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''Australopithecus garhi'' is a species of
australopithecine Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally ''Australopithecus'' ( cladistically including the genera ''Homo'', ''Paranthropus'', and '' Kenyanthropus''), and it typically include ...
from the
Bouri Formation The Bouri Formation is a sequence of sedimentary deposits that is the source of australopithecine and Homo (that is, hominin) fossils, artifacts, and bones of large mammals with cut marks from butchery with tools by early hominins. It is locate ...
in the
Afar Region The Afar Region (; aa, Qafar Rakaakayak; am, አፋር ክልል), formerly known as Region 2, is a regional state in northeastern Ethiopia and the homeland of the Afar people. Its capital is the planned city of Semera, which lies on the pav ...
of Ethiopia 2.6–2.5 million years ago (mya) during the Early Pleistocene. The first remains were described in 1999 based on several skeletal elements uncovered in the three years preceding. ''A. garhi'' was originally considered to have been a direct ancestor to ''
Homo ''Homo'' () is the genus that emerged in the (otherwise extinct) genus '' Australopithecus'' that encompasses the extant species ''Homo sapiens'' (modern humans), plus several extinct species classified as either ancestral to or closely relat ...
'' and the human line, but is now thought to have been an offshoot. Like other australopithecines, ''A. garhi'' had a brain volume of ; a jaw which jutted out (
prognathism Prognathism, also called Habsburg jaw or Habsburgs' jaw primarily in the context of its prevalence amongst members of the House of Habsburg, is a positional relationship of the mandible or maxilla to the skeletal base where either of the jaws pr ...
); relatively large
molars The molars or molar teeth are large, flat teeth at the back of the mouth. They are more developed in mammals. They are used primarily to grind food during chewing. The name ''molar'' derives from Latin, ''molaris dens'', meaning "millstone to ...
and
premolar The premolars, also called premolar teeth, or bicuspids, are transitional teeth located between the canine and molar teeth. In humans, there are two premolars per quadrant in the permanent set of teeth, making eight premolars total in the mouth ...
s; adaptations for both walking on two legs (
bipedalism Bipedalism is a form of terrestrial locomotion where an organism moves by means of its two rear limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from Latin ''bis'' 'double' ...
) and grasping while climbing (
arboreal Arboreal locomotion is the locomotion of animals in trees. In habitats in which trees are present, animals have evolved to move in them. Some animals may scale trees only occasionally, but others are exclusively arboreal. The habitats pose num ...
ity); and it is possible that, though unclear if, males were larger than females (exhibited
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most an ...
). One individual, presumed female based on size, may have been tall. ''A. garhi'' is the first pre-''Homo'' hominin postulated to have manufactured tools—using them in butchering—and may be counted among a growing body of evidence for pre-''Homo'' stone tool industries (the ability to manufacture tools was previously believed to have separated ''Homo'' from predecessors.) ''A. garhi'' possibly produced the
Oldowan The Oldowan (or Mode I) was a widespread stone tool archaeological industry (style) in prehistory. These early tools were simple, usually made with one or a few flakes chipped off with another stone. Oldowan tools were used during the Lower ...
industry which was previously considered to have been invented by the later '' H. habilis'', though this may have instead been produced by contemporary ''Homo''.


Taxonomy

The Ethiopian ''Australopithecus garhi'' was first described in 1999 by palaeoanthropologists
Berhane Asfaw Berhane Asfaw ( Amharic: በርሃነ አስፋው) (born August 22, 1954 in Gondar, Ethiopia) is an Ethiopian paleontologist of Rift Valley Research Service, who co-discovered human skeletal remains at Herto Bouri, Ethiopia later classified as ...
,
Tim D. White Tim D. White (born August 24, 1950) is an American paleoanthropologist and Professor of Integrative Biology at the University of California, Berkeley. He is best known for leading the team which discovered Ardi, the type specimen of ''Ardipithecu ...
,
Owen Lovejoy Owen Lovejoy (January 6, 1811 – March 25, 1864) was an American lawyer, Congregational minister, abolitionist, and Republican congressman from Illinois. He was also a "conductor" on the Underground Railroad. After his brother Elijah Lov ...
, Bruce Latimer, Scott Simpson, and
Gen Suwa Gen Suwa (born 1954) is a Japanese paleoanthropologist. He is known for his contributions to the understanding of the evolution of early hominids, including the discovery of a tooth from a hominid that was more than one million years older than the ...
based on fossils discovered in the Hatayae Beds of the
Bouri Formation The Bouri Formation is a sequence of sedimentary deposits that is the source of australopithecine and Homo (that is, hominin) fossils, artifacts, and bones of large mammals with cut marks from butchery with tools by early hominins. It is locate ...
in
Middle Awash The Middle Awash is a paleoanthropological research area in the Afar Region along the Awash River in Ethiopia's Afar Depression. It is a unique natural laboratory for the study of human origins and evolution and a number of fossils of the earlie ...
,
Afar Region The Afar Region (; aa, Qafar Rakaakayak; am, አፋር ክልል), formerly known as Region 2, is a regional state in northeastern Ethiopia and the homeland of the Afar people. Its capital is the planned city of Semera, which lies on the pav ...
, Ethiopia. The first hominin remains were discovered here in 1990—a partial
parietal bone The parietal bones () are two bones in the skull which, when joined at a fibrous joint, form the sides and roof of the cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, and four angles. It is named ...
(GAM-VP-1/2), left jawbone (GAM-VP-1/1), and left
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
(MAT-VP-1/1)—which are unassignable to a specific
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial n ...
. The first identifiable ''Australopithecus'' fossils–an adult
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
(BOU-VP-11/1)–were found on 17 November 1996 by T. Assebework. A partial skeleton (BOU-VP-12/1) was discovered 13 days later by White, comprising a mostly complete left
femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates wit ...
, right
humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...
,
radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...
, and
ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...
, and a partial
fibula The fibula or calf bone is a human leg, leg bone on the Lateral (anatomy), lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long ...
, foot, and jawbone. The
holotype specimen A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
, a partial skull (BOU-VP-12/130), was discovered on 20 November 1997 by Ethiopian palaeoanthropologist
Yohannes Haile-Selassie Yohannes Haile-Selassie Ambaye (born 23 February 1961) is an Ethiopian paleoanthropologist. An authority on pre-''Homo sapiens'' hominids, he particularly focuses his attention on the East African Rift and Middle Awash valleys.Mangels, John ( ...
. More skull fragments (BOU-VP-12/87) were recovered south of BOU-VP-12/1. On 17 November 1997, French palaeoanthropologist Alban Defleur discovered a complete mandible (BOU-VP-17/1) about north in the Esa Dibo
locality Locality may refer to: * Locality (association), an association of community regeneration organizations in England * Locality (linguistics) * Locality (settlement) * Suburbs and localities (Australia), in which a locality is a geographic subdivi ...
of the formation, and American palaeoanthropologist David DeGusta discovered a humerus (BOU-VP-35/1) north of BOU-VP-17/1. However, BOU-VP-11, -12, and -35 cannot conclusively be attributed to ''A. garhi''. The remains are dated to about 2.5 million years ago (mya) based on
argon–argon dating Argon–argon (or 40Ar/39Ar) dating is a radiometric dating method invented to supersede potassiumargon (K/Ar) dating in accuracy. The older method required splitting samples into two for separate potassium and argon measurements, while the newer ...
. When they were discovered, human evolution was obscured due to a paucity of remains from 3 to 2 mya, with the only
hominin The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
s from this timespan being identified from South Africa ('' A. africanus'') and
Lake Turkana Lake Turkana (), formerly known as Lake Rudolf, is a lake in the Kenyan Rift Valley, in northern Kenya, with its far northern end crossing into Ethiopia. It is the world's largest permanent desert lake and the world's largest alkaline lake ...
, Kenya (''
Paranthropus aethiopicus ''Paranthropus aethiopicus'' is an extinct species of robust australopithecine from the Late Pliocene to Early Pleistocene of East Africa about 2.7–2.3 million years ago. However, it is much debated whether or not ''Paranthropus'' is an invali ...
''). Likewise, the classification of
australopithecine Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally ''Australopithecus'' ( cladistically including the genera ''Homo'', ''Paranthropus'', and '' Kenyanthropus''), and it typically include ...
s and pre-''
Homo erectus ''Homo erectus'' (; meaning " upright man") is an extinct species of archaic human from the Pleistocene, with its earliest occurrence about 2 million years ago. Several human species, such as ''H. heidelbergensis'' and ''H. antecessor' ...
'' hominins has been the subject of much debate. The original describers considered ''A. garhi'' to be a descendant of the earlier ''
A. afarensis ''Australopithecus afarensis'' is an extinct species of australopithecine which lived from about 3.9–2.9 million years ago (mya) in the Pliocene of East Africa. The first fossils were discovered in the 1930s, but major fossil finds would not ...
'' which inhabited the same region, based mainly on dental similarities. Though they assigned the species to ''
Australopithecus ''Australopithecus'' (, ; ) is a genus of early hominins that existed in Africa during the Late Pliocene and Early Pleistocene. The genus ''Homo'' (which includes modern humans) emerged within ''Australopithecus'', as sister to e.g. ''Austral ...
'', the original describers believed it could represent an ancestor to ''Homo'', which, if the case, would possibly lead to reclassification as ''H. garhi''. Because the characteristics of ''A. garhi'' are unexpected for a human ancestor at this stage, the specific name, ''garhi'', means "surprise" in the local
Afar language The Afar language ( aa, Qafaraf, links=no; also known as ’Afar Af, Afaraf, Qafar af) is an Afroasiatic lang