A comparative study of the ocular skeleton of fossil and modern chondrichthyans

Many vertebrates have an ocular skeleton composed of cartilage and/or bone situated within the sclera of the eye. In this study we investigated whether modern and fossil sharks have an ocular skeleton, and whether it is conserved in morphology. We describe the scleral skeletal elements of three species of modern sharks and compare them to those found in fossil sharks from the Cleveland Shale (360 Mya). We also compare the elements to contemporaneous arthrodires from the same deposit. Surprisingly, the morphology of the skeletal support of the eye was found to differ significantly between modern and fossil sharks. All three modern shark species examined (spiny dogfish shark Squalus acanthias, porbeagle shark Lamna nasus and blue shark Prionace glauca) have a continuous skeletal element that encapsulates much of the eyeball; however, the tissue composition is different in each species. Histological and morphological examination revealed scleral cartilage with distinct tesserae in parts of the sclera of the porbeagle and blue shark, and more diffuse calcification in the dogfish. Strengthening of the scleral cartilage by means of tesserae has not been reported previously in the shark eye. In striking contrast, the ocular skeleton of fossil sharks comprises a series of individual elements that are arranged in a ring, similar to the arrangement in modern and fossil reptiles. Fossil arthrodires also have a multi-unit sclerotic ring but these are composed of fewer elements than in fossil sharks. The morphology of these elements has implications for the behaviour and visual capabilities of sharks that lived during the Devonian Period. This is the first time that such a dramatic variation in the morphology of scleral skeletal elements has been observed in a single lineage (Chondrichthyes), making this lineage important for broadening our understanding of the evolution of these elements within jawed vertebrates.