Frequency-dependent selection by tree squirrels: adaptive escape of nondormant white oaks

Frequency-dependent selection is theoretically predicted to have an important impact on predator-prey dynamics, but little is understood about this process in nature. We investigated oak-squirrel interactions to test the hypothesis that tree squirrels predate/disperse white oak acorns based on the frequency of their germination phenotypes (dormancy, i.e., delayed germination vs. nondormancy, i.e., immediately germinating), which in turn influences dispersal and establishment of these oaks. In Central China, we manipulated the relative frequency of dormant and nondormant acorns available to Pere David's Rock squirrel (Sciurotamias davidianus) with 2 white oaks (QAA, Quercus aliena var. acutesevata and QSB, Quercus serrata var. breviptiolata) that exhibit considerable variation in acorn germination between and within oak species throughout the fruiting season. We predicted that dispersal success (i.e., the probability of a viable acorn being hoarded) of each germination phenotype would vary with the phenotype frequency and hence show either positive or negative frequency-dependent selection. We found that acorn-embryo removal by squirrels may select for the dormant phenotype but is more likely to occur for early germinating acorns. We also found that both absolute and relative dispersal success of each germination phenotype increased with their frequency in the 2 white oaks, thus demonstrating positive frequency-dependent selection. Our results reveal that the frequency of germination phenotypes can account for as much as 40.5% and 21.4% of the observed dispersal success in QAA and QSB, respectively. We show that behavioral preferences of scatter-hoarding animals could result in frequency-dependent selection in nature, and such selection may help maintain variation in germination phenotypes in prey species.