Habitat fragmentation reduces plant fitness by disturbing pollination and modifying response to herbivory

In fragmented landscapes plant species are often confined to remnants of formerly more widespread habitats, with many of their populations being small and isolated. This study experimentally examined the effects of population size and isolation on pollination, herbivory and reproductive success in the forest herb Phyteuma spicatum (Campanulaceae). In an experiment in which population size and isolation were manipulated using plants from the same origin, population size positively affected pollinator visitation, but did not alter the generally high levels of herbivory. As a result, seed production was higher in large populations. Conversely, plants originating from 14 natural populations of varying size and degree of isolation did not differ in reproductive success when grown in the same environment, suggesting similar attractiveness to pollinators and reproductive potential. The intensity of herbivory, however, was higher in progeny of small populations, at least in terms of the proportion of biomass removed. In both experiments, there were no effects of population isolation. The results suggest (1) that small population size decreases reproductive success via direct negative effects on plant–pollinator interactions, (2) that this pattern is not offset by herbivory, but (3) that herbivory enforces fragmentation effects on pollination by further reducing the number of flowering individuals and (4) that habitat fragmentation may influence plant fitness by affecting plant response to herbivory. The effects of habitat fragmentation on plant populations in present-day landscapes are thus complex, illustrating the need for more integrated studies in conservation biology that take into account both mutualistic and antagonistic plant–animal interactions.