instead, Holochilus was considered to be a sigmodont, related to Sigmodon and Reithrodon, and Pseudoryzomys and Zygodontomys were considered to be members of Phyllotini, another large South American tribe. Although lacking mesoloph(id)s, these genera share other characters with oryzomyines, and a series of papers by Robert Voss and coworkers in the early 1990s established their membership in Oryzomyini.

In a 1993 paper, Voss and Carleton proposed the first cladistic