Substantial controversy surrounds the efficacy of biological control insects to reduce densities of Centaurea stoebe, a widespread, aggressive invasive plant in North America. We developed a graphical model to conceptualize the conditions required to explain the current contradictory findings, and then employed a series of manipulations to evaluate C. stoebe responses to herbivores. We manipulated soil nitrogen and competition in a field population and measured attack rates of a foliage and seed feeder (Larinus minutus), two gall flies (Urophora spp.), and a root feeder (Cyphocleonus achates), as well as their effects on the growth and reproduction of C. stoebe. Nitrogen limitation and competing vegetation greatly reduced C. stoebe growth. L. minutus most intensively reduced seed production in low-nitrogen soils, and removal of neighboring vegetation increased Larinus numbers per flower head and the percentage of flowers attacked by 15% and 11%, respectively. Cyphocleonus reduced flower production and aboveground biomass over two years, regardless of resources or competition. Our results, in conjunction with other published studies, demonstrate that positive, neutral, and negative plant growth responses to herbivory can be generated. However, under realistic field conditions and in the presence of multiple herbivores, our work repudiates earlier studies that indicate insect herbivores increase C. stoebe dominance.

No abstract

We measured seed germination and seedling survivorship of spotted knapweed, Centaurea stoebe, in a series of laboratory and field experiments to evaluate the efficacy of seed limitation as a management focus. This work was initiated 6 years after introduction of several biological control agents. The soil seed bank of the site used in this study contained a mean density of 5,848 seeds/m 2 (ranging from 0 to 16,364 seeds/m 2 ), and 92% of the seeds isolated from soils were shriveled, discolored, and/or partially decayed. Additionally, none of the intact seeds germinated, suggesting that the viable seed bank at our field study site has been exhausted. Centaurea stoebe seeds were planted into pots under a range of soil nitrogen (N) availability, with half of the pots containing a single density of previously established seedlings of a native cool-season grass, slender wheatgrass (Elymus trachycaulus). A watering regime mimicking local precipitation was applied. Spotted knapweed exhibited large biomass responses to N addition, but the presence of grasses suppressed the ability to exploit this N. Surprisingly, low soil N conditions improved knapweed survivorship in the presence of grasses. Nevertheless, recruitment and biomass were still far below the levels reached in the absence of competition. To evaluate the effect of density on successful recruitment, Centaurea stoebe seed was introduced into a meadow at three densities matching reduced levels of seed production under the constraints of seed predators. These densities were sown with or without a seed mixture of native species, into an existing plant community lacking C. stoebe, and seedling recruitment was recorded over 2.5 years. Across all plots and densities sown (568-2,272 seeds m -2 year -1 ), seedling recruitment was less than 1%. The invasion potential of spotted knapweed was greatly diminished when realistic levels of plant competition and biological control limit seed production. We therefore conclude that a combination of seed limitation and shortage of 'safe sites' within undisturbed vegetation can limit densities of C. stoebe.