The oldest articulated chondrichthyan from the Early Devonian period.

Chondrichthyans (including living sharks, skates, rays and chimaeras) have a fossil record of scales and dermal denticles perhaps dating back to the Late Ordovician period, about 455 million years ago. Their fossil tooth record extends to the earliest Devonian period, almost 418 million years ago, whereas the oldest known articulated shark remains date from the Early Devonian period, about 394 million years ago. Here we report the discovery of an articulated shark that is almost 409 million years old from the Early Devonian (early Emsian) period of New Brunswick, Canada. The specimen, identified as Doliodus problematicus (Woodward), sheds light on the earliest chondrichthyans and their interrelationships with basal jawed vertebrates. This species has been truly problematic. Previously known only from isolated teeth, it has been identified as an acanthodian and a chondrichthyan. This specimen is the oldest shark showing the tooth families in situ, and preserves one of the oldest chondrichthyan braincases. More notably, it shows the presence of paired pectoral fin-spines, previously unknown in cartilaginous fishes.