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''Australopithecus bahrelghazali'' is an extinct
species In biology, a species is the basic unit of Taxonomy (biology), 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 ...
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 ...
discovered in 1995 at Koro Toro, Bahr el Gazel, Chad, existing around 3.5 million years ago in the
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Central Africa Central Africa is a subregion of the African continent comprising various countries according to different definitions. Angola, Burundi, the Central African Republic, Chad, the Democratic Republic of the Congo, the Republic of the Congo, E ...
, and demonstrates that this group was widely distributed across Africa as opposed to being restricted to East and southern Africa as previously thought. The validity of ''A. bahrelghazali'' has not been widely accepted, in favour of classifying the specimens as ''
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 ...
'', a better known Pliocene australopithecine from East Africa, because of the anatomical similarity and the fact that ''A. bahrelghazali'' is known only from 3 partial jawbones and an isolated
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 ...
. The specimens inhabited a lakeside grassland environment with sparse tree cover, possibly similar to the modern
Okavango Delta The Okavango Delta (or Okavango Grassland; formerly spelled "Okovango" or "Okovanggo") in Botswana is a swampy inland delta formed where the Okavango River reaches a tectonic trough at an altitude of 930–1,000 m in the central part of the ...
, and similarly predominantly ate C4 savanna foods—such as grasses, sedges,
storage organ A storage organ is a part of a plant specifically modified for storage of energy (generally in the form of carbohydrates) or water. Storage organs often grow underground, where they are better protected from attack by herbivores. Plants that have ...
s, or rhizomes—and to a lesser degree also C3 forest foods—such as fruits, flowers, pods, or insects. However, the teeth seem ill-equipped to process C4 plants, so its true diet is unclear.


Research history

In 1995, two specimens were recovered from Koro Toro, Bahr el Gazel, Chad: KT12/H1 or "
Abel Abel ''Hábel''; ar, هابيل, Hābīl is a Biblical figure in the Book of Genesis within Abrahamic religions. He was the younger brother of Cain, and the younger son of Adam and Eve, the first couple in Biblical history. He was a shepherd ...
" (a jawbone preserving the premolars, canines, and the right second
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, w ...
) and KT12/H2 (an isolated first upper
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 ...
). They were discovered by the Franco-Chadian Paleoanthropological Mission, and reported by French palaeontologist
Michel Brunet Michel Brunet may refer to: * Michel Brunet (historian) Michel Brunet (July 24, 1917 in Montreal – September 4, 1985 in Montreal) was a Quebec historian and essayist. He received his B.A. and M.A. from the Université de Montréal and received ...
, French geographer Alain Beauvilain, French anthropologist
Yves Coppens Yves Coppens (9 August 1934 – 22 June 2022) was a French anthropologist. A graduate from the University of Rennes and Sorbonne, he studied ancient hominids and had multiple published works on this topic, and also produced a film. In October ...
, French palaeontologist Emile Heintz, Chadian geochemist engineer Aladji Hamit Elimi Ali Moutaye, and British palaeoanthropologist
David Pilbeam David Pilbeam (born 21 November 1940 in Brighton, Sussex, England) is the Henry Ford II Professor of the Social Sciences at Harvard University and curator of paleoanthropology at the Peabody Museum of Archaeology and Ethnology. He is a member ...
. Based on the wildlife assemblage, the remains were roughly dated to the middle to late
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Australopithecus 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 Ethiopia during that time period, barring more detailed anatomical comparisons. In 1996, they allocated it to a new species, ''A. bahrelghazali'', naming it after the region; Bahr el Gazel means "River of the Gazelles" in classical Arabic. They denoted KT12/H1 as the
holotype 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 seve ...
and KT12/H2 a
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). O ...
. Another jawbone was discovered at the K13 site in 1997, and a third from the KT40 site. In 2008, a
pelite A pelite (Greek: ''pelos'', "clay") or metapelite is a metamorphosed fine-grained sedimentary rock, i.e. mudstone or siltstone. The term was earlier used by geologists to describe a clay-rich, fine-grained clastic sediment or sedimentary rock, ...
(a type of
sedimentary rock Sedimentary rocks are types of rock that are formed by the accumulation or deposition of mineral or organic particles at Earth's surface, followed by cementation. Sedimentation is the collective name for processes that cause these particle ...
) recovered from the same sediments as Abel was radiometrically dated (using the 10Be/ 9Be ratio) to have been deposited 3.58 million years ago. However, Beauvilain responded that Abel was not found ''
in situ ''In situ'' (; often not italicized in English) is a Latin phrase that translates literally to "on site" or "in position." It can mean "locally", "on site", "on the premises", or "in place" to describe where an event takes place and is used in ...
'' but at the edge of a shallow gulley, and it is impossible to figure out from what
stratigraphic section A stratigraphic section is a sequence of layers of rocks in the order they were deposited. It is based on the principle of original horizontality, which states that layers of sediment are originally deposited horizontally under the action of ...
the specimen (or any other fossil from Koro Toro) was first deposited in, in order to accurately radiometrically date it. Nonetheless, Abel was redated in 2010 using the same methods to about 3.65 million years ago, and Brunet agreed with an age of roughly 3.5 million years ago. ''A. bahrelghazali'' was the first
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 ...
recovered from Central Africa, and disproved the earlier notion that they were restricted to east of the eastern branch of the
East African Rift The East African Rift (EAR) or East African Rift System (EARS) is an active continental rift zone in East Africa. The EAR began developing around the onset of the Miocene, 22–25 million years ago. In the past it was considered to be part of a ...
which formed in the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene Epoch made up of two stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million years ago) to 5.333 Ma. The ...
. Koro Toro is situated about from the Rift Valley, and the remains suggest australopithecines were widely distributed in grassland and woodland zones across the continent. The lack of other Central and West African australopithecines may be due to
sampling bias In statistics, sampling bias is a bias (statistics), bias in which a sample is collected in such a way that some members of the intended statistical population, population have a lower or higher sampling probability than others. It results in a bia ...
, as similarly aged fossil-bearing sediments are more or less unknown beyond East Africa. The ancestors of ''A. bahrelghazali'' may have left East Africa via the
Central African Shear Zone The Central African Shear Zone (CASZ) (or Shear System) is a wrench fault system extending in an ENE direction from the Gulf of Guinea through Cameroon into Sudan. The structure is not well understood. , there was still no general agreement about ...
. In 2014, the first australopithecine in the western branch of the East African Rift was reported in Ishango, Democratic Republic of the Congo. At present, the classification of ''Australopithecus'' and ''Paranthropus'' species is in disarray. ''Australopithecus'' is considered a grade taxon whose members are united by their similar physiology rather than how close they are to each other in the hominin family tree. In an attempt to resolve this, in 2003, Spanish writer Camilo José Cela Conde and evolutionary biologist Francisco J. Ayala proposed splitting off the genus "''Praeanthropus''" and including ''A. bahrelghazali'' alongside ''
Sahelanthropus ''Sahelanthropus tchadensis'' is an extinct species of the Homininae (African apes) dated to about , during the Miocene epoch. The species, and its genus ''Sahelanthropus'', was announced in 2002, based mainly on a partial cranium, nicknamed '' ...
'' (the only other fossil ape known from Chad), ''
A. anamensis ''Australopithecus anamensis'' is a hominin species that lived approximately between 4.2 and 3.8 million years ago and is the oldest known ''Australopithecus'' species, living during the Plio-Pleistocene era. Nearly one hundred fossil specimens ...
'', ''A. afarensis'', and '' A. garhi''. The validity of ''A. bahrelghazali'' has not been widely accepted given how few remains there are and how similar they are to ''A. afarensis''.


Anatomy

The teeth of KT12/H1 are quite similar to the jawbone of ''A. afarensis'', with large and incisor-like canines and bicuspid premolars (as opposed to molar-like premolars). Unlike ''A. afarensis'', the alveolar part of the jawbone where the tooth sockets are is almost vertical as opposed to oblique, possesses poorly developed superior transverse torus and moderate inferior torus (two ridges on the midline of the jaw on the tongue side), and thin enamel on the chewing surface of the premolars. Brunet and colleagues had listed the presence of 3 distinct tooth roots as a distinguishing characteristic, but the third premolar of the ''A. afarensis'' LH-24 specimen from
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 ...
, Ethiopia, was described in 2000 as having the same feature, which shows that premolar anatomy was highly variable for ''A. afarensis''. The
mandibular symphysis In human anatomy, the facial skeleton of the skull 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 p ...
(at the midline of the jaw) of KT40, especially, as well as KT12/H1 have the same dimensions as the symphysis of ''A. afarensis'', though theirs is relatively thick compared to the height.


Palaeoecology

Carbon isotope analysis indicates a diet of predominantly C4 savanna foods, such as grasses, sedges, underground
storage organ A storage organ is a part of a plant specifically modified for storage of energy (generally in the form of carbohydrates) or water. Storage organs often grow underground, where they are better protected from attack by herbivores. Plants that have ...
s (USOs), or rhizomes. There is a smaller C3 portion which may have comprised more typical ape food items such as fruits, flowers, pods, or insects. This indicates that, like contemporary and future australopiths, ''A. bahrelghazali'' was capable of exploiting whatever food was abundant in its environment, whereas most primates (including savanna chimps) avoid C4 foods. However, despite 55–80% of δ13C deriving from C4 sources similar to ''
Paranthropus boisei ''Paranthropus boisei'' is a species of australopithecine from the Early Pleistocene of East Africa about 2.5 to 1.15 million years ago. The holotype specimen, OH 5, was discovered by palaeoanthropologist Mary Leakey in 1959, and described ...
'' and the modern gelada (and considerably more than any tested ''A. afarensis'' population), ''A. bahrelghazali'' lacks the specialisations for such a diet. Because the teeth are not
hypsodont Hypsodont is a pattern of dentition with high-crowned teeth and enamel extending past the gum line, providing extra material for wear and tear. Some examples of animals with hypsodont dentition are cows and horses; all animals that feed on grit ...
, it could not have chewed large quantities of grass, and because the enamel is so thin, the teeth would not have been able to withstand the abrasive dirt particles of USOs. In regard to C4 sources, chimps and
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the co ...
s (which have even thinner enamel) consume plant medullas as a fallback food and sedges as an important energy and protein source; however a sedge-based diet likely could not have sustained ''A. bahrelghazali''. During the Pliocene around the expanded
Lake Chad Lake Chad (french: Lac Tchad) is a historically large, shallow, endorheic lake in Central Africa, which has varied in size over the centuries. According to the ''Global Resource Information Database'' of the United Nations Environment Programme ...
(or "Lake Mega-Chad"), insect
trace fossil A trace fossil, also known as an ichnofossil (; from el, ἴχνος ''ikhnos'' "trace, track"), is a fossil record of biological activity but not the preserved remains of the plant or animal itself. Trace fossils contrast with body fossils, ...
s indicate this was a well-vegetated region, and the abundance of rhizomes may suggest a seasonal climate with wet and dry seasons. Koro Toro has yielded several large mammals, including several
antelope The term antelope is used to refer to many species of even-toed ruminant that are indigenous to various regions in Africa and Eurasia. Antelope comprise a wastebasket taxon defined as any of numerous Old World grazing and browsing hoofed mamm ...
s, of which some were
endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found els ...
, the
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...
'' Loxodonta exoptata'', the
white rhinoceros The white rhinoceros, white rhino or square-lipped rhinoceros (''Ceratotherium simum'') is the largest extant species of rhinoceros. It has a wide mouth used for grazing and is the most social of all rhino species. The white rhinoceros consists ...
''
Ceratotherium praecox ''Diceros praecox'' is an extinct species of rhinoceros that lived in Africa during the Pliocene, around 4 million years ago. It is considered the direct ancestor of the living black rhinoceros ''(Diceros bicornis)''. Taxonomy ''Diceros praecox'' ...
'', the
pig The pig (''Sus domesticus''), often called swine, hog, or domestic pig when distinguishing from other members of the genus ''Sus'', is an omnivorous, domesticated, even-toed, hoofed mammal. It is variously considered a subspecies of ''Sus ...
'' Kolpochoerus afarensis'', a ''
Hipparion ''Hipparion'' (Greek, "pony") is an extinct genus of horse that lived in North America, Asia, Europe, and Africa during the Miocene through Pleistocene ~23 Mya—781,000 years ago. It lived in non-forested, grassy plains, shortgrass prairie or st ...
''
horse The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million ...
, a ''
Sivatherium ''Sivatherium'' ("Shiva's beast", from Shiva and ''therium'', Latinized form of Ancient Greek θηρίον -'' thēríon'') is an extinct genus of giraffids that ranged throughout Africa to the Indian subcontinent. The species ''Sivatherium giga ...
'', and a
giraffe The giraffe is a large African hoofed mammal belonging to the genus ''Giraffa''. It is the tallest living terrestrial animal and the largest ruminant on Earth. Traditionally, giraffes were thought to be one species, '' Giraffa cameloparda ...
. Some of these are also known from Pliocene East African sites, implying that animals could freely migrate between east and west of the Great African Rift. The K13 site features, in regard to
bovid The Bovidae comprise the biological family of cloven-hoofed, ruminant mammals that includes cattle, bison, buffalo, antelopes, and caprines. A member of this family is called a bovid. With 143 extant species and 300 known extinct species, ...
s, an abundance of
Reduncinae The bovid subfamily Reduncinae or tribe Reduncini is composed of nine species of antelope, all of which dwell in marshes, floodplains, or other well-watered areas, including the waterbucks and reedbucks. These antelopes first appear in the fossi ...
,
Alcelaphinae The subfamily Alcelaphinae or tribe Alcelaphini of the family Bovidae contains wildebeest, hartebeest, bonteboks, and several similar species. Depending on the classification, there are 6–10 species placed in four genera, although ''Beatragu ...
, and
Antilopinae The Antilopines are even-toed ungulates belonging to the subfamily Antilopinae of the family Bovidae. The members of tribe Antilopini include the gazelles, blackbucks, springboks, gerenuks, dibatags, and Central Asian gazelles, are often ...
, whereas
Tragelaphini The tribe Tragelaphini (sometimes referred to by some authors as "Strepsicerotini"), or the spiral-horned antelopes, are bovines that are endemic to sub-Sahara Africa. These include the bushbuck, kudus, and the elands. The scientific name is in ...
is much rarer, which indicates an open environment which was drier than Pliocene East African sites. In total, the area seems to have been predominantly grasslands with some tree cover. In addition, the area featured aquatic creatures, predominantly
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
, and also 10 other kinds of fish, the
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 ...
''
Hexaprotodon protamphibius ''Hexaprotodon'' is an extinct genus of hippopotamid known from Asia. The name ''Hexaprotodon'' means "six front teeth" as some of the fossil forms have three pairs of incisors.
'', an otter, a '' Geochelone''
tortoise Tortoises () are reptiles of the family Testudinidae of the order Testudines (Latin: ''tortoise''). Like other turtles, tortoises have a shell to protect from predation and other threats. The shell in tortoises is generally hard, and like ot ...
, a '' Trionyx'' softshell turtle, a
false gharial The false gharial (''Tomistoma schlegelii''), also known by the names Malayan gharial, Sunda gharial and tomistoma is a freshwater crocodilian of the family Gavialidae native to Peninsular Malaysia, Borneo, Sumatra and Java. It is listed as Vul ...
, and an
anatid The Anatidae are the biological family of water birds that includes ducks, geese, and swans. The family has a cosmopolitan distribution, occurring on all the world's continents except Antarctica. These birds are adapted for swimming, floating o ...
waterbird A water bird, alternatively waterbird or aquatic bird, is a bird that lives on or around water. In some definitions, the term ''water bird'' is especially applied to birds in freshwater ecosystems, although others make no distinction from seab ...
. These aquatic animals indicate Koro Toro had open-water lakes or streams with swampy grassy margins, connected to the Nilo-Sudan waterways (including the
Nile The Nile, , Bohairic , lg, Kiira , Nobiin language, Nobiin: Áman Dawū is a major north-flowing river in northeastern Africa. It flows into the Mediterranean Sea. The Nile is the longest river in Africa and has historically been considered ...
, Chari,
Niger ) , official_languages = , languages_type = National languagesSenegal Senegal,; Wolof: ''Senegaal''; Pulaar: 𞤅𞤫𞤲𞤫𞤺𞤢𞥄𞤤𞤭 (Senegaali); Arabic: السنغال ''As-Sinighal'') officially the Republic of Senegal,; Wolof: ''Réewum Senegaal''; Pulaar : 𞤈𞤫𞤲𞤣𞤢𞥄𞤲𞤣� ...
,
Volta Volta may refer to: Persons * Alessandro Volta (1745–1827), Italian physicist and inventor of the electric battery, count and eponym of the volt * Giovanni Volta (1928–2012), Italian Roman Catholic bishop * Giovanni Serafino Volta (1764–184 ...
, and
Gambia The Gambia,, ff, Gammbi, ar, غامبيا officially the Republic of The Gambia, is a country in West Africa. It is the smallest country within mainland AfricaHoare, Ben. (2002) ''The Kingfisher A-Z Encyclopedia'', Kingfisher Publicatio ...
Rivers). Koro Toro, during Mega-Chad events (which have been cyclical for the last 7 million years), may have been similar to the modern
Okavango Delta The Okavango Delta (or Okavango Grassland; formerly spelled "Okovango" or "Okovanggo") in Botswana is a swampy inland delta formed where the Okavango River reaches a tectonic trough at an altitude of 930–1,000 m in the central part of the ...
.


See also

*
African archaeology Africa has the longest record of human habitation in the world. The first hominins emerged 6-7 million years ago, and among the earliest anatomically modern human skulls found so far were discovered at Omo Kibish, Jebel Irhoud, and Florisbad. ...
* '' Ardipithecus ramidus'' * ''
Australopithecus anamensis ''Australopithecus anamensis'' is a hominin species that lived approximately between 4.2 and 3.8 million years ago and is the oldest known ''Australopithecus'' species, living during the Plio-Pleistocene era. Nearly one hundred fossil specimens ...
'' * ''
Australopithecus 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 ...
'' * '' Australopithecus deyiremeda'' * ''
Kenyanthropus ''Kenyanthropus'' is a hominin genus identified from the Lomekwi site by Lake Turkana, Kenya, dated to 3.3 to 3.2 million years ago during the Middle Pliocene. It contains one species, ''K. platyops'', but may also include the 2 million year ...
'' *
List of fossil sites This list of fossil sites is a worldwide list of localities known well for the presence of fossils. Some entries in this list are notable for a single, unique find, while others are notable for the large number of fossils found there. Many of t ...
(with link directory) *
List of human evolution fossils The following tables give an overview of notable finds of hominin fossils and remains relating to human evolution, beginning with the formation of the tribe Hominini (the divergence of the human and chimpanzee lineages) in the late Miocene, roug ...
(with images)


References


External links


''Australopithecus bahrelghazali'', Abel

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 ...
(August 2016). {{Taxonbar, from=Q310945 Australopithecus Fossil taxa described in 1995 Prehistoric Chad Pliocene primates