The role of the tymbal in cicada sound production.

1. The tymbal of Cyclochila australasiae consists of a biconvex membrane bearing alternating long and short ribs anteriorly and an irregularly shaped tymbal plate posteriorly. These sclerotised regions are coupled together by the surrounding highly flexible cuticle, which contains resilin. Dorsally, there is a thick pad of resilin, which functions as a spring, returning the tymbal to the out position and maintaining the stress on the long ribs. 2. Contraction of the tymbal muscle causes the tymbal plate to swing inwards, acting as a lever so that the surface of the tymbal moves through more than twice the distance of muscle shortening. This produces an inward movement and twisting of the dorsal ends of the long ribs, which then buckle in sequence, with each rib undergoing a sudden deformation from a convex to a V-shaped profile. Buckling takes place at the rib's weakest point, which is the narrow, highly sclerotised mid-region. 3. Inward buckling of the tymbal generates a loud click with a dominant frequency around 4 kHz. Resonances close to 4 kHz can be demonstrated in a buckled-in tymbal when driven by internal sound or by vibration at the tymbal plate. These resonances occur in sealed cicadas and those in which the abdominal air sac has been opened at both its anterior and posterior ends, which shows that the resonances are not due to the air sac; the tymbal itself is a resonant system. The maximum amplitude of tymbal vibration occurs at the V-shaped dimples in the centre of the long ribs. 4. When the tymbal plus abdominal air sac system is driven by vibration at the tymbal plate, the Q3dB of the sound radiated through the tympana is about 12.5, which is approximately the sum of those of the tymbal (Q=9.3) and of the air sac (Q=3.4) resonators. When the tymbal is not loaded by the air sac, i.e. in the sealed cicada and open cicada preparations, the Q3dB of its resonance is higher, between 13 and 20. 5. The click produced as the tymbal pops out is over 20 dB quieter than the in-click and has a dominant frequency around 6 kHz. When driven in the resting position, resonances are found close to 6 kHz but there is only a weak general vibration of the ribs and tymbal plate. When the tymbal is pushed in gradually, the resonant frequency changes from about 5.5 kHz to about 4.3 kHz as the tymbal buckles inwards. The left and right tymbals of the same insect may differ slightly in their acoustic properties. 6. As the tymbal buckles inwards, it displaces a volume of approximately 6 µl into the abdominal air sac volume of about 2 ml. The resulting sound pressure inside the air sac attains peak values of 155­159 dB SPL; the root mean square values are 141­144 dB SPL. The mean peak value just outside the tympana is 148.5 dB SPL. 7. Overall, the present work supports and extends our earlier model of cicada sound production: the tymbal click provides a coherent resonant source that drives the abdominal resonator, from which sound is radiated via the tympana. At the same time, the system provides the pressure transformation between muscle power and sound power that is desirable for efficient sound radiation.