Mineralized belemnoid cephalic cartilage from the late Triassic Polzberg Konservat-Lagerstatte (Austria)

Although hyaline cartilage is widely distributed in various invertebrate groups such as sabellid polychaetes, molluscs (cephalopods, gastropods) and a chelicerate arthropod group (horseshoe crabs), the enigmatic relationship and distribution of cartilage in taxonomic groups remains to be explained. It can be interpreted as a convergent trait in animal evolution and thus does not seem to be a vertebrate invention. Due to the poor fossil record of cartilaginous structures, occurrences of mineralized fossil cartilages are important for evolutionary biology and paleontology. Although the biochemical composition of recent cephalopod cartilage differs from vertebrate cartilage, histologically the cartilages of these animal groups resemble one another remarkably. In this study we present fossil material from the late Triassic Polzberg Konservat-Lagerstatte near Lunz am See (Lower Austria, Northern Calcareous Alps). A rich Carnian fauna is preserved here, whereby a morphogroup (often associated with belemnoid remains) of black, amorphous appearing fossils still remained undetermined. These multi-elemental, symmetrical fossils show remarkable similarities to recent cartilage. We examined the conspicuous micro- and ultrastructure of these enigmatic fossils by thin-sectioning and Scanning Electron Microscopy (SEM). The geochemical composition analyzed by Microprobe and Energy Dispersive X-ray Spectroscopy (SEM-EDX) revealed carbonization as the taphonomic pathway for this fossil group. Mineralization of soft tissues permits the 3D preservation of otherwise degraded soft tissues such as cartilage. We examined eighty-one specimens from the Polzberg locality and seven specimens from Cave del Predil (formerly Raibl, Julian Alps, Italy). The study included morphological examinations of these multi-elemental fossils and a focus on noticeable structures like grooves and ridges. The detected grooves are interpreted to be muscular attachment areas, and the preserved branched system of canaliculi is comparable to a channel system that is also present in recent coleoid cartilage. The new findings on these long-known enigmatic structures strongly point to the preservation of cephalic cartilage belonging to the belemnoid Phragmoteuthis bisinuata and its homologization to the cephalic cartilage of modern coleoids.