Euthycarcinoid arthropods (Cambrian–Triassic) were likely the first animals to transition from oceanic to freshwater and emergent environments. Although their basic bauplan is well known, they have a poor fossil record because their non-sclerotized exoskeleton was rarely preserved. Euthycarcinoids’ unusual morphology (varying numbers of body segments, seemingly dichotomous possession of either mandibles or a labrum, specialized or generalized limbs, and possession by some euthycarcinoid species of sternal pores—structures possibly analogous to coxal vesicles in myriapods) contribute to uncertainty regarding their relationship to other arthropod groups; while their poor fossil record masks the evolutionary transitions within and between the separate realms they inhabited (marine, freshwater, emergent). A new euthycarcinoid from a Permian polar proglacial lake is described herein that is morphologically unlike all other euthycarcinoids, and interpreted as being well adapted for a nekton-benthic lifestyle. Antarcticarcinus pagoda n. gen. n. sp. possesses a pair of large wing-like processes that project laterally from the preabdominal dorsal exoskeleton. A trace fossil from the overlying Mackellar Formation, cf. Orbiculichnus, which was previously interpreted as having been produced by insects taking off or landing on wet sediments, is reinterpreted herein as being produced by A. pagoda n. gen. n. sp. due to the high degree of morphological similarity between traces and body fossils. This occurrence indicates that euthycarcinoids were able to adapt to life in temperate freshwater environments, while possible subaerial adaptations hint at an ability to breathe air. Indeed, if euthycarcinoids could breathe air, Cambrian terrestrial forays and rapid transition (by the Ordovician) into freshwater environments might be explained.

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