The cranial morphology of the earliest known hominins in the genus Australopithecus remains unclear. The oldest species in this genus (Australopithecus anamensis, specimens of which have been dated to 4.2-3.9 million years ago) is known primarily from jaws and teeth, whereas younger species (dated to 3.5-2.0 million years ago) are typically represented by multiple skulls. Here we describe a nearly complete hominin cranium from Woranso-Mille (Ethiopia) that we date to 3.8 million years ago. We assign this cranium to A. anamensis on the basis of the taxonomically and phylogenetically informative morphology of the canine, maxilla and temporal bone. This specimen thus provides the first glimpse of the entire craniofacial morphology of the earliest known members of the genus Australopithecus. We further demonstrate that A. anamensis and Australopithecus afarensis differ more than previously recognized and that these two species overlapped for at least 100,000 years—contradicting the widely accepted hypothesis of anagenesis.