Metabolite and gene expression analysis reveal the molecular mechanism for petal colour variation in six Centaurea cyanus cultivars.

Centaurea cyanus is a popular garden plant native to Europe. Although their petals show abundant colour variations, the flavonoid profiling and the potential molecular mechanisms remain unclear. In the present study, we collected six cornflower cultivars with white, pink, red, blue, mauve and black petals. Ultra-performance liquid chromatography coupled with photodiode array and tandem mass spectrometry (UPLC-MS/MS) was used to investigate the comparative profiling of flavonoids both qualitatively and quantitatively. Ten anthocyanins, six flavones and two flavonols were separated and putatively identified. Except for white petals without any anthocyanins, both pink and red flowers contained pelargonidin derivatives, whereas blue, mauve and black petals accumulated cyanidins. The expression patterns of genes involved in the flavonoid biosynthesis were performed by real-time quantitative reverse transcription-PCR. The anthocyanin biosynthetic pathway in white petals was inhibited starting from flavanone 3-hydroxylase, resulting in the absence of anthocyanin accumulation. The open reading frame of flavonoid 3'-hydroxylase in pink and red petals was truncated; this led to loss of a haem binding site, a conserved motif in the cytochrome P450 family, and loss of conversion from dihydrokaempferol to dihydroquercetin. The significantly higher expression of structural genes corresponding to the hyper-accumulation of flavonoids in black petals may play an important role in black coloration. Remarkably, the mauve and blue petals accumulated the same cyanidin derivative but contained apigenin with different modifications on the 4' position, which may cause the coloration differences. The results obtained in this study will provide insights into the mechanisms of vivid colour diversities in cornflower.

Expansion history and environmental suitability shape effective population size in a plant invasion.

The margins of an expanding range are predicted to be challenging environments for adaptation. Marginal populations should often experience low effective population sizes (Ne ) where genetic drift is high due to demographic expansion and/or census population size is low due to unfavourable environmental conditions. Nevertheless, invasive species demonstrate increasing evidence of rapid evolution and potential adaptation to novel environments encountered during colonization, calling into question whether significant reductions in Ne are realized during range expansions in nature. Here we report one of the first empirical tests of the joint effects of expansion dynamics and environment on effective population size variation during invasive range expansion. We estimate contemporary values of Ne using rates of linkage disequilibrium among genome-wide markers within introduced populations of the highly invasive plant Centaurea solstitialis (yellow starthistle) in North America (California, USA), and within native Eurasian populations. As predicted, we find that Ne within the invaded range is positively correlated with both expansion history (time since founding) and habitat quality (abiotic climate). History and climate had independent additive effects with similar effect sizes, indicating an important role for both factors in this invasion. These results support theoretical expectations for the population genetics of range expansion, though whether these processes can ultimately arrest the spread of an invasive species remains an unanswered question.

Is population genetic structure of vascular plants shaped more by ecological or geographic factors? A study case on the Mediterranean endemic Centaurea filiformis (Asteraceae).

All known populations of the Sardinian endemic Centaurea filiformis Viv. (Asteraceae) were studied in order to understand the impact of both geographic and ecological factors on the genetic structuring of this species. Fourteen populations and 234 individuals were sampled. The demographic structure of the populations and the reproductive ecology were estimated in 28 plots. Population genetic analyses were based on SSR markers. Genetic structure was investigated by spatial Bayesian methods. Average densities of 0.51 individuals m-2 were detected, with a prevalence of adults. Ten species of pollinators were identified; C. filiformis ability to self-pollinate and myrmecochory were demonstrated experimentally. The populations displayed an average heterozygosity value of He  = 0.576 and high genetic differentiation (overall FST  = 0.218). Bayesian analysis suggests that five is the most probable number of gene pools of origin. A strong correlation between geographic distances and genetic distances among populations was highlighted. The demographic population structure of C. filiformis is dominated by adults, suggesting that it is a stable-regressive or senile species, investing more in local persistence than colonisation ability. Despite the scattered distribution, the populations studied do not present evidence of genetic erosion. The analysis of genetic differentiation reveals very high differentiation levels among populations, thus indicating that effective barriers exist against gene flow. A general conclusion is that population distribution results in a clear genetic structure for the populations studied, and that geography and not ecology is shaping the present distribution of this species.

Local pre-adaptation to disturbance and inbreeding-environment interactions affect colonisation abilities of diploid and tetraploid Centaurea stoebe.

Primary colonisation in invasive ranges most commonly occurs in disturbed habitats, where anthropogenic disturbance may cause physical damage to plants. The tolerance to such damage may differ between cytotypes and among populations as a result of differing population histories (adaptive differentiation between ruderal verus natural habitats). Moreover, founder populations often experience inbreeding depression, the effects of which may increase through physical damage due to inbreeding-environment interactions. We aimed to understand how such colonisation processes differ between diploid and tetraploid Centaurea stoebe populations, with a view to understanding why only tetraploids are invasive. We conducted a clipping experiment (frequency: zero, once or twice in the growing season) on inbred versus outbred offspring originating from 37 C. stoebe populations of varying cytotype, range and habitat type (natural versus ruderal). Aboveground biomass was harvested at the end of the vegetation period, while re-sprouting success was recorded in the following spring. Clipping reduced re-sprouting success and biomass, which was significantly more pronounced in natural than in ruderal populations. Inbreeding depression was not detected under benign conditions, but became increasingly apparent in biomass when plants were clipped. The effects of clipping and inbreeding did not differ between cytotypes. Adaptive differentiation in disturbance tolerance was higher among populations than between cytotypes, which highlights the potential of pre-adaptation in ruderal populations during early colonisation on anthropogenically disturbed sites. While the consequences of inbreeding increased through clipping-mediated stress, they were comparable between cytotypes, and consequently do not contribute to understanding the cytotype shift in the invasive range.

Competition does not come at the expense of colonization in seed morphs with increased size and dispersal.

PREMISE OF THE STUDY: Seed-level trade-offs of heterocarpic species remain poorly understood. We propose that seedlings emerging from seeds with a permanent pappus (dispersing seeds) are stronger competitors than those emerging from seeds without a pappus (nondispersing seeds) because dispersing seeds are larger and germinate faster than nondispersing seeds in Centaurea solstitialis. METHODS: We conducted a competition experiment with both seed morphs, in which we recorded emergence rate and proportion, estimated seed dispersal by wind (anemochory) and by mammals (exozoochory), and measured size and abundance of seed morphs. KEY RESULTS: We found that seedlings from pappus seeds had greater competitive abilities than those from non-pappus seeds. Similarly, pappus seedlings emerged at much faster rates and larger proportions than non-pappus seedlings. Pappus seeds were larger, were more numerous, and displayed improved exozoochory compared to non-pappus seeds. Anemochory was poor for both seed morphs. CONCLUSIONS: We found support for our hypothesis, raising in turn the possibility that competition and colonization are positively associated in seed morphs of heterocarpic species with enhanced exozoochory of larger seeds. These findings are not consistent with those from heterocarpic species with enhanced anemochory of smaller seeds or slower-germinating seeds. Our results additionally suggest that pappus and non-pappus seeds of C. solstitialis display a task-division strategy in which pappus morphs colonize and preempt unoccupied sites through improved dispersal and fast and large emergence of seedlings with increased competitive abilities, whereas non-pappus morphs promote site persistence through delayed germination and dormancy. This strategy may contribute to the success of C. solstitialis in highly variable environments.

Neighbour Origin and Ploidy Level Drive Impact of an Alien Invasive Plant Species in a Competitive Environment.

Our understanding of the potential mechanisms driving the spread and naturalization of alien plant species has increased over the past decades, but specific knowledge on the factors contributing to their increased impact in the introduced range is still urgently needed. The native European plant Centaurea stoebe occurs as two cytotypes with different life histories (monocarpic diploids, allo-polycarpic tetraploids). However, only tetraploids have been found in its introduced range in North America, where C. stoebe has become a most prominent plant invader. Here, we focus on the ploidy level of C. stoebe and origin of neighbouring community in explaining the high impact during the invasion of new sites in the introduced range. We conducted a mesocosm experiment under open-field conditions with the diploid (EU2x) and tetraploid (EU4x) cytotype of Centaurea stoebe from its native European (EU) range, and with the invasive tetraploid (NA4x) cytotype from the introduced North American (NA) range in competition with EU (old) or NA (new) neighbouring plant communities. In the presence of competition, the biomass of EU neighbouring community was reduced to a comparable level by all three geo-cytotypes of C. stoebe. In contrast, the biomass of the NA neighbouring community was reduced beyond when competing with tetraploid, but not with diploid C. stoebe. The fact that the biomass of all three geo-cytotypes of C. stoebe was correlated with the biomass of the EU neighbouring community, but not with that of the NA neighbouring community suggests that different mechanisms underlie the competitive interactions between C. stoebe and its old vs. new neighbouring communities, such as competition for the same limiting resources at home vs competition through novel allelo-chemicals or differential resource uptake strategies in the introduced range. We therefore caution to simply use the ecosystem impact assessed at home to predict impact in the introduced range.

Allogamy-Autogamy Switch Enhance Assortative Mating in the Allotetraploid Centaurea seridis L. Coexisting with the Diploid Centaurea aspera L. and Triggers the Asymmetrical Formation of Triploid Hybrids.

Hybridization between tetraploids and their related diploids is generally unsuccessful in Centaurea, hence natural formation of triploid hybrids is rare. In contrast, the diploid Centaurea aspera and the allotetraploid C. seridis coexist in several contact zones where a high frequency of triploid hybrids is found. We analyzed the floral biology of the three taxa to identify reproductive isolation mechanisms that allow their coexistence. Flowering phenology was recorded, and controlled pollinations within and between the three taxa were performed in the field. Ploidy level and germination of progeny were also assessed. There was a 50% flowering overlap which indicated a phenological shift. Diploids were strictly allogamous and did not display mentor effects, while tetraploids were found to be highly autogamous. This breakdown of self-incompatibility by polyploids is first described in Centaurea. The asymmetrical formation of the hybrid was also found: all the triploid intact cypselae came from the diploid mothers pollinated by the pollen of tetraploids. Pollen and eggs from triploids were totally sterile, acting as a strong triploid block. These prezygotic isolation mechanisms ensured higher assortative mating in tetraploids than in diploids, improving their persistence in the contact zones. However these mechanisms can also be the cause of the low genetic diversity and high genetic structure observed in C. seridis.

Bioactive Compounds in Wild, In vitro Obtained, Ex vitro Adapted, and Acclimated Plants of Centaurea davidovii (Asteraceae).

In vitro cultures were initiated from a single seed of Centaurea davidovii. Whole plantlets were regenerated and cultivated for several months on agar-solidified nutrient media differing by their composition: basal MS medium, MS medium supplemented with plant growth regulators, and liquid MS medium. Plantlets were ex vitro adapted and successfully acclimated to open-air conditions; flowering was observed in some individuals in the first summer, and mass flowering during the second summer. The contents of the total flavonoids and the total phenolic compounds were determined spectrophotometrically in the leaves of the in vitro plantlets cultured on different media, and then compared with those in the leaves of the wild plants and in the leaves of the acclimated plants of the field plot. The sesquiterpene lactone 8α-(5'-hydroxyangeloyl)-salonitenolide was determined by HPLC in leaf samples of C. davidovii wild plants, in vitro obtained plantlets and ex vitro acclimated plants in the greenhouse and on the experimental field plot. The composition of the nutrient medium influenced the contents of all studied bioactive substances. The highest concentrations of all tested secondary metabolites were detected in the leaves of the acclimated plants during mass flowering, the content of the lactone reaching 56.2 mg/g DW, which was several times more than in the other leaf samples. The obtained results revealed both the effectiveness of biotechnological methods for propagation and conservation of rare and endangered plant species, and the possibility to use C. davidovii plants ex vitro acclimated to field conditions as a source of secondary metabolites with potential biological activity.

Origin matters: diversity affects the performance of alien invasive species but not of native species.

At local scales, it has often been found that invasibility decreases with increasing resident plant diversity. However, whether resident community diversity similarly resists invasion by alien versus native species is seldom studied. We examined this issue by invading constructed native plant assemblages that varied in species and functional richness with invasive alien or native Asteraceae species. Assemblages were also invaded with spotted knapweed, Centaurea stoebe, a native European aster that has been previously used in diversity-invasibility experiments in North America. We also conducted a field survey to explore the generality of the patterns generated from our experimental study. Both experimental and observational work revealed that increasing diversity reduced the performance of alien but not native invaders. Centaurea stoebe invading its native community performed poorly regardless of resident diversity, whereas in a parallel, previously published study conducted in North America, C. stoebe easily invaded low-diversity but not high-diversity assemblages. Our results suggest that diversity is an attribute of resident communities that makes them more or less susceptible to invasion by novel invasive alien but not native plant species.

Biogeographic effects on early establishment of an invasive alien plant.

PREMISE OF THE STUDY: Biotic resistance is often studied in the context of how interactions between native biota and invading species influence the success of those invaders. Seldom, however, is the strength of "resistance" compared biogeographically, where the ability of a species to impede invader establishment is contrasted between an invader's native and introduced recipient community. METHODS: We conducted an experiment to examine how community diversity influences seedling recruitment of a plant invader where it is native and contrasted with results previously published from introduced ranges. In Switzerland, we created recipient communities that varied in species and functional richness and invaded them, or not, with seeds of Centaurea stoebe, a native European plant that has been previously used in an identical experiment in North America, where it is a prominent invader. KEY RESULTS: The biogeographic comparison revealed that the recipient community largely prevented C. stoebe seedling establishment at home (Switzerland), but not away (Montana, USA), and that diversity of the resident vegetation did not contribute to the effects observed in the introduced range. CONCLUSIONS: Our results provide evidence that differences in the biogeographic conditions and/or overall level of competition of resident community between the native and introduced range considerably suppresses seedling recruitment of the invasive plant, rather than resident diversity itself. In the case of C. stoebe, the surprisingly low establishment success in the experiments conducted in the native compared with the introduced range is likely to be influenced by the higher level of competition with resident community, by abiotic environmental conditions or interactions between these two factors in the native range. Release from factors suppressing seedling recruitment at home may contribute to the successful invasion of C. stoebe in North America.

Fragment size does not matter when you are well connected: effects of fragmentation on fitness of coexisting gypsophiles.

Most habitat fragmentation studies have focused on the effects of population size on reproductive success of single species, but studies assessing the effects of both fragment size and connectivity, and their interaction, on several coexisting species are rare. In this study, we selected 20 fragments along two continuous gradients of size and degree of isolation in a gypsum landscape in central Spain. In each fragment, we selected 15 individuals of each of three dominant gypsophiles (Centaurea hyssopifolia, Lepidium subulatum and Helianthemum squamatum, 300 plants per species, 900 plants in total) and measured several reproductive traits: inflorescence number, fruit set, seed set and seed mass. We hypothesised that plant fitness would be lower on small and isolated fragments due to an interaction between fragment size and connectivity, and that response patterns would be species-specific. Overall, fragment size had very little effect on reproductive traits compared to that of connectivity. We observed a positive effect of fragment connectivity on C. hyssopifolia fitness, mediated by the increased seed predation in plants from isolated fragments, resulting in fewer viable seeds per capitulum and lower seed set. Furthermore, seed mass was lower in plants from isolated fragments for both C. hyssopifolia and L. subulatum. In contrast, few reproductive traits of H. squamatum were affected by habitat fragmentation. We discuss the implications of species-specific responses to habitat fragmentation for the dynamics and conservation of gypsum plant communities. Our results highlight the complex interplay among plants and their mutualistic and antagonistic visitors, and reinforce the often-neglected role of habitat connectivity as a key component of the fragmentation process.

Contemporary evolution of plant reproductive strategies under global change is revealed by stored seeds.

Global change is expected to impose new selection pressures on natural populations. Phenotypic responses, such as earlier phenology in response to climate warming, have been repeatedly observed in the field. The recent pollinator decline is also expected to change selection on reproductive traits in flowering plants. However, it remains unclear whether short-term adaptation of plant reproductive strategies occurs in response to global change. In this study, we report the evolution of some important reproductive traits of the annual self-incompatible weed Centaurea cyanus. In a common garden experiment, we germinated stored seeds, sampled 18 years apart from the same location, in a region where warmer springs and indices of pollinator decline have been reported. Compared to the ancestral population (1992), our results showed that plants of the descendant population (2010) flowered earlier and also produced larger capitula with longer receptivity and a larger floral display. QST -FST comparisons indicated that natural selection has likely contributed to the evolution of some of the traits investigated. Lower FST within temporal samples than among spatial samples further suggests a limited role of gene flow from neighbouring populations. We therefore propose that trait shifts could partly be due to adaptation to global change.

Drought tolerance and plasticity in the invasive knapweed Centaurea stoebe s.l. (Asteraceae): effect of populations stronger than those of cytotype and range.

BACKGROUND AND AIMS: Spotted knapweed (Centaurea stoebe s.l., Asteraceae) is native to Europe, where it occurs as a diploid (2xEU) and tetraploid cytotype (4xEU), but so far only the tetraploid has been reported in the introduced range in North America (4xNA). In previous studies, significant range shifts have been found towards drier climates in 4xEU compared with 2xEU, and in 4xNA when compared with the native range. In addition, 4x plants showed thicker leaves and reduced specific leaf area compared with 2x plants, suggesting higher drought tolerance in 4x plants. It is thus hypothesized that the 4x cytotype might be better pre-adapted to drought than the 2x, and the 4xNA better adapted than the 4xEU due to post-introduction selection. METHODS: Plants of the three geocytotypes (2xEU, 4xEU and 4xNA ), each represented by six populations, were subjected to three water treatments over 6 weeks in a greenhouse experiment. Plasticity and reaction norms of above- and below-ground biomasses and their ratio, survival rate, stomatal conductance and carbon isotope discrimination were analysed using linear and generalized linear mixed effect models. KEY RESULTS AND CONCLUSIONS: Above-ground and total biomasses of European tetraploids were slightly less affected by drought than those of European diploids, and 4xEU plants maintained higher levels of stomatal conductance under moderate drought than 4xNA plants, thus supporting the pre-adaptation but not the post-introduction evolution hypothesis. Plasticity indexes for most of the traits were generally higher in 2xEU and 4xNA than in 4xEU plants, but these differences were not or were only marginally significant. Interestingly, the effect of population origin and its interaction with treatment was more important than the effects of geocytotype and range. Population means for the control treatment showed several significant associations either with latitude or some aspect of climatic data, suggesting evolution of local adaptations, especially within the 2xEU and 4xEU geocytotypes.

Fungal endophyte increases the allelopathic effects of an invasive forb.

Endophytic plant symbionts can have powerful effects on the way their hosts interact with pathogens, competitors, and consumers. The presence of endophytes in plants can alter food webs, community composition and ecosystem processes, suggesting that endophyte-plant symbioses may represent unique forms of extended phenotypes. We tested the impact of the fungal endophyte Alternaria alternata (phylotype CID 120) on the allelopathic effect of the invasive forb Centaurea stoebe when in competition with the North American native bunchgrass Koeleria macrantha in a greenhouse competition experiment. The allelopathic effect of C. stoebe on K. macrantha when infected with the fungal endophyte was more than twice that of endophyte-free C. stoebe. However, this allelopathic effect was a small part of the very large competitive effect of C. stoebe on K. macrantha in all treatments, likely because of the priority effects in our experimental design. To our knowledge, these results are the first experimental evidence for a symbiotic relationship between plants and fungal endophytes affecting allelopathic interactions between competing plants, and thus provide insight into the mechanisms by which fungal endophytes may increase the competitive ability of their hosts.

Pollinator interactions with yellow starthistle (Centaurea solstitialis) across urban, agricultural, and natural landscapes.

Pollinator-plant relationships are found to be particularly vulnerable to land use change. Yet despite extensive research in agricultural and natural systems, less attention has focused on these interactions in neighboring urban areas and its impact on pollination services. We investigated pollinator-plant interactions in a peri-urban landscape on the outskirts of the San Francisco Bay Area, California, where urban, agricultural, and natural land use types interface. We made standardized observations of floral visitation and measured seed set of yellow starthistle (Centaurea solstitialis), a common grassland invasive, to test the hypotheses that increasing urbanization decreases 1) rates of bee visitation, 2) viable seed set, and 3) the efficiency of pollination (relationship between bee visitation and seed set). We unexpectedly found that bee visitation was highest in urban and agricultural land use contexts, but in contrast, seed set rates in these human-altered landscapes were lower than in natural sites. An explanation for the discrepancy between floral visitation and seed set is that higher plant diversity in urban and agricultural areas, as a result of more introduced species, decreases pollinator efficiency. If these patterns are consistent across other plant species, the novel plant communities created in these managed landscapes and the generalist bee species that are favored by human-altered environments will reduce pollination services.

Biocontrol attack increases pollen limitation under some circumstances in the invasive plant Centaurea solstitialis.

Herbivore damage often deters pollinator visitation and many invasive plants in North America are pollinator-dependent. This has important implications for the biological control of invasive plants because it means that agents that deter pollinators may have a larger than expected impact on the plant. Yet interactions between pollinators and biocontrol agents are rarely evaluated. Centaurea solstitialis, one of the most problematic invasive species in California, is dependent on pollinators for reproduction. I factorially manipulated infection by a biocontrol pathogen and pollen supplementation to test for (1) pollen limitation in C. solstitialis, (2) whether infection increased pollen limitation, and (3) whether this varied across a soil moisture gradient. Plants growing on north-facing slopes where soil moisture was higher experienced mild pollen limitation in the absence of the pathogen and more pronounced pollen limitation when they were infected. Plants on drier south-facing slopes did not suffer from pollen limitation but instead appeared to suffer from resource limitation. Pathogen infection directly reduced seed set in C. solstitialis by 67-72%. On north-facing slopes, infection had an additional, indirect effect by increasing the degree of pollen limitation plants experienced. The trait that mediates this indirect pathogen-pollinator interaction is the number of inflorescences plants produced: infected plants made fewer inflorescences which led to greater pollen limitation. Although in the present study this outcome is dependent on abiotic factors that vary over small spatial scales, exploiting other invasive plants' dependence on pollinators by selecting agents that deter visitation may enhance agent impact.

Rapid evolution of an invasive weed.

Trade-offs between performance and the ability to tolerate abiotic and biotic stress have been suggested to explain both the success of invasive species and phenotypic differentiation between native and invasive populations. It is critical to sample broadly across both ranges and to account for latitudinal clines and maternal effects when testing this premise. Wild-collected Centaurea diffusa seeds were grown in benign and stressful conditions (drought, flooding, nutrient stress and simulated herbivory), to evaluate whether native and introduced individuals differ in performance or life history phenotypes. A second experiment used glasshouse-grown seeds to evaluate whether patterns remain comparable when maternal environment is consistent. Many traits differed between ranges, and in all cases but one, invasive individuals grew larger, performed better, or matured later. No trade-off in performance with herbivore defense was evident. Invasive populations may have been released from a trade-off between growth and drought tolerance apparent in the native range. Larger individuals with delayed maturity and greater reproductive potential have evolved in invasive populations, a pattern evident across broad population sampling, and after latitude and maternal environment were considered. Release from abiotic stress tolerance trade-offs may be important for the invasion success of Centaurea diffusa.

Dispersal pathways and genetic differentiation among worldwide populations of the invasive weed Centaurea solstitialis L. (Asteraceae).

The natural history of introduced species is often unclear due to a lack of historical records. Even when historical information is readily available, important factors of the invasions such as genetic bottlenecks, hybridization, historical relationships among populations and adaptive changes are left unknown. In this study, we developed a set of nuclear, simple sequence repeat markers and used these to characterize the genetic diversity and population structure among native (Eurasian) and non-native (North and South American) populations of Centaurea solstitialis L., (yellow starthistle). We used these data to test hypotheses about the invasion pathways of the species that were based on historical and geographical records, and we make inferences about historical relationships among populations and demographic processes following invasion. We confirm that the center of diversity and the native range of the species is likely the eastern Mediterranean region in the vicinity of Turkey. From this region, the species likely proceeded to colonize other parts of Europe and Asia via a slow, stepwise range expansion. Spanish populations were the primary source of seed to invade South America via human-mediated events, as was evident from historical records, but populations from the eastern Mediterranean region were also important. North American populations were largely derived from South America, but had secondary contributors. We suggest that the introduction history of non-native populations from disparate parts of the native range have allowed not just one, but multiple opportunities first in South America then again in North America for the creation of novel genotypes via intraspecific hybridization. We propose that multiple intraspecific hybridization events may have created especially potent conditions for the selection of a noxious invader, and may explain differences in genetic patterns among North and South America populations, inferred differences in demographic processes, as well as morphological differences previously reported from common garden experiments.

Dissecting impact of plant invaders: do invaders behave differently in the new range?

Knowledge from basic plant ecology suggests that impact of one plant species on another is driven by either competition for the same limiting resources, or by unique plant traits. These processes might be context specific, explaining a differential impact of exotic plant invaders in the native vs. introduced range. With the help of a conceptual framework, we aimed at identifying the relationship between invader biomass and impact in the invasive Centaurea stoebe by conducting pairwise competition experiments with 15 European (old) and 15 North American (new) neighboring species. Old neighbors grew larger and could use available soil moisture more efficiently for growth than new neighbors. Interestingly, biomass of C. stoebe explained a substantial amount of the variation in biomass of the coevolved neighbors, but not of the new "naive" neighbors. Thus, impact in the home range appears to be driven by competition for the same limiting resources, but by other factors in the introduced range, possibly by exploitation of resources that are not used by the new neighbors or by interference competition. This distinction has important consequences for the management of invasive species, as in our study ecosystem recovery is less likely after simple biomass reduction.

Invasive and non-invasive congeners show similar trait shifts between their same native and non-native ranges.

Differences in morphological or ecological traits expressed by exotic species between their native and non-native ranges are often interpreted as evidence for adaptation to new conditions in the non-native ranges. In turn this adaptation is often hypothesized to contribute to the successful invasion of these species. There is good evidence for rapid evolution by many exotic invasives, but the extent to which these evolutionary changes actually drive invasiveness is unclear. One approach to resolving the relationship between adaptive responses and successful invasion is to compare traits between populations from the native and non-native ranges for both exotic invaders and congeners that are exotic but not invasive. We compared a suite of morphological traits that are commonly tested in the literature in the context of invasion for three very closely related species of Centaurea, all of which are sympatric in the same native and non-native ranges in Europe and North America. Of these, C. solstitialis is highly invasive whereas C. calcitrapa and C. sulphurea are not. For all three species, plants from non-native populations showed similar shifts in key traits that have been identified in other studies as important putative adaptive responses to post-introduction invasion. For example, for all three species plants from populations in non-native ranges were (i) larger and (ii) produced seeds that germinated at higher rates. In fact, the non-invasive C. calcitrapa showed the strongest trait shift between ranges. Centaurea solstitialis was the only species for which plants from the non-native range increased allocation to defensive spines, and allocated proportionally less resources to reproduction, patterns contrary to what would be predicted by theory and other empirical studies to enhance invasion. Our results suggest caution when interpreting the commonly observed increase in size and reproductive capacity as factors that cause exotics to become invaders.