Spectrographic analysis showed that dholes produce sounds with two fundamental frequencies (components): the high-frequency and the low-frequency, which may occur simultaneously or separately. The fundamental frequency of the low-frequency component varied from 0.52 to 1.44 kHz, and that of the high-frequency component from 5.51 to 10.77 kHz. In calls where both the frequencies occurred simultaneously, they were not integer multiples of each other. They also had different frequency modulations and had additional combination frequency bands. These are features of biphonation. Our data showed biphonation occurs in a lot of dhole calls (20 to 92% of the calls, depending on the individual (n=14); average occurrence 44.3%, n=1317 sounds). The occurrence of biphonation did not differ between sexes and ages; however, occurrence of the high component only was significantly higher in subadult animals, whereas the occurrence of the low component only was significantly higher in adults. Based on the sound structures, we discuss probable mechanisms of sound production for both the components in the dhole. For the low component, the normal vocal folds oscillation mechanism is suggested. For the high component, four possible mechanisms of sound production are discussed. We conclude that the vortex-shedding mechanism is the most probable.
vocalization, nonlinear phenomena, biphonation, vocal membrane, whistle, Canidae, dhole, Cuon alpinus