title: Pangolin armor: Overlapping, structure, and mechanical properties of the keratinous scales.
abstract: The pangolin has a flexible dermal armor consisting of overlapping keratinous scales. Although they show potential for bioinspired flexible armor, the design principles of pangolin armor are barely known. Here we report on the overlapping organization, hierarchical structure (from the nano to the mesolevel), and mechanical response of scales from ground (Chinese) and arboreal (African tree) pangolins. Both scales exhibit the same overlapping organization, with each scale at the center of neighboring scales arranged in a hexagonal pattern. The scales have a cuticle of several layers of loosely attached flattened keratinized cells, while the interior structure exhibits three regions distinguished by the geometry and orientations of the keratinized cells, which form densely packed lamellae; each one corresponds to one layer of cells. Unlike most other keratinous materials, the scales show a crossed-lamellar structure (∼5μm) and crossed fibers (∼50μm). A nano-scale suture structure, observed for the first time, outlines cell membranes and leads to an interlocking interface between lamellae, thus enhancing the bonding and shear resistance. The tensile response of the scales shows an elastic limit followed by a short plateau prior to failure, with Young's modulus ∼1 GPa and tensile strength 60-100MPa. The mechanical response is transversely isotropic, a result of the cross lamellar structure. The strain rate sensitivity in the range of 10(-5)-10(-1)s(-1) region is found to be equal to 0.07-0.08, typical of other keratins and polymers. The mechanical response is highly dependent on the degree of hydration, a characteristic of keratins.
  
  
  
  Although many fish and reptiles have protective scales and carapaces, mammals are characteristically fast and light. The pangolin is one of the few mammal possessing a flexible dermal armor for protection from predators, such as lions. Here we study the arrangement of the scales as well as their hierarchical structure from the nano to the mesolevel and correlate it to the mechanical properties. The study reveals a unique structure consisting of crossed lamellae and interlocking sutures that provide exceptional performance and in-plane isotropy.
authors.affiliation: Materials Science and Engineering Program, University of California, San Diego, CA 92093, USA.
authors.forename: Bin
authors.initials: B
authors.lastname: Wang
authors.affiliation: Materials Science and Engineering Program, University of California, San Diego, CA 92093, USA; Department of Materials, ETH Zürich, Zürich 8093, Switzerland. Electronic address: wey005@eng.ucsd.edu.
authors.forename: Wen
authors.initials: W
authors.lastname: Yang
authors.affiliation: Materials Science and Engineering Program, University of California, San Diego, CA 92093, USA.
authors.forename: Vincent R
authors.initials: VR
authors.lastname: Sherman
authors.affiliation: Materials Science and Engineering Program, University of California, San Diego, CA 92093, USA; Department of Mechanical and Aerospace Engineering, University of California, San Diego, CA 92093, USA; Department of NanoEngineering, University of California, San Diego, CA 92093, USA. Electronic address: mameyers@eng.ucsd.edu.
authors.forename: Marc A
authors.initials: MA
authors.lastname: Meyers
mesh_terms: D000825:Animal Structures / Q000033:anatomy & histology* / Q000502:physiology*; D000818:Animals; D001696:Biomechanical Phenomena; D002462:Cell Membrane / Q000648:ultrastructure; D019245:Compressive Strength; D017709:Fractals; D006244:Hardness; D007633:Keratins / Q000737:chemistry*; D008322:Mammals / Q000033:anatomy & histology*; D013718:Tensile Strength
chemical_list: D007633:Keratins
keywords: Crossed lamellae & crossed fibers; Interlocking interface; Pangolin scale; Strain rate sensitivity; Suture-like cell membrane complex; Transverse isotropic