Spatial dynamics of Ilex aquifolium populations seed dispersal and seed bank: understanding the first steps of regeneration

The objective of this study was to analyse quantitatively the spatial distribution of holly (Ilex aquifolium L.) seed rain and seed bank, and to detect the relationships between these consecutive processes. We measured seed dispersal by birds and fallen fruits, and also density and viability of seed bank in two Ilex populations in central Spain. Analysis was made distinguishing the following microhabitats: holly woodland, edge of holly woodland, open grassland 10 m and 100 m from the woodland, fleshy fruit shrubs, dry fruit shrubs, and adjacent non-holly woodland. Spatial distribution of dispersed and in-soil seeds was measured by the clumping index. Seed rain and seed bank under holly woodlands were significantly higher than in the other microhabitats. Forest edges and fleshy fruit shrubs were the next microhabitats with the highest seed rain and seed bank density. Interannual and interlocality variations were not significant. The relative importance of the different dispersal methods varied between microhabitats, with a similar support of bird dispersed seeds and fallen fruits within the woodland and a greater influence of cattle dispersal in open areas. Seed spatial aggregation was significant in both dispersed seeds and soil seeds from holly woodlands and the edge of the forest. Aggregation under shrubs, grasslands and the adjacent forests evidenced a general random distribution of holly seeds (only in some cases clumping index was significant). Quantitative differences between seed rain and seed bank are important. Post-dispersal seed predation did not modify seed rain distribution, which was mirrored in the seed bank pattern. These two phases of holly regeneration had a heavy spatial influence, determined by the landscape structure and activity of the dispersal agents, that reflects a differential recruitment potential. Comparisons between both populations suggest that in the southern locality (Robregordo) holly has a weaker capacity to colonize open areas, and a stronger recruitment limitation due to propagule availability.