Abstract
Atractosteus spatula has been described as a living fossil (having existed for 100 Myr), retaining morphological characteristics of early ancestors such as the ability to breathe air and survive above water for hours. Its highly effective armour consists of ganoid scales. We analyse the protective function of the scales and identify key features which lead to their resistance to failure. Microstructural features include: a twisted cross-plied mineral arrangement that inhibits crack propagation in the external ganoine layer, mineral crystals that deflect cracks in the bony region in order to activate the strength of mineralized collagen fibrils, and saw-tooth ridges along the interface between the two scale layers which direct cracks away from the intrinsically weak interface. The macroscale geometry is additionally evaluated and it is shown that the scales retain full coverage in spite of minimal overlap between adjacent scales while conforming to physiologically required strain and maintaining flexibility via a process in which adjacent rows of scales slide and concurrently reorient.