Effects of frugivore preferences and habitat heterogeneity on seed rain: a multi-scale analysis

Seed rain mediated by frugivores is influenced by (1) the seed-deposition distances following fruit ingestion, (2) the disperser activity, as determined by its behaviour and habitat preferences, and (3) the structure of the habitat within the landscape. Here, we evaluated such components using the fleshy-fruited shrub Ephedra fragilis and the frugivorous Balearic lizard Podarcis lilfordi. We estimated seed-deposition patterns based on the displacements and habitat preferences of lizards, derived from visual surveys and telemetry data. The influence of variables potentially determining lizard habitat preference (i.e., height, slope, four measures of habitat abundance and four measures of habitat fragmentation) was evaluated at three spatial scales: 'home-range' (c. 2.5-10*10(3) m(2); telemetry data), 'within home-range' (c. 100 m(2); telemetry data) and 'microhabitat' (<100 m(2); visual survey). Cumulative lizard displacement (from each telemetric location to the initial capture point) saturated before the peak of seed defecation (seed-retention time), indicating that lizard home-range size and habitat preferences were the main determinants of the spread and shape of seed shadows. Shrub cover was positively correlated with habitat preference at the three scales of analysis, whereas slope was negatively correlated at the home-range scale. Model scenarios indicated that spatially-aggregated seed rain emerged when we incorporated the joint effect of habitat preference at the two largest (home-range and within home-range) scales. We conclude that, in order to predict seed rain in animal dispersed plants, it is important to consider the multi-scale effects of habitat preference by frugivores.