Australopithecus

Gracile australopith Fossil range: Pliocene
Australopithecus afarensis
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Class:	Mammalia
Order:	Primates
Family:	Hominidae
Genus:	Australopithecus R.A. Dart, 1925
Species
†A. afarensis †A. africanus †A. anamensis †A. bahrelghazali †A. garhi Formerly Australopithecus, Now Paranthropus †P. aethiopicus †P. robustus †P. boisei

Australopithecus (Latin australis "southern", Greek πίθηκος pithekos "ape") is a genus of extinct hominids, made up of the gracile australopiths, and formerly also included their larger relatives, the robust australopiths (which are now given their own genus). The genus Australopithecus is closely related to the human genus Homo, and may be ancestral to it.

Evolution

Map of the fossil sites of the early Australopithecines in Africa

Reconstruction of Australopithecus afarensis

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 3.7 million years. Until recently, the footprints have generally been classified as australopith because that had been the only form of pre-human known to have existed in that region at that time; however, some scholars have considered reassigning them to a yet unidentified very early species of the genus Homo.^{[citation needed]}

Australopithecus anamensis, Australopithecus afarensis and Australopithecus africanus are among the most famous of the extinct hominins. A. africanus used to be regarded as 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 A. afarensis or an even earlier form, possibly Kenyanthropus platyops), or both developed from a yet possibly unknown common ancestor independently.

According to the Chimpanzee Genome Project, both human (Ardipithecus, Australopithecus and Homo) and chimpanzee (Pan troglodytes and Pan paniscus) lineages diverged from a common ancestor about 5 to 6 million years ago, if we assume 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 diversion.) However, hominins discovered more recently are somewhat older than the molecular clock would theorize. Sahelanthropus tchadensis, commonly called "Toumai" is about 7 million years old and Orrorin tugenensis lived at least 6 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 an evolutionary 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. ^[1]

Morphology

Mrs. Pless, a female Australopithecus Africanus

The brains of most species of Australopithecus were roughly 35% of the size of that of a modern human brain. Most species of Australopithecus were diminutive and gracile, usually standing between 1.2 to 1.4 m tall (approx. 4 to 4.5 feet). In several variations of Australopithecus there is a considerable degree of sexual dimorphism, in this case males are larger than females. Modern hominids do not appear to display sexual dimorphism to the same degree — particularly, modern humans display a low degree of sexual dimorphism, with males being only 15% larger than females, on average. In Australopithecus, however, males can be up to 50% larger than females. New research suggests that sexual dimorphism may be less pronounced than this, but there is still debate on the subject. ^[2]

Species variations

Although opinions differ as to whether the species aethiopicus, boisei and robustus should be included within the genus Australopithecus, the current consensus in the scientific community is that they should be placed in a distinct genus, Paranthropus, which is believed to have developed from the ancestral Australopithecus line. Up 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. However, Paranthropus was morphologically distinct from Australopithecus, and its specialized morphology also implies that its behavior was quite different from that of its ancestors.

Evolutionary role

Cast of the skeleton of Lucy, an Australopithecus afarensis

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 modern apes, the bipedal stature is the key evidence which distinguishes the group from previous primates who are quadrupeds. The morphology of Australopithecus upsets what scientists previously believed, namely, that large brains preceded bipedalism. If A. afarensis was the definite hominid which left the footprints at Laetoli, it strengthens the notion that A. afarensis had a small brain but was a biped. Fossil evidence such as this has made it clear that bipedalism far predated large brains. However, it remains a matter of controversy how bipedalism first evolved millions of years ago (several concepts are still being studied). The advantages of bipedalism allowed hands to be free for grasping objects (e.g. carrying 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 evolution of bipedalism.

Skull of the Taung child

A recent study of primate evolution and morphology noted that all apes, both modern and fossil, show skeletal adaptations to upright posture of the trunk, and that fossils such as Orrorin tugenensis indicate bipedalism around 6 million years ago, around the time of the split between humans and chimpanzees indicated by genetic studies. This suggested that upright, straight-legged walking originally evolved as an adaptation to tree-dwelling. Studies of modern orangutans in Sumatra showed that these apes use four legs when walking on large stable branches, swing underneath slightly smaller branches, but are bipedal and keep their legs very straight when walking on multiple small 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.^[3]

It is suggested that the ancestors of gorillas and chimpanzees became more specialised in climbing vertical tree trunks, using a bent hip and bent knee posture which matches the knuckle-walking posture they use for ground travel. This was due to climate changes around 11 to 12 million years ago^{[citation needed]} that affected forests in East and Central Africa so that there were periods when openings prevented travel through the tree canopy, and at these times ancestral hominids could have adapted the upright walking behaviour for ground travel. Humans are closely related to these apes, and share features including wrist bones apparently strengthened for knuckle-walking.^[4] 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 two separate evolutions that happened after the last common ancestor with the human linerage.^[5] Further comparative analysis with other primates suggests these wrist bone adaptations support a palm based tree walking.^[5]

Radical changes in morphology took place before gracile australopiths evolved; the pelvis structure and feet are very similar to modern humans.^[6] The teeth have small canines, but australopiths generally evolved a larger post-canine dentition with thicker enamel.^[7]

Most species of Australopithecus were not any more adept at tool use than modern non-human 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. However, some have argued that A. garhi used stone tools due to a loose association of this species and butchered animal remains.

Diet

In a 1979 preliminary microwear study of Australopithecus fossil teeth, anthropologist Alan Walker theorized that robust australopiths were largely frugivorous.^[8] However, newer methods of studying fossils have suggested the possibility that Australopithecus was omnivorous. 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.^[9] Australopithecus mainly ate fruit, vegetables, and tubers.

History of Study

The first australopithecine to be discovered and documented was a fossil of a three year old Australopithecus afarensis which was discovered 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 who published his findings in Nature magazine in February 1925. Dart realised that the fossil contained a number of humanoid features, and so came to the conclusion that this was an early ancestor of humans. Ten years later, he and the Scottish paleontologist Robert Broom, set about to search for more early hominin specimens, and at several sites they found further A. afarensis remains as well as fossils of a species which Broom named Paranthropus (which would now be recognised as Australopithecus robustus). Initially, anthropologists were largely hostile to the idea that these discoveries were anything but apes, though this changed during the latter years of the 1940s.^[10]

The first australopithecine to be discovered in eastern Africa was a skull belonging to an Australopithecus bosei 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.^[10]

Notable specimens

• Laetoli footprints
• AL 129-1
• Lucy

• STS 5 (Mrs. Ples)
• STS 14
• STS 71
• Taung Child
• Selam

See also

• Aramis, Ethiopia
• List of fossil sites (with link directory)
• List of human evolution fossils (with images)
• Ardipithecus
• Homo Habilis

References

• Barraclough, G. (1989). Stone, N. (ed.). ed. Atlas of World History (3rd ed.). Times Books Limited. ISBN 0-7230-0304-1. 
• Leakey, Richard (1994). The Origins of Human Kind. New York: BasicBooks. ISBN 0-465-03135-8. 
• White, Tim D., et al. "Asa Issie, Aramis and the Origin of Australopithecus." Nature 440 (April 13, 2006), 883-89.

1. ^ Bower, Bruce (May 20, 2006). "Hybrid-Driven Evolution: Genomes show complexity of human-chimp split" (^{[dead link]} – ^{Scholar search}). Science News 169 (20): 308. http://www.sciencenews.org/articles/20060520/fob4.asp. 
2. ^ Beck, Roger B.; Linda Black, Larry S. Krieger, Phillip C. Naylor, Dahia Ibo Shabaka, (1999). World History: Patterns of Interaction. Evans ton, IL: McDougal Littell. ISBN 0-395-87274-X. 
3. ^ Thorpe SK, Holder RL, Crompton RH. (2007). Origin of human bipedalism as an adaptation for locomotion on flexible branches. Science. 316(5829):1328-31. PMID 17540902
4. ^ Richmond BG, Begun DR, Strait DS. (2001). Origin of human bipedalism: The knuckle-walking hypothesis revisited. Am J Phys Anthropol. Suppl 33:70-105. PMID 11786992
5. ^ ^{a b} Kivell TL, Schmitt D. (2009).Independent evolution of knuckle-walking in African apes shows that humans did not evolve from a knuckle-walking ancestor. Proc Natl Acad Sci U S A. Aug 25;106(34):14241-6. PMID 19667206 doi:10.1073/pnas.0901280106
6. ^ Lovejoy, C.O. (1988). "Evolution of Human walking". Scientific American. 259 (5): 82–89. 
7. ^ McHenry, H.M (2009). "Human Evolution". in Michael Ruse & Joseph Travis. Evolution: The First Four Billion Years. Cambridge, Massachusetts: The Belknap Press of Harvard University Press. pp. 261–265. ISBN 978-0-674-03175-3. 
8. ^ Billings, Tom. "Humanity's Evolutionary Prehistoric Diet and Ape Diets--continued, Part D)". http://www.beyondveg.com/nicholson-w/hb/hb-interview1d.shtml#microwear%20studies. 
9. ^ Billings, Tom. "Comparative Anatomy and Physiology Brought Up to Date--continued, Part 3B)". http://www.beyondveg.com/billings-t/comp-anat/comp-anat-3b.shtml. 
10. ^ ^{a b} Lewin, R. (1999). "The Australopithecines". Human Evolution: An Illustrated Introduction. Blackwell Science. pp. 112. 

External links

Wikibooks has a book on the topic of Introduction to Paleoanthropology

Wikispecies has information related to: Australopithecus

• Why australopithecines became bipedal
• Metadata and Virtual Models of Australopithecus Fossils on NESPOS