Cranial kinesis in palaeognathous birds.

Cranial kinesis in birds is induced by muscles located caudal on the cranium. These forces are transferred onto the moveable parts of the skull via the Pterygoid-Palatinum Complex (PPC). This bony structure therefore plays an essential role in cranial kinesis. In palaeognathous birds the morphology of the PPC is remarkably different from that of neognathous birds and is thought to be related to the specific type of cranial kinesis in palaeognaths known as central rhynchokinesis. We determined whether clear bending zones as found in neognaths are present in the upper bill of paleognaths, and measured bending forces opposing elevation of the upper bill. A static force model was used to calculate the opening forces that can be produced by some of the palaeognathous species. We found that no clear bending zones are present in the upper bill, and bending is expected to occur over the whole length of the upper bill. Muscle forces are more than sufficient to overcome bending forces and to elevate the upper bill. The resistance against bending by the bony elements alone is very low, which might indicate that bending of bony elements can occur during food handling when muscles are not used to stabilise the upper bill. Model calculations suggest that the large processi basipterygoidei play a role in stabilizing the skull elements, when birds have to resist external opening forces on the upper bill as might occur during tearing leafs from plants. We conclude that the specific morphology of the palaeognathous upper bill and PPC are not designed for active cranial kinesis, but are adapted to resist external forces that might cause unwanted elevation of the upper bill during feeding.