Extreme multisegmentation in a giant bivalved arthropod from the Cambrian Burgess Shale

The origin of mandibulate arthropods can be traced back to the Cambrian period to several carapace-bearing arthropod groups, but their morphological diversity is still not well characterized. Here, we describe Balhuticaris voltae, a bivalved arthropod from the 506-million-year-old Burgess Shale (Marble Canyon, British Columbia, Canada). This species has an extremely elongated and multisegmented body bearing ca. 110 pairs of homonomous biramous limbs, the highest number among Cambrian arthropods, and, at 245 mm, it represents one of the largest Cambrian arthropods known. Its unusual carapace resembles an arch; it covers only the frontalmost section of the body but extends ventrally beyond the legs. Balhuticaris had a complex sensory system and was probably an active swimmer thanks to its powerful paddle-shaped exopods and a long and flexible body. Balhuticaris increases the ecological and functional diversity of bivalved arthropods and suggests that cases of gigantism occurred in more arthropod groups than previously recognized.