''Australopithecus garhi'' is a species of
australopithecine 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, Artifact (archaeology), artifacts, and bones of large mammals with cut marks from butchery with tools by earl ...
in the
Afar Region
The Afar Region (; ; ), formerly known as Region 2, is a Regions of Ethiopia, regional state in northeastern Ethiopia and the homeland of the Afar people. Its capital is the planned city of Semera, which lies on the paved Awash, Ethiopia, Awash� ...
of Ethiopia 2.6–2.5 million years ago (mya) during the
Early Pleistocene
The Early Pleistocene is an unofficial epoch (geology), sub-epoch in the international geologic timescale in chronostratigraphy, representing the earliest division of the Pleistocene Epoch within the ongoing Quaternary Period. It is currently esti ...
. 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 a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'' 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 is a positional relationship of the mandible or maxilla to the skeletal base where either of the jaws protrudes beyond a predetermined imaginary line in the coronal plane of the skull.
In the case of ''mandibular'' prognathism (nev ...
); relatively large
molars
The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
and
premolar
The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s; adaptations for both walking on two legs (
bipedalism
Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
) 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 (scansorial), but others are exclusively arboreal. The hab ...
ity); and it is possible that, though unclear if, males were larger than females (exhibited
sexual dimorphism
Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
). 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 industry which was previously considered to have been invented by the later ''
H. habilis'', though this may have instead been produced by contemporary ''Homo''.
Anatomy
Like other australopithecines, ''A. garhi'' had a brain volume of about , a
sagittal crest
A sagittal crest is a ridge of bone running lengthwise along the midline of the top of the skull (at the sagittal suture) of many mammalian and reptilian skulls, among others. The presence of this ridge of bone indicates that there are excepti ...
running along the midline of the skull, and a
prognathic jaw (the jaw jutted out). Relatively, the postcanine teeth, the
molars
The molars or molar teeth are large, flat tooth, teeth at the back of the mouth. They are more developed in mammal, mammals. They are used primarily to comminution, grind food during mastication, chewing. The name ''molar'' derives from Latin, '' ...
and
premolar
The premolars, also called premolar Tooth (human), teeth, or bicuspids, are transitional teeth located between the Canine tooth, canine and Molar (tooth), molar teeth. In humans, there are two premolars per dental terminology#Quadrant, quadrant in ...
s, are massive (
post-canine megadontia), similar to or greater than those of other australopithecines and of the large-toothed ''
Paranthropus robustus
''Paranthropus robustus'' is a species of robust australopithecine from the Early and possibly Middle Pleistocene of the Cradle of Humankind, South Africa, about 2.27 to 0.87 (or, more conservatively, 2 to 1) million years ago. It has been ide ...
''.
Like the earlier ''A. afarensis'' from the same region, ''A. garhi'' had a humanlike humerus to femur ratio, and an apelike brachial index (lower to upper arm ratio) as well as curved
phalanges
The phalanges (: phalanx ) are digit (anatomy), digital bones in the hands and foot, feet of most vertebrates. In primates, the Thumb, thumbs and Hallux, big toes have two phalanges while the other Digit (anatomy), digits have three phalanges. ...
of the hand.
This is generally interpreted as adaptations for both walking on two legs (habitual
bipedalism
Bipedalism is a form of terrestrial locomotion where an animal moves by means of its two rear (or lower) Limb (anatomy), limbs or legs. An animal or machine that usually moves in a bipedal manner is known as a biped , meaning 'two feet' (from ...
) as well as for grasping while climbing in trees (
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 (scansorial), but others are exclusively arboreal. The hab ...
ity).
The BOU-VP-35/1 humerus specimen is notably larger than the humerus of the BOU-VP-12/1 specimen, which could potentially indicate size-specific
sexual dimorphism
Sexual dimorphism is the condition where sexes of the same species exhibit different Morphology (biology), morphological characteristics, including characteristics not directly involved in reproduction. The condition occurs in most dioecy, di ...
with males larger than females to a similar degree to what is postulated in ''A. afarensis'', but it is unclear if this does not represent normal size variation of the same sex as this is based on only two specimens. Nonetheless, on the basis of size, BOU-VP-12/130 is considered male and BOU-VP-17/1 female. Contemporary hominins from Kenya are about the same size as ''A. garhi''.
BOU-VP-17/1 may have been about tall.
[
''Australopithecus'' are thought to have had fast, apelike growth rates, lacking an extended childhood typical of modern humans. However, the legs of ''A. garhi'' are elongated, unlike those of other ''Australopithecus'', and, in humans, elongated limbs develop during the delayed adolescent growth spurt. This could mean that ''A. garhi'', compared to other ''Australopithecus'', either had a slower overall growth rate, or a more rapid leg growth rate.]
Taxonomy
The Ethiopian ''Australopithecus garhi'' was first described in 1999 by palaeoanthropologists Berhane Asfaw, Tim D. White, Owen Lovejoy, Bruce Latimer, Scott Simpson, and Gen Suwa 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, Artifact (archaeology), artifacts, and bones of large mammals with cut marks from butchery with tools by earl ...
in Middle Awash, Afar Region
The Afar Region (; ; ), formerly known as Region 2, is a Regions of Ethiopia, regional state in northeastern Ethiopia and the homeland of the Afar people. Its capital is the planned city of Semera, which lies on the paved Awash, Ethiopia, Awash� ...
, 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 known as a cranial suture, form the sides and roof of the neurocranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four bord ...
(GAM-VP-1/2), left jawbone (GAM-VP-1/1), and left humerus
The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
(MAT-VP-1/1)—which are unassignable to a specific genus
Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
. The first identifiable ''Australopithecus'' fossils–an adult ulna
The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
(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 (; : femurs or femora ), or thigh bone is the only long bone, bone in the thigh — the region of the lower limb between the hip and the knee. In many quadrupeds, four-legged animals the femur is the upper bone of the hindleg.
The Femo ...
, right humerus
The humerus (; : humeri) 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 (bone), radius and ulna, and consists of three sections. The humeral upper extrem ...
, radius
In classical geometry, a radius (: radii or radiuses) of a circle or sphere is any of the line segments from its Centre (geometry), center to its perimeter, and in more modern usage, it is also their length. The radius of a regular polygon is th ...
, and ulna
The ulna or ulnar bone (: ulnae or ulnas) is a long bone in the forearm stretching from the elbow to the wrist. It is on the same side of the forearm as the little finger, running parallel to the Radius (bone), radius, the forearm's other long ...
, and a partial fibula, foot, and jawbone. The holotype specimen
A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was Species description, formally described. It is either the single such physical example (or illus ...
, 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 Paleoanthropology, paleoanthropologist. An authority on pre-''Homo sapiens'' hominids, he particularly focuses his attention on the East African Rift and Middle Awash valleys. ...
. 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 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 Potassium-argon dating, potassiumargon (K/Ar) dating in accuracy. The older method required splitting samples into two for separate potassium and argon measur ...
. 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 (hominins) form a taxonomic tribe of the subfamily Homininae (hominines). They comprise two extant genera: ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos), and in standard usage exclude the genus '' Gorilla'' ( gorillas) ...
s from this timespan being identified from South Africa ('' A. africanus'') and Lake Turkana, Kenya (''Paranthropus aethiopicus
''Paranthropus aethiopicus'' is an Extinction, 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'' ...
''). Likewise, the classification of australopithecines and pre-''Homo erectus
''Homo erectus'' ( ) is an extinction, extinct species of Homo, archaic human from the Pleistocene, spanning nearly 2 million years. It is the first human species to evolve a humanlike body plan and human gait, gait, to early expansions of h ...
'' hominins has been the subject of much debate. The original describers considered ''A. garhi'' to be a descendant of the earlier '' A. afarensis'' which inhabited the same region, based mainly on dental similarities. Though they assigned the species to ''Australopithecus
''Australopithecus'' (, ; or (, ) is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera ''Homo'' (which includes modern humans), ''Paranthropus'', and ''Kenyanthropus'' evolved from some ''Aus ...
'', 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
Afar is an Afroasiatic language belonging to the Cushitic branch, primarily spoken by the Afar people, native to parts of Djibouti, Eritrea and Ethiopia. It is an official language in Ethiopia; and a national language in Djibouti and Eritre ...
. In 1999, American palaeoanthropologists David Strait and Frederick E. Grine concluded that ''A. garhi'' was instead an offshoot of the human line instead of an ancestor because ''A. garhi'' and ''Homo'' share no synapomorphies (traits unique to only them). In 2015, ''Homo'' was recorded from 2.8 mya, much earlier than ''A. garhi''.
Palaeoecology
The large teeth of ''Australopithecus'' species have historically been interpreted as having been adaptations for a diet of hard foods, but the durable teeth may instead have only served an important function during leaner times for harder fallback foods. That is, dental anatomy may not accurately portray normal ''Australopithecus'' diet, rather abnormal diet during times of famine.
Though it was not found with any tools, mammalian bones associated with the ''A. garhi'' remains exhibit cut and percussion marks made from stone tools: the left mandible
In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla).
The jawbone i ...
of an alcelaphine bovid with three successive, unambiguous cut marks presumably made while removing the tongue; a bovid tibia
The tibia (; : tibiae or tibias), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two Leg bones, bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outsi ...
with cut marks, chop marks, and impact scars from a hammerstone, possibly inflicted to harvest the bone marrow
Bone marrow is a semi-solid biological tissue, tissue found within the Spongy bone, spongy (also known as cancellous) portions of bones. In birds and mammals, bone marrow is the primary site of new blood cell production (or haematopoiesis). It i ...
; and a '' Hipparion'' (a horse) femur with cut marks consistent with dismemberment and filleting. As to why stone tools were not present, because the Hatayae locality was likely a featureless, grassy lake margin with so few raw materials for making stone tools, it is possible these hominins were creating and carrying tools some ways with them to butchering sites, intending to use them many times before discarding. It was previously believed that only ''Homo
''Homo'' () is a genus of great ape (family Hominidae) that emerged from the genus ''Australopithecus'' and encompasses only a single extant species, ''Homo sapiens'' (modern humans), along with a number of extinct species (collectively called ...
'' could manufacture tools;[ but it is also possible that the butcherers were not manufacturing tools and simply used naturally sharp rocks.]
At the nearby Gona site, where there is an abundance of raw materials, several Oldowan tools (an industry previously believed to have been invented by '' H. habilis'') were recovered from 1992 to 1994. The tools date to around 2.6–2.5 mya, the oldest evidence of manufacturing at the time, and since ''A. garhi'' was the only species identified in the vicinity at the time, this species was the best candidate for authorship. However, in 2015, the earliest remains of ''Homo'', LD 350-1, were discovered in Ledi-Geraru, also in the Afar Region, dating to 2.8–2.75 mya.[ More stone tools were found in 2019 dating to about 2.6 mya in Ledi-Geraru, predating the Gona artifacts, and these may be attributed to ''Homo''; the invention of sharp-edged Oldowan tools could actually be due to specific adaptations characteristic of ''Homo''.] Nonetheless, other australopithecines have been associated with stone tool manufacturing, such as the 2010 discovery of cut marks dating to 3.4 mya attributed to ''A. afarensis'', and the 2015 discovery of the Lomekwi
Lomekwi is an archaeological site located on the west bank of Turkana Lake in Kenya. It is an important milestone in the history of human archaeology. An archaeological team from Stony Brook University in the United States discovered traces o ...
culture from Lake Turkana dating to 3.3 mya possibly attributed to '' Kenyanthropus''.
See also
References
External links
Reconstruction
by Viktor Deak
*
*
Human Timeline (Interactive)
– Smithsonian, National Museum of Natural History
The National Museum of Natural History (NMNH) is a natural history museum administered by the Smithsonian Institution, located on the National Mall in Washington, D.C., United States. It has free admission and is open 364 days a year. With 4.4 ...
(August 2016).
*
{{Taxonbar, from=Q310527
Australopithecus
Pliocene primates
Mammals described in 1997
Fossil taxa described in 1997
Prehistoric Ethiopia
Pliocene mammals of Africa