The problematic Cambrian arthropod Tuzoia and the origin of mandibulates revisited

The origin of mandibulates, the hyperdiverse arthropod group that includes pancrustaceans and myriapods, dates back to the Cambrian. Bivalved arthropod groups such as hymenocarines have been argued to be early mandibulates, but many species are still poorly known, and their affinities remain uncertain. One of the most common and globally distributed Cambrian bivalved arthropods is Tuzoia. Originally described in 1912 from the Burgess Shale based on isolated carapaces, its full anatomy has remained largely unknown. Here, we describe new specimens of Tuzoia from the Canadian Burgess Shale (Wuliuan, Cambrian) showcasing exceptionally preserved soft tissues, allowing for the first comprehensive reconstruction of its anatomy, ecology and evolutionary affinities. The head bears antennae and differentiated cephalic appendages. The body is divided into a cephalothorax, a homonomous trunk bearing ca 10 pairs of legs with heptopodomerous endopods and enlarged basipods, and a tail fan with two pairs of caudal rami. These traits suggest that Tuzoia swam along the seafloor and used its spinose legs for predation or scavenging. Tuzoia is retrieved by a Bayesian phylogenetic analysis as an early mandibulate hymenocarine lineage, exemplifying the rapid diversification of this group in open marine environments during the Cambrian Explosion.