The effects of tracheal coiling on the vocalizations of cranes (Aves; Gruidae)

Summary

It is generally supposed that the elongated, often coiled tracheae of many species of birds are adaptations for the production of loud, penetrating calls. A corollary supposition is that the acoustic effects are produced by the resonant properties of the elongated tube, with the birds being analogized to a wind instrument. We have experimented with several species of cranes possessing different degrees of tracheal coiling. Regardless of the degree of coiling, all cranes can utter extremely loud calls using remarkably low driving pressures. Neither surgical modifications of the trachea nor changing the respiratory gases to helium-oxygen produced consistent changes of voice that could be unambiguously attributed to changes of tubal resonances. However, shortening the trachea markedly reduced vocal intensity, the degree of reduction being roughly proportional to the degree of shortening. Although some of that reduction may derive from an increased impedance mismatch at the external aperture of the tube, and some from a decreased radiation directly from the hard walls of the trachea, these explanations scarcely account for the dramatic effects we observed. We, therefore, hypothesize a more unusual mechanism: The tracheal coils that are embedded in the sternum serve a function analogous to the bridge of a stringed instrument, transmitting the vibrations of a tiny sound source to a large radiating surface, the sternum. The sternum then vibrates against the large internal air reservoir of the avian airsac system. As it has a complex shape, the sternum will have many resonances and will respond to many frequencies; as a solid oscillator, its resonances will not be greatly affected by low density gases. Hence, we suggest that cranes and other birds with enlarged windpipes are more properly analogized with a violin than a trombone.