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Scatterhoarding rodents often place caches in the open where pilferage rates are reduced, suggesting that they tradeoff higher risks of predation for more secure cache sites. We tested this hypothesis in two study systems by measuring predation risks inferred from measures of giving-up densities (GUDs) at known cache sites and other sites for comparison. Rodent GUDs were measured with small trays containing 3 L of fine sand mixed with sunflower seeds. In the first experiment, we relied on a 2-year seed dispersal study in a natural forest to identify caches of eastern gray squirrels (Sciurus carolinensis) and then measured GUDs at: (i) these caches; (ii) comparable points along logs and rocks where rodent activity was assumed highest; and (iii) a set of random points. We found that GUDs and, presumably, predation risks, were higher at both cache and random points than those with cover. At the second site, we measured GUDs of eastern gray squirrels in an open park system and found that GUDs were consistently lowest at the base of the tree compared to more open sites, where previous studies show caching by squirrels to be highest and pilferage rates by naïve competitors to be lowest. These results confirm that predation risks can influence scatterhoarding decisions but that they are also highly context dependent, and that the landscape of fear, now so well documented in the literature, could potentially shape the temporal and spatial patterns of seedling establishment and forest regeneration in systems where scatterhoarding is common.

As mutualists, seed dispersers may significantly affect mutualistic interactions and seedling recruitment of sympatric plants that share similar seed dispersers, but studies are rare. Here, we compared seed dispersal fitness in two co-occurring plant species (Armeniaca sibirica and Amygdalus davidiana) that inhabit warm temperate deciduous forest in northern China. We tested the hypothesis that seed trait-mediated selection by rodents may influence mutualistic interactions with rodents and then seedling establishment of co-occurring plant species. A. davidiana seeds are larger and harder (thick endocarps) than A. sibirica seeds, but they have similar levels of nutrients (crude fat, crude protein), caloric value and tannin. More A. sibirica seedlings are found in the field. Semi-natural enclosure tests indicated that the two seed species were both harvested by the same six rodent species, but that A. sibirica had mutualistic interactions (scatter hoarding) with four rodent species (Apodemus peninsulae, A. agrarius, Sciurotamias davidianus, Tamias sibiricus), and A. davidiana with only one (S. davidianus). Tagged seed dispersal experiments in the field indicated that more A. sibirica seeds were scatter-hoarded by rodents, and more A. sibirica seeds survived to the next spring and became seedlings. A. sibirica seeds derive more benefit from seed dispersal by rodents than A. davidiana seeds, particularly in years with limited seed dispersers, which well explained the higher seedling recruitment of A. sibirica compared with that of A. davidiana under natural conditions. Our results suggest that seed dispersers may play a significant role in seedling recruitment and indirect competition between co-occurring plant species.

Context. Mutualistic interactions between animals and plants shape the structure of plant-animal systems and, subsequently, affect plant-community structure and regeneration.Aims. To assess the effects of plant and rodent functional traits on the formation of mutualistic and predatory interactions regarding seed dispersal and predation in a warm-temperate forest.Methods. Seed scatter-hoarding and predation by six sympatric rodent species on seeds belonging to five sympatric tree species were tested under enclosure conditions. Key results. Functional traits (i.e. rodent body size and seed traits) are important to mutualism/predation in this seed-rodent system. The rodent-seed network is highly nested: large-sized rodents have mutualistic or predatory interactions with both large-and small-sized seed species, but small-sized rodents interacted with small-sized seed species only. Large seeds or seeds with hard coats enhanced mutualism and reduced predation.Conclusion. Body size of rodents and seed traits such as handling time and nutritional value are key factors in the formation of mutualistic and predatory interactions within seed-rodent systems.Implications. To promote seedling establishment in degenerated forests, introducing or protecting large-sized scatter hoarders and reducing the density of pure seed eaters are needed.Additional keywords: behavioural difference, plant-animal interactions, plant regeneration, seed dispersal.

Food hoarding and pilferage in rodents may be regulated by intense competition between sympatric species that have similar habitats, diets and activity, but studies exploring this remain rare. Here, we used semi-natural enclosures to investigate food-hoarding and cache pilferage interactions between sympatric Korean field mice (KFM) (Apodemus peninsulae) and Chinese white-bellied rats (CWR) (Niviventer confucianus). KFM and CWR have similar diets, habitat and nocturnal activity, but the smaller KFM larder and scatter hoards and larger CWR larder hoard only. We found that KFM harvest, larder-hoard and eat seeds at a greater intensity when CWR are present as an audience (present but cannot pilfer). KFM ate 11.5%, re-larder-hoarded 17.9% and re-scatter-hoarded 1.3% of their scatter-hoarded seeds, and ate 29.3% of their larder-hoarded seeds when CWR were present as pilferers. A total of 12.8% of the seeds scatter-hoarded and 50% of seeds directly put on the ground by KFM were pilfered by CWR. CWR did not alter hoarding intensity in the presence of KFM and their stores cannot be pilfered by KFM. These results indicate that large-sized rodent species (more dominant) significantly increase the hoarding intensity of small-sized species and show a unidirectional pilferage of seeds cached by small-sized species. The behavioural differences between these two species may reduce competition for resources and promote coexistence between sympatric rodents.

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