Caching territoriality and site preferences by a scatter-hoarder drive the spatial pattern of seed dispersal and affect seedling emergence

Abstract

1. For plants with seeds dispersed by scatter-hoarders, decision-making by animals when caching determines the spatial pattern of seed dispersal and lays the initial template for recruitment, driving the regeneration of many species. However, the mechanism by which animal behaviour shapes seed distributions in spatially complex landscapes is not well understood. We investigated caching territoriality and site preferences to determine the spatial pattern of seed caching at different scales and whether scatter-hoarding behaviour drives the spatial distribution of seedling emergence. 2. We used radio-tracking and automatic wildlife cameras to monitor holm oak (Quercus ilex) acorn caching by Eurasian magpies (Pica pica), who are effective scatter-hoarders in agroforestry systems. We assessed the effect of caching territories, distance to seed source, habitat, subhabitat, microsites, and caching material in the spatial pattern of acorn dispersal by magpies. In addition, we analysed the relationship between the density of cached acorns and of emerged seedlings in different habitats. 3. Breeding magpies cached the acorns inside their caching territories, where they preferred tilled areas over oak plantations and mostly avoided old fields. These differences in habitat preference were maximized at relatively short to medium dispersal distances, where most acorns were cached, and decreased or disappeared at long-distances. Within tree plantations, magpies preferred high plant-productivity sites over low productivity ones. At the finest spatial scale, magpies preferred structures built by animals, such as rabbit grit mounds and latrines and ant litter mounds, to cache the acorns. In many sites, magpies selected uncommon materials such as stones and litter to cover caches. In the subsequent spring, seedling emergence was positively correlated with acorn cache density. 4. Synthesis. Scatter-hoarding is a hierarchical process in which caching sites are selected using different criteria at different spatial scales driven by territoriality and site preferences. Territoriality constrained dispersal distance and the habitats available for acorn caching. Magpie territoriality therefore indirectly drives oak seedling emergence and can determine oak recruitment and forest regeneration.