Seed bank spatial structure in semi-arid environments: beyond the patch-bare area dichotomy

The prevalence of patchy structures in vegetation is a common feature in semi-arid ecosystems. Although the effect of patches on seed density is widely known, we still lack information on how patch features affect seed bank density and composition. Our aim was to answer two basic questions: (1) How do seed bank density and composition vary within and outside patch aboveground physical limits? and (2) Do patch characteristics affect soil seed bank density and composition? We sampled 50 shrub patches in a semi-arid gypsum system in Central Spain, measuring patch size, composition and structure, and seed bank at three locations per shrub (centre, edge and outside). We calculated the effect of interior patch location, patch composition and structure on seed density and composition. Patches acted both as seed sources, increasing seed density in neighbouring areas and as seed sinks by trapping seeds from bare areas. Patch structure (erect perennial cover) had the greatest effect on seed bank density, whereas patch size and microslope had the greatest influence on bare area density. Patch structure, composition and interior location explained the variation in seed bank composition. Patch effect extends to the surrounding bare matrix creating a seed bank gradient in density and composition. This effect is modulated by patch structure and composition and affects seed bank composition. Our results suggest that the spatial structure of gypsum community seed banks may act as a mechanism for a spatial storage effect contributing to the maintenance of high levels of diversity in semi-arid environments.