Does disturbance,competition or resource limitation underlie Hieracium lepidulum invasion in New Zealand? Mechanisms of establishment and persistence,and functional differentiation among invasive and native species

The processes underlying plant invasions have been the subject of much ecological research. Understanding mechanisms of plant invasions are difficult to elucidate from observations, yet are crucial for ecological management of invasions. Hieracium lepidulum, an asteraceous invader in New Zealand, is a species for which several explanatory mechanisms can be raised. Alternative mechanisms, including competitive dominance, disturbance of resident vegetation allowing competitive release or nutrient resource limitation reducing competition with the invader are raised to explain invasion. We tested these hypotheses in two field experiments which manipulated competitive, disturbance and nutrient environments in pre‐invasion and post‐invasion vegetation. H. lepidulum and resident responses to environmental treatments were measured to allow interpretation of underlying mechanisms of establishment and persistence. We found that H. lepidulum differed in functional response profile from native species. We also found that other exotic invaders at the sites were functionally different to H. lepidulum in their responses. These data support the hypothesis that different invaders use different invasion mechanisms from one another. These data also suggest that functional differentiation between invaders and native resident vegetation may be an important contributing factor allowing invasion. H. lepidulum appeared to have little direct competitive effect on post‐invasion vegetation, suggesting that competition was not a dominant mechanism maintaining its persistence. There was weak support for disturbance allowing initial establishment of H. lepidulum in pre‐invasion vegetation, but disturbance did not lead to invader dominance. Strong support for nutrient limitation of resident species was provided by the rapid competitive responses with added nutrients despite presence of H. lepidulum. Rapid competitive suppression of H. lepidulum once nutrient limitation was alleviated suggests that nutrient limitation may be an important process allowing the invader to dominate. Possible roles of historical site degradation and/or invader‐induced soil chemical/microbial changes in nutrient availability are discussed.     

Abiotic and biotic resistance to grass invasion in serpentine annual plant communities

Biological invasions severely impact native plant communities, causing dramatic shifts in species composition and the restriction of native species to spatially isolated refuges. Competition from resident species and the interaction between resource limitation and competition have been overlooked as mechanisms of community resistance in refugia habitats. We examined the importance of these factors in determining the resistance of California serpentine plant communities to invasion by three common European grasses, Avena barbata, Bromus diandrus, and Hordeum murinum. We added seeds of each of these grasses to plots subjected to six levels of resource addition (N, P, Ca, H₂O, all resources together, and a no-addition control) and two levels of competition (with resident community present or removed). Resource limitation and competition had strong effects on the biomass and reproduction of the three invaders. The addition of all resources together combined with the removal of the resident community yielded individual plants that were fourfold to 20-fold larger and sixfold to 20-fold more fecund than plants from control plots. Competitor removal alone yielded invaders that were twofold to sevenfold larger and twofold to ninefold more fecund. N addition alone or in combination with other resources led to a twofold to ninefold increase in the biomass and fecundity of the invaders. No other resource alone significantly affected native or invader performance, suggesting that N was the key limiting resource during our experiment. We found a significant interaction between abiotic and biotic resistance for Bromus, which experienced increased competitive suppression in fertilized plots. The threefold increase in resident biomass with N addition was likely responsible for this result. Our results confirm that serpentine plant communities are severely N limited, which, in combination with competition from resident species, promotes the resistance of these systems to invasions. Our work suggests that better understanding the relative sensitivities of invaders and residents to the physical environment is critical to predicting how abiotic and biotic factors interact to determine community resistance.     

Traits linked with species invasiveness and community invasibility vary with time,stage and indicator of invasion in a long‐term grassland experiment

Much uncertainty remains about traits linked with successful invasion – the establishment and spread of non‐resident species into existing communities. Using a 20‐year experiment, where 50 non‐resident (but mostly native) grassland plant species were sown into savannah plots, we ask how traits linked with invasion depend on invasion stage (establishment, spread), indicator of invasion success (occupancy, relative abundance), time, environmental conditions, propagule rain, and traits of invaders and invaded communities. Trait data for 164 taxa showed that invader occupancy was primarily associated with traits of invaders, traits of recipient communities, and invader‐community interactions. Invader abundance was more strongly associated with community traits (e.g. proportion legume) and trait differences between invaders and the most similar resident species. Annuals and invaders with high‐specific leaf area were only successful early in stand development, whereas invaders with conservative carbon capture strategies persisted long‐term. Our results indicate that invasion is context‐dependent and long‐term experiments are required to comprehensively understand invasions.     

Plant nitrogen and phosphorus limitation in 98 North American grassland soils

The availability of nutrients is a critical determinant of ecological dynamics in grasslands, but the relationships between soil resource availability and nutrient limitation across ecosystems are not clear. To better understand how soil nutrient availability determines nutrient limitation in vegetation, we grew the same species of grass (Schizachyrium scoparium) in 98 North American grassland soils and fertilized them factorially with nitrogen (N) and phosphorus (P). On average adding N, P, and the two nutrients together increased biomass relative to unfertilized plants by 81%, 22%, and 131%, respectively. Plants grown on low-P soils were not primarily limited by P. Instead, these plants were colimited by N and P, while plants grown on high-P soils were primarily limited by N and only secondarily limited by P. Limitation was not predicted by total soil N. The preponderance of colimitation between N and P on low-P soils suggests that low P availability alters the N cycle to constrain supplies to plants such that N and P are made available in proportion to their demand by plants.     

Soil fertility alters the nature of plant–resource interactions in invaded grassland communities

Resource competition theory suggests that the nature of diversity–resource–invasibility interactions will vary along fertility gradients, concurrent with changes in the relative availability of limiting above- versus below-ground resources. Experimental support for this contingency is lacking. Here, we manipulated resident diversity, baseline fertility, and the availabilities of light and soil nitrogen in grassland communities invaded by two functionally distinct non-native plant species (Lolium arundinaceum and Melilotus alba). We tested the hypotheses that increased resident diversity reduces community invasibility and dampens the effects of light and soil nitrogen pulses, and that the relative effects of light versus soil nitrogen additions on diversity–invasibility relationships depend on the baseline fertility of the study system. Our results reveal an overall weak negative effect of resident diversity on Lolium performance, but in contrast to our expectations, this diversity effect did not vary with light or soil nitrogen additions or with baseline fertility. However, the relative effects of above- versus below-ground resource additions on invader performance varied with baseline fertility as expected: Lolium responded most strongly to soil nitrogen additions in low-fertility mesocosms and most strongly to increased light availability in high-fertility mesocosms. In contrast to Lolium, nitrogen-fixing Melilotus was overall less responsive to diversity and resource manipulations. Together, these patterns do not lend support for the dependence of diversity–resource–invasibility relationships on either baseline fertility or invasive species identity, but they do highlight the dominant role of resources over diversity in determining invader performance, as well as the manner in which fertility alters the relative importance of above- versus below-ground resource pulses in promoting invasions.     

Nitrogen balance in the Trachypogon grasslands of Central Venezuela

The nitrogen balance of a Trachypogon grassland in Calabozo, Venezuela, is calculated for average conditions using biomass accumulation, nitrogen content, and turnover rates of organic matter. Burning Trachypogon grasslands results in losses of 8.5 kg N ha^?1 yr^?1, while rainfall inputs average 2.6 kg N ha^?1 yr^?1. Uptake of N by vegetation is 14.8 kg N ha^?1 yr^?1, but the total N required to build new tissue during a growing season is about 30 kg N ha^?1 yr^?1, so that about 50% of the nitrogen in the vegetation is recycled internally. Nitrogen lossesvia fire are probably balanced by biological N₂-fixation, but no data are available for N-fixation in these savannas. The calculations presented in this paper are based on few data and more measurements are needed to develop a conclusive picture of the N-balance of Trachypogon grasslands.     

The distribution and prevalence of helminths,coccidia and blood parasites in two competing species of gecko: implications for apparent competition

Across the Pacific the invading gecko species Hemidactylus frenatus has competitively displaced the resident gecko species Lepidodactylus lugubris in urban/surburban habitats. Do parasites enhance, inhibit, orhave no effect on this invasion? Parasites can confer an advantage to an invading species when the invader (1) introduces a new parasite to a resident species that has a greater detrimental effect on the resident than the invader, (2) is less susceptible to endemic parasites than the resident, and/or (3) increases the susceptibility of the resident to parasites. Conversely, parasites may protect a resident against invasion when endemic parasites have a greater impact on the invader than the resident. We screened more than one thousand H. frenatus and L. lugubris in areas of sympatry and allopatry from 28 islands and 5 sites on mainland Asia for a broad array of blood parasites, coccidia and helminths in order to evaluate the potential for parasites to affect their interaction. We found that 1) There were no parasites which appear to protect L. lugubris against invasion by H. frenatus. 2) H. frenatus does not introduce the same parasite to L. lugubris in every location where the two come in to contact, but probably has introduced different parasites in different locations. L. lugubris also seems to have introduced at least one parasite to H. frenatus. 3) The prevalence of parasite species shared by the two hosts is generally higher in H. frenatus; however, prevalence is determined by many factors and cannot be directly translated as susceptibility. We discuss the implications of this difference in prevalence for the Red Queen hypothesis. 4) The prevalence of the cestode Cylindrotaenia sp. is significantly higher in L. lugubris that are sympatric with H. frenatus than those which are allopatric.     

Litter quality,decomposition rates and saprotrophic mycoflora in Fallopia japonica (Houtt.) Ronse Decraene and in adjacent native grassland vegetation

Fallopia japonica succeeds in invading different ecosystems likely because of its huge biomass production. This biomass is characterized by low nutritional quality and low decomposition rates but knowledge on whether these features are correlated to microbial decomposers is still lacking. The aims of this work were: i) to determine litter decomposition rates of native grassland vegetation and F. japonica under different conditions in a year-round experiment; ii) to evaluate litter quality and/or site effect on the decomposition of the invader and native vegetation and iii) to characterize mycoflora isolated from F. japonica and native vegetation litter. The results showed that F. japonica litter decomposes 3–4 times slower than that of native grassland, mainly due to its low N content and consequently high C/N ratio both in leaves and stems. As decomposition proceeds C/N in F. japonica litter decreases to values approaching those of the grassland litter. Site had no effect on the decomposition rates of F. japonica and grassland litter. Total fungal load and composition differed between F. japonica and native litter, and also varied across sites. These results indicate that the successful invasive plant F. japonica affects the structure and functions of the invaded ecosystem through a huge production of low quality, slow-decomposing litter that selects saprotrophic fungi.     

Plant invasion alters trait composition and diversity across habitats

Increased globalization has accelerated the movement of species around the world. Many of these nonnative species have the potential to profoundly alter ecosystems. The mechanisms underpinning this impact are often poorly understood, and traits are often overlooked when trying to understand and predict the impacts of species invasions on communities. We conducted an observational field experiment in Canada's first National Urban Park, where we collected trait data for seven different functional traits (height, stem width, specific leaf area, leaf percent nitrogen, and leaf percent carbon) across an abundance gradient of the invasive Vincetoxicum rossicum in open meadow and understory habitats. We assessed invasion impacts on communities, and associated mechanisms, by examining three complementary functional trait measures: community‐weighted mean, range of trait values, and species’ distances to the invader in trait space. We found that V. rossicum invasion significantly altered the functional structure of herbaceous plant communities. In both habitats V. rossicum changed the community‐weighted means, causing invaded communities to become increasingly similar in their functional structure. In addition, V. rossicum also reduced the trait ranges for a majority of traits indicating that species are being deterministically excluded in invaded communities. Further, we observed different trends in the meadow and understory habitats: In the understory, resident species that were more similar to V. rossicum in multivariate trait space were excluded more, however this was not the case in the meadow habitat. This suggests that V. rossicum alters communities uniquely in each habitat, in part by creating a filter in which only certain resident species are able to persist. This filtering process causes a nonrandom reduction in species' abundances, which in turn would be expected to alter how the invaded ecosystems function. Using trait‐based frameworks leads to better understanding and prediction of invasion impacts. This novel framework can also be used in restoration practices to understand how invasion impacts communities and to reassemble communities after invasive species management.     

Comparative pollinator limitation of two non-native shrubs: do mutualisms influence invasions?

While interactions between invaders and resident species have received a great deal of attention recently, the role of mutualists in facilitating or constraining invasions is rarely considered. We investigated the reproductive ecology of two closely related, woody legumes, Cytisus scoparius (Scotch broom) and Genista monspessulana (French broom), invading the same sites. Both species are considered noxious non-native weeds in California, and are considered to be ecologically similar, but Genista has much smaller flowers than Cytisus. Neither species showed appreciable levels of autogamous selfing. When experimentally self-pollinated, Genista demonstrated less depression of fruit set and seed set relative to outcrossed flowers than did Cytisus. At two sites on the Marin peninsula, Calif., Genista flowers were consistently less likely to be pollinated than Cytisus flowers. Genista was significantly pollen limited at both sites, while Cytisus was pollen limited at only the site with lower visitation rates. In the three populations with demonstrable pollen limitation, we found a significant relationship between fruit production and natural pollinator visitation at the level of the individual plant. However, we did not find that overall patterns of fecundity were strongly predicted by differences in pollen limitation between species or between sites. While a previous study found a tight link between patterns of pollinator visitation and patterns of reproduction in Cytisus in Washington State, we conclude that a more complex and variable environment (in terms of resources, herbivores, and florivores) on the Marin Peninsula de-coupled the relationship between pollinators and fruit production in these invaders. Our results suggest that the role of mutualisms in promoting or constraining invasions is likely to vary considerably among invaded communities.     

Elevated success of multispecies bacterial invasions impacts community composition during ecological succession

Successful microbial invasions are determined by a species’ ability to occupy a niche in the new habitat whilst resisting competitive exclusion by the resident community. Despite the recognised importance of biotic factors in determining the invasiveness of microbial communities, the success and impact of multiple concurrent invaders on the resident community has not been examined. Simultaneous invasions might have synergistic effects, for example if resident species need to exhibit divergent phenotypes to compete with the invasive populations. We used three phylogenetically diverse bacterial species to invade two compositionally distinct communities in a controlled, naturalised in vitro system. By initiating the invader introductions at different stages of succession, we could disentangle the relative importance of resident community structure, invader diversity and time pre‐invasion. Our results indicate that multiple invaders increase overall invasion success, but do not alter the successional trajectory of the whole community.     

N/P imbalance as a key driver for the invasion of oligotrophic dune systems by a woody legume

Oligotrophic ecosystems, previously considered to be more resilient to invasive plants, are now recognised to be highly vulnerable to invasions. In these systems, woody legumes show belowground ecosystem engineering characteristics that enable invasion, however, the underlying processes are not well understood. Using a Portuguese primary dune ecosystem as an oligotrophic model system, belowground biomass pools, turnover rates and stoichiometry of a native (Stauracanthus spectabilis) and an invasive legume (Acacia longifolia) were compared and related to changes in the foliage of the surrounding native (Corema album) vegetation. We hypothesized that the invasive legume requires less phosphorus per unit of biomass produced and exhibits an enhanced nutrient turnover compared to the native vegetation, which could drive invasion by inducing a systemic N/P imbalance. Compared with the native legumes, A. longifolia plants had larger canopies, higher SOM levels and lower tissue P concentrations. These attributes were strongly related to legume influence as measured by increased foliar N content and less depleted δ¹⁵N signatures in the surrounding C. album vegetation. Furthermore, higher root N concentration and increased nutrient turnover in the rhizosphere of the invader were associated with depleted foliar P in C. album. Our results emphasize that while A. longifolia itself maintains an efficient phosphorus use in biomass production, at the same time it exerts a strong impact on the N/P balance of the native system. Moreover, this study highlights the engineering of a belowground structure of roots and rhizosphere as a crucial driver for invasion, due to its central role in nutrient turnover. These findings provide new evidence that, under nutrient‐limited conditions, considering co‐limitation and nutrient cycling in oligotrophic systems is essential to understand the engineering character of invasive woody legumes.     

Nutrient feedbacks to soil heterotrophic nitrogen fixation in forests

Multiple nutrient cycles regulate biological nitrogen (N) fixation in forests, yet long-term feedbacks between N-fixation and coupled element cycles remain largely unexplored. We examined soil nutrients and heterotrophic N-fixation across a gradient of 24 temperate conifer forests shaped by legacies of symbiotic N-fixing trees. We observed positive relationships among mineral soil pools of N, carbon (C), organic molybdenum (Mo), and organic phosphorus (P) across sites, evidence that legacies of symbiotic N-fixing trees can increase the abundance of multiple elements important to heterotrophic N-fixation. Soil N accumulation lowered rates of heterotrophic N-fixation in organic horizons due to both N inhibition of nitrogenase enzymes and declines in soil organic matter quality. Experimental fertilization of organic horizon soil revealed widespread Mo limitation of heterotrophic N-fixation, especially at sites where soil Mo was scarce relative to C. Fertilization also revealed widespread absence of P limitation, consistent with high soil P:Mo ratios. Responses of heterotrophic N-fixation to added Mo (positive) and N (negative) were correlated across sites, evidence that multiple nutrient controls of heterotrophic N-fixation were more common than single-nutrient effects. We propose a conceptual model where symbiotic N-fixation promotes coupled N, C, P, and Mo accumulation in soil, leading to positive feedback that relaxes nutrient limitation of overall N-fixation, though heterotrophic N-fixation is primarily suppressed by strong negative feedback from long-term soil N accumulation.     

Severity of impacts of an introduced species corresponds with regional eco‐evolutionary experience

Invasive plant impacts vary widely across introduced ranges. We tested the hypothesis that differences in the eco‐evolutionary experience of native communities with the invader correspond with the impacts of invasive species on native vegetation, with impacts increasing with ecological novelty. We compared plant species richness and composition beneath Pinus contorta to that in adjacent vegetation and other P. contorta stands across a network of sites in its native (Canada and USA) and non‐native (Argentina, Chile, Finland, New Zealand, Scotland, Sweden) ranges. At sites in North America and Europe, within the natural distribution of the genus Pinus, P. contorta was not associated with decreases in diversity. In the Southern Hemisphere, where there are no native Pinaceae, plant communities beneath P. contorta were less diverse than in other regions and compared to uninvaded native vegetation. Effects on native vegetation were particularly pronounced where P. contorta was a more novel life form and exhibited higher growth rates. Our results support the hypothesis that the eco‐evolutionary experience of the native vegetation, and thus the novelty of the invader, determines the magnitude of invader impacts on native communities. Understanding the eco‐evolutionary context of invasions will help to better understand and predict where invasion impacts will be greatest and to prioritize invasive species management.     

Your worst enemy could be your best friend: predator contributions to invasion resistance and persistence of natives

Native predators are postulated to have an important role in biotic resistance of communities to invasion and community resilience. Effects of predators can be complex, and mechanisms by which predators affect invasion success and impact are understood for only a few well-studied communities. We tested experimentally whether a native predator limits an invasive species’ success and impact on a native competitor for a community of aquatic insect larvae in water-filled containers. The native mosquito Aedes triseriatus alone had no significant effect on abundance of the invasive mosquito Aedes albopictus. The native predatory midge Corethrella appendiculata, at low or high density, significantly reduced A. albopictus abundance. This effect was not caused by trait-mediated oviposition avoidance of containers with predators, but instead was a density-mediated effect caused by predator-induced mortality. The presence of this predator significantly reduced survivorship of the native species, but high predator density also significantly increased development rate of the native species when the invader was present, consistent with predator-mediated release from interspecific competition with the invader. Thus, a native predator can indirectly benefit its native prey when a superior competitor invades. This shows the importance of native predators as a component of biodiversity for both biotic resistance to invasion and resilience of a community perturbed by successful invasion.     

Modelling invasibility in endogenously oscillating tree populations: timing of invasion matters

The timing of introduction of a new species into an ecosystem can be critical in determining the invasibility (i.e. the sensitivity to invasion) of a resident population. Here, we use an individual-based model to test how (1) the type of competition (symmetric versus asymmetric) and (2) seed masting influence the success of invasion by producing oscillatory dynamics in resident tree populations. We focus on a case where two species (one resident, one invader introduced at low density) do not differ in terms of competitive abilities. By varying the time of introduction of the invader, we show that oscillations in the resident population favour invasion, by creating “invasibility windows” during which resource is available for the invader due to transiently depressed resident population density. We discuss this result in the context of current knowledge on forest dynamics and invasions, emphasizing the importance of variability in population dynamics.     

Siderophores drive invasion dynamics in bacterial communities through their dual role as public good versus public bad

Microbial invasions can compromise ecosystem services and spur dysbiosis and disease in hosts. Nevertheless, the mechanisms determining invasion outcomes often remain unclear. Here, we examine the role of iron-scavenging siderophores in driving invasions of Pseudomonas aeruginosa into resident communities of environmental pseudomonads. Siderophores can be ‘public goods’ by delivering iron to individuals possessing matching receptors; but they can also be ‘public bads’ by withholding iron from competitors lacking these receptors. Accordingly, siderophores should either promote or impede invasion, depending on their effects on invader and resident growth. Using supernatant feeding and invasion assays, we show that invasion success indeed increased when the invader could use its siderophores to inhibit (public bad) rather than stimulate (public good) resident growth. Conversely, invasion success decreased the more the invader was inhibited by the residents’ siderophores. Our findings identify siderophores as a major driver of invasion dynamics in bacterial communities under iron-limited conditions.     

Responses of a native plant species from invaded and uninvaded areas to allelopathic effects of an invader

Invaders exert new selection pressures on the resident species, for example, through competition for resources or by using novel weapons. It has been shown that novel weapons aid invasion but it is unclear whether native species co‐occurring with invaders have adapted to tolerate these novel weapons. Those resident species which are able to adapt to new selective agents can co‐occur with an invader while others face a risk of local extinction. We ran a factorial common garden experiment to study whether a native plant species, Anthriscus sylvestris, has been able to evolve a greater tolerance to the allelochemicals exerted by the invader, Lupinus polyphyllus. Lupinus polyphyllus produces allelochemicals which potentially act as a novel, strong selective agent on A. sylvestris. We grew A. sylvestris seedlings collected from uninvaded (naïve) and invaded (experienced) sites growing alone and in competition with L. polyphyllus in pots filled with soil with and without activated carbon. Because activated carbon absorbs allelochemicals, its addition should improve especially naïve A. sylvestris performance in the presence of the invader. To distinguish the allelochemicals absorption and fertilizing effects of activated carbon, we grew plants also in a mixture of soil and fertilizer. A common garden experiment indicated that the performances of naïve and experienced A. sylvestris seedlings did not differ when grown with L. polyphyllus. The addition of activated carbon, which reduces interference by allelochemicals, did not induce differences in their performances although it had a positive effect on the aboveground biomass of A. sylvestris. Together, these results suggest that naïve and experienced A. sylvestris plants tolerated equally the invader L. polyphyllus and thus the tolerance has not occurred over the course of invasion.     

Community invasibility and invasion by non-native Fraxinus pennsylvanica trees in a degraded tropical forest

Whether invasion of introduced plant species may be aided by certain community properties is poorly understood for species-rich ecosystems, such as tropical montane forests. In Kenya, the non-native tree Fraxinus pennsylvanica has invaded degraded montane forests. We used generalized linear mixed models to examine the relative importance of different community properties to Fraxinus invasion after agricultural abandonment and in the secondary forest. Fraxinus invasion was positively related to plant community species diversity and the abundance of tree saplings, shrubs, ferns, and herbs in the abandoned fallows, but negatively related to the same community properties in the secondary forest. The number of Fraxinus recruits declined with declining propagule pressure in the fallows, but not in the secondary forest. Although adult and saplings of Fraxinus were positively related to community diversity in the fallows, Fraxinus appeared to decrease diversity in the secondary forest. These results show that the success of non-native species invasion and the effects of an invader on the resident community may depend both on properties and degree of disturbance of the community. Plant community diversity and evenness appeared to determine the invasion success by increasing invasibility of the abandoned fallows, but decreasing invasibility of the secondary forest. Our results from a tropical degraded forest emphasize the importance of including habitat characteristics when predicting both the potential of non-native plant invasion and effects of invasives on the particular community.     

Plant diversity increases resistance to invasion in the absence of covarying extrinsic factors

Biological invasion is a widespread, but poorly understood phenomenon. Elton's hypothesis, supported by theory, experiment, and anecdotal evidence, suggests that an important determinant of invasion success is resident biodiversity, arguing that high diversity increases the competitive environment of communities and makes them more difficult to invade. Observational studies of plant invasions, however, find little support for this hypothesis and argue strongly against it. Lack of control of extrinsic factors (e.g., disturbance, climate, or soil fertility) that covary with biodiversity and invasion in observational studies makes it difficult to determine if their findings truly refute Elton's hypothesis. We examined performance of Crepis tectorum (an invasive, annual composite weed) in experimental prairie grassland plots and greenhouse plant assemblages in which resident species richness was directly manipulated. Under these conditions, unlike observational studies, no covarying extrinsic factors could interfere with interpreting results. We found a strong inverse association between resident diversity and invader performance as predicted by Elton's hypothesis. Higher resident diversity increased crowding, decreased available light, and decreased available nutrients all of which increased the competitive environment of diverse plant assemblages and reduced C. tectorum success. Examination of individual resident species impacts on C. tectorum performance demonstrated that this diversity effect was not due to the sampling effect. These results suggest that both Elton's hypothesis and its competitive mechanism may operate in nature, but covarying extrinsic factors may obscure the negative impact of diversity on invader success.