Contrasting seed traits of co-existing seeds lead to a complex neighbor effect in a seed-rodent interaction

Scatter-hoarding rodents play important roles in seed dispersal and predation in many forest ecosystems. Existing studies have shown that the seed foraging preference of rodents is directly affected by seed traits and indirectly affected by the traits of other co-existing seeds nearby (i.e., neighbor effect). Plant seeds exhibit a combination of diverse seed traits, including seed size, chemical defense, and nutrient content. Therefore, it is difficult to evaluate the influence of each single seed trait on such neighbor effects. Here, by using artificial seeds, we investigated the impacts of contrasts in seed size, tannin content, and nutrient content on neighbor effects. We tracked 9000 tagged artificial seeds from 30 seed-seed paired treatments in a subtropical forest in southwest China. The contrast in seed size between paired seeds created obvious neighbor effects measured through three seed dispersal related indicators: the proportion of seeds being removed, the proportion of seeds cached, and the distance transported by rodents. However, the magnitudes and the signs of the neighbor effects differed among pairs, including both apparent mutualism and apparent competition, depending on the contrast in seed size between paired seeds. The contrasts of tannin and nutrient content between paired seeds showed relatively few neighbor effects. Our results suggest that the contrast in seed traits between the target seed and its neighboring seeds should be considered when studying rodent-seed interactions. Furthermore, we expect that similar complex neighbor effects may also exist in other plant-animal interactions, such as pollination and herbivory.