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Species may become invasive after introduction to a new range because phenotypic traits pre-adapt them to spread and become dominant. In addition, adaptation to novel selection pressures in the introduced range may further increase their potential to become invasive. The diploid Leucanthemum vulgare and the tetraploid L. ircutianum are native to Eurasia and have been introduced to North America, but only L. vulgare has become invasive. To investigate whether phenotypic differences between the two species in Eurasia could explain the higher abundance of L. vulgare in North America and whether rapid evolution in the introduced range may have contributed to its invasion success, we grew 20 L. vulgare and 21 L. ircutianum populations from Eurasia and 21 L. vulgare populations from North America under standardized conditions and recorded performance and functional traits. In addition, we recorded morphological traits to investigate whether the two closely related species can be clearly distinguished by morphological means and to what extent morphological traits have changed in L. vulgare post-introduction. We found pronounced phenotypic differences between L. vulgare and L. ircutianum from the native range as well as between L. vulgare from the native and introduced ranges. The two species differed significantly in morphology but only moderately in functional or performance traits that could have explained the higher invasion success of L. vulgare in North America. In contrast, leaf morphology was similar between L. vulgare from the native and introduced range, but plants from North America flowered later, were larger and had more and larger flower heads than those from Eurasia. In summary, we found litte evidence that specific traits of L. vulgare may have pre-adapted this species to become more invasive than L. ircutianum, but our results indicate that rapid evolution in the introduced range likely contributed to the invasion success of L. vulgare. [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]

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]

The taxonomy of the Centaurea stoebe complex is controversial. Diploid and tetraploid plants occur in its native European range, but to date only tetraploids have been recorded from its introduced range in North America. We examined morphological differentiation of C. stoebe using multivariate and univariate approaches to clarify the taxonomic status of the known cytotypes. We measured more than 40 morphological traits on plants originating from 78 populations, grown from seed under uniform glasshouse conditions. The ploidy of almost 300 plants from 2 native and 20 introduced populations from Canada was assessed to test for the absence of diploids from North America. Finally, we explored whether postintroduction processes have resulted in phenotypic changes in introduced plants which may have contributed to the invasion success of C. stoebe. Morphometric analyses showed a clear separation of 2x and 4x plants and thus supported recognition of both cytotypes as separate taxa. Differences in the life cycle, the number of florets, the shape of capitula, and the shape of young rosette leaves were the best discriminant characters. Only minor differences were found between native and introduced tetraploids. All plants from the introduced range except for one hexaploid were found to be tetraploid. Rare diploids from Canada were identified as Centaurea diffusa or Centaurea psamogenna. [ABSTRACT FROM AUTHOR]

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]

Abstract: Spotted knapweed (Centaurea stoebe) occurs from Western Asia to Western Europe both as diploid and tetraploid cytotypes, predominantly in single-cytotype populations with higher frequency of diploid populations. Interestingly, only tetraploids have been recorded so far from its introduced range in North America where they became highly invasive. We performed phylogenetic and network analyses of more than 40 accessions of the C. stoebe and C. paniculata groups and other related taxa using cloned internal transcribed spacer (ITS) and sequences of the chloroplast trnT–trnL and atpBrbcL regions to (i) assess the evolutionary origin of tetraploid C. stoebe s.l., and (ii) uncover the phylogeny of the C. stoebe group. Both issues have not been studied so far and thus remained controversial. Cloned ITS sequences showed the presence of two slightly divergent ribotypes occurring in tetraploid cytotype, while only one major ribotype was present in diploid C. stoebe s.str. This pattern suggests an allopolyploid origin of tetraploids with contribution of the diploid C. stoebe s.str. genome. Although we were not able to detect the second parental taxon, we hypothesize that hybridization might have triggered important changes in morphology and life history traits, which in turn may explain the colonization success of the tetraploid taxon. Bayesian relaxed clock estimations indicate a relatively recent – Pleistocene origin of the tetraploid C. stoebe s.l. Furthermore, our analyses showed a deep split between the C. paniculata and C. stoebe groups, and a young diversification of the taxa within the C. stoebe group. In contrast to nrDNA analyses, the observed pattern based on two cpDNA regions was inconclusive with respect to the origin and phylogeny of the studied taxa, most likely due to shared ancient polymorphism and frequent homoplasies. [Copyright &y& Elsevier]

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.

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., (© 2012 Blackwell Publishing Ltd/CNRS.)

One commonly accepted mechanism for biological invasions is that species, after introduction to a new region, leave behind their natural enemies and therefore increase in distribution and abundance. However, which enemies are escaped remains unclear. Escape from specialist invertebrate herbivores has been examined in detail, but despite the profound effects of generalist herbivores in natural communities their potential to control invasive species is poorly understood. We carried out parallel laboratory feeding bioassays with generalist invertebrate herbivores from the native (Europe) and from the introduced (North America) range using native and nonnative tetraploid populations of the invasive spotted knapweed, Centaurea stoebe. We found that the growth of North American generalist herbivores was far lower when feeding on C. stoebe than the growth of European generalists. In contrast, North American and European generalists grew equally well on European and North American tetraploid C. stoebe plants, lending no support for an evolutionary change in resistance of North American tetraploid C. stoebe populations against generalist herbivores. These results suggest that biogeographical differences in the response of generalist herbivores to novel plant species have the potential to affect plant invasions.