Discovery of the earliest-known tetrapod stapes.

The evolution of the middle ear is central to the discussion of how the first tetrapods adapted to life on land as well as their phylogeny. Here I report the discovery of the stapes of Acanthostega gunnari, from the Upper Devonian of east Greenland. This is the earliest tetrapod stapes so far described, and it throws new light on both these aspects of early tetrapod biology. It has been assumed that the common inheritance of all early tetrapods was a light, rod-like stapes associated with a temporal notch in the otic region that was thought to have supported a tympanum, or eardrum. The stapes would have conducted vibrations from the tympanum to the otic capsule. By contrast, the stapes of Acanthostega was stout with a broad distal ramus associated with the temporal notch. I suggest that the temporal notch of Acanthostega and other early tetrapods supported a spiracular opening rather than a tympanum, and that the stapes controlled palatal and spiracular movements in ventilation.