[All these photos and videos were taken last week at Gardens in the Wild in Framingham, Massachussets, the home of the New England Wildflower Society. Highly recommended! and the frogs are used to people, so they are much less skittish than they are around me in Maine. ]
The American Bullfrog, Lithobates catesbeianus, is a substantial chap. He can weigh up to 0.5Kg, more than 1lb. The bullfrog male’s mating call is low in pitch, with a frequency around 200-300Hz, and can be heard for up to one kilometer. Here is a recording (made in Maine a year or two ago.)
These calls are very very loud, and yet they call with their mouths closed. How do they do it?
When the male calls, he fills his bright yellow throat pouches with air. Each call lasts approximately 0.5 seconds: I have slowed down this video so you see this more clearly:
I had always thought that was why his calls were so loud.
But it turns out that is wrong. The inflated pouches are used as a sort of air reservoir so that air can be recycled to the lungs across the vocal folds where the sound is actually produced. Some sound is indeed then transmitted to the world through these pouches, but that is not the main source. The male has a greatly enlarged tympanum (ear-drum), the round patch behind the eye, as you can see in the photo.

It is this which hugely amplifies the sound energy, being responsible for some 98% of the energy actually transmitted. (Purgue 1997). Its resonating frequencies match those of the calls, so effectively it acts as a drum. (Purgue says you can see the eardrum move, but even when I slow my videos right down I can’t detect this!).
Even if I can’t see the eardrums move, what I can see is the whole body vibrations that are transmitted to the water: look at those ripples radiating outward as the frog calls.

Note also how much smaller the female’s ear-drum is: hers is used just to transmit sound to her inner ear, like ours.

There is always something new to learn: who knew that ear-drums could be a two-way sound system. They even look just like a high-end stereo speaker!

PS More on the generation of the initial sounds: (Ryan and Guerra 2014) ” In most frogs, air is expelled by contracting trunk muscles surrounding the lungs, which pushes the air through the larynx .. The incoming air causes the vibration of the vocal cords and the larynx itself. The air then enters the buccal cavity and passes through the vocal slits to inflate the vocal sac. One of the most conspicuous and near-universal traits of male frogs is the vocal sac. Its main function is to recycle air from the lungs to the vocal sac and back again. The vocal sac also radiates sound.”
PPS The kettle-drums used in classical music are more formally called timpani, essentially the plural of the same word as an ear-drum, tympanum. Hence my title.
PPPS It took about 250 photos to get those two still shots of the 0.5 second period when the vocal sac is inflated!