The edge regions in tergites of the desert isopod Hemilepistus reaumuri : the transition from hard cuticle to flexible a
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T.C. BIOLOGICAL AND BIOMIMETIC MATERIALS
The edge regions in tergites of the desert isopod Hemilepistus reaumuri: the transition from hard cuticle to flexible arthrodial membrane Franziska Ernst1 · Helge‑Otto Fabritius2,3 · Erika Griesshaber4 · Wolfgang W. Schmahl4 · Andreas Ziegler1 Received: 26 June 2020 / Accepted: 29 August 2020 / Published online: 17 September 2020 © The Author(s) 2020
Abstract The arthrodial membrane is a thin and flexible type of cuticle that inserts at the edge regions of neighbouring rigid skeletal elements creating a flexible connection. In the present study, we analyzed the structure, mineral composition, calcite organization and local stiffness and hardness of edge regions that form transitions to the arthrodial membranes in the tergites of the desert isopod Hemilepistus reaumuri. For the transitions to the arthrodial membrane, the results show an increase in the thickness of the epicuticle at cost of the distal exocuticle and a calcite layer, an increase in the ratio of phosphorus to calcium and a decrease in the local mechanical properties. The posterior edge region contains an unusually large stack of unidirectionally oriented parallel fibrils projecting to the lateral sides. At the edge, it turns down into a long ventral cuticle overlapping an anterior part of the neighbouring tergite. It forms a thin arched gap between the tergites that can help reducing water loss through the arthrodial membrane and protects the arthrodial membrane upon predation. A thick ventral ridge near the transition to the arthrodial membrane carrying bristles can prevent sand grains from access to the arthrodial membrane. From the dorsal cuticle to the transition to the arthrodial membrane, calcite units become larger and single crystalline turning their c-axes orientation perpendicular to the sagittal section plane. Comparison with edge regions of the beach isopod Tylos europaeus reveal common characteristics of the edge region, but also specific adaptations to the desert habitat of H. reaumuri. Keywords Biological–material functionalization · Crustacea · Cuticle · Amorphous calcium phosphate · Calcite co-orientation strength
1 Introduction
Electronic supplementary material The online version of this article (https://doi.org/10.1007/s00339-020-03961-0) contains supplementary material, which is available to authorized users. * Andreas Ziegler andreas.ziegler@uni‑ulm.de 1
Central Facility for Electron Microscopy, University of Ulm, Albert‑Einstein‑Allee 11, 89069 Ulm, Germany
2
Bionics and Materials Development, Hamm-Lippstadt University of Applied Sciences, Marker Allee 76‑78, 59063 Hamm, Germany
3
Department of Microstructure Physics and Alloy Design, Max-Planck-Institut für Eisenforschung GmbH, Max‑Planck‑Str. 1, 40237 Düsseldorf, Germany
4
Department of Earth and Environmental Sciences, LMU, Theresienstr. 41, 80333 Munich, Germany
The cuticle of Crustacea has a complex hierarchical organization which has attracted extensive interest during the last decade [1]. Its basic func
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