Nectocaridid ecology, diversity, and affinity: early origin of a cephalopod-like body plan

Nectocaridids are soft-bodied early to middle Cambrian organisms known from Burgess Shale-type deposits in Canada, China, and Australia. Originally described as unrelated species, they have recently been interpreted as a clade; their flexible tentacles, camera-type eyes, lateral fins, internal gills, axial cavity, and funnel point to a relationship with the cephalopods. However, aspects of this reinterpretation, including the relevance of the group to cephalopod evolution, have been called into question. Here, I examine new and existing nectocaridid material, including a large new form that may represent a sexual dimorph of Nectocaris pteryx. Differences between existing taxa largely represent taphonomic variation between sites and specimens-which provides further constraint on the organisms' anatomy. I revise the morphology of the tentacles and fins, and describe mouthparts and phosphatized gills for the first time. A mathematical analysis supports the presence of the earliest known camera-type eyes, and fluid mechanical considerations suggest that the funnel is optimized for efficient jet propulsion in a low Reynolds number flow regime. Nectocaridids closely resemble coleoid cephalopods, but a position deeper within Cephalopoda raises fewer stratigraphic challenges. Whether its coleoid-like construction reflects common ancestry or profound convergence, the Nectocaris body plan adds substantially to Cambrian disparity, demonstrating the rapid colonization of nektobenthic niches after the Cambrian explosion.