The interplay between shifts in biomass allocation and costs of reproduction in four grassland perennials under simulated successional change.

When perennial herbs face the risk of being outcompeted in the course of succession, they are hypothesized to either increase their biomass allocation to flowers and seeds or to invest more in vegetative growth. We tested these hypotheses in a 3-year garden experiment with four perennials (Hypochaeris radicata, Cirsium dissectum, Succisa pratensis and Centaurea jacea) by growing them in the midst of a tall tussock-forming grass (Molinia caerulea) that may successionally replace them in their natural habitat. In all species except for the short-lived H. radicata, costs of sexual reproduction were significant over the 3 years, since continuous bud removal enhanced total biomass or rosette number. To mimic succession we added nutrients, which resulted in a tripled grass biomass and higher death rates in the shorter-lived species. The simulated succession resulted also in a number of coupled growth responses in the survivors: enhanced plant size as well as elevated seed production. The latter was partly due to larger plant sizes, but mostly due to higher reproductive allocation, which in turn could be partly explained by lower relative somatic costs and by lower root-shoot ratios in the high-nutrient plots. Our results suggest that perennial plants can increase both their persistence and their colonization ability by simultaneously increasing their vegetative size and reproductive allocation in response to enhanced competition and nutrient influxes. These responses can be very important for the survival of a species in a metapopulation context.