Tentacular nature of the 'column' of the Cambrian diploblastic Xianguangia sinica

Unveiling the body architectures of Cambrian problematic fossils would provide novel insights into the radiation of metazoan body plans during the 'Cambrian Explosion' and the ancestral traits of major living animal clades. Xianguangia sinica, from the celebrated Chengjiang biota (similar to 518 Ma), is a typical Cambrian problematicum with disputable body architecture, particularly about its 'column' part. The contradictory interpretations of the 'column' of X. sinica have led to at least three hypotheses regarding its affinity in the diploblastic clade. Here we depict the detailed anatomy of the 'column' based on new, exquisitely preserved material. The 'column' of X. sinica is formed by 18 longsword-shaped tentacle-sheath complexes that can either close or be in a flowering state. There is no partitioned cavity internally when the 'column' is closed, invalidating the homology with the true column of living sea anemones. Each tentacle tapers distally and includes a distal flexible portion at about one-fourth the length of the tentacle. The proximal portion is stiff, bearing a set of paired dark stains, and is enveloped by a single outer sheath. Pinnules carrying a row of large cilia are fringed on both sides along the whole length of the tentacles. The body plan of X. sinica is accordingly revised as consisting of a calyx and 18 unique tentacle-sheath complexes that radially surround the mouth. Our result corroborates previous observations that suggest a close relationship between Xianguangia, Daihua and Dinomischus, all of which are here formally assigned to the family Dinomischidae, a monophyletic clade recovered in our phylogenetic analyses. Xianguangia sinica likely employs cilia-bearing pinnate tentacles for sieving particle matter down to 21 mu m, indicating that its nutrition source is suspended micro-planktonic organisms or other organic matter in the bottom water.