Toughening mechanisms of the elytra of the diabolical ironclad beetle

Rivera, Jesus; Hosseini, Maryam Sadat; Restrepo, David; Murata, Satoshi; Vasile, Drago; Parkinson, Dilworth Y.; Barnard, Harold; Arakaki, Atsushi; Zavattieri, Pablo; Kisailus, David

doi:10.1038/s41586-020-2813-8

Cited by 148 publications

(101 citation statements)

References 28 publications

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“…An extreme case of mechanical strengthening of the elytral body cover occurs in the "diabolically ironclad beetle" N o s o d e r m a d i a b o l i c u m L e C o n t e ( Z o p h e r i d a e , Tenebrionoidea) (Rivera et al 2020). Another unique and enhanced type of locking system has evolved in Pselaphinae, arguably a key feature in the evolution of this successful subfamily.…”

Section: Elytral Locking System and Flightlessnessmentioning

confidence: 99%

The specialized thoracic skeletomuscular system of the myrmecophile Claviger testaceus (Pselaphinae, Staphylinidae, Coleoptera)

et al. 2021

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External and internal structures of the thorax of the myrmecophile beetle Claviger testaceus (Clavigeritae, Pselaphinae) were examined and documented with state-of-the-art visualization techniques. Following a general trend in the omaliine lineage (Staphylinidae), the skeletal elements of the pro- and pterothorax in Claviger reach a maximum degree of compactness, with largely reduced inter- and intrasegmental sutures and skeletal elements linked with the flight apparatus. The musculature, especially metathoracic direct and indirect flight muscles, also shows a high degree of reduction. Two forms of wings were found among individuals of C. testaceus, both non-functional and representing an advanced stage of reduction. However, that wing vestiges are still present and the metanotum, only slightly reduced, suggests that loss of flight in this species is likely the result of a young evolutionary process. Several structures are linked with myrmecophilous habits: small body size facilitates transportation of beetles by ant workers and makes it easier to move inside nest tunnels; the remarkably compact body and mechanically robust appendages make the beetles less vulnerable to attacks by ant mandibles; the improved elytral interlocking mechanism and unusually expanded epipleura enhance the protection of vulnerable dorsal parts of the pterothorax and anterior abdomen; and glands associated with trichomes on the posterolateral elytral angle produce secretions attractive for ants. Various modifications of the thorax and anterior abdomen lead to an optimization of intimate associations with ants. The morphological syndrome enabling these beetles to cope with life in ant colonies evolved in several steps. This is suggested by an increasing solidification of the thoracic skeleton in related non-myrmecophilous groups and also by less modified related clavigerites;for instance, ant-associated tropical species are still able to fly.

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Section: Elytral Locking System and Flightlessnessmentioning

confidence: 99%

The specialized thoracic skeletomuscular system of the myrmecophile Claviger testaceus (Pselaphinae, Staphylinidae, Coleoptera)

et al. 2021

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“…They help to explain why both young thin- and thick-walled pistachio shell tissue showed a similar strain ( figure 4 a ) and fracture through the CW ( figure 5 c ). Interdigitated sutures together with laminated microstructures have recently been shown to be the most important toughening mechanisms in the elytra of the diabolical ironclad beetle [ 40 ]. In seahorse armour, interlocking joints of the bony plates achieve large deformations [ 41 , 42 ].…”

Section: Discussionmentioning

confidence: 99%

Twist and lock: nutshell structures for high strength and energy absorption

et al. 2021

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Nutshells achieve remarkable properties by optimizing structure and chemistry at different hierarchical levels. Probing nutshells from the cellular down to the nano- and molecular level by microchemical and nanomechanical imaging techniques reveals insights into nature's packing concepts. In walnut and pistachio shells, carbohydrate and lignin polymers assemble to form thick-walled puzzle cells, which interlock three-dimensionally and show high tissue strength. Pistachio additionally achieves high-energy absorption by numerous lobes interconnected via ball-joint-like structures. By contrast, the three times more lignified walnut shells show brittle LEGO-brick failure, often along the numerous pit channels. In both species, cell walls (CWs) show distinct lamellar structures. These lamellae involve a helicoidal arrangement of cellulose macrofibrils as a recurring motif. Between the two nutshell species, these lamellae show differences in thickness and pitch angle, which can explain the different mechanical properties on the nanolevel. Our in-depth study of the two nutshell tissues highlights the role of cell form and their interlocking as well as plant CW composition and structure for mechanical protection. Understanding these plant shell concepts might inspire biomimetic material developments as well as using walnut and pistachio shell waste as sustainable raw material in future applications.

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“…[16,17] Moreover, the appropriate orientation of phase field can enhance the fracture toughness without major compromise to stiffness and strength. [11,[18][19][20][21][22][23][24][25] Hence, the nacre with laminated structure is twice as hard and 1000 times as tough as its constituent phases, providing strength, hardness, and toughness. [10] Such investigations inspire us to design an oriented and gradient structure in the cocontinuous phase, which is highly expected to have excellent performance.…”

Section: Doi: 101002/macp202100012mentioning

confidence: 99%

Imparting Gradient and Oriented Characters to Cocontinuous Structure for Improving Integrated Performance

Sun

Song

et al. 2021

Macro Chemistry & Physics

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Inspired by the oriented, gradient, and heterogeneous characters of natural materials, an oriented and gradient cocontinuous structure is designed to obtain excellent mechanical performance via a gradient shear stress and temperature during melt processing. The investigation of hierarchical structure containing phase morphology, crystalline morphology, and lamellae in the cocontinuous structure can clarify the role of gradient shear stress and temperature in promoting gradient and oriented features. Given such oriented and gradient features in the cocontinuous structure, a notable enhancement in the tensile strength, tensile modulus, elongation at break, and impact strength is achieved in the model poly(butylene adipate‐ran‐terephthalate)/poly(butylene succinate) biodegradable cocontinuous blends. Compared with the isotropic cocontinuous structure, homogenous structure and discrete phase structure, oriented and gradient cocontinuous structure provides a promising balance of strength, hardness, and toughness. Local reinforcement is achieved by highly oriented cocontinuous phase, crystal, and lamellae in skin layer. Meanwhile, the toughness is achieved by weakly oriented cocontinuous phase. The tuning of cocontinuous structure with gradient and oriented features is suggested as a new pathway toward the development of “materials systems” with unprecedented mechanical performance.

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Toughening mechanisms of the elytra of the diabolical ironclad beetle

Cited by 148 publications

References 28 publications

The specialized thoracic skeletomuscular system of the myrmecophile Claviger testaceus (Pselaphinae, Staphylinidae, Coleoptera)

The specialized thoracic skeletomuscular system of the myrmecophile Claviger testaceus (Pselaphinae, Staphylinidae, Coleoptera)

Twist and lock: nutshell structures for high strength and energy absorption

Imparting Gradient and Oriented Characters to Cocontinuous Structure for Improving Integrated Performance

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