainly includes its neighbors and visitors in the community. The complexity of community assembly calls forth a quest for general principles, yet current results and theories on assembly rules differ widely. Using a grassland microcosm as a model system, we manipulated fertility, disturbance by defoliation, soil/microclimate, and arrival order of species belonging to two groups differing in functional attributes. We analyzed the outcome of community assembly dynamics in terms of species richness, invasibility, and species composition. The analyses revealed strong environmental control over species richness and invasibility. Species composition was mainly determined by the arrival order of species, indicating that historical contingency may change the outcome of community assembly. The probability for multiple equilibria appeared to increase with productivity and environmental stability. The importance of arrival order offers an explanation of the difficulties in predicting local occurrences of species in the field. In our experiment, variation in fertility and disturbance was controlling colonization with predictable effects on emergent community properties such as species richness. The key mechanism is suggested to be asymmetric competition, and our results show that this mechanism is relatively insensitive to the species through which it works. While our analyses indicate a positive and significant correlation between richness and invasibility, the significance disappears after accounting for the effect of the environment. The importance of arrival order (historical contingency) and environmental control supports the assumption of the unified neutral theory that different species within a trophic level can be considered functionally equivalent when it comes to community assembly. However, our results indicate that variation in asymmetric competition is the key factor determining the richness of the resulting communities, and this is far from neutral.     

三江源区不同建植年代人工草地群落演替与土壤养分变化

研究了了三源区不同建植期人工修复草地在不同演替阶段毒杂草[主要是甘肃马先蒿(Pedicularis kansuensis)]的入侵规律、数量特征,植物群落物种组成、生物苗和草地质最以及土壤养分、微生物活性的变化规律.结果表明,不同建植期人工修复草地植物群落的种类组成、植物功能群组成和群落数量特征存在显著差异.随着演替时间的推移,人工草地群落盖度、高度、物种数、生物最和多样性指数均表现出"V"字型变化规律,杂类草--甘肃马先蒿的数量特征变化尤为明显,在4 a的人工草地群落中开始局部入侵,在5～6 a的人工草地群落中大面积入侵,其入侵速度、入侵面积达到高峰期.土壤的含水量、容重、土壤中有机质、氮素和磷素在演替过程(7 a、9 a草地)中逐渐降低,到一定时期又逐步增加;随着演替的进行,不同建植期人工草地的土壤微牛物生物量碳和酶活性均呈"V"字型,变化.对于退化生态系统的恢复首先是植被恢复,其次是土壤肥力的恢复.土壤有机质等养分的积累、微生物活性的改善不仅能使土壤-植物复合系统的功能得以恢复,同时也能促进物种多样性的形成,有利于人工草地群落稳定性的提高.在试验区尽管植被恢复演替进行得比较缓慢,但从土壤发展的角度看,仍属进展演替.所以,在退化高寒草甸的恢复过程中,若降低和有效控制外界的干扰(如围栏封育),可为退化草地恢复提供繁殖体与土壤环境,实现人工草地逐步向恢复(正向)演替进行.图3表6参34     

An invader differentially affects leaf physiology of two natives across a gradient in diversity

Little is known about how exotics influence the ecophysiology of co-occurring native plants or how invader impact on plant physiology may be mediated by community diversity or resource levels. We measured the effect of the widespread invasive forb spotted knapweed (Centaurea maculosa) on leaf traits (leaf dry matter content, specific leaf area, leaf nitrogen percentage, leaf C:N ratios, and delta13C as a proxy for water use efficiency) of two co-occurring native perennial grassland species, Monarda fistulosa (bee balm) and Koeleria macrantha (Junegrass). The impact of spotted knapweed was assessed across plots that varied in functional diversity and that either experienced ambient rainfall or received supplemental water. Impact was determined by comparing leaf traits between identical knapweed-invaded and noninvaded assemblages. Virtually all M. fistulosa leaf traits were affected by spotted knapweed. Knapweed impact, however, did not scale with its abundance; the impact of knapweed on M. fistulosa was similar across heavily invaded low-diversity assemblages and lightly invaded high-diversity assemblages. In uninvaded assemblages, M. fistulosa delta13C, leaf nitrogen, and C:N ratios were unaffected by native functional group richness, whereas leaf dry matter content significantly increased and specific leaf area significantly decreased across the diversity gradient. The effects of spotted knapweed on K. macrantha were weak; instead native functional group richness strongly affected K. macrantha leaf C:N ratio, delta13C, and specific leaf area, but not leaf dry matter content. Leaf traits for both species changed in response to spotted knapweed or functional richness, and in a manner that may promote slower biomass accumulation and efficient conservation of resources. Taken together, our results show that an invader can alter native plant physiology, but that these effects are not a simple function of how many invaders exist in the community.     

Separating habitat invasibility by alien plants from the actual level of invasion

Habitats vary considerably in the level of invasion (number or proportion of alien plant species they contain), which depends on local habitat properties, propagule pressure, and climate. To determine the invasibility (susceptibility to invasions) of different habitats, it is necessary to factor out the effects of any confounding variables such as propagule pressure and climate on the level of invasion. We used 20 468 vegetation plots from 32 habitats in the Czech Republic to compare the invasibility of different habitats. Using regression trees, the proportion of alien plants, including archaeophytes (prehistoric to medieval invaders) and neophytes (recent invaders), was related to variables representing habitat properties, propagule pressure, and climate. The propagule pressure was expressed as the proportion of surrounding urban and industrial or agricultural land, human population density, distance from a river, and history of human colonization in the region. Urban and industrial land use had a positive effect on the proportion of both archaeophytes and neophytes. Agricultural land use, higher population density, and longer history of human impact positively affected the proportion of archaeophytes. Disturbed human-made habitats with herbaceous vegetation were most invaded by both groups of aliens. Neophytes were also relatively common in disturbed woody vegetation, such as broad-leaved plantations, forest clearings, and riverine scrub. These habitats also had the highest proportion of aliens after removing the effect of propagule pressure and climate, indicating that they are not only the most invaded, but also most invasible. These habitats experience recurrent disturbances and are rich, at least temporarily, in available nutrients, which supports the hypothesis that fluctuating resources are the major cause of habitat invasibility. The least invaded habitats were mires and alpine-subalpine grasslands and scrub. After removing the effect of propagule pressure and climate, some habitats actually invaded at an intermediate level had very low proportions of aliens. This indicates that these habitats (e.g., dry, wet, and saline grasslands, base-rich fens, and broad-leaved deciduous woodlands) are resistant to invasion.     

Forest succession suppressed by an introduced plant-fungal symbiosis

Microbial symbionts can affect plant nutrition, defensive chemistry, and biodiversity. Here we test the hypothesis that symbionts alter the speed and direction of plant succession in communities that are shifting from grasslands to forests. A widespread C3 grass introduced to the United States, Lolium arundinaceum (tall fescue), hosts a fungal endophyte that is toxic to herbivores. In replicated experimental grasslands, the presence of the endophyte in tall fescue reduced tree abundance and size, altered tree composition, and slowed plant species turnover. In addition, consumption of tree seedlings by voles (Microtus spp.) was 65% higher in plots with the endophyte at the one grassland site where these data were collected. Despite its negligible contribution to community biomass, a microbial symbiont suppressed tree establishment, posing an important constraint on the natural transition from grasslands to forests.     

Diversity-invasibility across an experimental disturbance gradient in Appalachian Forests

Research examining the relationship between community diversity and invasions by nonnative species has raised new questions about the theory and management of biological invasions. Ecological theory predicts, and small-scale experiments confirm, lower levels of nonnative species invasion into species-rich compared to species-poor communities, but observational studies across a wider range of scales often report positive relationships between native and nonnative species richness. This paradox has been attributed to the scale dependency of diversity-invasibility relationships and to differences between experimental and observational studies. Disturbance is widely recognized as an important factor determining invasibility of communities, but few studies have investigated the relative and interactive roles of diversity and disturbance on nonnative species invasion. Here, we report how the relationship between native and nonnative plant species richness responded to an experimentally applied disturbance gradient (from no disturbance up to clearcut) in oak-dominated forests. We consider whether results are consistent with various explanations of diversity-invasibility relationships including biotic resistance, resource availability, and the potential effects of scale (1 m2 to 2 ha). We found no correlation between native and nonnative species richness before disturbance except at the largest spatial scale, but a positive relationship after disturbance across scales and levels of disturbance. Post-disturbance richness of both native and nonnative species was positively correlated with disturbance intensity and with variability of residual basal area of trees. These results suggest that more nonnative plants may invade species-rich communities compared to species-poor communities following disturbance.     

Native plant diversity resists invasion at both low and high resource levels

Human modification of the environment is causing both loss of species and changes in resource availability. While studies have examined how species loss at the local level can influence invasion resistance, interactions between species loss and other components of environmental change remain poorly studied. In particular, the manner in which native diversity interacts with resource availability to influence invasion resistance is not well understood. We created experimental plant assemblages that varied in native species (1-16 species) and/or functional richness (defined by rooting morphology and phenology; one to five functional groups). We crossed these diversity treatments with resource (water) addition to determine their interactive effects on invasion resistance to spotted knapweed (Centaurea maculosa), a potent exotic invader in the intermountain West of the United States. We also determined how native diversity and resource addition influenced plant-available soil nitrogen, soil moisture, and light. Assemblages with lower species and functional diversity were more heavily invaded than assemblages with greater species and functional diversity. In uninvaded assemblages, experimental addition of water increased soil moisture and plant-available nitrogen and decreased light availability. The availability of these resources generally declined with increasing native plant diversity. Although water addition increased susceptibility to invasion, it did not fundamentally change the negative relationship between diversity and invasibility. Thus, native diversity provided strong invasion resistance even under high resource availability. These results suggest that the effects of local diversity can remain robust despite enhanced resource levels that are predicted under scenarios of global change.