Sahelanthropus Tchadensis
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''Sahelanthropus tchadensis'' is an
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
species of the
Homininae Homininae (), also called "African hominids" or "African apes", is a subfamily of Hominidae. It includes two tribes, with their extant as well as extinct species: 1) the tribe Hominini (with the genus ''Homo'' including modern humans and numerou ...
(African apes) dated to about , during the
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
epoch. The species, and its genus ''Sahelanthropus'', was announced in 2002, based mainly on a partial
cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, nicknamed ''Toumaï'', discovered in northern
Chad Chad (; ar, تشاد , ; french: Tchad, ), officially the Republic of Chad, '; ) is a landlocked country at the crossroads of North and Central Africa. It is bordered by Libya to the north, Sudan to the east, the Central African Republic ...
. ''Sahelanthropus tchadensis'' lived close to the time of the chimpanzee–human divergence, possibly related to ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopitheci ...
'', a species of Homininae that lived about one million years later. It may have been ancestral to both humans and chimpanzees (which would place it in the tribe
Hominini 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 ...
), or alternatively an early member of the tribe
Gorillini Gorillini is a taxonomic tribe containing three genera: ''Gorilla'' and the extinct ''Chororapithecus'' and ''Nakalipithecus ''Nakalipithecus nakayamai'' is an extinct species of great ape from Nakali, Kenya, from about 9.9–9.8 million ye ...
. In 2020, the
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 with ...
was analyzed, and it was found that ''Sahelanthropus'' was not
biped 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' a ...
al, casting some doubt on its position as a human ancestor, but this was refuted in 2022.


Taxonomy


Discovery

Funded by the Mission Paléoanthropologique Franco–Tchadienne, Ahounta Djimdoumalbaye, Fanoné Gongdibé, Mahamat Adoum, and Alain Beauvilain identified the first remains in the Toros-Menalla area (TM 266
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 subdivis ...
) in the
Djurab Desert The Djurab Desert (''Djourab'', ) is a desert in northern Chad. Part of the greater Sahara desert, it makes up much of the area of Chad's Borkou region. The Koro Toro settlement and maximum security prison is situated on the eastern boundary of ...
of northern Chad. By the time
Michel Brunet Michel Brunet may refer to: * Michel Brunet (historian) (1917–1985), Canadian historian * Michel Brunet (paleontologist) (born 1940), French paleontologist * Michel Brunet (figure skater) (born 1970), Canadian skater {{hndis, Brunet, Michel ...
and colleagues formally described the remains in 2002, a total of six specimens had been recovered: a nearly complete but heavily deformed skull, a fragment of the midline of the jaw with the
tooth socket Dental alveoli (singular ''alveolus'') are sockets in the jaws in which the roots of teeth are held in the alveolar process with the periodontal ligament. The lay term for dental alveoli is tooth sockets. A joint that connects the roots of the ...
s for an
incisor Incisors (from Latin ''incidere'', "to cut") are the front teeth present in most mammals. They are located in the premaxilla above and on the mandible below. Humans have a total of eight (two on each side, top and bottom). Opossums have 18, whe ...
and canine, a right third molar, a right first incisor, a right jawbone with the last
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 ...
to last molar, and a right canine. With the skull as 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 ...
, they were grouped into a new
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
and
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
as ''Sahelanthropus tchadensis'', the genus name referring to the
Sahel The Sahel (; ar, ساحل ' , "coast, shore") is a region in North Africa. It is defined as the ecoclimatic and biogeographic realm of transition between the Sahara to the north and the Sudanian savanna to the south. Having a hot semi-arid c ...
, and the species name to Chad. These, along with ''
Australopithecus bahrelghazali ''Australopithecus bahrelghazali'' is an extinct species of australopithecine discovered in 1995 at Koro Toro, Bahr el Gazel, Chad, existing around 3.5 million years ago in the Pliocene. It is the first and only australopithecine known from Ce ...
'', were the first discoveries of any fossil African
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the east ...
(outside the genus ''
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 relate ...
'') made beyond eastern and southern Africa. By 2005, a third premolar was recovered from the TM 266 locality, a lower jaw missing the region behind the second molar from the TM 292 locality, and a lower left jaw preserving the sockets for premolars and molars from the TM 247 locality. The skull was nicknamed "Toumaï", a name from the local
Daza language Daza (also known as Dazaga) is a Nilo-Saharan language spoken by the Daza people inhabiting northern Chad. The Daza are also known as the Gouran (Gorane) in Chad. Dazaga is spoken by around 380,000 people, primarily in the Djurab Desert region ...
meaning "hope of life", given to infants born just before the dry season. The nickname was suggested by the then-president of Chad,
Idriss Déby Idriss Déby Itno ' (18 June 1952 – 20 April 2021) was a Chadian politician and military officer who was the president of Chad from 1990 until his death in 2021. Déby was a member of the Bidayat clan of the Zaghawa ethnic group. A high-ranki ...
, in honour of one of his comrades-in-arms who had been killed in the coup d'état against his predecessor,
Hissène Habré Hissène Habré (Arabic: ''Ḥusaīn Ḥabrī'', Chadian Arabic: ; ; 13 August 1942 – 24 August 2021), also spelled Hissen Habré, was a Chadian politician and convicted war criminal who served as the 5th president of Chad from 1982 unt ...
. Toumaï also became a source of national pride, and Brunet announced the discovery before the Ministry of Foreign Affairs and a television audience in the capital of
N'Djamena N'Djamena ( ) is the capital and largest city of Chad. It is also a special statute region, divided into 10 districts or ''arrondissements''. The city serves as the centre of economic activity in Chad. Meat, fish and cotton processing are the ...
, "''l'ancêtre de l'humanité est Tchadien...Le berceau de l'humanité se trouve au Tchad. Toumaï est votre ancêtre''" ("The ancestor of humanity is Chadian...The cradle of humanity is in Chad. Toumaï is your ancestor.") Toumaï had been found with a femur, but this was stored with animal bones and shipped to the
University of Poitiers The University of Poitiers (UP; french: Université de Poitiers) is a public university located in Poitiers, France. It is a member of the Coimbra Group. It is multidisciplinary and contributes to making Poitiers the city with the highest studen ...
in 2003, where it was stumbled upon by graduate student Aude Bergeret the next year. She took the bone to the head of the Department of Geosciences, Roberto Macchiarelli, who considered it to be inconsistent with
biped 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' a ...
alism contra what Brunet ''et al.'' had earlier stated in their description analysing only the distorted skull. This was conspicuous because Brunet and his team had already explicitly stated Toumaï was associated with no limb bones, which could have proven or disproven their conclusions of locomotion. Because Brunet had declined to comment on the subject, Macchiarelli and Bergeret petitioned to present their preliminary findings during an annual conference organised by the Anthropological Society of Paris, which would be held at Poitiers that year. This was rejected as they had not formally published their findings yet. They were able to publish a full description in 2020, and concluded ''Sahelanthropus'' was not bipedal. In 2022, French primatologist Franck Guy and colleagues reported that a hominin left femur (TM 266-01-063), and a right (TM 266-01-358) and a left (TM 266-01-050)
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 ...
(forearm bone) were also discovered at the site in 2001, but were excluded originally from ''Sahelanthropus'' because they could not be reliably associated with the skull. They decided to include it because ''Sahelanthropus'' is the only hominin known from the site, and they concluded that the material is consistent with obligate bipedalism, the earliest evidence of such. All ''Sahelanthropus'' specimens, representing six to nine different adults, have been recovered within the area.


Taphonomy

Upon description, Brunet and colleagues were able to constrain the TM 266 locality to 7 or 6 million years ago (near the end of the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
) based on the animal assemblage, which made ''Sahelanthropus'' the earliest African ape at the time. In 2008, Anne-Elisabeth Lebatard and colleagues (which includes Brunet) attempted to radiometrically date using the 10Be/ 9Be ratio the sediments Toumaï was found near (dubbed the " anthracotheriid unit" after the commonplace '' Libycosaurus petrochii''). Averaging the ages of 28 samples, they reported an approximate date of 7.2–6.8 million years ago. Their methods were soon challenged by Beauvilain, who clarified that Toumaï was found on loose sediments at the surface rather than being "unearthed", and had probably been exposed to the harsh sun and wind for some time considering it was encrusted in an iron shell and
desert varnish Desert varnish or rock varnish is an orange-yellow to black coating found on exposed rock surfaces in arid environments. Desert varnish is approximately one micrometer thick and exhibits nanometer-scale layering. Rock rust and desert patina are ot ...
. This would mean it is unsafe to assume that the skull and nearby sediments were deposited at the same time, making such radiometric dating impossible. Further, the ''Sahelanthropus'' fossils lack white silaceous
cement A cement is a binder, a chemical substance used for construction that sets, hardens, and adheres to other materials to bind them together. Cement is seldom used on its own, but rather to bind sand and gravel ( aggregate) together. Cement mix ...
which is present on every other fossil in the site, which would mean they date to different time periods. Because the large mammal fossils were scattered across the area instead of concentrated like the ''Sahelanthropus'' fossils, the discoverers originally believed the ''Sahelanthropus'' fossils were dumped there by a palaeontologist or geologist, but later dismissed this because the skull was too complete to have been thrown away like that. In 2009, Beauvilain and argued that Toumaï was purposefully buried in a "grave", because the skull was also found with two parallel rows of large mammal fossils, seemingly forming a box. Because the "grave" is orientated in a northeast–southwest direction towards
Mecca Mecca (; officially Makkah al-Mukarramah, commonly shortened to Makkah ()) is a city and administrative center of the Mecca Province of Saudi Arabia, and the Holiest sites in Islam, holiest city in Islam. It is inland from Jeddah on the Red ...
, and all sides of the skull were exposed to the wind and were eroded (meaning the skull had somehow turned), they argued that Toumaï was first buried by nomads who identified the skull as human and collected nearby limb fossils (believing them to belong with the skull) and buried them, and was reburied again sometime after the 11th century by Muslims who reorientated the grave towards Mecca when the fossils were re-exposed.


Classification

When describing the species in 2002, Brunet ''et al.'' noted the combination of features which would be considered archaic or derived for a species on the human line (the subtribe Hominina), the latter namely bipedal locomotion and reduced
canine teeth In mammalian oral anatomy, the canine teeth, also called cuspids, dog teeth, or (in the context of the upper jaw) fangs, eye teeth, vampire teeth, or vampire fangs, are the relatively long, pointed teeth. They can appear more flattened howeve ...
, which they interpreted as evidence of its position near the
chimpanzee–human last common ancestor The chimpanzee–human last common ancestor (CHLCA) is the last common ancestor shared by the extant ''Homo'' (human) and '' Pan'' (chimpanzee and bonobo) genera of Hominini. Due to complex hybrid speciation, it is not currently possible to give ...
(CHLCA). This classification made ''Sahelanthropus'' the oldest Hominina, shifting the centre of origin for the
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
away from East Africa. They also suggested that ''Sahelanthropus'' could be a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and t ...
to the 5.5 to 4.5 million year old ''
Ardipithecus ''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 chimpa ...
'' and later Hominina. The classification of ''Sahelanthropus'' in Hominina, as well as ''Ardipithecus'' and the 6 million year old ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopitheci ...
'', was at odds with molecular analyses of the time which had placed the CHLCA between 6 and 4 million years ago based on a high
mutation rate In genetics, the mutation rate is the frequency of new mutations in a single gene or organism over time. Mutation rates are not constant and are not limited to a single type of mutation; there are many different types of mutations. Mutation rates ...
of about 70 mutations per generation. All these genera were anatomically too derived to represent a basal
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 ...
(the group containing chimps and humans), so molecular data would only permit their classification into more ancient and now-extinct lineages. This was overturned in 2012 by geneticists Aylwyn Scally and Richard Durbin who studied the genomes of children and their parents and found the mutation was actually half that, placing the CHLCA anywhere from 14 to 7 million years ago, though most geneticists and palaeoanthropologists use 8 to 7 million years ago. A further possibility is that Toumaï is not ancestral to either humans or chimpanzees at all, but rather an early representative of the
Gorillini Gorillini is a taxonomic tribe containing three genera: ''Gorilla'' and the extinct ''Chororapithecus'' and ''Nakalipithecus ''Nakalipithecus nakayamai'' is an extinct species of great ape from Nakali, Kenya, from about 9.9–9.8 million ye ...
lineage.
Brigitte Senut Brigitte Senut (27 January 1954, Paris) is a French paleoprimatologist and paleoanthropologist and a professor at the National Museum of Natural History, Paris. She is a specialist in the evolution of great apes and humans. Life and work Senu ...
and
Martin Pickford Martin Pickford was lecturer in the Chair of Paleoanthropology and Prehistory at the Collège de France and honorary affiliate at the Département Histoire de la Terre in the Muséum national d'histoire naturelle, Muséum national d'Histoire. ...
, the discoverers of ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopitheci ...
tugenensis'', suggested that the features of ''S. tchadensis'' are consistent with a female proto-
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four or fi ...
. Even if this claim is upheld the find would lose none of its significance, because at present, very few chimpanzee or gorilla ancestors have been found anywhere in Africa. Thus if ''S. tchadensis'' is an ancestral relative of the chimpanzees or gorillas, then it represents the earliest known member of ''their'' lineage. And ''S. tchadensis'' does indicate that the last common ancestor of humans and chimpanzees is unlikely to closely resemble extant chimpanzees, as had been previously supposed by some paleontologists. With the
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 ani ...
known to have existed in early hominins, the difference between ''
Ardipithecus ''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 chimpa ...
'' and ''Sahelanthropus'' may not be large enough to warrant a separate species for the latter.


Anatomy

Existing fossils include a relatively small
cranium The skull is a bone protective cavity for the brain. The skull is composed of four types of bone i.e., cranial bones, facial bones, ear ossicles and hyoid bone. However two parts are more prominent: the cranium and the mandible. In humans, the ...
, five pieces of
jaw The jaw is any opposable articulated structure at the entrance of the mouth, typically used for grasping and manipulating food. The term ''jaws'' is also broadly applied to the whole of the structures constituting the vault of the mouth and serv ...
, and some
teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, tear ...
, making up a head that has a mixture of derived and primitive features. A virtual reconstruction of the interior of the braincase indicated a cranial capacity of 378 cm3, similar to that of extant chimpanzees and approximately a third the size of modern human brains. The teeth, brow ridges, and facial structure differ markedly from those found in modern humans. Cranial features show a flatter face, U-shaped tooth rows, small
canines Canine may refer to: Zoology and anatomy * a dog-like Canid animal in the subfamily Caninae ** ''Canis'', a genus including dogs, wolves, coyotes, and jackals ** Dog, the domestic dog * Canine tooth, in mammalian oral anatomy People with the surn ...
, an anterior
foramen magnum The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblon ...
, and heavy brow ridges. The only known skull suffered a large amount of distortion during the time of fossilisation and discovery, as the cranium is dorsoventrally flattened, and the right side is depressed. In the original description in 2002, Brunet ''et al.'' said it "would not be unreasonable" to speculate that ''Sahelanthropus'' was capable of maintaining an upright posture while walking
biped 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' a ...
ally. Because they had not reported any limb bones or other post-cranial material (anything other than the skull), this was based on the reconstructed original orientation of the
foramen magnum The foramen magnum ( la, great hole) is a large, oval-shaped opening in the occipital bone of the skull. It is one of the several oval or circular openings (foramina) in the base of the skull. The spinal cord, an extension of the medulla oblon ...
(where the skull connects the spine), and their classification of ''Sahelanthropus'' into Hominina based on facial comparisons (one of the diagnostic characteristics of Hominina is bipedalism). This was soon disputed because the orientation of the foramen magnum is not an entirely conclusive piece of evidence in regard to the question of habitual posture, and the features used to classify ''Sahelanthropus'' into Hominina are not entirely unique to Hominina. In 2020, the femur had been formally described, and the study concluded it was not consistent with habitual bipedalism. However, recent postcranial evidence suggests characteristics consistent with habitual bipedalism and arboreal clambering. Ulnar morphologies including bone curvature, cross-section, and orientation combine to rule out quadrupedalism, and femoral morphologies allowing for higher compressive loads of bipedalism strongly suggest hominin characteristics in ''Sahelanthropus tchadensis''.


See also

* ''
Ardipithecus ''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 chimpa ...
'' *
Chimpanzee–human last common ancestor The chimpanzee–human last common ancestor (CHLCA) is the last common ancestor shared by the extant ''Homo'' (human) and '' Pan'' (chimpanzee and bonobo) genera of Hominini. Due to complex hybrid speciation, it is not currently possible to give ...
*
Human evolution Human evolution is the evolutionary process within the history of primates that led to the emergence of ''Homo sapiens'' as a distinct species of the hominid family, which includes the great apes. This process involved the gradual development of ...
*
List of human evolution fossils The following tables give an overview of notable finds of Hominini, hominin fossils and Skeleton, remains relating to human evolution, beginning with the formation of the tribe Hominini (the divergence of the Chimpanzee–human last common ancest ...
''(with images)'' * ''
Orrorin ''Orrorin tugenensis'' is a postulated early species of Homininae, estimated at and discovered in 2000. It is not confirmed how ''Orrorin'' is related to modern humans. Its discovery was used to argue against the hypothesis that australopitheci ...
''


References


Further reading

* * * *


External links


Fossil Hominids: Toumai


* ttps://web.archive.org/web/20080705163738/http://www.livescience.com/humanbiology/ap_050406_chad_bones.html New Findings Bolster Case for Ancient Human Ancestor
''Sahelanthropus tchadensis'', Toumaï
Detailed composition of the Franco-Chadian palaeoanthropological Mission, the sahara scientific missions, the discovery's context, controversy about the misplacement of a molar, the minimum number of individuals, the geology of the site, was Toumaï buried ? and research to date the skull,...
Sahelanthropus
news reporting by John Hawks
''S. tchadensis'' reconstruction

Sahelanthropus tchadensis
Origins – Exploring the Fossil Record – Bradshaw Foundation
Human Timeline (Interactive)
Smithsonian,
National Museum of Natural History The National Museum of Natural History 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. In 2021, with 7 ...
(August 2016). {{Taxonbar, from=Q192383, from2=Q15103944 Hominini Homininae Miocene primates of Africa Fossil taxa described in 2002 Prehistoric Chad