Euthycarcinoids record the transition of stem group myriapods from marine (Apankura) to freshwater environments, and their trace fossil record attests to amphibious nearshore (intertidal and coastal dune) habits by the mid-Cambrian (34–36). Our findings from H. rhyniensis reveal that some characters broadly associated with terrestrialization in myriapods, such as the mouthparts being encapsulated in a preoral chamber, are not innovations of the terrestrial crown group but rather evolved in aquatic environments.