Appendicular anatomy of the artiopod Emeraldella brutoni from the middle Cambrian (Drumian) of western Utah

The non-biomineralized artiopod Emeraldella brutoni Stein, Church & Robison, from the middle Cambrian (Drumian) Wheeler Formation in Utah represents the only confirmed occurrence of the genus Emeraldella outside of the stratigraphically older (Wuliuan) Burgess Shale Konservat-Lagerstätte in British Columbia. The hitherto known sole specimen of this species is preserved in dorsal view and lacks critical information on the ventral appendages. Here, we redescribe E. brutoni based on a new completely articulated specimen that illustrates the appendage organization in exceptional detail. The main body consists of a cephalic region covered by a semicircular head shield, a trunk including 10 tergites with expanded pleurae plus a cylindrical terminal segment, and a long articulated tailspine. The head carries a pair of elongate and flexible antennae, a pair of lateral eyes, and three pairs of post-antennal appendages. We report the presence of eyes in Emeraldella for the first time. The first post-antennal limb solely consists of an endopod with well-developed paired spiniform endites. The remaining cephalic appendages and those associated with all but the last trunk segments possess exopods terminating in paddle-shaped, distal lobes fringed with robust setae. The cylindrical terminal segment bears a pair of posteriorly oriented caudal flaps reminiscent of trunk exopods, and a styliform, possibly uniarticulate tailspine longer than the main body. The new data on E. brutoni suggests an exopodal origin for the paired caudal structures in Vicissicaudata, and improve our understanding of the fundamental organization of this major clade within Artiopoda.