Spatial Pattern and Scale Influence Invader Demographic Response to Simulated Precipitation Change in an Annual Grassland Community.

It is important to predict which invasive species will benefit from future changes in climate, and thereby identify those invaders that need particular attention and prioritization of management efforts. Because establishment, persistence, and spread determine invasion success, this prediction requires detailed demographic information. Explicit study of the impact of pattern on demographic response is particularly important for species that are naturally patchy, such as the invasive grass, Aegilops triuncialis. In the northern California Coast Range, where climate change may increase or decrease mean annual rainfall, we conducted a field experiment to understand the interaction of climate change and local-scale patterning on the demography of A. triuncialis. We manipulated precipitation (reduced, ambient, or augmented), seed density, and seeding pattern. Demographic and environmental data were collected for three years following initial seeding. Pattern and scale figure prominently in the demographic response of A. triuncialis to precipitation manipulation. Pattern interacts with precipitation and seeding density in its influence on per-plant seed output. Although per-plot seed production was highest when seeds were not aggregated, per-plant seed output was higher in aggregated patches. Results suggest aggregation of invasive A. triuncialis reduces the detrimental impact of interspecific competition in its invaded community, and that interspecific competition per se has a stronger impact than intraspecific competition.

Climate structures genetic variation across a species' elevation range: a test of range limits hypotheses.

Gene flow may influence the formation of species range limits, and yet little is known about the patterns of gene flow with respect to environmental gradients or proximity to range limits. With rapid environmental change, it is especially important to understand patterns of gene flow to inform conservation efforts. Here we investigate the species range of the selfing, annual plant, Mimulus laciniatus, in the California Sierra Nevada. We assessed genetic variation, gene flow, and population abundance across the entire elevation-based climate range. Contrary to expectations, within-population plant density increased towards both climate limits. Mean genetic diversity of edge populations was equivalent to central populations; however, all edge populations exhibited less genetic diversity than neighbouring interior populations. Genetic differentiation was fairly consistent and moderate among all populations, and no directional signals of contemporary gene flow were detected between central and peripheral elevations. Elevation-driven gene flow (isolation by environment), but not isolation by distance, was found across the species range. These findings were the same towards high- and low-elevation range limits and were inconsistent with two common centre-edge hypotheses invoked for the formation of species range limits: (i) decreasing habitat quality and population size; (ii) swamping gene flow from large, central populations. This pattern demonstrates that climate, but not centre-edge dynamics, is an important range-wide factor structuring M. laciniatus populations. To our knowledge, this is the first empirical study to relate environmental patterns of gene flow to range limits hypotheses. Similar investigations across a wide variety of taxa and life histories are needed.

Contemporary evolution of an invasive grass in response to elevated atmospheric CO(2) at a Mojave Desert FACE site.

Elevated atmospheric CO2 has been shown to rapidly alter plant physiology and ecosystem productivity, but contemporary evolutionary responses to increased CO2 have yet to be demonstrated in the field. At a Mojave Desert FACE (free-air CO2 enrichment) facility, we tested whether an annual grass weed (Bromus madritensis ssp. rubens) has evolved in response to elevated atmospheric CO2 . Within 7 years, field populations exposed to elevated CO2 evolved lower rates of leaf stomatal conductance; a physiological adaptation known to conserve water in other desert or water-limited ecosystems. Evolution of lower conductance was accompanied by reduced plasticity in upregulating conductance when CO2 was more limiting; this reduction in conductance plasticity suggests that genetic assimilation may be ongoing. Reproductive fitness costs associated with this reduction in phenotypic plasticity were demonstrated under ambient levels of CO2 . Our findings suggest that contemporary evolution may facilitate this invasive species' spread in this desert ecosystem.

Niche partitioning between close relatives suggests trade-offs between adaptation to local environments and competition.

Niche partitioning among close relatives may reflect trade-offs underlying species divergence and coexistence (e.g., between stress tolerance and competitive ability). We quantified the effects of habitat and congeneric species interactions on fitness for two closely related herbaceous plant species, Mimulus guttatus and Mimulus laciniatus, in three common habitat types within their sympatric range. Drought stress strongly reduced survival of M. guttatus in fast-drying seeps occupied by M. laciniatus, suggesting that divergent habitat adaptation maintains this niche boundary. However, neither seedling performance nor congeneric competition explained the absence of M. laciniatus from shady streams where M. guttatus thrives. M. laciniatus may be excluded from this habitat by competition with other species in the community or mature M. guttatus. Species performance and competitive ability were similar in sympatric meadows where plant community stature and the growing season length are intermediate between seeps and streams. Stochastic effects (e.g., dispersal among habitats or temporal variation) may contribute to coexistence in this habitat. Habitat adaptation, species interactions, and stochastic mechanisms influence sympatric distributions for these recently diverged species.

Trade-offs, spatial heterogeneity, and the maintenance of microbial diversity.

Specialization and concomitant trade-offs are assumed to underlie the non-neutral coexistence of lineages. Trade-offs across heterogeneous environments can promote diversity by preventing competitive exclusion. However, the importance of trade-offs in maintaining diversity in natural microbial assemblages is unclear, as trade-offs are frequently not detected in artificial evolution experiments. Stressful conditions associated with patches of heavy-metal enriched serpentine soils provide excellent opportunities for examining how heterogeneity may foster genetic diversity. Using a spatially replicated design, we demonstrate that rhizobium bacteria symbiotic with legumes inhabiting contrasting serpentine and nonserpentine soils exhibit a trade-off between a genotype's nickel tolerance and its ability to replicate rapidly. Furthermore, we detected adaptive divergence in rhizobial assemblages across soil type heterogeneity at multiple sites, suggesting that this trade-off may promote the coexistence of phenotypically distinct bacterial lineages. Trade-offs and adaptive divergence may be important factors maintaining the tremendous diversity within natural assemblages of bacteria.

Within- and trans-generational plasticity affects the opportunity for selection in barbed goatgrass (Aegilops triuncialis).

PREMISE OF THE STUDY: Environments are composed of selective agents, and environments may also modify the efficacy of these agents. Environments affect the rate of maximum evolutionary change by influencing variation in relative fitness (i.e., the opportunity for selection, or I). Within- and transgenerational plastic environmental responses may affect I, speeding or slowing processes of local adaptation. • METHODS: We determined whether environmental factors affected the opportunity for selection (I) in Aegilops triuncialis (barbed goatgrass) by measuring I as a within- and transgenerational plastic response to two maternal glasshouse environments (serpentine/dry and loam/moist). We also determined whether this species' two most common genetic lineages (determined by DNA microsatellite length polymorphism) varied in response to glasshouse treatments. • KEY RESULTS: Opportunity for selection was less for plants grown in the dry serpentine environment than for plants grown in the moist loam environment. This response varied between genetic lineages. The east lineage exhibited a within-generation response to the dry serpentine environment. For both seed mass and average seed weight in this lineage, the opportunity for selection was lower in dry serpentine than in moist loam. The west lineage had a transgenerational response to the dry serpentine such that the opportunity for selection for seed number and seed mass was lower for plants produced by mothers grown in dry serpentine than for plants produced by mothers in moist loam. • CONCLUSIONS: Phenotypic variation in relative fitness is constrained by the dry serpentine environment, which leads to lower evolvability in this environment. Within- and transgenerational effects of the environment may slow local adaptation to serpentine soils.

Evolution of root plasticity responses to variation in soil nutrient distribution and concentration.

Root plasticity, a trait that can respond to selective pressure, may help plants forage for nutrients in heterogeneous soils. Agricultural breeding programs have artificially selected for increased yield under comparatively homogeneous soil conditions, potentially decreasing the capacity for plasticity in crop plants like barley (Hordeum vulgare). However, the effects of domestication on the evolution of root plasticity are essentially unknown. Using a split container approach, we examined the differences in root plasticity among three domestication levels of barley germplasm (wild, landrace, and cultivar) grown under different concentrations and distribution patterns of soil nutrients. Domestication level, nutrient concentration, and nutrient distribution interactively affected average root diameter; differential root allocation (within-plant plasticity) was greatest in wild barley (Hordeum spontaneum), especially under low nutrient levels. Correlations of within-plant root plasticity and plant size were most pronounced in modern cultivars under low-nutrient conditions. Barley plants invested more resources to root systems when grown in low-nutrient soils and allocated more roots to higher-nutrient locations. Root plasticity in barley is scale dependent and varies with domestication level. Although wild barley harbors a greater capacity for within-plant root plasticity than domesticated barley, cultivars exhibited the greatest capacity to translate within-plant plasticity into increased plant size.

Mutualism and adaptive divergence: co-invasion of a heterogeneous grassland by an exotic legume-rhizobium symbiosis.

Species interactions play a critical role in biological invasions. For example, exotic plant and microbe mutualists can facilitate each other's spread as they co-invade novel ranges. Environmental context may influence the effect of mutualisms on invasions in heterogeneous environments, however these effects are poorly understood. We examined the mutualism between the legume, Medicago polymorpha, and the rhizobium, Ensifer medicae, which have both invaded California grasslands. Many of these invaded grasslands are composed of a patchwork of harsh serpentine and relatively benign non-serpentine soils. We grew legume genotypes collected from serpentine or non-serpentine soil in both types of soil in combination with rhizobium genotypes from serpentine or non-serpentine soils and in the absence of rhizobia. Legumes invested more strongly in the mutualism in the home soil type and trends in fitness suggested that this ecotypic divergence was adaptive. Serpentine legumes had greater allocation to symbiotic root nodules in serpentine soil than did non-serpentine legumes and non-serpentine legumes had greater allocation to nodules in non-serpentine soil than did serpentine legumes. Therefore, this invasive legume has undergone the rapid evolution of divergence for soil-specific investment in the mutualism. Contrary to theoretical expectations, the mutualism was less beneficial for legumes grown on the stressful serpentine soil than on the non-serpentine soil, possibly due to the inhibitory effects of serpentine on the benefits derived from the interaction. The soil-specific ability to allocate to a robust microbial mutualism may be a critical, and previously overlooked, adaptation for plants adapting to heterogeneous environments during invasion.

Gene flow increases fitness at the warm edge of a species' range.

According to theory, gene flow to marginal populations may stall or aid adaptation at range limits by swamping peripheral populations with maladaptive gene flow or by enhancing genetic variability and reducing inbreeding depression, respectively. We tested these contrasting predictions by manipulating patterns of gene flow of the annual plant, Mimulus laciniatus, at its warm range limit. Gene flow was experimentally applied by using crosses within warm-limit populations (selfed and outcrossed), between warm-limit populations, and between warm-limit and central range populations across two elevational transects. We measured the fitness of offspring in a common garden at the warm-edge species range limit. All sources of gene flow increased seedling emergence at the range limit, suggesting local inbreeding depression at both range limit populations; however, lifetime reproductive success only increased significantly when pollen originated from another warm-limit population. Center-to-warm-edge gene flow was maladaptive by delaying time to development at this warm, fast-drying range limit, whereas edge-to-edge gene flow hastened emergence time and time to reproduction. By empirically testing theory on the effects of gene flow on the formation of geographic range limits, we find benefits of gene flow among populations to be greatest when gene flow is between populations occupying the same range limit. Our results emphasize the overlooked importance of gene flow among populations occurring near the same range limit and highlight the potential for prescriptive gene flow as a conservation option for populations at risk from climate change.

Artifactual increase in journal self-citation.

INTRODUCTION: After submission of a manuscript to a peer-reviewed anesthesia journal, several authors were asked to cite additional references from the journal to which they submitted. We hypothesized that there were differences among the anesthesiology journals in both the total number of self-citations and the proportion of self-citations to the total number of references in each manuscript for the years 2005 and 2010. METHODS: We conducted a review of a sample of manuscripts from 2005 and 2010 to examine the number and rate of self-citations. As a secondary analysis, we reviewed impact factor (IF), rate of self-referencing, and contribution of self-citations to IF in the population of manuscripts published in 8 anesthesia journals between 2000 and 2009 using the ISI Journal Citation Reports. RESULTS: The number (P < 0.0001) and rate (P < 0.0001) of self-citations among the different journals were significantly different in 2005, with similar results for 2010 in the number (P < 0.0001) and rate (P = 0.0002) of self-citations. The mean range of number of self-citations ranged from 0.45 (95% confidence interval [CI], 0.06 to 0.84) to 3.95 (95% CI, 2.2 to 5.7) in 2005 and from 0.25 (95% CI, -0.05 to 0.55) to 4.5 (95% CI, 2.2 to 6.9) in 2010. On a per-journal basis, no difference in the number of self-citations was noted between 2005 and 2010. Analysis of the ISI Journal Citation Reports from 2000 to 2009 suggested a general decline in the contribution of self-cites to the IF over time for the aggregate journals (Spearman correlation coefficient (Rs) -0.25 (95% CI, -0.45 to -0.03), P = 0.02), with the exception of the journal in question (Rs = 0.59 (95% CI, -0.1 to 0.88), P = 0.05). Positive correlations were found between self-cited rate and IF (Rs 0.52, 95% CI, 0.34 to 0.66, P < 0.0001), percentage of self-cites to years used in IF calculation and IF (Rs 0.41, 95% CI, 0.21 to 0.58, P < 0.0001), and δ-IF and IF (Rs 0.89, 95% CI, 0.84 to 0.93, P < 0.0001). CONCLUSION: Although the number and rate of self-citations differed among anesthesia journals, the contribution of self-citation to IF has declined over time for most anesthesia journals. These results suggest periodic reassessment may be important to ensure that the publication process remains transparent and impartial to bias.

Fitness variation and local distribution limits in an annual plant population.

Understanding how genetic variation shapes species' distributions involves examining how variation is distributed across a species' range as well as how it responds to underlying environmental heterogeneity. We examined patterns of fitness variation across the local distribution of an annual composite (Lasthenia fremontii) spanning a small-scale inundation gradient in a California vernal pool wetland. Using seeds collected from the center and edge of a population, paternal half-sib families were generated and transplanted back to the center and edge of the original population. All transplants were adapted to the conditions at the center of the population. The effect of the environment on the opportunity for selection depended on the model of selection assumed. Under a model of hard selection, variance in absolute fitness was lower among transplants at the edge of the population than at the center. Under a model of soft selection, the variance in relative fitness was similar between center and edge microhabitats. Given that this population is likely well-mixed, differences in habitat quality between center and edge microhabitats will likely cause selection at the center of the population to dominate the evolutionary trajectory of this population.

Patterns of introduction and adaptation during the invasion of Aegilops triuncialis (Poaceae) into Californian serpentine soils.

Multiple introductions can play a prominent role in explaining the success of biological invasions. One often cited mechanism is that multiple introductions of invasive species prevent genetic bottlenecks by parallel introductions of several distinct genotypes that, in turn, provide heritable variation necessary for local adaptation. Here, we show that the invasion of Aegilops triuncialis into California, USA, involved multiple introductions that may have facilitated invasion into serpentine habitats. Using microsatellite markers, we compared the polymorphism and genetic structure of populations of Ae. triuncialis invading serpentine soils in California to that of accessions from its native range. In a glasshouse study, we also compared phenotypic variation in phenological and fitness traits between invasive and native populations grown on loam soil and under serpentine edaphic conditions. Molecular analysis of invasive populations revealed that Californian populations cluster into three independent introductions (i.e. invasive lineages). Our glasshouse common garden experiment found that all Californian populations exhibited higher fitness under serpentine conditions. However, the three invasive lineages appear to represent independent pathways of adaptation to serpentine soil. Our results suggest that the rapid invasion of serpentine habitats in California may have been facilitated by the existence of colonizing Eurasian genotypes pre-adapted to serpentine soils.

The role of adaptive trans-generational plasticity in biological invasions of plants.

High-impact biological invasions often involve establishment and spread in disturbed, high-resource patches followed by establishment and spread in biotically or abiotically stressful areas. Evolutionary change may be required for the second phase of invasion (establishment and spread in stressful areas) to occur. When species have low genetic diversity and short selection history, within-generation phenotypic plasticity is often cited as the mechanism through which spread across multiple habitat types can occur. We show that trans-generational plasticity (TGP) can result in pre-adapted progeny that exhibit traits associated with increased fitness both in high-resource patches and in stressful conditions. In the invasive sedge, Cyperus esculentus, maternal plants growing in nutrient-poor patches can place disproportional number of propagules into nutrient-rich patches. Using the invasive annual grass, Aegilops triuncialis, we show that maternal response to soil conditions can confer greater stress tolerance in seedlings in the form of greater photosynthetic efficiency. We also show TGP for a phenological shift in a low resource environment that results in greater stress tolerance in progeny. These lines of evidence suggest that the maternal environment can have profound effects on offspring success and that TGP may play a significant role in some plant invasions.

Not all ski slopes are created equal: disturbance intensity affects ecosystem properties.

In mountain regions around the world, downhill ski areas represent a significant source of anthropogenic disturbance while also providing recreation and revenue. Ski-run creation always results in some level of disturbance, but disturbance intensity varies greatly with construction method. Ski runs may be established either by clearing (cutting and removing tall vegetation) or by clearing and then machine-grading (leveling the soil surface with heavy equipment). To quantify how these different intensities of initial disturbance affect ecosystem properties, we extensively surveyed vegetation, soils, and environmental characteristics on cleared ski runs, graded ski runs, and adjacent reference forests across seven large downhill ski resorts in the northern Sierra Nevada, USA. We found that the greater disturbance intensity associated with grading resulted in greater impacts on all ecosystem properties considered, including plant community composition and diversity, soil characteristics relating to processes of nutrient cycling and retention, and measures of erosion potential. We also found that cleared ski runs retained many ecological similarities to reference forests and might even offer some added benefits by possessing greater plant species and functional diversity than either forests or graded runs. Because grading is more damaging to multiple indicators of ecosystem function, clearing rather than grading should be used to create ski slopes wherever practical.

Understanding the consequences of seed dispersal in a heterogeneous environment.

Plant distributions are in part determined by environmental heterogeneity on both large (landscape) and small (several meters) spatial scales. Plant populations can respond to environmental heterogeneity via genetic differentiation between large distinct patches, and via phenotypic plasticity in response to heterogeneity occurring at small scales relative to dispersal distance. As a result, the level of environmental heterogeneity experienced across generations, as determined by seed dispersal distance, may itself be under selection. Selection could act to increase or decrease seed dispersal distance, depending on patterns of heterogeneity in environmental quality with distance from a maternal home site. Serpentine soils, which impose harsh and variable abiotic stress on non-adapted plants, have been partially invaded by Erodium cicutarium in northern California, USA. Using nearby grassland sites characterized as either serpentine or non-serpentine, we collected seeds from dense patches of E. cicutarium on both soil types in spring 2004 and subsequently dispersed those seeds to one of four distances from their maternal home site (0, 0.5, 1, or 10 m). We examined distance-dependent patterns of variation in offspring lifetime fitness, conspecific density, soil availability, soil water content, and aboveground grass and forb biomass. ANOVA revealed a distinct fitness peak when seeds were dispersed 0.5 m from their maternal home site on serpentine patches. In non-serpentine patches, fitness was reduced only for seeds placed back into the maternal home site. Conspecific density was uniformly high within 1 m of a maternal home site on both soils, whereas soil water content and grass biomass were significantly heterogeneous among dispersal distances only on serpentine soils. Structural equation modeling and multigroup analysis revealed significantly stronger direct and indirect effects linking abiotic and biotic variation to offspring performance on serpentine soils than on non-serpentine soils, indicating the potential for soil-specific selection on seed dispersal distance in this invasive species.

Factors driving distribution limits in an annual plant community.

Studies examining plant distribution patterns across environmental gradients have generally focused on perennial-dominated systems, and we know relatively little about the processes structuring annual communities. Here, the ecological factors determining local distribution patterns of five dominant annual species distributed across micro-topographic gradients in ephemeral California wetlands are examined. Over two growing seasons in three vernal pools, patterns of inundation and above-ground biomass were characterized across the microtopographic gradient, population boundaries for five dominant species were documented and a reciprocal transplant experiment and neighbor removal treatment were conducted to test the relative effects of within-pool elevation, competition and seed dispersal on plant performance. Despite large differences in inundation time between growing seasons, above-ground biomass and the elevation of population boundaries remained consistent. The predicted 'optimal' depth for each species shifted between years, but competition and recruitment limitation restricted species' abilities to track these conditions. The distributions of the focal taxa are primarily driven by differential responses to environmental conditions associated with different microtopographic positions along pool inundation gradients, and are reinforced by competition and dispersal constraints. The relative importance of competition, other environmental factors and dispersal patterns appear to contrast with results obtained in systems dominated by perennial plants.

Multiple origins promote the ecological amplitude of allopolyploid Aegilops (Poaceae).

Polyploidy has been ubiquitous in plant evolution and is thought to be an important engine of biodiversity that facilitates speciation, adaptation, and range expansion. Polyploid species can exhibit higher ecological tolerance than their progenitor species. For allotetraploid species, this higher tolerance is often attributed to the existence of heterosis resulting from entire genome duplication. However, multiple origins of allopolyploid species may further promote their ecological success by providing genetic variability in ecological traits underlying local adaptation and range expansion. Here we show in a group of allopolyploid species in the genus Aegilops that range size and abundance are correlated with the number of inferred origins. We found that allopolyploid Aegilops spp. contain multiple chloroplast haplotypes, each identical to haplotypes of the diploid progenitor species, indicating multiple origins as the major source of variation. The number of inferred origins in each allopolyploid species was correlated to the total area occupied by the allopolyploid and the tendency for the species to be common. Additionally, we found differences in ecological tolerance among independent origins in Aegilops triuncialis. These results strongly support the hypothesis that the introduction of genetic variability by multiple origins can increase the ecological amplitude and evolutionary success of allopolyploid species.

Functional ecology of ecotypic differentiation in the Californian serpentine sunflower (Helianthus exilis).

* Here, we examined phenotypic differences between locally adapted serpentine and riparian populations of the serpentine sunflower Helianthus exilis from northern California, USA. * Within a common environment, plants from serpentine and riparian sites were grown in regular potting soil or serpentine soil. Physiology, morphology, phenology and fitness-related traits were measured. * Overall, riparian plants grew more rapidly, attained a larger final size, produced larger leaves, and smaller flowering heads. Riparian plants also invested less in root biomass and were more water-use-efficient than the serpentine plants. Serpentine and riparian plants also differed in leaf concentrations of boron, magnesium, sodium and molybdenum. * These ecotypic differences suggest contrasting adaptive strategies to cope with either edaphic stress in serpentine sites or intense above-ground competition at riparian sites. There was a significant population origin x soil type crossing interaction in one fitness trait (average dry weight) that mirrored local adaptation previously documented for these riparian and serpentine ecotypes. However, because all other fitness traits did not exhibit this crossing interaction in our common garden study, it is possible that phenotypic differences underlying local adaptation may be amplified in the field as a result of biotic and abiotic interactions.

Local adaptation, patterns of selection, and gene flow in the Californian serpentine sunflower (Helianthus exilis).

The traditional view of the species as the fundamental unit of evolution has been challenged by observations that in heterogeneous environments, gene flow may be too restricted to overcome the effects of local selection. Whether a species evolves as a cohesive unit depends critically on the dynamic balance between homogenizing gene flow among populations and potentially disruptive local adaptation. To examine this evolutionary balance between "global" gene flow and local selection, we studied northern Californian populations of Helianthus exilis, the serpentine sunflower, within a mosaic of contrasting serpentine and nonserpentine areas that differ considerably in soil chemistry and water availability. Local adaptation to riparian and serpentine habitats was studied in Helianthus exilis along with an analysis of gene flow patterns among populations within these habitats. Local adaptation was assessed in H. exilis during 2002 and 2003 using reciprocal transplant experiments at multiple locations within serpentine and riparian habitats. Effects of competition and germination date on the expression of local adaptation were also examined within the reciprocal transplant experiments. Local adaptation was detected in both years at the local site level and at the level of habitat. The analysis of the transplanted populations indicated that the patterns of selection differed considerably between riparian and serpentine sites. Differential survivorship occurred in serpentine habitats, whereas selection on reproductive output predominated in riparian habitats. Local adaptation was expressed only in the absence of competition. Local adaptation in terms of survivorship was most strongly expressed in treatments with delayed seed germination. Microsatellite markers were used to quantify population genetic parameters and examine the patterns of gene flow among sampled populations. Analysis of molecular markers revealed a system of population patches that freely exchange genes with each other. Strong selection seems to maintain ecotypic variation within this endemic sunflower species, while extensive gene flow among populations prevents local speciation between serpentine and riparian ecotypes.

Variation in resource availability changes the impact of invasive thistles on native bunchgrasses.

The threat posed by invasive nonnative plants to native plant populations is one of the largest challenges facing both conservation biology and restoration ecology. California has been highly impacted by invaders, although many relict stands of native plants are found on shallow, rocky soils with limited resources. The abiotic conditions of these sites may strongly influence the performance of an invasive plant and its effect on resident native species. In addition, the maturity of native plants in these sites may modulate an invader's impact; larger, well-established plants may be better able to resist invaders. In this study we examined how the impact of an invasive thistle (Centaurea solstitialis) on a native perennial bunchgrass (Nassella pulchra) changed in response to variation in soil depth, soil water availability, and bunchgrass maturity. We measured plant performance in terms of survival, growth, reproduction, and predawn water potential. We found that soil depth, water availability, and bunchgrass maturity acted in concert to influence the impact of the invasive thistle on the native bunchgrass. Both species performed better in deep soils, especially during dry years. The combination of shallow soil and low water availability reduced C. solstitialis performance and ameliorated its negative effect on N. pulchra growth and reproduction. Higher water availability resulted in a stronger negative effect of C. solstitialis on N. pulchra in both shallow and deep soils. However, as N. pulchra matured and increased in size, we saw a steady decline in C. solstitialis growth and reproductive output. Higher water availability increased the performance of C. solstitialis in shallow soils. C. solstitialis may thus have a stronger impact on N. pulchra and be more able to invade relict stands of N. pulchra in shallow soils during high-rainfall years. However, established stands of N. pulchra appear to be more resistant to invasion by C. solstitialis as N. pulchra plants grow older and larger.