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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. [ABSTRACT FROM AUTHOR]

Various factors have been shown contributing to the ecosystem impact of invasive alien plants, but their relative importance remains unclear. We focused on the effects of neighbouring plant community and soil biota as these biotic factors have been repeatedly put forward to explain invasion success (e.g. as components of the novel weapons and of the biotic release hypothesis)., To assess their relative importance in explaining the high impact of Centaurea stoebe during the invasion of new sites in the introduced range, we conducted a greenhouse experiment with both European ( EU) and North American ( NA) tetraploid C. stoebe competing with/without EU vs. NA neighbouring community and with the two neighbouring communities growing without C. stoebe. Plants were grown in sterilized commercial soil inoculated with naïve soil (from rhizosphere of plants other than C. stoebe) originating either from the home EU or the introduced NA range and half of which was sterilized to remove soil biota., In the competition pots, relative competitive ability (difference between the relative growth rates of C. stoebe and neighbouring community) and impact level (biomass of the neighbouring community relative to that in non-competition pots) of C. stoebe was significantly higher when grown with the NA than with the EU neighbouring community, although growth rates of EU and NA neighbouring communities did not differ in the non-competition pots. Both soil origin and C. stoebe origin had no effect on these processes. Soil sterilization increased growth of both C. stoebe and neighbouring communities, but had only a moderate effect on impact level and type, and no effect on the relative competitive ability of C. stoebe., These results suggest that during the colonization of new sites in North American grasslands, the impact of C. stoebe is strongly driven by reduced competitive ability of NA neighbours compared with EU neighbours, while altered biotic soil conditions in the introduced range and post-introduction evolutionary changes in the invader are of less importance. This differential impact appears to be due to inherently different mechanisms underlying the competitive interactions between EU and NA neighbouring communities when grown with C. stoebe. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Aim The spotted knapweed ( Centaurea stoebe), a plant native to south-east and central Europe, is highly invasive in North America. We investigated the spatio-temporal climatic niche dynamics of the spotted knapweed in North America along two putative eastern and western invasion routes. We then considered the patterns observed in the light of historical, ecological and evolutionary factors. Location Europe and North America. Methods The niche characteristics of the east and west invasive populations of spotted knapweed in North America were determined from documented occurrences over 120 consecutive years (1890-2010). For this investigation, the 2.5 and 97.5 percentiles of values along temperature and precipitation gradients, as given by the two first axes of a principal components analysis ( PCA), were calculated. We additionally measured the climatic dissimilarity between invaded sites and the native niche using a multivariate environmental similarity surface ( MESS) analysis. Results Along both invasion routes, the species established in regions with climatic conditions that were similar to those in the native niche. An initial spread in ruderal habitats always preceded spread in (semi-)natural habitats. In the east, the niche gradually increased over time until it reached limits similar to the native niche. Conversely, in the west the niche abruptly expanded after an extended time lag into climates not occupied in the native range; only the native cold niche limit was conserved. Main conclusions Our study reveals that different niche dynamics have taken place during the eastern and western invasions. This pattern indicates different combinations of historical, ecological and evolutionary factors in the two ranges. We hypothesize that the lack of a well-developed transportation network in the west at the time of the introduction of spotted knapweed confined the species to a geographically and climatically isolated region. The invasion of dry rangelands may have been favoured during the agricultural transition in the 1930s by release from natural enemies, local adaptation and less competitive vegetation, but further experimental and molecular studies are needed to explain these contrasting niche patterns fully. Our study illustrates the need and benefit of applying large-scale, temporally explicit approaches to understanding biological invasions. [ABSTRACT FROM AUTHOR]

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 "naïve" 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. [ABSTRACT FROM AUTHOR]

* Premise of the study: Because seeds have essential functions in the life cycle of plants, even subtle changes in their characteristics may have important demographic consequences. In this study, we examined whether potential changes in seed characteristics as a result of polyploidy or postintroduction evolution may have contributed to the invasion of Centaurea stoebe (Asteraceae). This plant occurs as diploid and tetraploid cytotypes in its native range in Europe, whereas only tetraploids have been found and become invasive in North America. Specific comparisons among these three "geo-cytotypes" allow us to explore hypotheses of preadaptation resulting from polyploidy (European diploids vs. European tetraploids) and postintroduction evolution (European tetraploids vs. North American tetraploids). * Methods: Using seeds collected from plants of each geo-cytotype grown in a common maternal environment, we compared seed mass, morphology (achene and pappus size), dispersal potential (falling velocity, seed roughness), survival, germination, and seedling emergence in a combination of laboratory, greenhouse and field experiments. * Key results: We found increased seed mass in North American tetraploids compared with European tetraploids. Seed morphology and dispersal potential were largely similar in all geo-cytotypes. Seed survival under field conditions was higher in native and invasive tetraploids compared with diploids. Germination in the laboratory was similar among all geo-cytotypes, but seedling emergence under field conditions was higher in invasive tetraploids than in the other geo-cytotypes. * Conclusions: Our findings suggest that a combination of preadaptation due to polyploidy (increased seed survival) and further postintroduction evolution in North American tetraploids (increased seed mass and seedling emergence) may have contributed to their invasion. [ABSTRACT FROM AUTHOR]

Biogeographic experiments that test how multiple interacting factors influence exotic plant abundance in their home and recipient communities are remarkably rare. We examined the effects of soil fungi, disturbance and propagule pressure on seed germination, seedling recruitment and adult plant establishment of the invasive Centaurea stoebe in its native European and non-native North American ranges., Centaurea stoebe can establish virtual monocultures in parts of its non-native range, but occurs at far lower abundances where it is native. We conducted parallel experiments at four European and four Montana ( USA) grassland sites with all factorial combinations of ± suppression of soil fungi, ±disturbance and low versus high knapweed propagule pressure [100 or 300 knapweed seeds per 0.3 m × 0.3 m plot (1000 or 3000 per m2)]. We also measured germination in buried bags containing locally collected knapweed seeds that were either treated or not with fungicide., Disturbance and propagule pressure increased knapweed recruitment and establishment, but did so similarly in both ranges. Treating plots with fungicides had no effect on recruitment or establishment in either range. However, we found: (i) greater seedling recruitment and plant establishment in undisturbed plots in Montana compared to undisturbed plots in Europe and (ii) substantially greater germination of seeds in bags buried in Montana compared to Europe. Also, across all treatments, total plant establishment was greater in Montana than in Europe., Synthesis. Our results highlight the importance of simultaneously examining processes that could influence invasion in both ranges. They indicate that under 'background' undisturbed conditions, knapweed recruits and establishes at greater abundance in Montana than in Europe. However, our results do not support the importance of soil fungi or local disturbances as mechanisms for knapweed's differential success in North America versus Europe. [ABSTRACT FROM AUTHOR]

The evolution of optimal life history strategies is central for the fitness of organisms in a given environment. Besides divergent selection, other factors may determine regional patterns of differentiation and contribute to life history evolution. In a common-garden environment over three years, we examined life history differentiation across different eco-geographical regions and effects of a specialist root-insect herbivore in the widespread European plant Centaurea stoebe, spotted knapweed. This plant occurs as two cytotypes with contrasting life cycles: monocarpic diploids and polycarpic tetraploids. In addition, the tetraploid cytotype has more recently become invasive in North America. We found significant regional differentiation in traits related to the timing of reproduction and reproductive allocation, but contrasting patterns in diploids and tetraploids. In diploids the degree of regional differentiation was higher compared to native tetraploids, and in the latter compared to invasive tetraploids. Furthermore, a pronounced shift in environmental conditions between the native and introduced range could have contributed to the differentiation between native and invasive tetraploids. The study also revealed the potential of the root-mining insect herbivore Agapeta zoegana, used as a biological control organism, to increase plant performance (presumably through overcompensatory growth), especially in the polycarpic tetraploids, and more so in the introduced populations. These findings suggest that patterns of regional differentiation in C. stoebe may be partly determined by divergent selection, but also strongly modulated by life cycle differences among geo-cytotypes. Furthermore, our study highlights the importance in applying a comprehensive and long-term approach when studying regional differentiation in plants. [ABSTRACT FROM AUTHOR]

Biological invasions are inherently demographic processes, but trait differences between native and introduced genotypes are rarely linked to population growth rates. Native European Centaurea stoebe occurs as two cytotypes with different life histories (monocarpic diploids, polycarpic tetraploids); however, only tetraploids have been found in its introduced range in North America. In a common garden experiment using artificial populations, we compared the demographic performance of the three geo-cytotypes in the presence and absence of a specialist herbivore using periodic matrix models. We found no difference in population growth rate between the two European cytotypes and no significant effects of herbivory in all geo-cytotypes. However, there was a pronounced increase in population growth rate for North American compared with European tetraploids due to increased seed production and juvenile establishment. These results suggest that genetic drift or rapid evolution, rather than pre-adaptation through polyploidy may explain the invasion success of tetraploids. [ABSTRACT FROM AUTHOR]

Background and Aims In a mixed-ploidy population, strong frequency-dependent mating will lead to the elimination of the less common cytotype, unless prezygotic barriers enhance assortative mating. However, such barriers favouring cytotype coexistence have only rarely been explored. Here, an assessment is made of the mechanisms involved in formation of mixed-ploidy populations and coexistence of diploid plants and their closely related allotetraploid derivates from the Centaurea stoebe complex (Asteraceae). Methods An investigation was made of microspatial and microhabitat distribution, life-history and fitness traits, flowering phenology, genetic relatedness of cytotypes and intercytotype gene flow (cpDNA and microsatellites) in six mixed-ploidy populations in Central Europe. Key Results Diploids and tetraploids were genetically differentiated, thus corroborating the secondary origin of contact zones. The cytotypes were spatially segregated at all sites studied, with tetraploids colonizing preferentially drier and open microhabitats created by human-induced disturbances. Conversely, they were rare in more natural microsites and microsites with denser vegetation despite their superior persistence ability (polycarpic life cycle). The seed set of tetraploid plants was strongly influenced by their frequency in mixed-ploidy populations. Triploid hybrids originated from bidirectional hybridizations were extremely rare and almost completely sterile, indicating a strong postzygotic barrier between cytotypes. Conclusions The findings suggest that tetraploids are later immigrants into already established diploid populations and that anthropogenic activities creating open niches favouring propagule introductions were the major factor shaping the non-random distribution and habitat segregation of cytotypes at fine spatial scale. Establishment and spread of tetraploids was further facilitated by their superior persistence through the perennial life cycle. The results highlight the importance of non-adaptive spatio-temporal processes in explaining microhabitat and microspatial segregation of cytotypes. [ABSTRACT FROM AUTHOR]

Abstract: A caged field experiment was used to determine how Centaurea stoebe L. phenostage (rosette, single-stem, multiple-stem) and ploidy level (diploid=2× and tetraploid=4×) influence oviposition and feeding of two biological control agents, Agapeta zoegana (Lep.: Cochylidae) and Cyphocleonus achates (Col.: Curculionidae). Ploidy level did not influence oviposition patterns of A. zoegana but rosette and one-stem plants had significantly more eggs than multiple-stem (4×) plants. Differences in oviposition levels did not translate into differences in larval densities, but 2× plants (particularly large one-stem plants) had significantly more larvae than 4× plants. There was a significant positive correlation between numbers of larvae and root diameter. Ploidy level and phenostage both had a significant effect on C. achates feeding damage, with adults feeding more frequently on multiple-stem plants. No C. achates larvae were observed when the roots were dissected. Furthermore, the generalist herbivore Arion lusitanicus, naturally present in the garden plots, was predominantly associated with young rosette plants, a stage at which survival rate is acknowledged to be the most important determinant of knapweed density. These results indicate that the combined damage caused by A. zoegana and C. achates, superimposed on damage caused by generalist herbivores in the local community, could provide effective control for C. stoebe. [Copyright &y& Elsevier]

1. Introduced plants have the potential to rapidly evolve traits of ecological importance that may add to their innate potential to become invasive. During invasions, selection may favour genotypes that are already pre-adapted to conditions in the new habitat and, over time, alter the characteristics of subsequent generations. 2. Spotted knapweed ( Centaurea stoebe) occurs in two predominantly spatially separated cytotypes in its native range (Europe–Western Asia), but currently only the tetraploid form has been confirmed in the introduced range (North America), where it is invasive. We used several common garden experiments to examine, across multiple populations, whether tetraploids and diploids from the native range differ in life cycle, leaf traits and reproductive capacity and if such differences would explain the predominance of tetraploids and their advance into new habitats in the introduced range. We also compared the same traits in tetraploids from the native and introduced range to determine whether any rapid adaptive changes had occurred since introduction that may have enhanced invasive potential of the species in North America. 3. We found tetraploids had lower specific leaf area, less lamina dissection and fewer, narrower leaves than diploids. Diploids exhibited a monocarpic life cycle and produced few if any accessory rosettes. Diploids produced significantly more seeds per capitulum and had more capitula per plant than tetraploids. In contrast, the vast majority of European tetraploids continued to flower in both seasons by regenerating from multiple secondary rosettes, demonstrating a predominantly polycarpic life cycle. 4. During early growth tetraploids from North America achieved greater biomass than both tetraploids and diploids from the native range but this did not manifest as larger above-ground biomass at maturity. In North American tetraploids there was also evidence of a shift towards a more strictly polycarpic life cycle, less leaf dissection, greater carbon investment per leaf, and greater seed production per capitulum. 5. Synthesis. Our results suggest that the characteristics of tetraploid C. stoebe pre-adapted them (compared to diploid conspecifics) for spread and persistence of the species into habitats in North America characterized by a more continental climate. After the species’ introduction, small but potentially important shifts in tetraploid biology have occurred that may have contributed significantly to successful invasion. [ABSTRACT FROM AUTHOR]

Polyploidy is often assumed to increase the spread and thus the success of alien plant species, but few empirical studies exist. We tested this hypothesis with Centaurea maculosa Lam., a species native to Europe and introduced into North America approximately 120 years ago where it became highly invasive. We analyzed the ploidy level of more than 2000 plants from 93 native and 48 invasive C. maculosa populations and found a pronounced shift in the relative frequency of diploid and tetraploid cytotypes. In Europe diploid populations occur in higher frequencies than tetraploids and only four populations had both cytotypes, while in North America diploid plants were found in only one mixed population and thus tetraploids clearly dominated. Our results showed a pronounced shift in the climatic niche between tetraploid populations in the native and introduced range toward drier climate in North America and a similar albeit smaller shift between diploids and tetraploids in the native range. The field data indicate that diploids have a predominately monocarpic life cycle, while tetraploids are often polycarpic. Additionally, the polycarpic life-form seems to be more prevalent among tetraploids in the introduced range than among tetraploids in the native range. Our study suggests that both ploidy types of C. maculosa were introduced into North America, but tetraploids became the dominant cytotype with invasion. We suggest that the invasive success of C. maculosa is partly due to preadaptation of the tetraploid cytotype in Europe to drier climate and possibly further adaptation to these conditions in the introduced range. The potential for earlier and longer seed production associated with the polycarpic life cycle constitutes an additional factor that may have led to the dominance of tetraploids over diploids in the introduced range. [ABSTRACT FROM AUTHOR]