A large new leanchoiliid from the Burgess Shale and the influence of inapplicable states on stem arthropod phylogeny

Characterized by atypical frontalmost appendages, leanchoiliids are early arthropods whose phylogenetic placement has been much debated. Morphological interpretations have differed, some of which concern critical characters such as the number of eyes and head appendages, but methodological approaches also have diverged. Here, we describe a new leanchoiliid, Yawunik kootenayi gen. et sp. nov., based on 42 specimens from the newly discovered Marble Canyon locality of the Burgess Shale (Kootenay National Park, British Columbia; middle Cambrian). This new morphotype demonstrates the presence of a four-segmented head in leanchoiliids, along with two small antero-median eyes in addition to lateral eyes. Yawunik is characterized by a 12-segmented trunk and a carinate, lanceolate telson adorned with minute spines. The ‘great appendages’ of the animal bear teeth on their two distal rami, which would have enhanced their ability to grasp prey. Attitudes of specimens, resulting from burial at multi...