Abstract

Members of the Chondrichthyes (Elasmobranchii and Holocephali) are distinguished by their largely cartilaginous endoskeletons, which comprise an uncalcified core overlain by a mineralized layer; in the Elasmobranchii (sharks, skates, rays) most of this mineralization takes the form of calcified polygonal tiles known as tesserae. In recent years, these skeletal tissues have been described in ever increasing detail in sharks and rays, but those of Holocephali (chimaeroids) have been less well-studied, with conflicting accounts as to whether or not tesserae are present. During embryonic ontogeny in holocephalans, cervical vertebrae fuse to form a structure called the synarcual. The synarcual mineralizes early and progressively, anteroposteriorly and dorsoventrally, and therefore presents a good skeletal structure in which to observe mineralized tissues in this group. Here, we describe the development and mineralization of the synarcual in an adult and stage 36 elephant shark embryo (Callorhinchus milii). Small, discrete, but irregular blocks of cortical mineralization are present in stage 36, similar to what has been described recently in embryos of other chimaeroid taxa such as Hydrolagus, while in Callorhinchus adults, the blocks of mineralization are more irregular, but remain small. This differs from fossil members of the holocephalan crown group (Edaphodon), as well as from stem group holocephalans (e.g., Symmorida, Helodus, Iniopterygiformes), where tesserae are notably larger than in Callorhinchus and show similarities to elasmobranch tesserae, for example with respect to polygonal shape.

Highlights

  • During ontogeny, most vertebrate skeletons are initially composed predominantly of hyaline cartilage and largely replaced by bone via endochondral ossification (Hall, 1975, 2005)

  • We report the presence of a layer of mineralization in the Callorhinchus embryo, maintained in adults, comprising small, irregularly shaped units, but lacking many of the characteristics of tesserae in the elasmobranchs

  • General Morphology The axial skeleton of chondrichthyans typically includes a series of cartilages dorsal and ventral to the notochord, and in the elasmobranchs, centra associated with the notochord (e.g., Dean, 1895; Gadow and Abbott, 1895; Goodrich, 1930; Compagno, 1977; Criswell et al, 2017b)

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Summary

Introduction

Most vertebrate skeletons are initially composed predominantly of hyaline cartilage and largely replaced by bone via endochondral ossification (Hall, 1975, 2005). Mineralization of the Callorhinchus Vertebral Column of hyaline-like cartilage, with elasmobranchs developing a comparatively thin outer layer of cortical mineralization over most of their skeleton during ontogeny (Hall, 2005; Egerbacher et al, 2006; Dean et al, 2009, 2015; Seidel et al, 2016, 2019b; Atake et al, 2019; Debiais-Thibaud, 2019) This mineralized tissue begins as small separated islets near the cartilage surface, which gradually grow via mineral accretion to fill the intervening spaces, eventually forming a cortex of abutting polygonal tiles called tesserae (Dean and Summers, 2006; Dean et al, 2009, 2015; Seidel et al, 2016, 2019b). This mosaic of uncalcified cartilage, tesserae and perichondrium is called tessellated cartilage and comprises most of the cranial and postcranial skeleton (Kemp and Westrin, 1979; Dean and Summers, 2006; Seidel et al, 2016, 2017a)

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