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.     

Centaurea solstitialis Invasion Success Is Influenced by Nassella pulchra Size

Replacement of perennial grasses with non‐native annual grasses in California's Central Valley grasslands and foothills has increased deep soil water availability. Yellow starthistle (Centaurea solstitialis), a deep‐rooted invasive thistle, can use this water to invade annual grasslands. Native perennial bunchgrasses, such as Purple needlegrass (Nassella pulchra), also use deep soil water, so there is an overlap in resource use between N. pulchra and C. solstitialis. Restoration of N. pulchra to annual grasslands could result in strong competitive interactions between N. pulchra and C. solstitialis, which may reduce survival, growth, and reproduction of the invader. The strength of this competitive interaction can increase as N. pulchra plants mature, increase in size, and develop more extensive root systems. We studied how the size of N. pulchra affected the success of C. solstitialis invasion over 2 years. We allowed C. solstitialis seed to fall naturally into plots containing N. pulchra plants. For each plot, we measured the number of C. solstitialis seedlings and mature plants, as well as C. solstitialis biomass and seedhead production. In both years of the study, C. solstitialis number, biomass, and seedhead production declined significantly as N. pulchra size increased. However, even C. solstitialis grown with the largest N. pulchra plants produced some seed, especially during the higher rainfall year. We conclude that restoration plantings with larger, established N. pulchra plants will be more resistant to invasion by C. solstitialis than young N. pulchra plantings, but site management must continue as long as a C. solstitialis seed source is present.     

Back to the basics: using density series to test regulation versus limitation for invasive plants

Numerous hypotheses have been invoked to explain invasion in plant communities. Here, we use a fundamental tool from plant ecology, density series in situ, to explore whether a reasonable starting point for highly successful invasive plant species is to consider regulation (biotic effects) and limitation (environmental constraints). To explore the utility of density series to understanding invasion we present a case study using C. solstitialis, a perfect candidate since it is a prolific seed producer and invasive in many grasslands globally. Using seed addition into existing vegetation in three grasslands with densities of up to 6500 seeds per m², we found no evidence for regulation via intra or interspecific interference but large differences amongst sites. This strongly suggests that in this species limitations imposed by the environment are the only constraints. Hence, we propose that an excellent starting point for ecologists studying invasion should involve back to the basics experiments on novel species to determine whether other hypotheses need be invoked.     

Geographic mosaics of plant–soil microbe interactions in a global plant invasion

Aim Our aim in this study was to document the global biogeographic variation in the effects of soil microbes on the growth of Centaurea solstitialis (yellow starthistle; Asteraceae), a species that has been introduced throughout the world, but has become highly invasive only in some introduced regions. Location  To assess biogeographic variation in plant–soil microbe interactions, we collected seeds and soils from native Eurasian C. solstitialis populations and introduced populations in California, Argentina and Chile. Methods To test whether escape from soil‐borne natural enemies may contribute to the success of C. solstitialis, we compared the performance of plants using seeds and soils collected from each of the biogeographic regions in greenhouse inoculation/sterilization experiments. Results  We found that soil microbes had pervasive negative effects on plants from all regions, but these negative effects were significantly weaker in soils from non‐native ranges in Chile and California than in those from the non‐native range in Argentina and the native range in Eurasia. Main conclusions The biogeographic differences in negative effects of microbes in this study conformed to the enemy‐release hypothesis (ERH) overall, but the strong negative effect of soil biota in Argentina, where C. solstitialis is invasive, and weaker effects in Chile where it is not, indicated that different factors influencing invasion are likely to occur in large scale biogeographic mosaics of interaction strengths.     

The seedbanks of the Breckland heaths and heath grasslands, eastern England, and their relationship to the vegetation and the effects of management

The vegetation and the associated germinable seedbank of a range of heath and grass heath communities in Breckland, eastern England, were compared. The composition of vegetation and seedbank were similar for heather-dominated communities due to the predominance of Calluna vulgaris . Recent rotovation of previously heather-dominated communities invaded by Deschampsia flexuosa reduced the number of emerging C. vulgaris seedlings by more than 80%. The composition of seedbanks of calcicolous grasslands differed considerably from the vegetation, with a significant contribution from many ruderal species not present in the vegetation and an absence of important species such as Pilosella officinarum. Acidic grasslands were intermediate between calcicolous grasslands and heather-dominated heathlands, with seedbanks characterized by high densities of Rumex acetosella and a number of ruderal species that were absent or rare in the vegetation. The composition of the seedbank of more mesotrophic grasslands suggests that the imposition of some form of disturbance may recreate more open grasslands typical of Breckland.     

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.     

Distribution,density and impact of invasive plants in Lake Nakuru National Park,Kenya

Invasive plants have invaded swathes of grasslands in Lake Nakuru National Park thus necessitating the Park management to institute measures to control them. Despite this, information on the status and impact of invasive plants in these grasslands is lacking. Six grassland types were identified and assigned random numbers. Five study sites were then randomly selected from each grassland type. The Point‐Centre Quarter method was used to determine the distribution and density of seven major invasive plants, whilst their impact was assessed by comparing biomass and crude protein content of important forage grasses in invaded and noninvaded grasslands. Results show that the distribution of invasive plants did not differ among the grasslands (F_{5, 30} = 1.47, P = 0.229) but their densities differed among the plants (F_{6, 30} = 20.99, P < 0.001). The mean biomass in invaded and noninvaded grasslands (22 g m^?2 versus 37.2 g m^?2) was significantly different (t‐test, P < 0.05), while crude protein content of grasses in invaded (7.73, % DM) and noninvaded (9.3, % DM) patches was not significantly different (t‐test; P > 0.05). Our results suggest that invasive plants lower grass production and hence might reduce food availability to grazers.     

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.     

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.     

Multi-scale impacts of crested wheatgrass invasion in mixed-grass prairie

Most of North America’s northern Great Plains have been cultivated for crop production, leaving remnants of natural mixed-grass prairie fragmented and threatened by alien plant invasions. The region’s most widespread alien perennial forage crop, crested wheatgrass (Agropyron cristatum sensu amplo), has invaded native grassland and raised concerns regarding its ecological impact. To evaluate impacts at multiple scales of organization, adjacent invaded and uninvaded mixed-grass prairie were sampled at eight widely separated locations. At the population level, native C₃ mid-grasses and forbs were less abundant in invaded grasslands, while native C₃ and C₄ short-grass abundance was not different. At community and landscape levels, diversity was lower in invaded grasslands largely because of lower forb species richness and cover, and crested wheatgrass dominance of both cover (14% basal cover) and seedbank (404 seeds m⁻²). At the ecosystem level, both vegetation and litter biomass were greater in invaded grasslands, however, below ground organic matter (roots and litter), soil organic carbon, total nitrogen and phosphorus were not different. Crested wheatgrass invasion of mixed-grass prairie was associated with lower diversity within and among plant communities, and appears to simplify the composition of mixed-grass prairie landscapes. Hypotheses for crested wheatgrass dominance and persistence following invasion are suggested.     

Selective and Non-Selective Control of Invasive Plants: The Short-Term Effects of Growing-Season Prescribed Fire, Herbicide, and Mowing in Two Texas Prairies

Conservation of North American grasslands is hampered by the impact of invasive herbaceous species. Selective control of these plants, although desirable, is complicated by the shared physiology and phenology of the invader and the native components of the invaded plant community. Fortunately, there is evidence that some management practices, such as prescribed fire, herbicide, and mowing, can cause differential responses in native and invasive grassland species. However, timing of treatment is critical, and fire has been shown to increase rates of invasion when implemented during the dormant season. Bothriochloa ischaemum, an introduced C4 Eurasian grass is an increasing problem in grasslands, particularly in southern and central regions of North America. To date, there has been little success in effective selective control. Two invaded grassland sites representative of Blackland Prairie and Edwards Plateau ecoregions were subjected to two growing‐season prescribed fire treatments, single and double herbicide applications, and single and double mowing treatments. Mowing had no effect on either B. ischaemum or other dominant species at either site one‐year posttreatment. However, growing‐season fire and herbicide were both effective at reducing the abundance of B. ischaemum, with other codominant species responding either negatively to herbicide or neutrally or positively to fire. The vulnerability of B. ischaemum to growing‐season fire may be associated with the ecology of its native range. The negative growth response to growing‐season fire, combined with its lower implementation costs, indicates that this method warrants further investigation as a selective management tool for other problematic species in invaded grasslands.     

Bacterial Communities Associated with <Emphasis Type="Italic">Chenopodium album</Emphasis> and <Emphasis Type="Italic">Stellaria media</Emphasis> Seeds from Arable Soils

The bacterial community compositions in Chenopodium album and Stellaria media seeds recovered from soil (soil weed seedbank), from bulk soil, and from seeds harvested from plants grown in the same soils were compared. It was hypothesized that bacterial communities in soil weed seedbanks are distinct from the ones present in bulk soils. For that purpose, bacterial polymerase chain reaction denaturing gradient gel electrophoresis (PCR–DGGE) fingerprints, made from DNA extracts of different soils and seed fractions, were analyzed by principal component analysis. Bacterial fingerprints from C. album and S. media seeds differed from each other and from soil. Further, it revealed that bacterial fingerprints from soil-recovered and plant-harvested seeds from the same species clustered together. Hence, it was concluded that microbial communities associated with seeds in soil mostly originated from the mother plant and not from soil. In addition, the results indicated that the presence of a weed seedbank in arable soils can increase soil microbial diversity. Thus, a change in species composition or size of the soil weed seedbank, for instance, as a result of a change in crop management, could affect soil microbial diversity. The consequence of increased diversity is yet unknown, but by virtue of identification of dominant bands in PCR–DGGE fingerprints as Lysobacter oryzae (among four other species), it became clear that bacteria potentially antagonizing phytopathogens dominate in C. album seeds in soil. The role of these potential antagonists on weed and crop plant growth was discussed.     

Spatial variation of seedling distribution in an east Mediterranean pine woodland at the beginning of post-fire succession

Most of the area in pine woodlands is occupied by perennial seeders that regenerate from seeds in the first winter after the fire and by annuals. Control of the germination in the regenerating vegetation after wildfire is therefore a primary ecological component of the post-fire succession in this ecosystem. The aim of the study presented here was to determine the distribution of Pinus, Cistus and other plants seeds around burned Pinus halepensis trees, and to measure the conditions related to seed germination in the upper soil layers in the same locations. The study was carried out in a 50-year old planted Pinus halepensis woodland that was burned down by a wildfire in July 1995. The variation of seedbank density was determined by collecting samples under the canopies of burned trees and in a nearby open area. Pine seedbank density decreased and that of Cistus and annuals increased with increasing distance from the burned trunks. Most pine seeds were present in the ash layer while those of the other plants were in the soil. In situ germination experiments showed that seedling density decreased with distance from the burned trunks while the proportion of pines in the seedling population increased. This was a result of seedbank variation and germination inhibition by the high pH conditions caused by the ash. The establishment of sparse pine seedling under the dead tree canopies insured their rapid development without interference by other plants and played a key role in the regeneration and stability of the pine woodland community. The concomitant mass germination of the perennial seeders in the rest of the area prevented invasion by annuals.     

Allelopathic effects of three plant invaders on germination of native species: a field study

The ability of some invasive plant species to produce biochemical compounds toxic to native species, called allelopathy, is thought to be one of the reasons for their success when introduced to a novel range, an idea known as the Novel Weapons Hypothesis. However, support for this hypothesis mainly comes from bioassays and experiments conducted under controlled environments, whereas field evidence is rare. In a field experiment, we investigated whether three plant species invasive in Europe, Solidago gigantea, Impatiens glandulifera and Erigeron annuus, inhibit the germination of native species through allelopathy more than an adjacent native plant community. At three sites for each invasive species, we compared the germination of native species that were sown on invaded and non-invaded plots. Half of these plots were amended with activated carbon to reduce the influence of potential allelopathic compounds. The germination of sown seeds and of seeds from the seedbank was monitored over a period of 9 weeks. Activated carbon generally enhanced seed germination. This effect was equally pronounced in invaded and adjacent non-invaded plots, indicating that invasive species do not suppress germination more than a native plant community. In addition, more seeds germinated from the seedbank on invaded than on non-invaded soil, probably due to previous suppression of germination by the invasive species. Our field study does not provide evidence for the Novel Weapons Hypothesis with respect to the germination success of natives. Instead, our results suggest that if invasive species release allelopathic compounds that suppress germination, they do so to a similar degree as the native plant community.     

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.     

Soil biota composition and the performance of a noxious weed across its invaded range

The success of invasive plant species is driven, in part, by feedback with soil ecosystems. Yet, how variation in belowground communities across latitudinal gradients affects invader distributions remains poorly understood. To determine the effect of soil communities on the performance of the noxious weed Cirsium arvense across its invaded range, we grew seedlings for 40 days in soils collected across a 699 km linear distance from both inside and outside established populations. We also described the mesofaunal and bacterial communities across all soil samples. We found that C. arvense typically performed better when grown in soils sourced from northern populations than from southern locations where it has a longer invasion history. We also found evidence that C. arvense performed best in soils sourced from outside invaded patches, although this was not consistent across all sites. The bacterial community showed a significant increase in the magnitude of compositional change in invaded sites at higher latitudes, while the mesofaunal community showed the opposite pattern. Bacterial community composition was significantly correlated with C. arvense performance, although mesofaunal community composition was not. Our results demonstrate that the interactions between an invasive plant and associated soil communities change across the invaded range, and the bacterial community in particular may affect variation in plant performance. Observed patterns may be caused by C.arvense presence and time since invasion allowing for an accumulation of species‐specific pathogens in southern soils, while the naïveté of northern soils to invasion results in a more responsive bacterial community. Although these interactions are difficult to predict, such effects could possibly facilitate the establishment of this exotic species to novel locations.     

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.     

Interspecific variation in persistence of buried weed seeds follows trade‐offs among physiological,chemical, and physical seed defenses

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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.     

A comparison of native and invasive populations of three clonal plant species in Germany and New Zealand

Aim Our aim was to test for changes in growth patterns of three clonally growing plant species (Achillea millefolium, Hieracium pilosella and Hypericum perforatum) between native and invaded regions. We addressed the hypotheses that with differing important life‐history traits, invasive populations perform better than native populations, and that this expected better performance is linked to weakened trade‐offs between individual growth and sexual and clonal reproduction. Location Germany and New Zealand. Methods We conducted field surveys for the three above‐mentioned species in both native German and invasive New Zealand populations, and collected data at both population and individual levels. Results At the population level, the proportion of flowering plants, population size and population density were all higher in invasive populations. Similarly, at the individual level, the number of stolons per plant, stolon–biomass ratio and population crowdedness (local plant density in a specified area around a target plant) were significantly higher in New Zealand. Plant height did not differ between countries, and plant biomass was lower in New Zealand than in Germany for Achillea millefolium and Hypericum perforatum. These two species showed significant trade‐offs between individual growth and sexual and clonal reproduction. Achillea millefolium exhibited a weakened trade‐off in its invaded range, where the same proportion of flowering plants was sustained at much higher levels of population crowdedness than in its native range. Main conclusions The apparent invasion success of the three study species is generally due to better overall performance in their respective invaded ranges. In respect of both Achillea millefolium and Hypericum perforatum, this is driven primarily by increased vegetative reproduction. In contrast, Hieracium pilosella seems to benefit more from increased sexual reproduction in its invaded range. Shifts in trade‐offs as a general trend seem to be of minor importance.