Growth and competitive effects of Centaurea stoebe populations in response to simulated nitrogen deposition.

Increased resource availability can promote invasion by exotic plants, raising concerns over the potential effects of global increases in the deposition of nitrogen (N). It is poorly understood why increased N favors exotics over natives. Fast growth may be a general trait of good invaders and these species may have exceptional abilities to increase growth rates in response to N deposition. Additionally, invaders commonly displace locals, and thus may have inherently greater competitive abilities. The mean growth response of Centaurea stoebe to two N levels was significantly greater than that of North American (NA) species. Growth responses to N did not vary among C. stoebe populations or NA species. Without supplemental N, NA species were better competitors than C. stoebe, and C. stoebe populations varied in competitive effects. The competitive effects of C. stoebe populations increased with N whereas the competitive effects of NA species decreased, eliminating the overall competitive advantage demonstrated by NA species in soil without N added. These results suggest that simulated N deposition may enhance C. stoebe invasion through increasing its growth and relative competitive advantage, and also indicate the possibility of local adaptation in competitive effects across the introduced range of an invader.

Novel weapons and invasion: biogeographic differences in the competitive effects of Centaurea maculosa and its root exudate (+/-)-catechin.

Recent studies suggest that the invasive success of Centaurea maculosa may be related to its stronger allelopathic effects on native North American species than on related European species, one component of the "novel weapons" hypothesis. Other research indicates that C. maculosa plants from the invasive range in North America have evolved to be larger and better competitors than conspecifics from the native range in Europe, a component of the "evolution of increased competitive ability" hypothesis. These hypotheses are not mutually exclusive, but this evidence sets the stage for comparing the relative importance of evolved competitive ability to inherent competitive traits. In a competition experiment with a large number of C. maculosa populations, we found no difference in the competitive effects of C. maculosa plants from North America and Europe on other species. However, both North American and European C. maculosa were much better competitors against plants native to North America than congeners native to Romania, collected in areas where C. maculosa is also native. These results are consistent with the novel weapons hypothesis. But, in a second experiment using just one population from North America and Europe, and where North American and European species were collected from a broader range of sites, competitive interactions were weaker overall, and the competitive effects of C. maculosa were slightly stronger against European species than against North American species. Also consistent with the novel weapons hypothesis, (+/-)-catechin had stronger effects on native North American species than on native European species in two experiments. Our results suggest that the regional composition of the plant communities being invaded by C. maculosa may be more important for invasive success than the evolution of increased size and competitive ability.

Novel weapons: invasive plant suppresses fungal mutualists in America but not in its native Europe.

Why some invasive plant species transmogrify from weak competitors at home to strong competitors abroad remains one of the most elusive questions in ecology. Some evidence suggests that disproportionately high densities of some invaders are due to the release of biochemicals that are novel, and therefore harmful, to naive organisms in their new range. So far, such evidence has been restricted to the direct phytotoxic effects of plants on other plants. Here we found that one of North America's most aggressive invaders of undisturbed forest understories, Alliaria petiolata (garlic mustard) and a plant that inhibits mycorrhizal fungal mutualists of North American native plants, has far stronger inhibitory effects on mycorrhizas in invaded North American soils than on mycorrhizas in European soils where A. petiolata is native. This antifungal effect appears to be due to specific flavonoid fractions in A. petiolata extracts. Furthermore, we found that suppression of North American mycorrhizal fungi by A. petiolata corresponds with severe inhibition of North American plant species that rely on these fungi, whereas congeneric European plants are weakly affected. These results indicate that phytochemicals, benign to resistant mycorrhizal symbionts in the home range, may be lethal to naïve native mutualists in the introduced range and indirectly suppress the plants that rely on them.

Concentrations of the Allelochemical (+/-)-catechin IN Centaurea maculosa soils.

The phytotoxin (+/-)-catechin has been proposed to mediate invasion and autoinhibition by the Eurasian plant Centaurea maculosa (spotted knapweed). The importance of (+/-)-catechin to C. maculosa ecology depends in part on whether sufficient catechin concentrations occur at appropriate times and locations within C. maculosa soil to influence neighboring plants. Previous research on catechin in C. maculosa soils has yielded conflicting results, with some studies finding high soil catechin concentrations and other, more recent studies finding little or no catechin in field soils. Here, we report the most extensive study of soil catechin concentrations to date. We examined soil catechin concentrations in 402 samples from 11 C. maculosa sites in North America sampled in consecutive months over 1 yr, excluding winter months. One site was sampled on seven dates, another was sampled twice, and the remaining nine sites were each sampled once on a range of sampling dates. Methods used were similar to those with which we previously measured high soil catechin concentrations. We detected catechin only in the site that was sampled on seven dates and only on one sampling date in that site (May 16 2006), but in all samples collected on that date. The mean soil catechin concentration on that date was 0.65 +/- 0.45 (SD) mg g(-1), comparable to previously reported high concentrations. There are a number of possible explanations for the infrequency with which we detected soil catechin in this work compared to previous studies. Differences in results could reflect spatial and temporal variation in catechin exudation or degradation, as we examined different sites in a different year from most previous studies. Also, large quantities of catechin were detected in blanks for two sampling periods in the present study, leading us to discard those data. This contamination suggests that previous reports of high catechin concentrations that did not include blanks should be viewed with caution. Our results suggest that pure catechin is only rarely present in C. maculosa bulk soils. Thus, although catechin may play a role in C. maculosa invasion, the infrequency of soil catechin that we determined in this study suggests that we cannot be as certain of its role as previous reports of high soil catechin concentrations suggested.

Invasive plant suppresses the growth of native tree seedlings by disrupting belowground mutualisms.

The impact of exotic species on native organisms is widely acknowledged, but poorly understood. Very few studies have empirically investigated how invading plants may alter delicate ecological interactions among resident species in the invaded range. We present novel evidence that antifungal phytochemistry of the invasive plant, Alliaria petiolata, a European invader of North American forests, suppresses native plant growth by disrupting mutualistic associations between native canopy tree seedlings and belowground arbuscular mycorrhizal fungi. Our results elucidate an indirect mechanism by which invasive plants can impact native flora, and may help explain how this plant successfully invades relatively undisturbed forest habitat.

Oxalate contributes to the resistance of Gaillardia grandiflora and Lupinus sericeus to a phytotoxin produced by Centaurea maculosa.

Centaurea maculosa Lam. is a noxious weed in western North America that produces a phytotoxin, (+/-)-catechin, which is thought to contribute to its invasiveness. Areas invaded by C. maculosa often result in monocultures of the weed, however; in some areas, North American natives stand their ground against C. maculosa and show varying degrees of resistance to its phytotoxin. Two of these resistant native species, Lupinus sericeus Pursh and Gaillardia grandiflora Van Houtte, were found to secrete increased amounts of oxalate in response to catechin exposure. Mechanistically, we found that oxalate works exogenously by blocking generation of reactive oxygen species in susceptible plants and reducing oxidative damage generated in response to catechin. Furthermore, field experiments show that L. sericeus indirectly facilitates native grasses in grasslands invaded by C. maculosa, and this facilitation can be correlated with the presence of oxalate in soil. Addition of exogenous oxalate to native grasses and Arabidopsis thaliana (L.) Heynh grown in vitro alleviated the phytotoxic effects of catechin, supporting the field experiments and suggesting that root-secreted oxalate may also act as a chemical facilitator for plant species that do not secrete the compound.

Soil biota and exotic plant invasion.

Invasive plants are an economic problem and a threat to the conservation of natural systems. Escape from natural enemies might contribute to successful invasion, with most work emphasizing the role of insect herbivores; however, microbial pathogens are attracting increased attention. Soil biota in some invaded ecosystems may promote 'exotic' invasion, and plant-soil feedback processes are also important. Thus, relatively rare species native to North America consistently demonstrate negative feedbacks with soil microbes that promote biological diversity, whereas abundant exotic and native species demonstrate positive feedbacks that reduce biological diversity. Here we report that soil microbes from the home range of the invasive exotic plant Centaurea maculosa L. have stronger inhibitory effects on its growth than soil microbes from where the weed has invaded in North America. Centaurea and soil microbes participate in different plant-soil feedback processes at home compared with outside Centaurea's home range. In native European soils, Centaurea cultivates soil biota with increasingly negative effects on the weed's growth, possibly leading to its control. But in soils from North America, Centaurea cultivates soil biota with increasingly positive effects on itself, which may contribute to the success of this exotic species in North America.