Three-Dimensionally Preserved Appendages in an Early Cambrian Stem-Group Pancrustacean

Pancrustaceans boast impressive diversity, abundance, and ecological impact in the biosphere throughout the Phanerozoic [1]. Molecular clock estimates suggest an early Cambrian divergence for pancrustaceans [2, 3]. Despite the wealth of Palaeozoic exceptional fossiliferous deposits [4-7], the early evolution of Pancrustacea remains elusive given the difficulty of recognizing synapomorphies between Cambrian forms and extant representatives. Although early studies suggested crustacean affinities for Cambrian bivalved euarthropods [8-11], this view has fallen out of favor by recent reappraisals of their morphology [12-16]. The best evidence for total-group pancrustaceans comes from Cambrian microfossils preserved as three-dimensional phosphatic replicates in Orsten-type assemblages [4, 17-19] or as "small carbonaceous fossils" (SCFs) [20, 21]. Although these taphonomic windows capture minute morphology enabling detailed comparisons with extant representatives, these microfossils are limited to larval stages (Orsten) or recalcitrant fragmentary remains (SCFs) restricting their phylogenetic precision [5, 12, 19, 20, 22, 23]. We employed X-ray computed tomography [24] to reveal the three-dimensionally appendage morphology of the Chengjiang bivalved euarthropod Ercaicunia multinodosa [25] from the early Cambrian of China. E. multinodosa possesses characters uniquely shared with extant crustaceans, including differentiated tritocerebral antennae and epipodite-bearing biramous trunk appendages. Similarities between E. multinodosa with clypecaridids [9], waptiids [16] and hymenocarines [11, 14] suggest that these euarthropods may also possess similarly differentiated appendages, but these details are obstructed by the limits of preservation of compacted macrofossils. E. multinodosa illuminates the early evolution of pancrustacean appendage differentiation and represents the oldest unequivocal crown-group mandibulate known from complete macrofossils [22].