Waptia fieldensis Walcott, a mandibulate arthropod from the middle Cambrian Burgess Shale

Waptia fieldensis Walcott, 1912 is one of the iconic animals from the middle Cambrian Burgess Shale biota that had lacked a formal description since its discovery at the beginning of the twentieth century. This study, based on over 1800 specimens, finds that W. fieldensis shares general characteristics with pancrustaceans, as previous authors had suggested based mostly on its overall aspect. The cephalothorax is covered by a flexible, bivalved carapace and houses a pair of long multisegmented antennules, palp-bearing mandibles, maxillules, and four pairs of appendages with five-segmented endopods-the anterior three pairs with long and robust enditic basipods, the fourth pair with proximal annulations and lamellae. The post-cephalothorax has six pairs of lamellate and fully annulated appendages which appear to be extensively modified basipods rather than exopods. The front part of the body bears a pair of stalked eyes with the first ommatidia preserved in a Burgess Shale arthropod, and a median 'labral' complex flanked by lobate projections with possible affinities to hemi-ellipsoid bodies. Waptia confirms the mandibulate affinity of hymenocarines, retrieved here as part of an expanded Pancrustacea, thereby providing a novel perspective on the evolutionary history of this hyperdiverse group. We construe that Waptia was an active swimming predator of soft prey items, using its anterior appendages for food capture and manipulation, and also potentially for clinging to epibenthic substrates.