Scale-dependent effects of habitat fragmentation on hawthorn pollination, frugivory, and seed predation.

Habitat fragmentation is a major cause of functional disruption in plant-animal interactions. The net effect on plant regeneration is, however, controversial because a given landscape change can simultaneously hamper mutualism and attenuate antagonism. Furthermore, fragmentation effects may emerge at different spatial scales, depending on the size of the foraging range of the different interacting animals. We studied pollination by insects, frugivory by birds acting as seed dispersers, and postdispersal seed predation by rodents in 60 individual hawthorn (Crataegus monogyna Jacq.) trees in relation to structural fragmentation in the surrounding habitat. We evaluated fragmentation at three spatial scales by measuring the percentage of forest cover in three concentric areas around each tree of, respectively, 10-m, 20- to 50-m, and 50- to 100-m radius. The number of developing pollen tubes per flower style and fruit set decreased in proportion to the decrease of forest cover. Similarly, the magnitude of frugivory in focal trees was negatively affected by habitat loss. In contrast, seed predation was higher under plants in highly fragmented contexts. The effect of fragmentation was additive in terms of reducing the potential of plant regeneration. Moreover, the functional scale of response to habitat loss differed among interactions. Fragmentation effects on pollination emerged at the largest scale, whereas seed predation was mostly affected at the intermediate scale. In contrast to expectations from the larger foraging range of birds, fragmentation effects on frugivory mainly operated at the finest scale, favored by the ability of birds to cope hierarchically with spatial heterogeneity at different scales. Given that two opposing demographic forces (frugivory and seed predation) would be potentially affected by fine-scale features, we propose structural scale as the primary spatial dimension of fragmentation effects on the process of plant regeneration.