The rostral portion of the skull of a new genus and species of rodent, Armintomys tullbergi, from the earliest middle Eocene of the Wind River Basin (Wyoming) provides the geologically oldest known record of the hystricomorphous zygomasseteric structure. Armintomys also preserves the oldest known occurrence of incisor enamel that is transitional from pauciserial to uniserial. Other dental characters include: anteriorly grooved incisor, small premolars, and relatively primitive sciuravidlike molars. Analysis of this unique combination of characters implies that Armintomys is the oldest known myomorph rodent and the only known representative of a new family. Armintomyidae, which is referred, with question, to the myomorph superfamily Dipodoidea. Armintomys is more primitive, especially in premolar retention and structure, than the Bridgerian zapodid Elymys from Nevada, but adds to evidence from the latter for an early origin and radiation of dipodoid rodents.