The early Pliocene African hominoid Ardipithecus ramidus was diagnosed as a having a unique phylogenetic relationship with the Australopithecus + Homo clade based on nonhoning canine teeth, a foreshortened cranial base, and postcranial characters related to facultative bipedality. However, pedal and pelvic traits indicating substantial arboreality have raised arguments that this taxon may instead be an example of parallel evolution of human-like traits among apes around the time of the chimpanzee-human split. Here we investigated the basicranial morphology of Ar. ramidus for additional clues to its phylogenetic position with reference to African apes, humans, and Australopithecus. Besides a relatively anterior foramen magnum, humans differ from apes in the lateral shift of the carotid foramina, mediolateral abbreviation of the lateral tympanic, and a shortened, trapezoidal basioccipital element. These traits reflect a relative broadening of the central basicranium, a derived condition associated with changes in tympanic shape and the extent of its contact with the petrous. Ar. ramidus shares with Australopithecus each of these human-like modifications. We used the preserved morphology of ARA-VP 1/500 to estimate the missing basicranial length, drawing on consistent proportional relationships in apes and humans. Ar. ramidus is confirmed to have a relatively short basicranium, as in Australopithecus and Homo. Reorganization of the central cranial base is among the earliest morphological markers of the Ardipithecus + Australopithecus + Homo clade.A s the confluence of the neural, locomotor, and masticatory systems, the cranial base has been the site of profound structural change in human evolution. The modern human basicranium differs from that of our closest living relatives, the great apes, in numerous aspects of shape and morphological detail (1-4). In humans, the foramen magnum and occipital condyles are more anteriorly located, the midline basicranial axis is relatively short anteroposteriorly and strongly "flexed" internally, and the bilateral structures marking vascular and neural pathways through the central part of the base are more widely separated. This organization alters the relationships between the petrous and tympanic parts of the temporal bone. These phylogenetically derived features are already seen in the earliest known skulls of Australopithecus, ca. 3.0-3.4 Ma (5, 6).The cranium of Ardipithecus ramidus, an early Pliocene (4.4 Ma) hominoid from Ethiopia, was shown to have a relatively anterior foramen magnum on a short basicranium, corroborating evidence of nonhoning canine teeth and terrestrial bipedality for phylogenetic attribution of this taxon. These sets of derived characters are shared uniquely with the Australopithecus + Homo clade (7-10). At the same time, pelvic and pedal characters indicate that Ar. ramidus also retained considerable arboreal capabilities (11)(12)(13)(14). Despite the evidence for a unique phylogenetic relationship with the Australopithecus + Homo clade, it has been a...