The skeletal completeness of the Palaeozoic chondrichthyan fossil record.

Chondrichthyes (sharks, rays, ratfish and their extinct relatives) originated and diversified in the Palaeozoic but are rarely preserved as articulated or partly articulated remains because of their predominantly cartilaginous endoskeletons. Consequently, their evolutionary history is perceived to be documented predominantly by isolated teeth, scales and fin spines. Here, we aim to capture and analyse the quality of the Palaeozoic chondrichthyan fossil record by using a variation of the skeletal completeness metric, which calculates how complete the skeletons of individuals are compared to estimates of their original entirety. Notably, chondrichthyan completeness is significantly lower than any published vertebrate group: low throughout the Silurian and Permian but peaking in the Devonian and Carboniferous. Scores increase to a range similar to pelycosaurs and parareptiles only when taxa identified solely from isolated teeth, scales and spines are excluded. We argue that environmental influences probably played an important role in chondrichthyan completeness. Sea level significantly negatively correlates with chondrichthyan completeness records and resembles patterns already evident in records of ichthyosaurs, plesiosaurs and sauropodomorphs. Such observed variations in completeness highlight the impact of different sampling biases on the chondrichthyan fossil record and the need to acknowledge these when inferring patterns of chondrichthyan macroevolution.