Reproductive effort and herbivory timing in a perennial herb: fitness components at the individual and population levels.

We experimentally investigated how pollinator- and herbivore-induced changes influence the performance of the long-lived herb Primula veris. Eight treatments that corresponded to natural factors normally affecting this species were designed to enhance or reduce reproductive success and resource availability (flower removal, supplementary pollination, defoliation). During the experimental season and in the following year we quantified responses in terms of survival, growth, and seed production of reproductive plants. Matrix population models were used to calculate population growth rate using the demographic parameters recorded in permanent plots and respective treatment groups. Seed production was not limited by pollen availability, and we found no evidence of a cost of reproduction. Leaf removal had either no effect or a negative effect on future performance, depending on the timing of removal. Defoliation early in the season reduced current seed production and future growth, whereas removal during fruit development affected performance in the following year. Demographic models suggest that leaf damage has a smaller negative impact than flower removal on overall performance in this population. Our results suggest that the source-sink paths vary over the season and that the timing of herbivory may influence the extent to which effects are carried over to subsequent reproductive seasons.