Resistance to Centaurea solstitialis invasion from annual and perennial grasses in California and Argentina

A common explanation for Centaurea solstitialis invasion in California is that it occupies an ??empty niche?? created by the replacement of native perennial grasses by exotic annual grasses and concomitant increases in soil water availability. This hypothesis, however, cannot explain C. solstitialis invasion into perennial-dominated grasslands of central Argentina. We assessed invasibility of annual versus perennial grass communities in these regions through parallel field experiments where we created grass plots and, after one year of establishment, measured effects on water and light, and added C. solstitialis seeds in two successive trials. Additionally, we removed vegetation around naturally occurring C. solstitialis in both regions, and examined the performance of Californian and Argentinean C. solstitialis individuals when growing under common conditions simulating climate in California and Argentina. In California, both grass types offered high resistance to C. solstitialis invasion, water was generally greater under perennials than annuals, and light was similarly low beneath both types. In Argentina, invasibility was generally greater in annual than perennial plots, water was similar between groups, and light was much greater beneath annuals. Removal experiments showed that competition from annual grasses in California and perennial grasses in Argentina greatly reduce C. solstitialis performance. Additionally, Californian and Argentinean individuals did not exhibit genetic differentiation in studied traits. Our results suggest that dominant plant functional groups in both California and Argentina offer substantial resistance to C. solstitialis invasion. The success of this species might be tightly linked to a remarkable ability to take advantage of disturbance in both regions.     

Range-expanding populations of a globally introduced weed experience negative plant-soil feedbacks

Background

Biological invasions are fundamentally biogeographic processes that occur over large spatial scales. Interactions with soil microbes can have strong impacts on plant invasions, but how these interactions vary among areas where introduced species are highly invasive vs. naturalized is still unknown. In this study, we examined biogeographic variation in plant-soil microbe interactions of a globally invasive weed, Centaurea solstitialis (yellow starthistle). We addressed the following questions (1) Is Centaurea released from natural enemy pressure from soil microbes in introduced regions? and (2) Is variation in plant-soil feedbacks associated with variation in Centaurea''s invasive success?

Methodology/Principal Findings

We conducted greenhouse experiments using soils and seeds collected from native Eurasian populations and introduced populations spanning North and South America where Centaurea is highly invasive and noninvasive. Soil microbes had pervasive negative effects in all regions, although the magnitude of their effect varied among regions. These patterns were not unequivocally congruent with the enemy release hypothesis. Surprisingly, we also found that Centaurea generated strong negative feedbacks in regions where it is the most invasive, while it generated neutral plant-soil feedbacks where it is noninvasive.

Conclusions/Significance

Recent studies have found reduced below-ground enemy attack and more positive plant-soil feedbacks in range-expanding plant populations, but we found increased negative effects of soil microbes in range-expanding Centaurea populations. While such negative feedbacks may limit the long-term persistence of invasive plants, such feedbacks may also contribute to the success of invasions, either by having disproportionately negative impacts on competing species, or by yielding relatively better growth in uncolonized areas that would encourage lateral spread. Enemy release from soil-borne pathogens is not sufficient to explain the success of this weed in such different regions. The biogeographic variation in soil-microbe effects indicates that different mechanisms may operate on this species in different regions, thus establishing geographic mosaics of species interactions that contribute to variation in invasion success.     

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

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

The importance of factors controlling species abundance and distribution varies in native and non‐native ranges

How variation in factors controlling species abundance and distribution between native and non‐native ranges compares to that within ranges remains poorly understood. We used a globally distributed ruderal, Centaurea solstitialis (Centaurea), to explore the possibility that the importance of those factors exhibits great variation between and within ranges. To test our hypothesis, we established seed addition experiments with soil disturbance (turnover and control) and biocide (fungicides, insecticide, and control) treatments in two regions within native (the Caucasus and south‐western Turkey) and non‐native (the western United States – US – and central Argentina) distributions. Also, we estimated the rate of vegetation recovery after disturbance (resilience) and related it to Centaurea density in experimental plots. Disturbance strongly increased Centaurea density in all regions. Density was similar between the native Caucasus and non‐native Argentina and much greater in those regions than in the native Turkey and non‐native US in biocide‐free plots. Fungicides had positive effects on density in the US and negative ones in the Caucasus and Argentina, resulting in no differences between those three regions and greater density in the US than Turkey. Insecticide applications promoted Centaurea density in Turkey and Argentina, but inter‐regional comparisons of density in treated plots were comparable to those in biocide‐free plots. Overall, plants were smaller and less fecund in Turkey than the other regions, except the US. The greatest fungal attack was documented in Turkey, and herbivory was stronger there and in Argentina than in the Caucasus and US. The resilience of the local community explained a large proportion of variation in Centaurea density. These results support our hypothesis, and reveal that the speed at which competition is re‐gained after disturbance may influence global variation in Centaurea abundance. Because many ruderals exhibit native and non‐native distributions, our results are likely to be generalized to other systems.     

Invasive species grows faster,competes better,and shows greater evolution toward increased seed size and growth than exotic non-invasive congeners

Comparisons of introduced exotics that invade and those that do not can yield important insights into the ecology of invasions. Centaurea solstitialis, C. calcitrapa, and C. sulphurea are closely related, share a similar life history and were each introduced to western North America from Southern Europe ~100–200 years ago. However, of these three species, only C. solstitialis has become invasive. We collected seeds from different populations for each of the three species both in the native range of Spain and the non-native range of California, measured individual seed mass, and grew plants from these seeds in a greenhouse experiment in Montana. The invasive C. solstitialis had the smallest seeds and seedlings of the three congeners. However, in contrast to its non-invasive congeners, C. solstitialis had the highest relative growth rates when grown in competition. C. solstitialis was also the only species to show significant differences in traits between populations from different ranges, with plants from the non-native range of California demonstrating greater competitive resistance, larger seed size, and larger seedling mass than plants from the native range in Spain. This suggests that C. solstitialis may be evolving toward larger seed and seedling sizes in this non-native range. Relative growth rate showed no inter-regional variation for any species, but was higher for C. solstitialis than its congeners when in competition, and thus may interact with the evolution of larger seeds and plant mass in ways that contribute to the extraordinary invasive success of this species.     

Species interactions contribute to the success of a global plant invader

Biological invasions are ubiquitous ecological phenomena that often impact native ecosystems. Some introduced species have evolved traits that enhance their ability to compete and dominate in recipient communities. However, it is still unknown if introduced species can evolve traits that may enhance their species interactions to fuel invasion success. We tested whether Centaurea solstitialis (yellow starthistle) from introduced populations have greater performance than native counterparts, and whether they generate more beneficial plant-soil interactions. We used common garden and plant-soil feedback experiments with soils and seeds from native Eurasian and introduced Californian populations. We found that performance of Centaurea did not differ among source genotypes, implying that the success of this invasive species is not due to evolutionary changes. However, Centaurea grew significantly larger in soils from introduced regions than from native regions, indicating a reduction in natural enemy pressure from native populations. We conclude that species interactions, not evolution, may contribute to Centaurea’s invasion success in introduced populations.     

Defoliation of Centaurea solstitialis Stimulates Compensatory Growth and Intensifies Negative Effects on Neighbors

Efforts to arrest the spread of invasive weeds with herbivory may be hindered by weak effects of the herbivores or strong compensatory responses of the invaders. We conducted a greenhouse experiment to study the effects of defoliation and soil fungi on competition between the invasive weed Centaurea solstitialis and C. solstitialis and Avena barbata, a naturalized Eurasian annual grass, and Nassella pulchra, a native California bunchgrass. Surprisingly, considering the explosive invasion of grasslands by C. solstitialis, Avena and Nassella were strong competitors and reduced the invader’s biomass by 80.2% and 80.1% over all defoliation and soil fungicide treatments, respectively. However, our experiments were conducted in artificial environments where competition was probably accentuated. When fungicide was applied to the soil, the biomass of C. solstitialis was reduced in all treatment combinations, but reduction in the biomass of the invader had no corollary impact on the grasses. There was no overall effect of defoliation on the final biomass of C. solstitialis as the invader compensated fully for severe clipping. In fact, the directional trend of the clipping effect was +6.4% over all treatments after eight weeks. A significant neighbor × soil fungicide × clipping effect suggested that the compensatory response was the strongest without soil fungicide and when C. solstitialis was alone (+ 19%). Our key finding was that the compensatory response of C. solstitialis in all treatments was associated with an increase in the weed’s negative effects on Nassella and Avena – there was a significant decrease in the total biomass of both grasses and the reproductive biomass of Avena in pots with clipped C. solstitialis. Our results were obtained in controlled conditions that may have been conducive to compensatory growth, but they suggest the existence of mechanisms that may allow C. solstitialis, like other Centaurea species, to resist herbivory.     

The Role of Light and Soil Moisture in Plant Community Resistance to Invasion by Yellow Starthistle (Centaurea solstitialis)

To resist establishment by an invasive plant, a community may require one or more species functionally similar to the invader in their resource acquisition pattern. In this study, communities consisting of native winter annual forbs, non‐native annual grasses, native perennials, or a combination of the two native communities were established with and without Centaurea solstitialis to determine the effect of soil moisture and light availability on plant community invasion resistance. The annual plant communities were unable to resist invasion by C. solstitialis. In the native winter annual forb community, senescence in late spring increased light penetration (>75%) to the soil surface, allowing seeded C. solstitialis to quickly establish and dominate the plots. In addition, native annual forbs utilized only shallow soil moisture, whereas C. solstitialis used shallow and deep soil moisture. In communities containing native perennials, only Elymus glaucus established well and eventually dominated the plots. During the first 2 years of establishment, water use pattern of perennial communities was similar to native annual forbs and resistance to invasion was associated with reduced light availability during the critical stages of C. solstitialis establishment. In later years, however, water use pattern of perennial grass communities was similar or greater than C. solstitialis‐dominated plots. These results show that Central Valley grasslands that include E. glaucus resist C. solstitialis invasion by a combination of light suppression and soil water competition. Spatiotemporal resource utilization patterns, and not just functional similarity, should be considered when developing restoration strategies to resist invasion by many non‐native species.     

Germination responses of an invasive species in native and non-native ranges

Studying germination in the native and non‐native range of a species can provide unique insights into processes of range expansion and adaptation; however, traits related to germination have rarely been compared between native and non‐native populations. In a series of common garden experiments, we explored whether differences in the seasonality of precipitation, specifically, summer drought vs summer rain, and the amount and variation of annual and seasonal precipitation affect the germination responses of populations of an annual ruderal plant, Centaurea solstitialis, from its native range and from two non‐native regions with different climates. We found that seeds from all native populations, irrespective of the precipitation seasonality of the region in which they occurred, and non‐native populations from regions with dry summers displayed similarly high germination proportions and rates. In contrast, genotypes from the non‐native region with predominantly summer rain exhibited much lower germination fractions and rates. Also, percent germination was strongly correlated with variation in precipitation in winter, the season that follows germination for C. solstitialis. Specifically, germination was lower for native and non‐native populations experiencing greater variation in winter precipitation. This correlation, however, was greatly influenced by the non‐native region with summer rain, which also exhibited the greatest variation in winter precipitation among studied regions. These results suggest that rather than general climatic patterns, the degree of risk experienced at early developmental stages could exert an important control over the germination strategy of C. solstitialis populations in both native and non‐native ranges. Also, these findings reveal a largely unique germination response in C. solstitialis genotypes growing in the non‐native region with summer rain and high variation in winter precipitation. Our work raises the possibility that rapid adaptive changes in germination strategies may contribute to the success of globally distributed invaders.     

Non‐native conditions favor non‐native populations of invasive plant: demographic consequences of seed size variation?

Trait differences between native and non‐native populations may explain the greater abundance and impact of some organisms in their non‐native ranges than in their native ranges. Here, we conducted reciprocal common gardens in southwestern Turkey (home) and central Argentina (away) to explore the hypothesis that the greater success of the invasive ruderal Centaurea solstitialis in Argentina than Turkey is partially explained by differences between home and away populations. Unusual among common gardens, our experimental design included seed additions to explicitly evaluate population level responses, as well as disturbance and no‐disturbance treatments. We documented seed mass in native and non‐native populations, and during the experiment, we periodically measured density, plant size, and herbivory. After six months, we determined the establishment of plants for populations from both origins in both home and away common gardens. Seed mass was two times larger for Argentinean than Turkish populations. Density, plant size and final establishment were also greater for plants from Argentinean than from Turkish populations, but only in the common garden in Argentina. In Turkey, no differences between population origins were detected for these variables. Herbivory was similar for populations from both origins in both common gardens. As expected, disturbance generally increased plant performance in both regions. Our results suggest that increased seed size in non‐native populations may have demographic consequences under non‐native conditions that can contribute to the invasive success of C. solstitialis. This is the first reciprocal common garden that supports the idea that seed size variation contributes to demographic differences for an invasive species between native and non‐native distributions, but our findings further suggest that seed size effects on demography depend on the ecological context in which population processes occur.     

No evidence for root-mediated allelopathy in Centaurea solstitialis, a species in a commonly allelopathic genus

Phytotoxicity bioassays and pot experiments using activated carbon both suggest that Centaurea solstitialis (yellow star-thistle) does not rely on phytotoxic root exudates for invasion of California grasslands. Pot experiments in which five native species were grown in the presence/absence of C. solstitialis and in the presence/absence of activated carbon (fully crossed design) showed that C. solstitialis competitively suppressed native species, but did not inhibit them through allelochemicals. In separate experiments examining the role of root exudates in invasion success, treatment with crude root exudates and chloroform-extracted root exudates from C. solstitialis reduced growth of the model plant Arabidopsis thaliana. However, high concentrations of the exudates (50%, v/v or 500 μg mL⁻¹) were required to inhibit A. thaliana growth and did not result in A. thaliana mortality, suggesting the presence of only a weak growth inhibitor. Moreover, high concentrations of C. solstitialis crude root exudates did not affect the growth of five native grass species often displaced by C. solstitialis invasions in California grasslands. Finally, root exudates collected from C. solstitialis had weaker effects on a native California root parasite, Triphysaria versicolor, than root exudates collected from Zea mays, a species not renowned for its competitive or invasive capabilities. Our results suggest that, while C. solstitialis might possibly “be persuaded to yield a product that is toxic to one species or another” (Population biology of plants, Academic, 1977), we find no evidence that allelopathic root exudates play a role in the competitive success of this invasive.     

Traits correlate with invasive success more than plasticity: A comparison of three Centaurea congeners

The importance of phenotypic plasticity for successful invasion by exotic plant species has been well studied, but with contradictory and inconclusive results. However, many previous studies focused on comparisons of native and invasive species that co‐occur in a single invaded region, and thus on species with potentially very different evolutionary histories. We took a different approach by comparing three closely related Centaurea species: the highly invasive C. solstitialis, and the noninvasive but exotic C. calcitrapa and C. sulphurea. These species have overlapping distributions both in their native range of Spain and in their non‐native range of California. We collected seeds from 3 to 10 populations from each region and species and grew them in common garden greenhouse conditions to obtain an F1 generation in order to reduce maternal effects. Then, F1 seeds were grown subjected to simulated herbivory, variation in nutrient availability, and competition, to explore plasticity in the responses to these conditions. We found little variation in phenotypic plasticity among species and regions, but C. solstitialis plants from California produced more biomass in competition than their Spanish conspecifics. This species also had the highest relative growth rates when in competition and when grown under low nutrient availability. Noninvasive congeners produced intermediate or opposite patterns.     

Linking the pattern to the mechanism: How an introduced mammal facilitates plant invasions

Non‐native mammals that are disturbance agents can promote non‐native plant invasions, but to date there is scant evidence on the mechanisms behind this pattern. We used wild boar (Sus scrofa) as a model species to evaluate the role of non‐native mammals in promoting plant invasion by identifying the degree to which soil disturbance and endozoochorous seed dispersal drive plant invasions. To test if soil disturbance promotes plant invasion, we conducted an exclosure experiment in which we recorded emergence, establishment and biomass of seedlings of seven non‐native plant species planted in no‐rooting, boar‐rooting and artificial rooting patches in Patagonia, Argentina. To examine the role of boar in dispersing seeds we germinated viable seeds from 181 boar droppings and compared this collection to the soil seed bank by collecting a soil sample adjacent to each dropping. We found that both establishment and biomass of non‐native seedlings in boar‐rooting patches were double those in no‐rooting patches. Values in artificial rooting patches were intermediate between those in boar‐rooting and no‐rooting treatments. By contrast, we found that the proportion of non‐native seedlings in the soil samples was double that in the droppings, and over 80% of the germinated seeds were native species in both samples. Lastly, an effect size test showed that soil disturbance by wild boar rather than endozoochorous dispersal facilitates plant invasions. These results have implications for both the native and introduced ranges of wild boar, where rooting disturbance may facilitate community composition shifts.     

Soil conditioning effects of Phragmites australis on native wetland plant seedling survival

Interactions between introduced plants and soils they colonize are central to invasive species success in many systems. Belowground biotic and abiotic changes can influence the success of introduced species as well as their native competitors. All plants alter soil properties after colonization but, in the case of many invasive plant species, it is unclear whether the strength and direction of these soil conditioning effects are due to plant traits, plant origin, or local population characteristics and site conditions in the invaded range. Phragmites australis in North America exists as a mix of populations of different evolutionary origin. Populations of endemic native Phragmites australis americanus are declining, while introduced European populations are important wetland invaders. We assessed soil conditioning effects of native and non‐native P. australis populations on early and late seedling survival of native and introduced wetland plants. We further used a soil biocide treatment to assess the role of soil fungi on seedling survival. Survival of seedlings in soils colonized by P. australis was either unaffected or negatively affected; no species showed improved survival in P. australis‐conditioned soils. Population of P. australis was a significant factor explaining the response of seedlings, but origin (native or non‐native) was not a significant factor. Synthesis: Our results highlight the importance of phylogenetic control when assessing impacts of invasive species to avoid conflating general plant traits with mechanisms of invasive success. Both native (noninvasive) and non‐native (invasive) P. australis populations reduced seedling survival of competing plant species. Because soil legacy effects of native and non‐native P. australis are similar, this study suggests that the close phylogenetic relationship between the two populations, and not the invasive status of introduced P. australis, is more relevant to their soil‐mediated impact on other plant species.     

Shifts in priming partly explain impacts of long‐term nitrogen input in different chemical forms on soil organic carbon storage

Input of labile organic carbon can enhance decomposition of extant soil organic carbon (SOC) through priming. We hypothesized that long‐term nitrogen (N) input in different chemical forms alters SOC pools by altering priming effects associated with N‐mediated changes in plants and soil microbes. The hypothesis was tested by integrating field experimental data of plants, soil microbes and two incubation experiments with soils that had experienced 10 years of N enrichment with three chemical forms (ammonium, nitrate and both ammonium and nitrate) in an alpine meadow on the Tibetan Plateau. Incubations with glucose–¹³C addition at three rates were used to quantify effects of exogenous organic carbon input on the priming of SOC. Incubations with microbial inocula extracted from soils that had experienced different long‐term N treatments were conducted to detect effects of N‐mediated changes in soil microbes on priming effects. We found strong evidence and a mechanistic explanation for alteration of SOC pools following 10 years of N enrichment with different chemical forms. We detected significant negative priming effects both in soils collected from ammonium‐addition plots and in sterilized soils inoculated with soil microbes extracted from ammonium‐addition plots. In contrast, significant positive priming effects were found both in soils collected from nitrate‐addition plots and in sterilized soils inoculated with soil microbes extracted from nitrate‐addition plots. Meanwhile, the abundance and richness of graminoids were higher and the abundance of soil microbes was lower in ammonium‐addition than in nitrate‐addition plots. Our findings provide evidence that shifts toward higher graminoid abundance and changes in soil microbial abundance mediated by N chemical forms are key drivers for priming effects and SOC pool changes, thereby linking human interference with the N cycle to climate change.     

The small‐scale spatiotemporal pattern of the seedbank and vegetation of a highly invasive weed,Centaurea solstitialis: strength in numbers

The dynamics of invasive plant populations are intriguing and informative of the importance of population and community‐level processes. A dominant approach to understanding and describing invasion has been the development of unique hypotheses to explain invasion. However, here we directly explore the relevance of the small‐scale, spatiotemporal pattern in seedbanks and plants of the highly invasive weed, Centaurea solstitialis, to determine whether pattern can be used to contrast predictions associated with the simple ecological hypotheses of seed versus microsite limitations. At three invaded grasslands in California, highly invaded (> 20 adult plants present), invaded (< 10 adults), and uninvaded (no C. solstitialis plants) sites were selected. The spatial pattern of the seedbank was assessed using fine‐scale, 2 cm diameter contiguous cores and geostatistical statistics, and the number of C. solstitialis seeds in the seedbank was recorded in addition to the total community seedbank density. Three of the four critical predictions associated with the seed limitation hypothesis were clearly supported as an explanation for the patterns of C. solstitialis invasion observed in the field. The density of C. solstitialis seeds decreased from high to low extents of invasion, there was no relationship between the community seedbank and C. solstitialis seeds, and the distances between C. solstitialis plants was inversely related to the density of C. solstitialis seeds. However, both the persistent and transient seedbanks of C. solstitialis were spatially aggregated with autocorrelation up to 12 cm² which suggests that aggregation is a consistent attribute of this species in the seedbank regardless of extent of invasion. This basic pattern‐based approach clearly detected an ecological signal of invasive seedbank dynamics and is thus a useful tool for subsequent studies of invasions in grasslands.     

Functionally Similar Species Confer Greater Resistance to Invasion: Implications for Grassland Restoration

Plant community functional composition can be manipulated in restored ecosystems to reduce the establishment potential of invading species. This study was designed to compare invasion resistance among communities with species functionally similar or dissimilar to yellow starthistle (Centaurea solstitialis), a late‐season annual. A field experiment was conducted in the Central Valley of California with six experimental plant communities that included (1) six early‐season native annual forbs (AF); (2) five late‐season native perennials and one summer annual forb (NP); (3) a combination of three early‐season native annual forbs and three late‐season native perennials (FP); (4) six early‐season non‐native annual grasses (AG); (5) monoculture of the late‐season native perennial grass Elymus glaucus (EG); and (6) monoculture of the late‐season native perennial Grindelia camporum (GC). Following establishment, C. solstitialis seed was added to half of the plots, and a monoculture of C. solstitialis (CS) was established as a control. Over a 5‐year period, the AF and AG communities were ineffective at preventing C. solstitialis invasion. Centaurea solstitialis cover remained less than 10% in the FP and NP communities, except in year 1. By the fourth year, E. glaucus cover was greater than 50% in NP and FP communities and had spread to all other communities (e.g., 27% cover in CS in year 5). Communities containing E. glaucus, which is functionally similar to C. solstitialis, better resisted invasion than communities lacking a functional analog. In contrast, G. camporum, which is also functionally similar to C. solstitialis, failed to survive. Consequently, species selection for restored communities must consider not only functional similarity to the invader but also establishment success, competitiveness, and survivorship.     

Restoration of Native Perennials in a California Annual Grassland after Prescribed Spring Burning and Solarization

Grasslands dominated by exotic annual grasses have replaced native perennial vegetation types in vast areas of California. Prescribed spring fires can cause a temporary replacement of exotic annual grasses by native and non‐native forbs, but generally do not lead to recovery of native perennials, especially where these have been entirely displaced for many years. Successful reintroduction of perennial species after fire depends on establishment in the postfire environment. We studied the effects of vegetation changes after an April fire on competition for soil moisture, a key factor in exotic annual grass dominance. As an alternative to fire, solarization effectively kills seeds of most plant species but with a high labor investment per area. We compared the burn to solarization in a study of establishment and growth of seeds and transplants of the native perennial grass Purple needlegrass (Nassella pulchra) and coastal sage species California sagebrush (Artemisia californica). After the fire, initial seed bank and seedling densities and regular percent cover and soil moisture (0–20 cm) data were collected in burned and unburned areas. Burned areas had 96% fewer viable seeds of the dominant annual grass, Ripgut brome (Bromus diandrus), leading to replacement by forbs from the seed bank, especially non‐native Black mustard (Brassica nigra). In the early growing season, B. diandrus dominating unburned areas consistently depleted soil moisture to a greater extent between rains than forbs in burned areas. However, B. diandrus senesced early, leaving more moisture available in unburned areas after late‐season rains. Nassella pulchra and A. californica established better on plots treated with fire and/or solarization than on untreated plots. We conclude that both spring burns and solarization can produce conditions where native perennials can establish in annual grasslands. However, the relative contribution of these treatments to restoration appears to depend on the native species being reintroduced, and the long‐term success of these initial restoration experiments remains to be determined.     

Virulence of oomycete pathogens from Phragmites australis‐invaded and noninvaded soils to seedlings of wetland plant species

Soil pathogens affect plant community structure and function through negative plant–soil feedbacks that may contribute to the invasiveness of non‐native plant species. Our understanding of these pathogen‐induced soil feedbacks has relied largely on observations of the collective impact of the soil biota on plant populations, with few observations of accompanying changes in populations of specific soil pathogens and their impacts on invasive and noninvasive species. As a result, the roles of specific soil pathogens in plant invasions remain unknown. In this study, we examine the diversity and virulence of soil oomycete pathogens in freshwater wetland soils invaded by non‐native Phragmites australis (European common reed) to better understand the potential for soil pathogen communities to impact a range of native and non‐native species and influence invasiveness. We isolated oomycetes from four sites over a 2‐year period, collecting nearly 500 isolates belonging to 36 different species. These sites were dominated by species of Pythium, many of which decreased seedling survival of a range of native and invasive plants. Despite any clear host specialization, many of the Pythium species were differentially virulent to the native and non‐native plant species tested. Isolates from invaded and noninvaded soils were equally virulent to given individual plant species, and no apparent differences in susceptibility were observed between the collective groups of native and non‐native plant species.     

Biogeographical variation in community response to root allelochemistry: novel weapons and exotic invasion

Centaurea diffusa is one of the most destructive invasive weeds in the western USA and allelopathy appears to contribute to its invasiveness ( Callaway & Aschehoug 2000 ). Here we identify a chemical from the root exudates of C. diffusa, 8‐hydroxyquinoline, not previously reported as a natural product, and find that it varies biogeographically in its natural concentration and its effect as an allelochemical. 8‐Hydroxyquinoline is at least three times more concentrated in C. diffusa‐invaded North American soils than in this weed's native Eurasian soils and has stronger phytotoxic effects on grass species from North America than on grass species from Eurasia. Furthermore, experimental communities built from North American plant species are far more susceptible to invasion by C. diffusa than communities built from Eurasian species, regardless of the biogeographical origin of the soil biota. Sterilization of North American soils suppressed C. diffusa more than sterilization of Eurasian soils, indicating that North American soil biota may also promote invasion by C. diffusa. Eurasian plants and soil microbes may have evolved natural resistance to 8‐hydroxyquinoline while North American plants have not, suggesting a remarkable potential for evolutionary compatibility and homeostasis among plants within natural communities and a mechanism by which exotic weeds destroy these communities.