Australopithecus (/ˌɒstrələˈpɪθɪkəs, -loʊ-/ OS-trə-lo-PITH-i-kəs;[1] from Latin australis, meaning 'southern', and Greek πίθηκος (pithekos), meaning 'ape', informal australopithecine or australopith (although the term australopithecine has a broader meaning as a member of the subtribe Australopithecina, [2][3] which includes this genus as well as the Paranthropus, Kenyanthropus,[4] Ardipithecus,[4] and Praeanthropus genera) [5] is a 'genus' of hominins. From paleontological and archaeological evidence, the genus Australopithecus apparently evolved in eastern Africa around 4 million years ago before spreading throughout the continent and eventually becoming extinct two million years ago. Australopithecus is not literally extinct (in the sense of having no living descendants) as the Kenyanthropus, Paranthropus and Homo genera probably emerged as sister of a late Australopithecus species such as A. Africanus and/or A. Sediba. During that time, a number of australopithecine species emerged, including Australopithecus afarensis, A. africanus, A. anamensis, A. bahrelghazali, A. deyiremeda (proposed), A. garhi, and A. sediba.

For some hominid species of this time – A. robustus, A. boisei and A. aethiopicus – some debate exists whether they truly constitute members of the genus Australopithecus. If so, they would be considered 'robust australopiths', while the others would be 'gracile australopiths'. However, if these more robust species do constitute their own genus, they would be under the genus name Paranthropus, a genus described by Robert Broom when the first discovery was made in 1938, which makes these species P. robustus, P. boisei and P. aethiopicus.

Australopithecus species played a significant part in human evolution, the genus Homo being derived from Australopithecus at some time after three million years ago. In addition, they were the first hominids to possess certain genes, known as the duplicated SRGAP2, which increased the length and ability of neurons in the brain.[6] One of the australopith species evolved into the genus Homo in Africa around two million years ago (e.g. Homo habilis), and eventually modern humans, H. sapiens sapiens.[7]

In January 2019, scientists reported that Australopithecus sediba is distinct from, but shares anatomical similarities to, both the older Australopithecus africanus, and the younger Homo habilis.[8]

Temporal range: 4.5–1.977 Ma
Early PlioceneEarly Pleistocene
Australopithecusafarensis reconstruction
Australopithecus afarensis reconstruction, San Diego Museum of Man
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Family: Hominidae
Subfamily: Homininae
Tribe: Hominini
Subtribe: Australopithecina
Genus: Australopithecus
R.A. Dart, 1925
Type species
Australopithecus africanus
Dart, 1925

Also called Praeanthropus

Cladistically included genera (traditionally sometimes excluded):


Gracile australopiths shared several traits with modern apes and humans, and were widespread throughout Eastern and Northern Africa around 3.5 million years ago. The earliest evidence of fundamentally bipedal hominids can be observed at the site of Laetoli in Tanzania. This site contains hominid footprints that are remarkably similar to those of modern humans and have been dated to as old as 3.6 million years.[9] The footprints have generally been classified as australopith, as they are the only form of prehuman hominins known to have existed in that region at that time.

Map of the fossil sites of the early hominids (4.4-1M BP)
Map of the fossil sites of the early australopithecines in Africa

Australopithecus anamensis, A. afarensis, and A. africanus are among the most famous of the extinct hominins. A. africanus was once considered to be ancestral to the genus Homo (in particular Homo erectus). However, fossils assigned to the genus Homo have been found that are older than A. africanus. Thus, the genus Homo either split off from the genus Australopithecus at an earlier date (the latest common ancestor being either A. afarensis or an even earlier form, possibly Kenyanthropus), or both developed from a yet possibly unknown common ancestor independently.

According to the Chimpanzee Genome Project, the human (Ardipithecus, Australopithecus and Homo) and chimpanzee (Pan troglodytes and Pan paniscus) lineages diverged from a common ancestor about five to six million years ago, assuming a constant rate of evolution. It is theoretically more likely for evolution to happen more slowly, as opposed to more quickly, from the date suggested by a gene clock (the result of which is given as a youngest common ancestor, i.e., the latest possible date of divergence.) However, hominins discovered more recently are somewhat older than the presumed rate of evolution would suggest.[10]

Sahelanthropus tchadensis, commonly called "Toumai", is about seven million years old and Orrorin tugenensis lived at least six million years ago. Since little is known of them, they remain controversial among scientists since the molecular clock in humans has determined that humans and chimpanzees had a genetic split at least a million years later. One theory suggests that the human and chimpanzee lineages diverged somewhat at first, then some populations interbred around one million years after diverging.[10]


The brains of most species of Australopithecus were roughly 35% of the size of a modern human brain. Most species of Australopithecus were diminutive and gracile, usually standing 1.2 to 1.4 m (3 ft 11 in to 4 ft 7 in) tall. In several variations is a considerable degree of sexual dimorphism, males being larger than females.[11]

According to one scholar, A. Zihlman, Australopithecus body proportions closely resemble those of bonobos (Pan paniscus),[12] leading evolutionary biologists such as Jeremy Griffith to suggest that bonobos may be phenotypically similar to Australopithecus.[13] Furthermore, thermoregulatory models suggest that Australopithecus species were fully hair covered, more like chimpanzees and bonobos, and unlike humans.[14]

Modern humans do not display the same degree of sexual dimorphism as Australopithecus appears to have. In modern populations, males are on average a mere 15% larger than females, while in Australopithecus, males could be up to 50% larger than females. New research suggests, however, that australopithecines exhibited a lesser degree of sexual dimorphism than these figures suggest, but the issue is not settled.[11]

Species variations

Opinions differ as to whether the species A. aethiopicus, A. boisei, and A. robustus should be included within the genus Australopithecus, and no current consensus exists as to whether they should be placed in a distinct genus, Paranthropus, which is suggested to have developed from the ancestral Australopithecus line. Until the last half-decade, the majority of the scientific community included all the species shown in the box at the top of this article in a single genus. The postulated genus Paranthropus was morphologically distinct from Australopithecus, and its specialized morphology implies that its behaviour may have been quite different from that of its ancestors, although it has been suggested that the distinctive features of A. aethiopicus, A. boisei, and A. robustus may have evolved independently.

Evolutionary role

The fossil record seems to indicate that Australopithecus is the common ancestor of the distinct group of hominids now called Paranthropus (the "robust australopiths"), and most likely the genus Homo, which includes modern humans. Although the intelligence of these early hominids was likely no more sophisticated than in modern apes, the bipedal stature is the key element that distinguishes the group from previous primates, which were quadrupeds. The morphology of Australopithecus upset what scientists previously believed — namely, that strongly increased brain size had preceded bipedalism.

If A. afarensis was the definite hominid that left the footprints at Laetoli, that strengthens the notion that A. afarensis had a small brain, but was a biped. Fossil evidence such as this makes it clear that bipedalism far predated large brains. However, it remains a matter of controversy as to how bipedalism first emerged (several concepts are still being studied). The advantages of bipedalism were that it left the hands free to grasp objects (e.g., carry food and young), and allowed the eyes to look over tall grasses for possible food sources or predators. However, many anthropologists argue that these advantages were not large enough to cause the emergence of bipedalism.

A recent study of primate evolution and morphology noted that all apes, both modern and fossil, show skeletal adaptations to erect posture of the trunk, and that fossils such as Orrorin tugenensis indicate bipedalism around six million years ago, around the time of the split between humans and chimpanzees indicated by genetic studies. This suggested that erect, straight-legged walking originated as an adaptation to tree-dwelling. Studies of modern orangutans in Sumatra have shown that these apes use four legs when walking on large, stable branches, and swing underneath slightly smaller branches, but are bipedal and keep their legs very straight when walking on multiple flexible branches under 4 cm diameter, while also using their arms for balance and additional support. This enables them to get nearer to the edge of the tree canopy to get fruit or cross to another tree.[15]

The ancestors of gorillas and chimpanzees are suggested to have become more specialised in climbing vertical tree trunks, using a bent hip and bent knee posture that matches the knuckle-walking posture they use for ground travel. This was due to climate changes around 11 to 12 million years ago that affected forests in East and Central Africa, so periods occurred when openings prevented travel through the tree canopy, and at these times, ancestral hominids could have adapted the erect walking behaviour for ground travel. Humans are closely related to these apes, and share features including wrist bones apparently strengthened for knuckle-walking.[16]

However, the view that human ancestors were knuckle-walkers is now questioned since the anatomy and biomechanics of knuckle-walking in chimpanzees and gorillas are different, suggesting that this ability evolved independently after the last common ancestor with the human lineage.[17] Further comparative analysis with other primates suggests that these wrist-bone adaptations support a palm-based tree walking.[17]

Radical changes in morphology took place before gracile australopiths evolved; the pelvis structure and feet are very similar to modern humans.[18] The teeth have small canines, but australopiths generally evolved a larger postcanine dentition with thicker enamel.[19]

Most species of Australopithecus were not any more adept at tool use than modern nonhuman primates, yet modern African apes, chimpanzees, and most recently gorillas, have been known to use simple tools (i.e. cracking open nuts with stones and using long sticks to dig for termites in mounds), and chimpanzees have been observed using spears (not thrown) for hunting.

For a long time, no known stone tools were associated with A. afarensis, and paleoanthropologists commonly thought that stone artifacts only dated back to about 2.5 million years ago.[20] However, a 2010 study suggests the hominin species ate meat by carving animal carcasses with stone implements. This finding pushes back the earliest known use of stone tools among hominins to about 3.4 million years ago.[21]

Some have argued that A. garhi used stone tools due to a loose association of this species and butchered animal remains.


Australopithecines have thirty two teeth, like modern humans, but with an intermediate formation; between the great apes and humans. Their molars were parallel, like those of great apes, and they had a slight pre-canine diastema. But, their canines were smaller, like modern humans, and with the teeth less interlocked than in previous hominins. In fact, in some australopithecines the canines are shaped more like incisors.[22]

The molars of Australopithicus fit together in much the same way human's do, with low crowns and four low, rounded cusps used for crushing. They have cutting edges on the crests.[22]

Robust australopithecines (like A. boisei and A. robustus) had larger cheek, or buccal, teeth than the smaller – or gracile – species (like A. afarensis and A. africanus). It is possible that they had more tough, fibrous plant material in their diets while the smaller species of Australopithecus had more meat. But it is also possibly due to their generally larger build requiring more food. Their larger molars do support a slightly different diet, including some hard food.[22]

Australopithecines also had thick enamel, like those in genus Homo, while other great apes have markedly thinner enamel. One explanation for the thicker enamel is that these hominins were living more on the ground than in the trees and were foraging for tubers, nuts, and cereal grains. They would also have been eating a lot of gritty dirt with the food, which would wear at enamel, so thicker enamel would be advantageous. Or, it could simply indicate a change in diet. Robust australopithecines wore their molar surfaces down flat, unlike the more gracile species, who kept their crests, which certainly seems to suggest a different diet. The gracile Australopithecus had larger incisors, which indicates tearing and more meat in the diet, likely scavenged. The wear patterns on the tooth surfaces support a largely herbivorous diet.[22]

When we examine the buccal microwear patterns on the teeth of A. afarensis and A. anamensis, we see that A. afarensis did not consume a lot of grasses or seeds, but rather ate fruits and leaves, but A. anamensis did eat grasses and seeds in addition to fruits and leaves.[23]


Artistic interpretation of Australopithecus afarensis

In a 1979 preliminary microwear study of Australopithecus fossil teeth, anthropologist Alan Walker theorized that robust australopiths were largely frugivorous.[24] Australopithecus species mainly ate fruit, vegetables, small lizards, and tubers. Much research has focused on a comparison between the South African species A. africanus and Paranthropus robustus. Early analyses of dental microwear in these two species showed, compared to P. robustus, A. africanus had fewer microwear features and more scratches as opposed to pits on its molar wear facets.[25]

These observations have been interpreted as evidence that P. robustus may have fed on hard and brittle foods, such as some nuts and seeds.[25] More recently, new analyses based on three-dimensional renderings of wear facets have confirmed earlier work, but have also suggested that P. robustus ate hard foods primarily as a fallback resource, while A. africanus ate more mechanically tough foods.[26] A recent study looking at enamel fractures suggests A. africanus actually ate more hard foods than P. robustus, with double the frequency of antemortem chips.[27]

In 1992, trace-element studies of the strontium/calcium ratios in robust australopith fossils suggested the possibility of animal consumption, as they did in 1994 using stable carbon isotopic analysis.[28]

In 2005, fossils of animal bones with butchery marks dating 2.6 million years old were found at the site of Gona, Ethiopia. Fossil evidence indicates meat consumption by at least three species of hominins occurring around 2.6-2.5 Mya: A. africanus, A. garhi, and P. aethiopicus.[29]

In 2010, fossils of butchered animal bones dated 3.4 million years old were found in Ethiopia, close to regions where australopith fossils were found.[30]

A study in 2018 found non-carious cervical lesions, caused by acid erosion, on the teeth of A. africanus suggesting the individual ate a lot of acidic fruits.[31]

History of study

The type specimen for genus Australopithecus was discovered in 1924, in a lime quarry by workers at Taung, South Africa. The specimen was studied by the Australian anatomist Raymond Dart, who was then working at the University of the Witwatersrand in Johannesburg. The fossil skull was from a three-year-old bipedal primate that he named Australopithecus africanus. The first report was published in Nature in February 1925. Dart realised that the fossil contained a number of humanoid features, and so, he came to the conclusion that this was an early ancestor of humans.[32] Later, Scottish paleontologist Robert Broom and Dart set about to search for more early hominin specimens, and at several sites they found more A. africanus remains, as well as fossils of a species Broom named Paranthropus (which would now be recognised as P. robustus). Initially, anthropologists were largely hostile to the idea that these discoveries were anything but apes, though this changed during the late 1940s.[32]

The first australopithecine discovered in eastern Africa was a skull belonging to an A. boisei that was excavated in 1959 in the Olduvai Gorge in Tanzania by Mary Leakey. Since then, the Leakey family have continued to excavate the gorge, uncovering further evidence for australopithecines, as well as for Homo habilis and Homo erectus.[32] The scientific community took 20 years to widely accept Australopithecus as a member of the family tree.

Then, in 1997, an almost complete Australopithecus skeleton with skull was found in the Sterkfontein caves of Gauteng, South Africa. It is now called "Little Foot" and it is around 3.7 million years old. It was named Australopithecus prometheus[33][34] which has since been placed within A. africanus. Other fossil remains found in the same cave in 2008 were named Australopithecus sediba, which lived 1.9 million years ago. A. africanus probably evolved into A. sediba, which some scientists think may have evolved into H. erectus,[35] though this is heavily disputed.


A taxonomy of the Australopithecus within the great apes is assessed as follows, with Paranthropus and Homo emerging among the Australopithecus.[36] The genus Australopithecus with conventional definitions is assessed to be highly paraphyletic, i.e. it is not a natural group, and the genera Kenyanthropus, Paranthropus and Homo are included.[37][38][39] The exact phylogeny within Australopithecus is still highly controversial. Approximate radiation dates of daughter clades is shown in Millions of years ago (Mya). Sahelanthropus, Orrorin, and Ardipithecus, possibly sisters to Australopithecus, are not shown here.

Hominoidea (20.4 Mya)

Hylobatidae (gibbons)

Hominidae (15.7)

Ponginae (orangutans)

Homininae  (8.8)

Gorillini (gorillas)

Hominini  (6.3)

Panina (chimpanzees)

Hominina (4)

A. anamensis

A. afarensis

A. garhi

A. deyiremeda (†3.3)

A. africanus

Homo (2.5)

Homo habilis Habilis Skull.png

Paranthropus (†1.2)

Kenyanthropus platyops

Homo rudolfensis Rudolfensis Skull.png


Homo ergaster Ergaster Skull.png

Homo erectus Erectus Skull.png


Homo antecessor ssp. Antecessor Skull.png


Homo heidelbergensis(†0.7)

Homo neanderthalensis (†0.25)Neanderthalensis Skull.png

Homo sapiens Sapiens Skull.png

Inconsistent taxonomy

Even though Australopithecus is classified as a "genus", several other genera appear to have emerged in it: Homo, Kenyanthropus and Paranthropus. This genus is thus regarded as an entrenched paraphyletic wastebasket taxon.[40][41][42] Resolving this into monophyletic groupings requires extensive renaming of species in the binomial nomenclature. Possibilities are to rename Homo sapiens to Australopithecus sapiens[43] (or even Pan sapiens[44][45]), or to rename all the Australopithecus species.[46]

Notable specimens

  • KT-12/H1, an A. bahrelghazali mandibular fragment, discovered 1995 in Sahara, Chad
  • AL 129-1, an A. afarensis knee joint, discovered 1973 in Hadar, Ethiopia
  • Karabo, a juvenile male A. sediba, discovered in South Africa
  • Laetoli footprints, preserved hominin footprints in Tanzania
  • Lucy, a 40%-complete skeleton of a female A. afarensis, discovered 1974 in Hadar, Ethiopia
  • Selam, remains of a three-year-old A. afarensis female, discovered in Dikika, Ethiopia
  • STS 5 (Mrs. Ples), the most complete skull of an A. africanus ever found in South Africa
  • STS 14, remains of an A. africanus, discovered 1947 in Sterkfontein, South Africa
  • STS 71, skull of an A. africanus, discovered 1947 in Sterkfontein, South Africa
  • Taung Child, skull of a young A. africanus, discovered 1924 in Taung, South Africa


Plaque marking the discovery of Australopithecus in Tanzania

The spot where the first Australopithecus boisei was discovered in Tanzania.

Mrs Ples

Original skull of Mrs. Ples, a female A. africanus

Taung child - Skin and Muscles

Taung Child by Cicero Moraes, Arc-Team, Antrocom NPO, Museum of the University of Padua.

Cast of the skeleton of Lucy, an A. afarensis

Australopithecus africanus - Cast of taung child

Skull of the Taung child

See also


  2. ^ Wood & Richmond 2000.
  3. ^ Briggs & Crowther 2008, p. 124.
  4. ^ a b Wood 2010.
  5. ^ Cela-Conde & Ayala 2003.
  6. ^ Reardon, Sara (2012), "The Humanity Switch", New Scientist (AU/NZ), 12 May 2012 No. 2864, pp. 10–11. ISSN 1032-1233
  7. ^ Toth, Nicholas and Schick, Kathy (2005). "African Origins" in The Human Past: World Prehistory and the Development of Human Societies (Editor: Chris Scarre). London: Thames and Hudson. Page 60. ISBN 0-500-28531-4
  8. ^ Dartmouth College (17 January 2019). "Understanding our early human ancestors: Australopithecus sediba". EurekAlert!. Retrieved 21 January 2019.
  9. ^ David A. Raichlen; Adam D. Gordon; William E. H. Harcourt-Smith; Adam D. Foster; Wm. Randall Haas Jr (2010). Rosenberg, Karen (ed.). "Laetoli Footprints Preserve Earliest Direct Evidence of Human-Like Bipedal Biomechanics". PLoS ONE. 5 (3): e9769. Bibcode:2010PLoSO...5.9769R. doi:10.1371/journal.pone.0009769. PMC 2842428. PMID 20339543.
  10. ^ a b Bower, Bruce (May 20, 2006). "Hybrid-Driven Evolution: Genomes show complexity of human-chimp split". Science News. 169 (20): 308–309. doi:10.2307/4019102. JSTOR 4019102.
  11. ^ a b Beck, Roger B.; Linda Black; Larry S. Krieger; Phillip C. Naylor; Dahia Ibo Shabaka (1999). World History: Patterns of Interaction. McDougal Littell. ISBN 978-0-395-87274-1.
  14. ^ David-Barrett, T.; Dunbar, R.I.M. (2016). "Bipedality and Hair-loss Revisited: The Impact of Altitude and Activity Scheduling". Journal of Human Evolution. 94: 72–82. doi:10.1016/j.jhevol.2016.02.006. PMC 4874949. PMID 27178459.
  15. ^ Thorpe, SK; Holder, RL; Crompton, RH. (2007). "Origin of human bipedalism as an adaptation for locomotion on flexible branches". Science. 316 (5829): 1328–31. Bibcode:2007Sci...316.1328T. doi:10.1126/science.1140799. PMID 17540902.
  16. ^ Richmond, BG; Begun, DR; Strait, DS (2001). "Origin of human bipedalism: The knuckle-walking hypothesis revisited". American Journal of Physical Anthropology. Suppl 33: 70–105. doi:10.1002/ajpa.10019. PMID 11786992.
  17. ^ a b Kivell, TL; Schmitt, D. (Aug 2009). "Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor". Proceedings of the National Academy of Sciences of the United States of America. 106 (34): 14241–6. Bibcode:2009PNAS..10614241K. doi:10.1073/pnas.0901280106. PMC 2732797. PMID 19667206.
  18. ^ Lovejoy, C. O. (1988). "Evolution of Human walking". Scientific American. 259 (5): 82–89. Bibcode:1988SciAm.259e.118L. doi:10.1038/scientificamerican1188-118. PMID 3212438.
  19. ^ McHenry, H. M. (2009). "Human Evolution". In Michael Ruse; Joseph Travis (eds.). Evolution: The First Four Billion Years. Cambridge, Massachusetts: The Belknap Press of Harvard University Press. pp. 261–265. ISBN 978-0-674-03175-3.
  22. ^ a b c d Kay, R.F., 1985, 'DENTAL EVIDENCE FOR THE DIET OF AUSTRALOPITHECUS', Annual Review of Anthropology, 14, pp. 315-341.
  23. ^ Martínez, L., Estebaranz-Sánchez, F., Galbany, J., & Pérez-Pérez, A., 2016, 'Testing Dietary Hypotheses of East African Hominines Using Buccal Dental Microwear Data', PLOS ONE, 11, pp. 1-25.
  24. ^ Billings, Tom. "Humanity's Evolutionary Prehistoric Diet and Ape Diets--continued, Part D)". Archived from the original on 8 January 2007. Retrieved 2007-01-06.
  25. ^ a b Grine FE (1986). "Dental evidence for dietary differences in Australopithecus and Paranthropus - a quantitative-analysis of permanent molar microwear". Journal of Human Evolution. 15 (8): 783–822. doi:10.1016/S0047-2484(86)80010-0.
  26. ^ Scott RS, Ungar PS, Bergstrom TS, Brown CA, Grine FE, Teaford MF, Walker A (2005). "Dental microwear texture analysis shows within-species diet variability in fossil hominins". Nature. 436 (7051): 693–695. Bibcode:2005Natur.436..693S. doi:10.1038/nature03822. PMID 16079844.
  27. ^ "(PDF) Behavioral inferences from the high levels of dental chipping in Homo naledi". ResearchGate. Retrieved 2019-01-10.
  28. ^ Billings, Tom. "Comparative Anatomy and Physiology Brought Up to Date--continued, Part 3B)". Archived from the original on 15 December 2006. Retrieved 2007-01-06.
  29. ^ Nature. "Evidence for Meat-Eating by Early Humans".
  30. ^ Nature (2010). "Butchering dinner 3.4 million years ago". Nature. doi:10.1038/news.2010.399.
  31. ^ "Root grooves on two adjacent anterior teeth of Australopithecus africanus". International Journal of Paleopathology. 22: 163–167. 2018-09-01. doi:10.1016/j.ijpp.2018.02.004. ISSN 1879-9817.
  32. ^ a b c Lewin, Roger (1999). "The Australopithecines". Human Evolution: An Illustrated Introduction. Blackwell Science. pp. 112–113. ISBN 0632043091.
  33. ^ BRUXELLES L., CLARKE R. J., MAIRE R., ORTEGA R., et STRATFORD D. – 2014. - Stratigraphic analysis of the Sterkfontein StW 573 Australopithecus skeleton and implications for its age. Journal of Human Evolution,
  34. ^ "New stratigraphic research makes Little Foot the oldest complete Australopithecus".
  35. ^ Celia W. Dugger; John Noble Wilford (April 8, 2010). "New Hominid Species Discovered in South Africa". The New York Times.
  36. ^ Saylor, Beverly Z.; Scott, Gary; Levin, Naomi E.; Deino, Alan; Alene, Mulugeta; Ryan, Timothy M.; Melillo, Stephanie M.; Gibert, Luis; Haile-Selassie, Yohannes (2015). "New species from Ethiopia further expands Middle Pliocene hominin diversity". Nature. 521 (7553): 483–488. doi:10.1038/nature14448. ISSN 1476-4687.
  37. ^ Villmoare, Brian (2018-01-30). "Early Homo and the role of the genus in paleoanthropology". American Journal of Physical Anthropology. 165: 72–89. doi:10.1002/ajpa.23387. ISSN 0002-9483. PMID 29380889.
  38. ^ "(PDF) 2 @BULLET Enhanced cognitive capacity as a contingent fact of hominid phylogeny". ResearchGate. Retrieved 2019-01-12.
  39. ^ "Cowen: History of Life, 5th Edition - Student Companion Site". Wiley. p. 20/5. Retrieved 2019-01-12.
  40. ^ Kimbel, William H. (2015), Henke, Winfried; Tattersall, Ian (eds.), "The Species and Diversity of Australopiths", Handbook of Paleoanthropology, Springer Berlin Heidelberg, pp. 2071–2105, doi:10.1007/978-3-642-39979-4_50, ISBN 9783642399787, retrieved 2019-03-24
  41. ^ Fleagle, John G. (2013-03-08). Primate Adaptation and Evolution. Academic Press. p. 364. ISBN 9780123786333.
  42. ^ Schwarz, J.H. (2004). "Barking up the wrong ape--australopiths and the quest for chimpanzee characters in hominid fossils". Collegium antropologicum. 28 Suppl 2: 87–101.
  43. ^ Flegr, Jaroslav (2013-11-27). "Why Drosophila is not Drosophila any more, why it will be worse and what can be done about it?". Zootaxa. 3741 (2): 295–300. doi:10.11646/zootaxa.3741.2.8. ISSN 1175-5334.
  44. ^ Pietrzak-Franger, Dr Monika; Schaff, Prof Dr Barbara; Voigts, Prof Dr Eckart (2014-02-28). Reflecting on Darwin. Ashgate Publishing, Ltd. p. 118. ISBN 9781472414090.
  45. ^ Gribbin, John (2009-08-27). Science: A History: A History. Penguin Books Limited. ISBN 9780141042220.
  46. ^ Hawks, John (2017-03-20). "The plot to kill Homo habilis". Medium. Retrieved 2019-03-24.

Further reading

  • Barraclough, G. (1989). Stone, N. (ed.). Atlas of World History (3rd ed.). Times Books Limited. ISBN 978-0-7230-0304-5..
  • Leakey, Richard (1994). The Origins of Human Kind. New York: BasicBooks. ISBN 978-0-465-03135-1..
  • White, Tim D.; WoldeGabriel, Giday; Asfaw, Berhane; Ambrose, S; Beyene, Y; Bernor, RL; Boisserie, JR; Currie, B; Gilbert, H; Haile-Selassie, Y; Hart, WK; Hlusko, LJ; Howell, FC; Kono, RT; Lehmann, T; Louchart, A; Lovejoy, CO; Renne, PR; Saegusa, H; Vrba, ES; Wesselman, H; Suwa, G (2006). "Asa Issie, Aramis and the Origin of Australopithecus". Nature. 440 (7086): 883–889. Bibcode:2006Natur.440..883W. doi:10.1038/nature04629. PMID 16612373..
  • Gibbons, Ann (2006). The first human. New York: Doubleday. p. 306. ISBN 978-0385512268.
  • Reader, John (2011). Missing links: in search of human origins. New York: Oxford University Press. p. 538. ISBN 978-0-19-927685-1.
  • Tattersall, Ian (2012). Masters of the Planet, the search for our human origins. Palgrave-Macmillan. pp. 1–79. ISBN 978-0-230-10875-2.

External links


Australopithecines are generally Australopithecus (cladistically including the Homo, Paranthropus, and Kenyanthropus genera), and it typically includes the earlier Ardipithecus. All these related species are now sometimes collectively classified as a subtribe of the Hominini tribe called Australopithecina. They are the extinct, close relatives of humans and, 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 (see Hominidae; terms "hominids" and hominins).

The terms australopithecine, et al., come from a former classification as members of a distinct subfamily, the Australopithecinae. Members of Australopithecus are sometimes referred to as the gracile australopithecines, while Paranthropus are called the "robust australopithecines".The australopithecines occurred in the Plio-Pleistocene era and were bipedal, and they were dentally similar to humans, but with a brain size not much larger than that of modern apes, with lesser encephalization than in the genus Homo. Humans (genus Homo) may have descended from australopithecine ancestors and the genus Ardipithecus is a possible ancestor of the australopithecines.

Australopithecus afarensis

Australopithecus afarensis (Latin: "Southern ape from Afar") is an extinct hominin that lived between 3.9 and 2.9 million years ago in Africa. A. afarensis was slenderly built, like the younger Australopithecus africanus. A. afarensis is thought to be more closely related to the genus Homo (which includes the modern human species Homo sapiens), whether as a direct ancestor or a close relative of an unknown ancestor, than any other known primate from the same time. Some researchers include A. afarensis in the genus Praeanthropus.The most famous fossil is the partial skeleton named Lucy (3.2 million years old) found by Donald Johanson and colleagues, who, in celebration of their find, repeatedly played the Beatles song "Lucy in the Sky with Diamonds".

Australopithecus africanus


Australopithecus anamensis

Australopithecus anamensis is a hominin species that lived approximately four million years ago. Nearly one hundred fossil specimens are known from Kenya and Ethiopia, representing over 20 individuals.

It is accepted that A. anamensis is ancestral to A. afarensis and continued an evolving lineage.

Fossil evidence determines that Australopithecus anamensis is the earliest hominin species in the Turkana Basin.

Australopithecus bahrelghazali

Australopithecus bahrelghazali is a hominin species that inhabited the Bahr el Ghazal region in Central Africa around 3.6 million years ago. It was discovered in 1995 by a Franco-Chadian team led by the paleontologist Michel Brunet. All four fossils that have been found since 1995 are jaw (mandibular or maxillar) fragments.

A. bahrelghazali is the only australopithecine species found in Central Africa. It is also of great importance as it is the first fossil to show that there is a geographical "third window", that is, beyond East Africa and South Africa, of early hominin evolution. The locality is roughly 2,500 km (1,600 mi) west of the East African Great Rift Valley, making it far removed from what is broadly thought to be the "cradle" area of human evolution.

Australopithecus deyiremeda

Australopithecus deyiremeda is a proposed species of early hominin among those who lived about 3.5–3.3 million years ago in northern Ethiopia, around the same time and place as several discovered specimens of Australopithecus afarensis, including the well-known "Lucy".

The discoverers believe A. deyiremeda is a new species. If true, some fossils identified as A. afarensis may more properly belong to A. deyiremeda. Some anthropologists have suggested that the identification of A. deyiremeda as a new species requires more evidence than has been obtained thus far.

Australopithecus garhi

Australopithecus garhi is a 2.5-million-year-old gracile australopithecine species whose fossils were discovered in 1996 by a paleontologist research team led by Berhane Asfaw and Tim White.The remains are suggested as representing the transitional stage between the Australopithecus and Homo genera.

Australopithecus sediba

Australopithecus sediba is a species of Australopithecus of the early Pleistocene, identified based on fossil remains dated to about 2 million years ago.

The species is known from six skeletons discovered in the Malapa Fossil Site at the Cradle of Humankind World Heritage Site in South Africa, including a juvenile male (MH1 also called "Karabo", the holotype), an adult female (MH2, the paratype), an adult male, and three infants.

The fossils were found together at the bottom of the Malapa Cave, where they apparently fell to their death, and have been dated to between 1.980 and 1.977 million years ago.Over 220 fragments from the species have been recovered to date. The partial skeletons were initially described in two papers in the journal Science by American and South African paleoanthropologist Lee R. Berger from the University of the Witwatersrand, Johannesburg and colleagues as a newly discovered species of early human ancestor called Australopithecus sediba ("sediba" meaning "natural spring" or "well" in the Sotho language). MH1 is disarticulated and 34% complete if skeletal elements known to be in an unprepared block are included (59.6% if small elements are excluded) while MH2 is 45.6% complete (again 59.6% excluding small elements) and exhibits partial articulation. A later paper concluded Australopithecus sediba is distinct from, but shares anatomical similarities to, both the older Australopithecus africanus and the younger Homo habilis.Australopithecus sediba may have lived in savannas but ate fruit and other foods from the forest—behavior similar to modern-day savanna chimpanzees. The conditions in which the individuals were buried and fossilized were extraordinary, permitting the extraction of plant phytoliths from dental plaque.

Cradle of Humankind

The Cradle of Humankind is a paleoanthropological site about 50 km (31 mi) northwest of Johannesburg, South Africa, in the Gauteng province. Declared a World Heritage site by UNESCO in 1999, the site currently occupies 47,000 hectares (180 sq mi) and contains a complex of limestone caves. The registered name of the site in the list of World Heritage sites is Fossil Hominid Sites of South Africa.

The Sterkfontein Caves were the site of the discovery of a 2.3-million-year-old fossil Australopithecus africanus (nicknamed "Mrs. Ples"), found in 1947 by Robert Broom and John T. Robinson. The find helped corroborate the 1924 discovery of the juvenile Australopithecus africanus skull known as the "Taung Child", by Raymond Dart, at Taung in the North West Province of South Africa, where excavations still continue.

Nearby the site, but not in the site, the Rising Star Cave system contains the Dinaledi Chamber (chamber of stars), in which were discovered fifteen fossil skeletons of an extinct species of hominin, provisionally named Homo naledi.

Sterkfontein alone has produced more than a third of early hominid fossils ever found prior to 2010. The Dinaledi Chamber contains over 1,500 H. naledi fossils, the most extensive discovery of a single hominid species ever found in Africa.


Homo (Latin: homō, "human being") is the genus which 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 related to modern humans (depending on a species), most notably Homo erectus and Homo neanderthalensis.

The genus is taken to emerge with the appearance of Homo habilis, just over two million years ago. Genus Homo, together with the genus Paranthropus is probably sister to A. africanus in the genus Australopithecus, which itself had previously split from the lineage of Pan, the chimpanzees.Homo erectus appeared about two million years ago and, in several early migrations, it spread throughout Africa (where it is dubbed Homo ergaster) and Eurasia.


An adaptive and successful species, Homo erectus persisted for more than a million years, and gradually diverged into new species by around 500,000 years ago.Homo sapiens (anatomically modern humans) emerges close to 300,000 to 200,000 years ago, most likely in Africa, and Homo neanderthalensis emerged at around the same time in Europe and Western Asia.

H. sapiens dispersed from Africa in several waves, from possibly as early as 250,000 years ago, and certainly by 130,000 years ago, the so-called Southern Dispersal beginning about 70,000 years ago leading to the lasting colonisation of Eurasia and Oceania by 50,000 years ago.

Both in Africa and Eurasia, H. sapiens met with and interbred with archaic humans. Separate archaic (non-sapiens) human species are thought to have survived until around 40,000 years ago (Neanderthal extinction), with possible late survival of hybrid species as late as 12,000 years ago (Red Deer Cave people).

Homo gautengensis

Homo gautengensis is a hominin species proposed by biological anthropologist Darren Curnoe in 2010. The species is composed of South African hominin fossils previously attributed to Homo habilis, Homo ergaster, or, in some cases, Australopithecus, and is argued by Curnoe to be the earliest species in the genus Homo.

Homo habilis

Homo habilis is a proposed archaic species of Homo, which lived between roughly 2.1 and 1.5 million years ago, during the Gelasian and early Calabrian stages of the Pleistocene geological epoch.The type specimen is OH 7, discovered in 1960 at Olduvai Gorge in Tanzania, associated with the Oldowan lithic industry; the fossils were identified as a separate species of Homo with the proposed binomial name of H. habilis ("handy man") in 1964. In its appearance and morphology, H. habilis is intermediate between Australopithecus and the somewhat younger Homo erectus and its classification in the genus Homo has been the subject of controversial debate since its original proposal. A main argument for its classification as the first Homo ("human") species was its use of flaked stone tools. However, evidence for earlier tool use (3.39 million years ago) by undisputed members of Australopithecus has been found in the 1990s.In January 2019, scientists reported that Australopithecus sediba is distinct from, but shares anatomical similarities to, both the older Australopithecus africanus, and the younger Homo habilis.


Kenyanthropus platyops is a 3.5 to 3.2-million-year-old (Pliocene) hominin fossil discovered in Lake Turkana, Kenya in 1999 by Justus Erus, who was part of Meave Leakey's team.Leakey (2001) proposes that the fossil represents an entirely new hominin species and genus, while others classify it as a species of Australopithecus, Australopithecus platyops, others as a species of Homo, Homo platyops, and yet others interpret it as an individual of Australopithecus afarensis.

Archaeological discoveries in Lomekwi in 2015, identifying possibly the oldest known evidence of hominin use of tools to date, have indicated that Kenyanthropus platyops may have been the earliest tool-users known.

Lucy (Australopithecus)

Lucy is the common name of AL 288-1, several hundred pieces of bone fossils representing 40 percent of the skeleton of a female of the hominin species Australopithecus afarensis. In Ethiopia, the assembly is also known as Dinkinesh, which means "you are marvelous" in the Amharic language. Lucy was discovered in 1974 in Africa, near the village Hadar in the Awash Valley of the Afar Triangle in Ethiopia, by paleoanthropologist Donald Johanson of the Cleveland Museum of Natural History.The Lucy specimen is an early australopithecine and is dated to about 3.2 million years ago. The skeleton presents a small skull akin to that of non-hominin apes, plus evidence of a walking-gait that was bipedal and upright, akin to that of humans (and other hominins); this combination supports the view of human evolution that bipedalism preceded increase in brain size. A 2016 study proposes that Australopithecus afarensis was also to a large extent tree-dwelling, though the extent of this is debated."Lucy" acquired her name from the song "Lucy in the Sky with Diamonds" by The Beatles, which was played loudly and repeatedly in the expedition camp all evening after the excavation team's first day of work on the recovery site. After public announcement of the discovery, Lucy captured much public interest, becoming a household name at the time.

Lucy became famous worldwide, and the story of her discovery and reconstruction was published in a book by Johanson. Beginning in 2007, the fossil assembly and associated artifacts were exhibited publicly in an extended six-year tour of the United States; the exhibition was called Lucy's Legacy: The Hidden Treasures of Ethiopia. There was discussion of the risks of damage to the unique fossils, and other museums preferred to display casts of the fossil assembly. The original fossils were returned to Ethiopia in 2013, and subsequent exhibitions have used casts.


Paranthropus (from Greek παρα, para "beside"; άνθρωπος, ánthropos "human") is a genus of extinct hominins that lived between 2.6 and 1.1 million years ago. Also known as robust australopithecines, they were bipedal hominids probably descended from the gracile australopithecine hominids (Australopithecus) 2.7 million years ago.The genus is characterised by robust craniodental anatomy, including gorilla-like sagittal cranial crests which suggest strong muscles of mastication, and broad, grinding herbivorous teeth. However, Paranthropus skulls lack the transverse cranial crests of modern gorillas. The low rate of enamel fractures on Paranthropus teeth suggest this characteristic craniodental anatomy evolved to cope with a diet of tough vegetation, rather than hard foods.

Paranthropus aethiopicus

Paranthropus aethiopicus or Australopithecus aethiopicus is an extinct species of hominin, one of the robust australopithecines.

Paranthropus boisei

Paranthropus boisei or Australopithecus boisei was an early hominin, described as the largest of the genus Paranthropus (robust australopithecines). It lived in Eastern Africa during the Pleistocene epoch from about 2.4 until about 1.4 million years ago.

Paranthropus robustus


Paranthropus robustus is generally dated to have lived between 2.0 and 1.2 million years ago. It had large jaws and jaw muscles with the accompanying sagittal crest, and post-canine teeth that were adapted to serve in the dry environment they lived in. The post-canine teeth also commonly display pitting enamel hypoplasia, thought to be caused by a genetic condition, amelogenesis imperfecta, and was likely common due to instability in crucial gene(s) after evolving such large teeth.

Taung Child

The Taung Child (or Taung Baby) is the fossilised skull of a young Australopithecus africanus. It was discovered in 1924 by quarrymen working for the Northern Lime Company in Taung, South Africa. Raymond Dart described it as a new species in the journal Nature in 1925.

The Taung skull is in repository at the University of Witwatersrand. Dean Falk, a specialist in brain evolution, has called it "the most important anthropological fossil of the twentieth century."


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