Phylogenetic diversity is a better predictor of wetland community resistance to Alternanthera philoxeroides invasion than species richness.

Highly biodiversity communities have been shown to better resist plant invasions through complementarity effects. Species richness (SR) is a widely used biodiversity metric but lacks explanatory power when there are only a few species. Communities with low SR can have a wide variety of phylogenetic diversities (PD), which might allow for a better prediction of invasibility. We assessed the effect of diversity reduction of a wetland community assemblage typical of the Beijing area on biotic resistance to invasion of the exotic weed Alternanthera philoxeroides and compared the reduction in SR and PD in predicting community invasibility. The eight studied resident species performed similarly when grown alone and when grown in eight-species communities together with the invasive A. philoxeroides. Variation partitioning showed that PD contributed more to variation in both A. philoxeroides traits and community indicators than SR. All A. philoxeroides traits and community indicators, except for evenness index, showed a linear relationship with PD. However, only stem length of A. philoxeroides differed between the one- and two-species treatments, and the diversity index of the communities differed between the one- and two-species treatments and between the one- and four-species treatments. Our results showed that in natural or semi-natural wetlands with relatively low SR, PD may be a better predictor of invasibility than SR. When designing management strategies for mitigating A. philoxeroides invasion, deliberately raising PD is expected to be more efficient than simply increasing species number.

Evidence of climatic niche shift during biological invasion.

Niche-based models calibrated in the native range by relating species observations to climatic variables are commonly used to predict the potential spatial extent of species' invasion. This climate matching approach relies on the assumption that invasive species conserve their climatic niche in the invaded ranges. We test this assumption by analysing the climatic niche spaces of Spotted Knapweed in western North America and Europe. We show with robust cross-continental data that a shift of the observed climatic niche occurred between native and non-native ranges, providing the first empirical evidence that an invasive species can occupy climatically distinct niche spaces following its introduction into a new area. The models fail to predict the current invaded distribution, but correctly predict areas of introduction. Climate matching is thus a useful approach to identify areas at risk of introduction and establishment of newly or not-yet-introduced neophytes, but may not predict the full extent of invasions.

Low genetic differentiation among seasonal cohorts in Senecio vulgaris as revealed by amplified fragment length polymorphism analysis.

Common groundsel, Senecio vulgaris (Asteraceae), is a highly selfing semelparous ephemeral weed that belongs to the few plant species in central Europe capable of growing, flowering and fruiting all year round. In temperate climates, flowering S. vulgaris cohorts were found to appear up to three times per year. Using amplified fragment length polymorphism (AFLP) molecular markers we examined temporal genetic differentiation among spring, summer and autumn cohorts at each of seven sites located in two regions in Switzerland. Strong genetic differentiation among cohorts may indicate the existence of seasonal races of S. vulgaris, reproductively isolated by nonoverlapping flowering phenologies. Analysis of molecular variance (amova) revealed that < 2.5% of the AFLP variation resided among cohorts within sites, whereas there was significant genetic differentiation among plants from different sites (15.6%) and among individuals within cohorts (81.9%). Significant genetic differentiation was also observed between the two regions. Isolation-by-distance was found on a regional scale, but not on a local scale. Gene flow was estimated to be approximately 15-fold higher among cohorts within sites than among sites. We further found, on average, similar levels of genetic diversity within the three seasonal cohorts. The results of this study demonstrate that season of growth represents a weak barrier for genetic exchange among S. vulgaris populations and does not affect molecular variance. Therefore, there is no evidence for the existence of seasonally specialized races of S. vulgaris. We discuss some implications of the results for the biological control of S. vulgaris using a native rust fungus.

Does natural selection promote population divergence? A comparative analysis of population structure using amplified fragment length polymorphism markers and quantitative traits.

Divergent natural selection is considered an important force in plant evolution leading to phenotypic differentiation between populations exploiting different environments. Extending an earlier greenhouse study of population differentiation in the selfing annual plant Senecio vulgaris, we estimated the degree of population divergence in several quantitative traits related to growth and life history and compared these estimates with those based on presumably neutral molecular markers (amplified fragment length polymorphisms; AFLPs). This approach allowed us to disentangle the effects of divergent selection from that of other evolutionary forces (e.g. genetic drift). Five populations were examined from each of two habitat types (ruderal and agricultural habitats). We found a high proportion of total genetic variance to be among populations, both for AFLP markers (phiST = 0.49) and for quantitative traits (range of QST: 0.26-0.77). There was a strong correlation between molecular and quantitative genetic differentiation between pairs of populations (Mantel's r = 0.59). However, estimates of population differentiation in several quantitative traits exceeded the neutral expectation (estimated from AFLP data), suggesting that divergent selection contributed to phenotypic differentiation, especially between populations from ruderal and agricultural habitats. Estimates of within-population variation in AFLP markers and quantitative genetic were poorly correlated, indicating that molecular marker data may be of limited value to predict the evolutionary potential of populations of S. vulgaris.

Genetic structure of the annual weed Senecio vulgaris in relation to habitat type and population size.

Throughout the world, the highly selfing annual common groundsel, Senecio vulgaris (Asteraceae) is a common weed. Recently, it has also colonized ecological compensation areas in agro-ecosystems. We investigated the genetic structure of S. vulgaris using random amplified polymorphic DNA (RAPD) profiles of 80 plants from nine populations representing three habitat types in two regions in Switzerland. RAPD variation among regions (19.8%), among populations within regions (19.2%) and within populations (61.1%) was highly significant (ANOVA; P < 0.001). Gene flow estimated from the observed differentiation among populations (PhiST = 0.382) was low (assuming Wright's island model, Nem = 0.404). Genetic distances between pairs of populations were significantly correlated with geographical distances (Mantel test; r = 0.37, P < 0.03). Molecular variance obtained with AMOVA was lowest in the small populations in compensation areas (1.13), intermediate in vineyard populations (2.49), all located in northern Switzerland and highest in the larger vegetable field populations from western Switzerland (3.41; P < 0.05). Overall, there was a positive correlation of molecular variance and population size (P < 0.05), as expected under genetic drift. However, molecular variance was negatively correlated with population size among populations in ecological compensation areas, suggesting that selection was also important. We also applied triazine herbicide to leaves of three offspring of each of the 80 plants. Plants from populations of compensation areas showed higher mean levels and reduced variation in the resistance to triazine herbicide than plants from vineyards and vegetable fields. This suggests that compensation areas were colonized from adjacent corn fields, in which there has been selection for herbicide resistance. We discuss the implications of our results for the biological control of S. vulgaris.

Phenotypic plasticity of Senecio vulgaris from contrasting habitat types: growth and pyrrolizidine alkaloid formation.

The growth-differentiation balance hypothesis (GDB), which postulates a physiological trade-off between growth and differentiation (morphological and chemical), has been tested almost exclusively for carbon-based secondary metabolites. Little attention has been paid to N-based compounds. In this study we aimed to test the predictions of the GDB hypothesis under field conditions for growth and pyrrolizidine alkaloid (PA) formation in Senecio vulgaris. We conducted a reciprocal transplant experiment at two sites differing widely in their nutrient supply. These included a conventionally managed vineyard (V) and a strip of local wild flowers between crop fields, which was established to promote species diversity in agroecosystems (C). No fertilizer or pesticides are allowed in such ecological compensation areas. In C, we expected lower growth but higher PA formation than in V. Due to differentiated selection regimes in the two habitat types with regard to nutrient (nitrogen) availability in the soil, we also expected different N-allocation patterns for the genotypes of the two collection sites. Plants of V produced more biomass and were taller than the plants of C. The relatively poor nitrogen conditions in C favored an earlier differentiation towards generative organs. In plants of C, higher concentrations of PAs were found than in plants of V. There exists a close negative correlation between growth and PA formation, indicating a trade-off. The origin of the plant material had only a little effect on PA formation. The observed phenotypic reaction of PA formation in S. vulgaris in the two habitats fits quite well the predictions of GDB theory. It is shown that this general response is overlaid by physiological factors leading to a pattern of PA accumulation, which is not readily predictable by nonmechanistic theories.

Adaptation of Senecio vulgaris (Asteraceae) to ruderal and agricultural habitats.

Adaptation of the annual plant Senecio vulgaris to ruderal and agricultural habitats was investigated. We expected S. vulgaris to be adapted to the agricultural habitat through nutrient-specific differentiation of relatively few genotypes responding to the generally high homogenous nutrient levels at the agricultural habitat caused by constant fertilization. To assess adaptation of S. vulgaris, vegetative and reproductive responses of seed families from various populations of agricultural and ruderal habitats, grown in the greenhouse at high and low nutrient levels, were compared. Data were analyzed with a three-level nested ANOVA based on the levels habitat, population, and family. A significant habitat effect indicated that S. vulgaris from ruderal and agricultural habitats were genetically different with plants from the agricultural habitat having larger leaves and a higher reproduction. A significant habitat by nutrient effect showed a stronger response of reproduction to nutrients at the agricultural habitat, suggesting that genetic differentiation among habitats is nutrient-specific. Contrary to expectations, only the agricultural habitat showed genetic diversity of S. vulgaris. Results suggest that nutrient levels at the agricultural habitat are more heterogeneous as generally proposed leading to a relatively high genetic variation.

Suitability of two root-mining weevils for the biological control of scentless chamomile, Tripleurospermum perforatum, with special regard to potential non-target effects.

The biology and host range of the two root-mining weevils Diplapion confluens Kirby and Coryssomerus capucinus (Beck), two potential agents for the biological control of scentless chamomile Tripleurospermum perforatum (Mérat) Laínz, were studied in the field in southern Germany and eastern Austria, and in a common garden and under laboratory conditions in Delémont, Switzerland from 1993 to 1999. Both weevils were univoltine, and females started to lay eggs in early spring. Diplapion confluens had three and C. capucinus five instars. Larvae of both species were found in the field from mid-April until the end of July; later instars preferentially fed in the vascular cylinder of the shoot base, root crown or root. Although larvae of both species occupy the same temporal and spatial niche within their host plants, they occurred at all investigated field sites together, and showed a similar distribution within sites. No negative or positive interspecific association was detected. Host-specificity tests including no-choice, single-choice, and multiple-choice tests under confined conditions, as well as tests under field conditions with natural and augmented insect densities revealed that both herbivores were specific to plant species in the tribe Anthemideae. However, their development to mature larva or adult on several cultivated plants, as well as on one plant species native to North America, rendered them unsuitable for field release in North America. It was concluded that to investigate non-target effects reliably, host-specificity tests with biological control agents should be carried out under a variety of conditions, particularly with augmented insect densities, as are expected to occur naturally after release.

Complex relation between triazine-susceptible phenotype and genotype in the weed Senecio vulgaris may be caused by chloroplast DNA polymorphism.

The weed Senecio vulgaris acquired high levels of resistance to triazine herbicides soon after the latter's introduction. As in most weeds, triazine resistance is conferred by a point mutation in the chloroplast psbA gene that negatively affects the fitness of its carrier. To assess levels of triazine resistance in S. vulgaris field populations, we adopted a PCR-RFLP-based molecular diagnostic test recently developed for the triazine resistance-conferring region of the psbA gene of other weeds, including Brassica napus, Chenopodium spp. and Amaranthus spp., and compared these molecular results to the phenotypic response after triazine application. A highly significant linear correlation was found between phytotoxic symptoms and biomass reduction. Variability in phenotypic response was not only found between populations or inbred lines of S. vulgaris but also within replicates of the same inbred line. No clear relationship, however, was found between the DNA restriction pattern and the phenotypic response to triazine application, thereby throwing doubt on the use of such molecular diagnostic tests to track triazine resistance in S. vulgaris. Our results indicate that the chloroplast genome of S. vulgaris is polymorphic and that the level of polymorphism may be variable within single leaves of individual plants. We discuss the possible genetic basis of this polymorphism and its consequence for the acquisition and inheritance of chloroplast-based traits.

Influence of cnicin, a sesquiterpene lactone ofCentaurea maculosa (Asteraceae), on specialist and generalist insect herbivores.

The sesquiterpene lactone cnicin was extracted fromCentaurea maculosa andCentaurea vallesiaca. We examined its effects on the ovipositional response and larval development of generalist and specialist insect herbivores associated withC. maculosa. For the oviposition trials, three plant species (C. maculosa, Achillea millefolium, andCichorium intybus), half of which were sprayed with 3% of cnicin, were exposed to the specialist mothsStenodes straminea, Agapeta zoegana, andPterolonche inspersa in field cages. All three species significantly preferredC. maculosa to other plants andP. inspersa significantly preferred cnicin-sprayed plants to untreated plants for oviposition. Tested over all species, cnicin significantly increased the number of eggs laid on a given plant. A larval diet test examined the toxicity of cnicin for larvae of the generalist noctuid mothSpodoptera littoralis. Cnicin concentrations of 3% and 6% were lethal and 1% and 0.5% seriously inhibited growth and development. The larvae of theC. maculosa specialistStenodes straminea survived at 6% cnicin, but none of the pupae hatched.Agapeta zoegana was able to survive at 1% and 3% cnicin. Both specialists had difficulties with the artificial diet, but weight increase and survival was not further reduced when cnicin was present compared with on the control diet. In conclusion, cnicin influenced host recognition by the specialist species, and larvae of the generalist did not survive on natural levels of cnicin. Growth and survival of the specialist were not influenced by cnicin but were considerably hampered on artificial diet.