The high toughness and work to fracture of hierarchical composites, like antler bone, involve structural 13 mechanisms at the molecular, nano-and micro scales, which are not completely explored. A key 14 characteristic of the high energy absorption of such materials is the large hysteresis during cyclic loading, but 15 its origin remains unknown. In situ synchrotron X-ray diffraction tests during tensile loading of antler bone 16 showed heterogeneous fibrillar deformation and hysteresis. To explain the origin of these mechanisms from 17 the nanostructure of antler bone, here we develop a class of finite-element fibril models whose predictions 18 are compared to experimental data across a range of potential composite architectures. We demonstrate 19 that the key structural motif enabling a match to experimental data is an axially staggered arrangement of 20 stiff mineralized collagen fibrils coupled with weak, damageable interfibrillar interfaces. 21 22 between structural properties at different scales 3. Bio-composites, such as bone, shells 29 and nacre, represent an excellent example of how the design at lower hierarchical scales 30 confers higher mechanical properties than the single constituents 4. Although the stiffness 31 of these biocomposites is comparable to that of the basic constituent at the nanoscale, 32 their toughness results hugely increased. For instance, in bone and shell, the toughness of 33 the mineral constituents is << 1MPa*m 1/2 while the toughness of their macrostructure 34 varies, respectively, in a range of 2-7 MPa*m 1/2 and 3-7 MPa*m 1/2. 35 36 Bone, as shown in Figure 1, at the nanometre scale length is a composite of stiff inorganic 37 hydroxyapatite platelets interleaved with a softer organic matrix, made principally of type I 38 tropocollagen proteins 5. This sub-structure, together with an intrafibrillar phase of 39 noncollageneous proteins and mineral, forms mineralized fibrils that are arranged into 40 aggregate structures at higher levels and larger length scales, such as fibril arrays and 41 lamellae 1. The structural aspects of this architecture served as inspiration for bioinspired 42 materials that replicate the nanometre scale fibril-matrix 6-10 and intrafibrillar 11 structure, or 43 at micrometre scales 12,13. Nonetheless, the mechanical interactions between the 44 constituent units and the higher length scales remain a matter of active research. In 45 particular, previous studies focused on how the hierarchical architecture brings functionally 46 desirable properties such as high toughness 14 , energy absorption and fatigue resistance 15. 47