Australopithecus bahrelghazali

Last updated

Australopithecus bahrelghazali
Temporal range: Piacenzian 3.6/3.5–3  Ma
O
S
D
C
P
T
J
K
Pg
N
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Genus: Australopithecus
Species:
A. bahrelghazali
Binomial name
Australopithecus bahrelghazali
Brunet et al., 1995
Synonyms

Praeanthropus 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 Central Africa, 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 , 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 specimens inhabited a lakeside grassland environment with sparse tree cover, possibly similar to the modern Okavango Delta, and similarly predominantly ate C4 savanna foods—such as grasses, sedges, storage organs, 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.

Contents

Research history

Bahr el Ghazal, Chad ; Australopithecus bahrelghazali 1995 discovery map.png
Koro-Toro.jpg
Location of Koro Toro in Africa (left) and within Chad with Lake Mega-Chad outlined (right)

In 1995, two specimens were recovered from Koro Toro, Bahr el Gazel, Chad: KT12/H1 or "Abel" (a jawbone preserving the premolars, canines, and the right second incisor) and KT12/H2 (an isolated first upper premolar). They were discovered by the Franco-Chadian Paleoanthropological Mission, and reported by French palaeontologist Michel Brunet, French geographer Alain Beauvilain, French anthropologist Yves Coppens, French palaeontologist Emile Heintz, Chadian geochemist engineer Aladji Hamit Elimi Ali Moutaye, and British palaeoanthropologist David Pilbeam. Based on the wildlife assemblage, the remains were roughly dated to the middle to late Pliocene 3.5–3 million years ago; consequently, the describers decided to preliminarily assign the remains to Australopithecus afarensis , which inhabited Ethiopia during that time period, barring more detailed anatomical comparisons. [1] 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 and KT12/H2 a paratype. [2] Another jawbone was discovered at the K13 site in 1997, [3] and a third from the KT40 site. [4]

In 2008, a pelite (a type of sedimentary rock) recovered from the same sediments as Abel was radiometrically dated (using the 10Be/9Be ratio) to have been deposited 3.58 million years ago. [5] However, Beauvilain responded that Abel was not found in situ but at the edge of a shallow gulley, and it is impossible to figure out from what stratigraphic section the specimen (or any other fossil from Koro Toro) was first deposited in, in order to accurately radiometrically date it. [6] Nonetheless, Abel was redated in 2010 using the same methods to about 3.65 million years ago, [7] and Brunet agreed with an age of roughly 3.5 million years ago. [8]

A. bahrelghazali was the first australopithecine recovered from Central Africa, and disproved the earlier notion that they were restricted to east of the eastern branch of the East African Rift which formed in the Late Miocene. Koro Toro is situated about 2,500 km (1,600 mi) 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, as similarly aged fossil-bearing sediments are more or less unknown beyond East Africa. [1] The ancestors of A. bahrelghazali may have left East Africa via the Central African Shear Zone. [9] In 2014, the first australopithecine in the western branch of the East African Rift was reported in Ishango, Democratic Republic of the Congo. [10]

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 (the only other fossil ape known from Chad), A. anamensis , A. afarensis, and A. garhi . [11]

The validity of A. bahrelghazali has not been widely accepted given how few remains there are and how similar they are to A. afarensis. [12]

African hominin timeline(in mya)
View references
H. sapiensH. nalediH. rhodesiensisH. ergasterAu. sedibaP. robustusP. boiseiH. rudolfensisH. habilisAu. garhiP. aethiopicusLD 350-1K. platyopsAu. bahrelghazaliAu. deyiremedaAu. africanusAu. afarensisAu. anamensisAr. ramidusAr. kadabbaAustralopithecus bahrelghazali

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. [1] 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, Ethiopia, was described in 2000 as having the same feature, which shows that premolar anatomy was highly variable for A. afarensis. [13] The mandibular symphysis (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. [4]

Palaeoecology

Carbon isotope analysis indicates a diet of predominantly C4 savanna foods, such as grasses, sedges, underground storage organs (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. [14] However, despite 55–80% of δ13C deriving from C4 sources similar to Paranthropus boisei 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, 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 bonobos (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. [9]

Koro Toro may have been similar to the modern Okavango Delta (above) Okavango delta - Botswana - panoramio.jpg
Koro Toro may have been similar to the modern Okavango Delta (above)

During the Pliocene around the expanded Lake Chad (or "Lake Mega-Chad"), insect trace fossils indicate this was a well-vegetated region, and the abundance of rhizomes may suggest a seasonal climate with wet and dry seasons. [15] Koro Toro has yielded several large mammals, including several antelopes, of which some were endemic, the elephant Loxodonta exoptata , the white rhinoceros Ceratotherium praecox , the pig Kolpochoerus afarensis , a Hipparion horse, a Sivatherium , and a giraffe. 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. [1] The K13 site features, in regard to bovids, an abundance of Reduncinae, Alcelaphinae, and Antilopinae, whereas Tragelaphini is much rarer, which indicates an open environment which was drier than Pliocene East African sites. [16] In total, the area seems to have been predominantly grasslands with some tree cover. [14] In addition, the area featured aquatic creatures, predominantly catfish, and also 10 other kinds of fish, the hippo Hexaprotodon protamphibius , an otter, a Geochelone tortoise, a Trionyx softshell turtle, a false gharial, and an anatid waterbird. 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, Chari, Niger, Senegal, Volta, and Gambia Rivers). [17] 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. [8]

See also

Related Research Articles

<i>Ardipithecus</i> Extinct genus of hominins

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 chimpanzees, the relation of this genus to human ancestors and whether it is a hominin is now a matter of debate. Two fossil species are described in the literature: A. ramidus, which lived about 4.4 million years ago during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago. Initial behavioral analysis indicated that Ardipithecus could be very similar to chimpanzees, however more recent analysis based on canine size and lack of canine sexual dimorphism indicates that Ardipithecus was characterised by reduced aggression, and that they more closely resemble bonobos.

<i>Kenyanthropus</i> Oldest-known tool-making hominin

Kenyanthropus is a genus of extinct hominin 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 old Homo rudolfensis, or K. rudolfensis. Before its naming in 2001, Australopithecus afarensis was widely regarded as the only australopithecine to exist during the Middle Pliocene, but Kenyanthropus evinces a greater diversity than once acknowledged. Kenyanthropus is most recognisable by an unusually flat face and small teeth for such an early hominin, with values on the extremes or beyond the range of variation for australopithecines in regard to these features. Multiple australopithecine species may have coexisted by foraging for different food items, which may be reason why these apes anatomically differ in features related to chewing.

<i>Orrorin</i> Postulated early hominin discovered in Kenya

Orrorin tugenensis is a postulated early species of Homininae, estimated at 6.1 to 5.7 million years ago and discovered in 2000. It is not known how Orrorin are related to modern humans. Its discovery was used to argue against the hypothesis that australopithecines are human ancestors, although this remains the most prevalent hypothesis of human evolution as of 2012.

<i>Australopithecus</i> Genus of hominin ancestral to modern humans

Australopithecus is a genus of early hominins that existed in Africa during the Pliocene and Early Pleistocene. The genera Homo, Paranthropus, and Kenyanthropus evolved from some Australopithecus species. Australopithecus is a member of the subtribe Australopithecina, which sometimes also includes Ardipithecus, though the term "australopithecine" is sometimes used to refer only to members of Australopithecus. Species include A. garhi, A. africanus, A. sediba, A. afarensis, A. anamensis, A. bahrelghazali and A. deyiremeda. Debate exists as to whether some Australopithecus species should be reclassified into new genera, or if Paranthropus and Kenyanthropus are synonymous with Australopithecus, in part because of the taxonomic inconsistency.

<i>Sahelanthropus</i> Extinct hominid from Miocene Africa

Sahelanthropus tchadensis is an extinct species of the hominid dated to about 7 million years ago, during the Miocene epoch. The species, and its genus Sahelanthropus, was announced in 2002, based mainly on a partial cranium, nicknamed Toumaï, discovered in northern Chad.

<i>Australopithecus afarensis</i> Extinct hominid from the Pliocene of East Africa

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 take place until the 1970s. From 1972 to 1977, the International Afar Research Expedition—led by anthropologists Maurice Taieb, Donald Johanson and Yves Coppens—unearthed several hundreds of hominin specimens in Hadar, Ethiopia, the most significant being the exceedingly well-preserved skeleton AL 288-1 ("Lucy") and the site AL 333. Beginning in 1974, Mary Leakey led an expedition into Laetoli, Tanzania, and notably recovered fossil trackways. In 1978, the species was first described, but this was followed by arguments for splitting the wealth of specimens into different species given the wide range of variation which had been attributed to sexual dimorphism. A. afarensis probably descended from A. anamensis and is hypothesised to have given rise to Homo, though the latter is debated.

<i>Australopithecus africanus</i> Extinct hominid from South Africa

Australopithecus africanus is an extinct species of australopithecine which lived between about 3.3 and 2.1 million years ago in the Late Pliocene to Early Pleistocene of South Africa. The species has been recovered from Taung, Sterkfontein, Makapansgat, and Gladysvale. The first specimen, the Taung child, was described by anatomist Raymond Dart in 1924, and was the first early hominin found. However, its closer relations to humans than to other apes would not become widely accepted until the middle of the century because most had believed humans evolved outside of Africa. It is unclear how A. africanus relates to other hominins, being variously placed as ancestral to Homo and Paranthropus, to just Paranthropus, or to just P. robustus. The specimen "Little Foot" is the most completely preserved early hominin, with 90% of the skeleton intact, and the oldest South African australopith. However, it is controversially suggested that it and similar specimens be split off into "A. prometheus".

<i>Australopithecus anamensis</i> Extinct hominin from Pliocene east Africa

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.

<i>Paranthropus aethiopicus</i> Extinct species of hominin of East Africa

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 invalid grouping and is synonymous with Australopithecus, so the species is also often classified as Australopithecus aethiopicus. Whatever the case, it is considered to have been the ancestor of the much more robust P. boisei. It is debated if P. aethiopicus should be subsumed under P. boisei, and the terms P. boisei sensu lato and P. boisei sensu stricto can be used to respectively include and exclude P. aethiopicus from P. boisei.

<i>Australopithecus garhi</i> Extinct hominid from the Afar Region of Ethiopia 2.6–2.5 million years ago

Australopithecus garhi is a species of australopithecine from the Bouri Formation in the Afar Region 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 and the human line, but is now thought to have been an offshoot. Like other australopithecines, A. garhi had a brain volume of 450 cc (27 cu in); a jaw which jutted out (prognathism); relatively large molars and premolars; adaptations for both walking on two legs (bipedalism) and grasping while climbing (arboreality); and it is possible that, though unclear if, males were larger than females. One individual, presumed female based on size, may have been 140 cm tall.

<i>Paranthropus boisei</i> Extinct species of hominin of East Africa

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 at Olduvai Gorge, Tanzania and described by her husband Louis a month later. It was originally placed into its own genus as "Zinjanthropus boisei", but is now relegated to Paranthropus along with other robust australopithecines. However, it is also argued that Paranthropus is an invalid grouping and synonymous with Australopithecus, so the species is also often classified as Australopithecus boisei.

<span class="mw-page-title-main">Hominini</span> Tribe of mammals

The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera Homo (humans) and Pan and in standard usage excludes the genus Gorilla (gorillas).

<span class="mw-page-title-main">Australopithecine</span> Extinct subtribe of the Hominini tribe, and members of the human clade

Australopithecina or Hominina is a subtribe in the tribe Hominini. The members of the subtribe are generally Australopithecus, and it typically includes the earlier Ardipithecus, Orrorin, Sahelanthropus, and Graecopithecus. All these closely related species are now sometimes collectively termed australopiths or homininians. They are the extinct, close relatives of modern humans and, together with the extant genus Homo, comprise the human clade. Members of the human clade, i.e. the Hominini after the split from the chimpanzees, are now called Hominina.

Abel (KT-12/H1) is the name given to the only specimen ever discovered of Australopithecus bahrelghazali. Abel was found in January 1995 in Chad in the Kanem Region by the paleontologist Michel Brunet, who named the fossil "Abel" in memory of his close friend Abel Brillanceau, who had died of malaria in 1989.

<span class="mw-page-title-main">Michel Brunet (paleontologist)</span> French paleontologist and professor

Michel Brunet is a French paleontologist and a professor at the Collège de France. In 2001 Brunet announced the discovery in Central Africa of the skull and jaw remains of a late Miocene hominid nicknamed Toumaï. These remains may predate the earliest previously known hominid remains, Lucy, by over three million years; however, this conclusion is the subject of a significant controversy.

David Pilbeam 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 of the National Academy of Sciences. He received his Ph.D. from Yale University.

The Djurab Desert is a desert in northern Chad. Part of the greater Sahara desert, it makes up much of the area of Chad's Borkou region.

Australopithecus deyiremeda is an extinct species of australopithecine from Woranso–Mille, Afar Region, Ethiopia, about 3.5 to 3.3 million years ago during the Pliocene. Because it is known only from three partial jawbones, it is unclear if these specimens indeed represent a unique species or belong to the much better-known A. afarensis. A. deyiremeda is distinguished by its forward-facing cheek bones and small cheek teeth compared to those of other early hominins. It is unclear if a partial foot specimen exhibiting a dextrous big toe can be assigned to A. deyiremeda. A. deyiremeda lived in a mosaic environment featuring both open grasslands and lake- or riverside forests, and anthropologist Fred Spoor suggests it may have been involved in the Kenyan Lomekwi stone-tool industry typically assigned to Kenyanthropus. A. deyiremeda coexisted with A. afarensis, and they may have exhibited niche partitioning to avoid competing with each other for the same resources, such as by relying on different fallback foods during leaner times.

Tchadailurus is a genus of machairodontine felid from the late Miocene of Chad, Africa.

<i>Afrocygnus</i> Extinct genus of birds

Afrocygnus is an extinct genus of swan, which lived during the Late Miocene, and perhaps up to the Late Pliocene, in what is today North Africa. The only genus of swan known in Africa, aside from fragmentary Pleistocene remains found in East Africa and from occasional observations of vagrant European swans along the Mediterranean coast, it lived in what was during the Miocene a damp wetland spanning from Libya to Chad, alongside the Antracothere Libycosaurus and the early Homininae Sahelanthropus. The genus is considered as the sister taxon of the extant genus Cygnus. Fossils of the genus have been uncovered in the Sahabi Formation of Cyrenaica in Libya, and in the Toros-Menalla locality in the Djurab Desert of Northern Chad.

References

  1. 1 2 3 4 Brunet, M.; Beauvilain, A.; Coppens, Y.; Heintz, É.; Moutaye, A. H. E; Pilbeam, D. (1995). "The first australopithecine 2,500 kilometres west of the Rift Valley (Chad)". Nature. 378 (6554): 273–275. Bibcode:1995Natur.378..273B. doi:10.1038/378273a0. PMID   7477344. S2CID   4365908.
  2. Brunel, Michel; Beauvilain, A.; Coppens, Yves; Heintz, É.; Moutaye, A. H. E; Pilbeam, D. (1996). "Australopithecus bahrelghazali, une nouvelle espèce d'Hominidé ancien de la région de Koro Toro (Tchad)" [Australopithecus bahrelghazali, a new species of fossil hominid from Koro Toro (Chad)](PDF). Comptes Rendus des Séances de l'Académie des Sciences. 322: 907–913.
  3. Brunet, M.; Beauvilain, A.; et al. (1997). "Tchad : un nouveau site à Hominidés Pliocène". Comptes rendus des séances de l'Académie des Sciences. 324: 341–345.
  4. 1 2 Guy, F.; Mackaye, H.-T.; et al. (2008). "Symphyseal shape variation in extant and fossil hominoids, and the symphysis of Australopithecus bahrelghazali". Journal of Human Evolution. 55 (1): 37–47. doi:10.1016/j.jhevol.2007.12.003. PMID   18222528.
  5. Lebatard, A.-E.; Bourlès, D. L.; Duringer, P.; Jolivet, M.; Braucher, R.; Carcaillet, J.; Schuster, M.; Arnaud, N.; Monié, P.; Lihoreau, F.; Likius, A.; Taisso Mackaye, H.; Vignaud, P.; Brunet, M. (2008). "Cosmogenic nuclide dating of Sahelanthropus tchadensis and Australopithecus bahrelghazali: Mio-Pliocene hominids from Chad". Proceedings of the National Academy of Sciences. 105 (9): 3226–3231. Bibcode:2008PNAS..105.3226L. doi: 10.1073/pnas.0708015105 . PMC   2265126 . PMID   18305174.
  6. Beauvilain, A. (2008). "The contexts of discovery of Australopithecus bahrelghazali (Abel) and of Sahelanthropus tchadensis (Toumaï): unearthed, embedded in sandstone, or surface collected?" (PDF). South African Journal of Science. 104 (5–6): 165–168.
  7. Lebetard, A.-E.; Bourlès, D. L.; Braucher, R.; et al. (2010). "Application of the authigenic 10Be/9Be dating method to continental sediments: Reconstruction of the Mio-Pleistocene sedimentary sequence in the early hominid fossiliferous areas of the northern Chad Basin". Earth and Planetary Science Letters. 297 (1–2): 57–70. Bibcode:2010E&PSL.297...57L. doi:10.1016/j.epsl.2010.06.003.
  8. 1 2 Brunet, M. (2010). "Two new Mio-Pliocene Chadian hominids enlighten Charles Darwin's 1871 prediction". Philosophical Transactions of the Royal Society B. 365 (1556): 3315–3321. doi:10.1098/rstb.2010.0069. PMC   2981960 . PMID   20855305.
  9. 1 2 Macho, G. A. (2015). "Pliocene hominin biogeography and ecology". Journal of Human Evolution. 87: 78–86. doi:10.1016/j.jhevol.2015.06.009. PMID   26198846.
  10. Crevecoeur, I.; Skinner, M. M.; Bailey, S. E.; et al. (2014). "First Early Hominin from Central Africa (Ishango, Democratic Republic of Congo)". PLOS ONE. 9 (1): e84652. Bibcode:2014PLoSO...984652C. doi: 10.1371/journal.pone.0084652 . PMC   3888414 . PMID   24427292.
  11. Cela-Conde, C. J.; Ayala, F. J. (2003). "Genera of the human lineage". Proceedings of the National Academy of Sciences. 100 (13): 7684–7689. Bibcode:2003PNAS..100.7684C. doi: 10.1073/pnas.0832372100 . PMC   164648 . PMID   12794185.
  12. Spoor, F.; Leakey, M. G.; O'Higgins, P. (2016). "Middle Pliocene hominin diversity: Australopithecus deyiremeda and Kenyanthropus platyops". Philosophical Transactions of the Royal Society B. 371 (1698): 20150231. doi:10.1098/rstb.2015.0231. PMC   4920288 . PMID   27298462.
  13. White, T. D.; Suwa, G.; Simpson, S.; Asfaw, B. (2000). "Jaws and teeth of Australopithecus afarensis from Maka, Middle Awash, Ethiopia". American Journal of Physical Anthropology. 111 (1): 66. doi:10.1002/(SICI)1096-8644(200001)111:1<45::AID-AJPA4>3.0.CO;2-I. PMID   10618588.
  14. 1 2 Lee-Thorp, J.; Likius, A.; Mackaye, H. T.; et al. (2012). "Isotopic evidence for an early shift to C4 resources by Pliocene hominins in Chad". Proceedings of the National Academy of Sciences. 109 (50): 20369–20372. doi: 10.1073/pnas.1204209109 . PMC   3528505 . PMID   23150583. S2CID   37279343.
  15. Schuster, M.; Duringer, P.; Ghienne, J.-F.; et al. (2009). "Chad Basin: Paleoenvironments of the Sahara since the Late Miocene". Comptes Rendus Geoscience. 341 (8–9): 603–611. Bibcode:2009CRGeo.341..603S. doi:10.1016/j.crte.2009.04.001. S2CID   133162696.
  16. Geraads, D.; Brunet, M.; Mackaye, H. T.; Vignaud, P. (2000). "Pliocene Bovidae (Mammalia) from the Koro Toro Australopithecine sites, Chad" (PDF). Journal of Vertebrate Paleontology. 21 (2): 335–346. doi:10.1671/0272-4634(2001)021[0335:PBMFTK]2.0.CO;2. S2CID   55934117.
  17. Otero, O.; Pinton, A.; Mackaye, H. T.; Likius, A.; Vignaud, P.; Brunet, M. (2010). "The early/late Pliocene ichthyofauna from Koro-Toro, Eastern Djurab, Chad". Geobios. 43 (2): 241–255. Bibcode:2010Geobi..43..241O. doi:10.1016/j.geobios.2009.10.003.