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Updated: Apr 11, 2026

Characterization Of Multi-layered Fish Scales Atractosteus spatula Using Nanoindentation, X-ray CT, FTIR, and SEM
Published on: July 10, 2014
The armored carapace of the boxfish
Wen Yang1, Steven E Naleway1, Michael M Porter1
1Materials Science and Engineering Program, University of California, San Diego, La Jolla, CA 92093-0411, USA.
Abstract:
The boxfish (Lactoria cornuta) has a carapace consisting of dermal scutes with a highly mineralized surface plate and a compliant collagen base. This carapace must provide effective protection against predators as it comes at the high cost of reduced mobility and speed. The mineralized hydroxyapatite plates, predominantly hexagonal in shape, are reinforced with raised struts that extend from the center toward the edges of each scute. Below the mineralized plates are non-mineralized collagen fibers arranged in through-the-thickness layers of ladder-like formations. At the interfaces between scutes, the mineralized plates form suture-like teeth structures below which the collagen fibers bridge the gap between neighboring scutes. These sutures are unlike most others as they have no bridging Sharpey's fibers and appear to add little mechanical strength to the overall carapace. It is proposed that the sutured interface either allows for accommodation of the changing pressures of the boxfish's ocean habitat or growth, which occurs without molting or shedding. In both tension and punch testing the mineralized sutures remain relatively intact while most failures occur within the collagen fibers, allowing for the individual scutes to maintain their integrity. This complex structure allows for elevated strength of the carapace through an increase in the stressed area when attacked by predators in both penetrating and crushing modes.
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