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Abstract A crucial step in the evolution of Euarthropoda (chelicerates, myriapods, pancrustaceans) was the transition between fossil groups that possessed frontal appendages innervated by the first segment of the brain (protocerebrum), and living groups with a protocerebral labrum and paired appendages innervated by the second brain segment (deutocerebrum). Appendage homologies between the groups are controversial. Here we describe two specimens of opabiniid-like euarthropods, each bearing an anterior proboscis (a fused protocerebral appendage), from the Middle Ordovician Castle Bank Biota, Wales, UK. Phylogenetic analyses support a paraphyletic grade of stem-group euarthropods with fused protocerebral appendages and a posterior-facing mouth, as in the iconic Cambrian panarthropodOpabinia. These results suggest that the labrum may have reduced from an already-fused proboscis, rather than a pair of arthropodized appendages. If some shared features between the Castle Bank specimens and radiodonts are considered convergent rather than homologous, phylogenetic analyses retrieve them as opabiniids, substantially extending the geographic and temporal range of Opabiniidae. 

Once considered ‘weird wonders’ of the Cambrian, the emblematic Burgess Shale animals Anomalocaris and Opabinia are now recognized as lower stem-group euarthropods and have provided crucial data for constraining the polarity of key morphological characters in the group. Anomalocaris and its relatives (radiodonts) had worldwide distribution and survived until at least the Devonian. However, despite intense study, Opabinia remains the only formally described opabiniid to date. Here we reinterpret a fossil from the Wheeler Formation of Utah as a new opabiniid, Utaurora comosa nov. gen. et sp. By visualizing the sample of phylogenetic topologies in treespace, our results fortify support for the position of U. comosa beyond the nodal support traditionally applied. Our phylogenetic evidence expands opabiniids to multiple Cambrian stages. Our results underscore the power of treespace visualization for resolving imperfectly preserved fossils and expanding the known diversity and spatio-temporal ranges within the euarthropod lower stem group.