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''Chororapithecus'' is an extinct
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 ...
from 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 paved ...
of Ethiopia roughly 8 million years ago during 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 ...
, comprising one species, ''C. abyssinicus''. It is known from 9 isolated teeth discovered in a 2005–2007 survey of the Chorora Formation. The teeth are indistinguishable from those of
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 ...
s in terms of absolute size and relative proportions, and it has been proposed to be an early member of
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 ...
. However, this is controversial given the paucity of remains, and notable anatomical differences between ''Chororapithecus'' and gorilla teeth. The Kenyan
ape Apes (collectively Hominoidea ) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and as well as Europe in prehistory), which together with its siste ...
''
Nakalipithecus ''Nakalipithecus nakayamai'' is an extinct species of great ape from Nakali, Kenya, from about 9.9–9.8 million years ago during the Late Miocene. It is known from a right jawbone with 3 molar tooth, molars and from 11 isolated teeth, and the ...
'' has been proposed to be an ancestor of ''Chororapithecus'' or at least closely related. If correct, they would be the only identified fossil members of any modern non-
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
great ape lineage, and would push the gorilla–human
last common ancestor In biology and genetic genealogy, the most recent common ancestor (MRCA), also known as the last common ancestor (LCA) or concestor, of a set of organisms is the most recent individual from which all the organisms of the set are descended. The ...
from 8 million years ago (identified by molecular analysis) to 10 million years ago. The teeth are adapted for processing tough plant fibres as well as hard, brittle food, and the formation is thought to represent a forested lakeside habitat.


Taxonomy

''Chororapithecus'' teeth were discovered 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 paved ...
, Ethiopia, in a 2005–2007 survey in the Beticha
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 ...
of the Chorora Formation, hence the name, and the formation itself is named after the Chorora village about south of the locality. The specific name, ''abyssinicus'', is in reference to
Abyssinia The Ethiopian Empire (), also formerly known by the exonym Abyssinia, or just simply known as Ethiopia (; Amharic and Tigrinya: ኢትዮጵያ , , Oromo: Itoophiyaa, Somali: Itoobiya, Afar: ''Itiyoophiyaa''), was an empire that historica ...
, the former name of Ethiopia. The ape was described in 2007 by anthropologists
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 th ...
, Reiko Kono, Shigehiro Katoh,
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 ...
, and
Yonas Beyene Yonas Beyene is an Ethiopian archaeologist. He is known for his works on the Paleolithic archaeology of Konso and the Middle Awash, Ethiopia. Recently, he prepared the Nomination Files that eventually led to the registration of thKonso Cultural L ...
. The remains represent at least 3, perhaps over 6, different individuals. 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 ...
, CHO-BT 4, is a right upper second molar, and the
paratype In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype nor a syntype). Of ...
s are a left lower canine, 3 right upper third molars, a left lower third molar, a left lower first molar, and a left and a right lower molar fragment, making for a total of 9 isolated teeth. The discoverers noted the teeth have some of the same adaptations for shearing food as those of
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 ...
s, and classified it as the first fossil member and the only other
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 ...
of the
tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...
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 ...
. Because the ''Chororapithecus'' teeth have several specializations not shared with those of gorillas (they exhibit a
derived Derive may refer to: * Derive (computer algebra system), a commercial system made by Texas Instruments * ''Dérive'' (magazine), an Austrian science magazine on urbanism *Dérive, a psychogeographical concept See also * *Derivation (disambiguatio ...
condition compared to the presumed last common ancestor, LCA), they did not consider it as ancestral to the gorilla. However, the discovers also conceded it is possible that ''Chororapithecus'' and gorillas instead
convergently evolved Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
the same teeth due to a similar diet. If ''Chororapithecus'' is not a gorillin, it may be a
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
hominine 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 ...
or not a hominine at all. The teeth were originally dated to 10.5–10 million years ago (mya), and the discoverers then concluded that the gorilla–human LCA existed about 12 mya, but they were re-dated to about 8 mya. If ''Chororapithecus'' was indeed a
stem Stem or STEM may refer to: Plant structures * Plant stem, a plant's aboveground axis, made of vascular tissue, off which leaves and flowers hang * Stipe (botany), a stalk to support some other structure * Stipe (mycology), the stem of a mushro ...
gorilla, the latter date is more consistent with the timing of 8 mya for the LCA according to molecular data. Based on the revised date and similarly large premolar size, the 10 million year old Kenyan ''Nakalipithecus'' has been proposed to have been the ancestor to ''Chororapithecus'', which would move the LCA to 10 mya if correct. Nonetheless, because there are so few remains known, its relations to modern
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 ...
s is unclear. It was the first extinct ape to have been proposed to be a member of the gorilla lineage. It is debated if great apes evolved in Africa or Eurasia given the abundance of early fossil apes species in the latter and the paucity in the former, despite all modern great apes except the
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
being known from Africa. The first Miocene African great ape was discovered in 1997, ''
Samburupithecus ''Samburupithecus'' is an extinct primate that lived in Kenya during the middle to late Miocene. The one species in this genus, ''Samburupithecus kiptalami'', is known only from a maxilla fragment dated to discovered in 1982 and formally descri ...
'', and the only others known are ''
Nakalipithecus ''Nakalipithecus nakayamai'' is an extinct species of great ape from Nakali, Kenya, from about 9.9–9.8 million years ago during the Late Miocene. It is known from a right jawbone with 3 molar tooth, molars and from 11 isolated teeth, and the ...
'' and ''Chororapithecus''. If ''Chororapithecus'' is indeed an early gorilla, then it would point to an African origin for great apes. However, earlier Eurasian
dryopithecine Dryopithecini is an extinct tribe of Eurasian and African great apes that are believed to be close to the ancestry of gorillas, chimpanzees and humans. Members of this tribe are known as dryopithecines. Taxonomy * Tribe Dryopithecini† ** ''Ke ...
s may be early hominines.


Anatomy

The teeth, both in absolute size and relative proportions, are the same as in gorillas, and the molars range in size between the largest and smallest of what is normally seen in adult gorillas. Like in gorillas, the upper molars have a long
protocone A cusp is a pointed, projecting, or elevated feature. In animals, it is usually used to refer to raised points on the crowns of teeth. The concept is also used with regard to the leaflets of the four heart valves. The mitral valve, which has two ...
crest, and the lower molars have a correspondingly long
trigonid 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, ...
crest, which increase shearing efficacy. Compared to gorillas, the cusp tips are relatively peripheral, are not well pronounced, and the enamel is thicker especially at the side cusps where the tooth borders other teeth. This causes a wide basin on the middle of the molar. This is reminiscent of the condition seen in
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 ...
(humans and chimps). The upper molars are elongated and narrow, and also have a gorilla-like enamel-
dentine Dentin () (American English) or dentine ( or ) (British English) ( la, substantia eburnea) is a calcified tissue of the body and, along with enamel, cementum, and pulp, is one of the four major components of teeth. It is usually covered by ena ...
juncture.


Palaeobiology

The teeth exhibit adaptations for shearing, but the low cusp height indicates the teeth likely folded and pulverised tough plant fibres rather than cut through them as gorillas and other
folivore In zoology, a folivore is a herbivore that specializes in eating leaves. Mature leaves contain a high proportion of hard-to-digest cellulose, less energy than other types of foods, and often toxic compounds.Jones, S., Martin, R., & Pilbeam, D. (1 ...
s with higher cusps do. The thick enamel is more similar to that of
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genus ...
s, and may indicate the consumption of hard, brittle foods. Orangutan molars are probably adapted for eating hard fruits and nuts. ''Chororapithecus'' and ''
Oreopithecus ''Oreopithecus'' (from the Greek , and , , meaning "hill-ape") is an extinct genus of hominoid primate from the Miocene epoch whose fossils have been found in today's Tuscany and Sardinia in Italy. It existed nine to seven million years ago in ...
'' are the only known folivorous Miocene apes. The Chorora Formation represents a braided river system, possibly a forested area alongside a lake in a forest-savanna mosaic environment. It is the only
sub-Saharan Sub-Saharan Africa is, geographically, the area and regions of the continent of Africa that lies south of the Sahara. These include West Africa, East Africa, Central Africa, and Southern Africa. Geopolitically, in addition to the African co ...
vertebrate-bearing formation spanning from 9–7.4 mya, and records the earliest known occurrences of
cercopithecine The Cercopithecinae are a subfamily of the Old World monkeys, which comprises roughly 71 species, including the baboons, the macaques, and the vervet monkeys. Most cercopithecine monkeys are limited to sub-Saharan Africa, although the macaques r ...
monkeys,
hippo The hippopotamus ( ; : hippopotamuses or hippopotami; ''Hippopotamus amphibius''), also called the hippo, common hippopotamus, or river hippopotamus, is a large semiaquatic mammal native to sub-Saharan Africa. It is one of only two extant ...
s, and
rabbit Rabbits, also known as bunnies or bunny rabbits, are small mammals in the family Leporidae (which also contains the hares) of the order Lagomorpha (which also contains the pikas). ''Oryctolagus cuniculus'' includes the European rabbit speci ...
s in Africa. At the Beticha locality, the large vertebrate assemblage is: colobine and cercopithecine monkeys, a hippo, the pig ''
Nyanzachoerus ''Nyanzachoerus'' is an extinct genus of the pig family ( Suidae) belonging to the subfamily Tetraconodontinae. The several species of ''Nyanzachoerus'' lived in Africa from the Miocene to Pliocene The Pliocene ( ; also Pleiocene) is th ...
'', the antelope-like siveratheriine and ''
Palaeotragus ''Palaeotragus'' ("ancient goat") is a genus of very large, primitive, okapi-like giraffids from the Miocene of Africa and Eurasia. ''Palaeotragus primaevus'' is the older species, being found in early to mid-Miocene strata, while ''P. germaini' ...
'', a bosalephine antelope, wild cattle, the horse ''
Cormohipparion ''Cormohipparion'' is an extinct genus of horse belonging to the tribe Hipparionini that lived in North America during the late Miocene to Pliocene ( Hemphillian to Blancan in the NALMA classification). This ancient species of horse grew up to l ...
'' and an unidentified
equine Equinae is a subfamily of the family Equidae, which have lived worldwide (except Indonesia and Australia) from the Hemingfordian stage of the Early Miocene (16 million years ago) onwards. They are thought to be a monophyletic grouping.B. J. MacFa ...
, a rhino, what may be the elephant ''
Stegotetrabelodon ''Stegotetrabelodon'' is an extinct genus of primitive Elephantinae, elephantid with gomphothere-like anatomical features from the Late Miocene to Early Pliocene of Africa and Eurasia and the Arabian peninsula. The type species is ''S. syrticus'' ...
'', a percrocutid
hyena Hyenas, or hyaenas (from Ancient Greek , ), are feliform carnivoran mammals of the family Hyaenidae . With only four extant species (each in its own genus), it is the fifth-smallest family in the Carnivora and one of the smallest in the clas ...
, and a large cat. Because horses and
rodent Rodents (from Latin , 'to gnaw') are mammals of the order Rodentia (), which are characterized by a single pair of continuously growing incisors in each of the upper and lower jaws. About 40% of all mammal species are rodents. They are na ...
s are much less common in the Beticha locality than the type locality of the formation, Beticha may have been a comparatively more forested and closed environment.


See also


References


External links

*
Researchers find fossils of 10 million-year-old ape
at
Reuters Reuters ( ) is a news agency owned by Thomson Reuters Corporation. It employs around 2,500 journalists and 600 photojournalists in about 200 locations worldwide. Reuters is one of the largest news agencies in the world. The agency was estab ...
{{good article Fossil taxa described in 2007 Miocene primates of Africa Monotypic prehistoric primate genera Prehistoric apes Gorillas Prehistoric primate genera