Ardipithecus is a genus of an extinct hominine that lived during the Late Miocene and Early Pliocene epochs in the Afar Depression, Ethiopia. Originally described as one of the earliest ancestors of humans after they diverged from the chimpanzees, the relation of this genus to human ancestors and whether it is a hominin is now a matter of debate.[1] Two fossil species are described in the literature: A. ramidus, which lived about 4.4 million years ago[2] during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago (late Miocene).[3] Initial behavioral analysis indicated that Ardipithecus could be very similar to chimpanzees,[1] however more recent analysis based on canine size and lack of canine sexual dimorphism indicates that Ardipithecus was characterised by reduced aggression,[4] and that they more closely resemble bonobos.[5]
A. ramidus was named in September 1994. The first fossil found was dated to 4.4 million years ago on the basis of its stratigraphic position between two volcanic strata: the basal Gaala Tuff Complex (G.A.T.C.) and the Daam Aatu Basaltic Tuff (D.A.B.T.).[6] The name Ardipithecus ramidus stems mostly from the Afar language, in which Ardi means "ground/floor" and ramid means "root". The pithecus portion of the name is from the Greek word for "ape".[7]
Like most hominids, but unlike all previously recognized hominins, it had a grasping hallux or big toe adapted for locomotion in the trees. It is not confirmed how many other features of its skeleton reflect adaptation to bipedalism on the ground as well. Like later hominins, Ardipithecus had reduced canine teeth and reduced canine sexual dimorphism.[8]
In 1992–1993 a research team headed by Tim White discovered the first A. ramidus fossils—seventeen fragments including skull, mandible, teeth and arm bones—from the Afar Depression in the Middle Awash river valley of Ethiopia. More fragments were recovered in 1994, amounting to 45% of the total skeleton. This fossil was originally described as a species of Australopithecus, but White and his colleagues later published a note in the same journal renaming the fossil under a new genus, Ardipithecus. Between 1999 and 2003, a multidisciplinary team led by Sileshi Semaw discovered bones and teeth of nine A. ramidus individuals at As Duma in the Gona area of Ethiopia's Afar Region.[9] The fossils were dated to between 4.35 and 4.45 million years old.[10]
Ardipithecus ramidus had a small brain, measuring between 300 and 350 cm3. This is slightly smaller than a modern bonobo or female common chimpanzee brain, but much smaller than the brain of australopithecines like Lucy (~400 to 550 cm3) and roughly 20% the size of the modern Homo sapiens brain. Like common chimpanzees, A. ramidus was much more prognathic than modern humans.[11]
The teeth of A. ramidus lacked the specialization of other apes, and suggest that it was a generalized omnivore and frugivore (fruit eater) with a diet that did not depend heavily on foliage, fibrous plant material (roots, tubers, etc.), or hard and or abrasive food. The size of the upper canine tooth in A. ramidus males was not distinctly different from that of females. Their upper canines were less sharp than those of modern common chimpanzees in part because of this decreased upper canine size, as larger upper canines can be honed through wear against teeth in the lower mouth. The features of the upper canine in A. ramidus contrast with the sexual dimorphism observed in common chimpanzees, where males have significantly larger and sharper upper canine teeth than females.[12] Of the living apes, bonobos have the smallest canine sexual dimorphism, although still greater than that displayed by A. ramidus.[13]