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Australopithecus afarensis
Fossil range: Pliocene
Cast of the remains of "Lucy"
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Hominidae
Subfamily: Homininae
Genus: Australopithecus
Species: A. afarensis
Binomial name
Australopithecus afarensis
Johanson & White, 1978

Australopithecus afarensis is an extinct hominid which lived between 3.8 and 2.9 million years ago.[citation needed] A. afarensis was slenderly built, like the younger Australopithecus africanus. It is thought that A. afarensis was ancestral to both the genus Australopithecus and the genus Homo, which includes the modern human species, Homo sapiens. The most famous fossil is the partial skeleton named Lucy (3.2 million years old) by Donald Johanson and colleagues, after they played the Beatles song Lucy in the Sky with Diamonds over and over in celebration of their find.[1][2][3]

Contents

Localities

Australopithecus afarensis fossils have only been discovered within northern Africa. Despite Laetoli being the type locality for A. afarensis, the most extensive remains assigned to this species are found in Hadar, Afar Region of Ethiopia, including the above-mentioned "Lucy" partial skeleton and the "First Family" found at the A.L. 333 locality. Other localities bearing A. afarensis remains include Omo, Maka, Fejej and Belohdelie in Ethiopia, and Koobi Fora and Lothagam in Kenya.

Physical characteristics

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Craniodental features and brain size

Australopithecus afarensis skull reconstruction, displayed at Museum of Man, San Diego, California.

Compared to the modern and extinct great apes, A. afarensis has reduced canines and molars, although they are still relatively larger than in modern humans. A. afarensis also has a relatively small brain size (~380–430 cm³) and a prognathic (i.e. projecting anteriorly) face.

The image of a bipedal hominin with a small brain and primitive face was quite a revelation to the paleoanthropological world at the time. This was due to the earlier belief that an increase in brain size was the first major hominin adaptive shift. Before the discoveries of A. afarensis in the 1970s, it was widely thought that an increase in brain size preceded the shift to bipedal locomotion. This was mainly because the oldest known hominins at the time had relatively large brains (e.g. KNM-ER 1470, Homo rudolfensis, which was found just a few years before Lucy and had a cranial capacity of ~800 cm³).

There is considerable debate regarding the locomotor behaviour of A. afarensis. Some believe that A. afarensis was almost exclusively bipedal, while others believe that the creatures were partly arboreal. The anatomy of the hands, feet and shoulder joints in many ways favour the latter interpretation. The curvature of the finger and toe bones (Phalanges) approaches that of modern-day apes, and is most likely reflective of their ability to efficiently grasp branches and climb. The presence of a wrist-locking mechanism might suggest that they engaged in knuckle-walking.[4] However, these conclusions have been questioned on the basis of close analysis of knuckle-walking and the comparison of wrist bones in different species of primates.[5] The shoulder joint is also oriented more cranially (i.e. towards the skull) than in modern humans. Combined with the relatively long arms A. afarensis are thought to have had, this is thought by many to be reflective of a heightened ability to use the arm above the head in climbing behaviour. Furthermore, scans of the skulls reveal a canal and bony labyrinth morphology, which some suggest is not conducive to proper bipedal locomotion.[citation needed]

Lucy skeleton reconstruction. Cleveland Natural History Museum.

However, there are also a number of traits in the A. afarensis skeleton which strongly reflect bipedalism, to the extent that some researchers have suggested that bipedality evolved long before A. afarensis.[6] In overall anatomy, the pelvis is far more human-like than ape-like. The iliac blades are short and wide, the sacrum is wide and positioned directly behind the hip joint, and there is clear evidence of a strong attachment for the knee extensors. While the pelvis is not wholly human-like (being markedly wide with flared with laterally orientated iliac blades), these features point to a structure that can be considered radically remodeled to accommodate a significant degree of bipedalism in the animals' locomotor repertoire. Importantly, the femur also angles in toward the knee from the hip. This trait would have allowed the foot to have fallen closer to the midline of the body, and is a strong indication of habitual bipedal locomotion. Along with humans, present day orangutans and spider monkeys possess this same feature. The feet also feature adducted big toes, making it difficult if not impossible to grasp branches with the hindlimbs. The loss of a grasping hindlimb also increases the risk of an infant being dropped or falling, as primates typically hold onto their mothers while the mother goes about her daily business. Without the second set of grasping limbs, the infant cannot maintain as strong a grip, and likely had to be held with help from the mother. The problem of holding the infant would be multiplied if the mother also had to climb trees. Bones of the foot also indicate bipedality.[7][8]

A reconstruction of a female Australopithecus afarensis

Computer simulations using dynamic modelling of the skeleton's inertial properties and kinematics have indicated that A. afarensis was able to walk in the same way modern humans walk, with a normal erect gait or with bent hips and knees, but could not walk in the same way as chimpanzees. The upright gait would have been much more efficient than the bent knee and hip walking, which would have taken twice as much energy.[9][10] It appears probable that A. afarensis was quite an efficient bipedal walker over short distances, and the spacing of the footprints at Laetoli indicates that they were walking at 1.0 m/s or above, which matches human small-town walking speeds.[11]

It is commonly thought that upright bipedal walking evolved from knuckle-walking with bent legs, in the manner used by chimpanzees and gorillas to move around on the ground, but fossils such as Orrorin tugenensis indicate bipedalism around 5 to 8 million years ago, in the same general period when genetic studies suggest the lineage of chimpanzees and humans diverged. Modern apes and their fossil ancestors show skeletal adaptations to an upright posture used in tree climbing, and it has been proposed that upright, straight-legged walking originally evolved as an adaptation to tree-dwelling. Studies of modern orangutans in Sumatra have shown these apes using four legs when walking on large stable branches and when swinging underneath slightly smaller branches, but are bipedal and maintain their legs very straight when using multiple small flexible branches under 4 cm. in 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 grasp fruit or cross to another tree.

Climate changes around 11 to 12 million years ago affected forests in East and Central Africa, establishing periods where openings prevented travel through the tree canopy, and during these times ancestral hominids could have adapted the upright walking behaviour for ground travel, while the ancestors of gorillas and chimpanzees became more specialised in climbing vertical tree trunks or lianas with a bent hip and bent knee posture, ultimately leading them to use the related knuckle-walking posture for ground travel. This would lead to A. afarensis usage of upright bipedalism for ground travel, while still having arms well adapted for climbing smaller trees. However, chimpanzees and gorillas are the closest living relatives to humans, and share anatomical features including a fused wrist bone which may also suggest knuckle-walking by human ancestors.[12][13][14] Other studies suggest that an upright spine and a primarily vertical body plan in primates dates back to Morotopithecus bishopi in the Early Miocene of 21.6 million years ago.[15][16] the earliest humanlike primates. Known from fossil remains found in Africa, australopithecines, or australopiths, represent the group from which the ancestors of modern humans emerged. As generally used, the term australopithecines covers all early human fossils dated from about 7 million to 2.5 million years ago, and some of those dated from 2.5 million to 1.4 million years ago. The group became extinct after that time.

At some time before about 2.5 million years ago, a currently unknown australopithecine species gave rise to the ancestor of our own genus Homo. The earliest members of Homo resembled late australopiths, and coexisted with them for more than a million years, but the Homo species had larger brains.[citation needed]

Australopiths evolved from apes and had a combination of apelike and humanlike traits. Their faces protruded like those of apes, and they probably had a similar amount of body hair. However, several traits distinguished australopithecines from the apes. First, australopiths are believed to have walked upright on two legs most of the time—a practice known as bipedalism—instead of using all four limbs for locomotion. In addition, australopiths had smaller canine teeth than those of apes. Studies of australopith hand bones suggest that at least one species could have made the earliest stone tools around 2.5 million years ago.[citation needed] However, some scientists believe only members of the larger-brained genus Homo would have had the mental capacity and hand manipulation skills needed to make these tools (see Stone Age).

Scientists have identified many australopithecine species. Some of these are classified in the genus Australopithecus (meaning “southern ape”). Other australopithecines belong to different genera, including Sahelanthropus, Orrorin, Ardipithecus, Kenyanthropus, and, by some classifications, Paranthropus.

Social characteristics

Australopithecus afarensis reconstruction

It is difficult to reconstruct the social behaviour of extinct fossil species. However, the social structure is likely to be comparable to some extent to that of modern apes and monkeys, given the average difference in body size between males and females (known as sexual dimorphism). Although there is considerable debate over how large the degree of sexual dimorphism was between males and females of A. afarensis, it is likely that males were relatively larger than females. If observations on the relationship between sexual dimorphism and social group structure from modern great apes are applied to A. afarensis then these creatures most likely lived in small family groups containing a single dominant male and a number of breeding females.[17]

There are no known stone tools associated with A. afarensis, and the present archeological record of stone artifacts only dates back to approximately 2.5 million years ago.[18]

Lineage questions

In 1977, Donald Johanson and his colleague Tim White carried out detailed morphological studies on their finds to date, including both Lucy and the "First Family" fossils. They compared the fossils to chimpanzee, gorilla and modern human specimens, and casts of extinct hominid fossils, with particular attention to jaws and dental arcades, and found that their fossils were somewhere between humans and apes, possibly closer to apes, though with essentially human bodies. They reached the conclusion that it could not be classified in the genus Homo and should be in the genus Australopithecus as the new species Australopithecus afarensis. They believed that this extinct hominid would prove to be ancestral to Australopithecus robustus and Australopithecus africanus which is thought to be an ancestor of Homo that includes the modern human species, Homo sapiens,[19] and this conclusion was widely accepted.[18] However, in 2006 scientists Yoel Rak, Avishag Ginzburg, and Eli Geffen carried out a morphological analysis which found that the mandibular ramus (jawbone) of australopithecus afarensis specimen A. L. 822-1 discovered in 2002 is robust , and from further studies they concluded that "australopithecus afarensis" is more likely an ancestor of only the robust australopithecines branch of the hominid evolutionary tree, and not an ancestor of the Australopithecus africanus branch, so not a direct ancestor of man. They concluded that Ardipithecus ramidus discovered by White and colleagues in the 1990s is a more likely ancestor of the human clade.[20][21]

Notable Fossils

Type specimen

The type specimen for A. afarensis is LH 4, an adult mandible from the site of Laetoli, Tanzania.

AL 129-1

The first A. afarensis knee joint was discovered in November 1973 by Donald Johanson as part of a team involving Maurice Taieb, Yves Coppens and Tim White in the Middle Awash of Ethiopia's Afar Depression.

AL 288-1 (Lucy)

Picture of Lucy remains replica, Museo Nacional de Antropología, Mexico City

The first A. afarensis skeleton was discovered on November 24, 1974 near Hadar in Ethiopia by Tom Gray in the company of Donald Johanson, as part of a team involving Maurice Taieb, Yves Coppens and Tim White in the Middle Awash of Ethiopia's Afar Depression.

Site 333

Michael Bush, one of Don Johanson's students, made another major discovery in 1975: near Lucy, on the other side of the hill, he found the "First Family", including 200 fragments of A. afarensis. The site of the findings is now known as "site 333", by a count of fossil fragments uncovered, such as teeth and pieces of jaw. 13 individuals were uncovered and all were adults, with no injuries caused by carnivores. All 13 individuals seemed to have died at the same time, thus Johanson concluded that they might have been killed instantly from a flash flood.

Selam

Selam or DIK 1-1

On September 20, 2006, Scientific American magazine presented the findings of a dig in Dikika, Ethiopia, a few miles from the place where Lucy was found. The recovered skeleton of a 3-year-old A. afarensis girl comprises almost the entire skull and torso, and most parts of the limbs. The features of the skeleton suggest adaptation to walking upright (bipedalism) as well as tree-climbing, features that match the skeletal features of Lucy and fall midway between human and humanoid ape anatomy. "Baby Lucy" has officially been named "Selam" (meaning peace in most Ethiopian languages).[22]

Others

Related work

Further findings at Afar, including the many hominin bones in site 333, produced more bones of concurrent date, and led to Johanson and White's eventual argument that the Koobi Fora hominins were concurrent with the Afar hominins. In other words, Lucy was not unique in evolving bipedalism and a flat face.

Recently, an entirely new species has been discovered, called Kenyanthropus platyops, however the cranium KNM WT 40000 has a much distorted matrix making it hard to distinguish (however a flat face is present). This had many of the same characteristics as Lucy, but is possibly an entirely different genus.

Another species, called Ardipithecus ramidus, was found by White and colleagues in the 1990s. This was fully bipedal, yet appears to have been contemporaneous with a woodland environment. Scientists have not yet been able to draw an estimation of the cranial capacity of Ar. ramidus as only small jaw and leg fragments have been discovered thus far.

See also

References

  1. ^ Johanson & Maitland 1981, pp. 283–297
  2. ^ Johanson, D.C. (2009). "Lucy (Australopithecus afarensis)". in Michael Ruse & Joseph Travis. Evolution: The First Four Billion Years. Cambridge, Massachusetts: The Belknap Press of Harvard University Press. pp. 693–697. ISBN 978-0-674-03175-3. 
  3. ^ Wood, B.A. (1994). "Evolution of australopithecines". in Jones, S., Martin, R., & Pilbeam, D.. The Cambridge Encyclopedia of Human Evolution. Cambridge, U.K.: Cambridge University Press. p. 234. ISBN 0-521-3270-3.  Also ISBN-0-521-46786-1 (paperback)
  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. ^ 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. ^ Weiss, M.L., & Mann, A.E. (1985). Human Biology and Behaviour: An anthropological perspective (4th ed). Boston: Little Brown. ISBN 0-673-39013-6. 
  8. ^ Latimer, B., & Lovejoy, C.O. (1989). "The Calcaneus of Australopithecus afarensis and its implications for the Evolution of Bipedality". American Journal of Physical Anthropology 78 (3): 369–386. 
  9. ^ "BBC - Science & Nature - The evolution of man". Mother of man - 3.2 million years ago. http://www.bbc.co.uk/sn/prehistoric_life/human/human_evolution/mother_of_man1.shtml. Retrieved 2007-11-01. 
  10. ^ "PREMOG - Research". How Lucy walked. Primate Evolution & Morphology Group (PREMOG), the Department of Human Anatomy and Cell Biology, the School of Biomedical Sciences at the University of Liverpool. 18 May 2007. http://www.liv.ac.uk/premog/premog-research.htm#backevidence. Retrieved 2007-11-01. 
  11. ^ "PREMOG - Supplementry Info". The Laetoli Footprint Trail: 3D reconstruction from texture; archiving, and reverse engineering of early hominin gait. Primate Evolution & Morphology Group (PREMOG), the Department of Human Anatomy and Cell Biology, the School of Biomedical Sciences at the University of Liverpool. 18 May 2007. http://www.liv.ac.uk/premog/premog-sup-info-Laetoli.htm. Retrieved 2007-11-01. 
  12. ^ Ian Sample, science correspondent (June 1, 2007). "New theory rejects popular view of man's evolution - Research - EducationGuardian.co.uk". http://education.guardian.co.uk/higher/research/story/0,,2093002,00.html. Retrieved 2007-11-05. 
  13. ^ "BBC NEWS - Science/Nature - Upright walking 'began in trees'". 31 May 2007. http://news.bbc.co.uk/1/hi/sci/tech/6709627.stm. Retrieved 2007-11-05. 
  14. ^ Thorpe S.K.S.; Holder R.L., and Crompton R.H. (24 May 2007). "PREMOG - Supplementry Info". Origin of Human Bipedalism As an Adaptation for Locomotion on Flexible Branches. Primate Evolution & Morphology Group (PREMOG), the Department of Human Anatomy and Cell Biology, the School of Biomedical Sciences at the University of Liverpool. http://www.liv.ac.uk/premog/premog-sup-info-SCIENCE.htm. Retrieved 2007-11-01. 
  15. ^ Aaron G. Filler (December 24, 2007). "Redefining the word “Human” – Do Some Apes Have Human Ancestors? : OUPblog". http://blog.oup.com/2007/12/human/#more-1425. Retrieved 2007-12-27. 
  16. ^ Aaron G. Filler (October 10, 2007). "PLoS ONE: Homeotic Evolution in the Mammalia: Diversification of Therian Axial Seriation and the Morphogenetic Basis of Human Origins". http://www.plosone.org/article/fetchArticle.action?articleURI=info:doi/10.1371/journal.pone.0001019. Retrieved 2007-12-27. 
  17. ^ Wood, B.A.(1994) p.239
  18. ^ a b Jones, S., Martin, R., & Pilbeam, D., ed (1994). The Cambridge Encyclopedia of Human Evolution. Cambridge: Cambridge University Press. ISBN 0-521-3270-3.  Also ISBN-0-521-46786-1 (paperback)
  19. ^ Johanson & Maitland 1981, pp. 265–266, 278–279, 283–297
  20. ^ Yoel Rak; Avishag Ginzburg and Eli Geffen (April 17, 2007). "From the Cover: Gorilla-like anatomy on Australopithecus afarensis mandibles suggests Au. afarensis link to robust australopiths -- Rak et al. 104 (16): 6568 -- Proceedings of the National Academy of Sciences". http://www.pnas.org/cgi/content/full/104/16/6568. Retrieved 2007-10-19. 
  21. ^ Yoel Rak; Avishag Ginzburg and Eli Geffen (April 17, 2007). "From the Cover: Gorilla-like anatomy on Australopithecus afarensis mandibles suggests Au. afarensis link to robust australopiths -- Rak et al. 104 (16): 6568 Data Supplement - HTML Page - index.htslp -- Proceedings of the National Academy of Sciences". http://www.pnas.org/cgi/content/full/0606454104/DC1. Retrieved 2007-10-19. 
  22. ^ Wong, Kate (2006-09-20). "Lucy's Baby: An extraordinary new human fossil comes to light". Scientific American. http://www.sciam.com/article.cfm?chanID=sa003&articleID=00076C1D-62D1-1511-A2D183414B7F0000. 
  • BBC - Dawn of Man (2000) by Robin Mckie| ISBN 0-7894-6262-1
  • Barraclough, G. (1989). Stone, N. (ed.). ed. Atlas of World History (3rd edition ed.). Times Books Limited. ISBN 0-7230-0304-1. 
  • Australopithecus afarensis from The Human Origins Program at the Smithsonian Institution
  • Delson, E., I. Tattersall, J.A. Van Couvering & A.S. Brooks (eds.), ed (2000). Encyclopedia of human evolution and prehistory (2nd Edition ed.). Garland Publishing, New York. ISBN 0-8153-1696-8. 
  • Johanson, D.C.; Maitland, A.E. (1981). Lucy: The Beginning of Humankind. St Albans: Granada. ISBN 0-586-08437-1. 

External links


Wikispecies

Up to date as of January 23, 2010

From Wikispecies

Australopithecus afarensis

Taxonavigation

Main Page
Cladus: Eukaryota
Supergroup: Unikonta
Cladus: Opisthokonta
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Deuterostomia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclassis: Tetrapoda
Classis: Mammalia
Subclassis: Theria
Infraclassis: Placentalia
Ordo: Primates
Subordo: Haplorrhini
Infraordo: Simiiformes
Parvordo: Catarrhini
Superfamilia: Hominoidea
Familia: Hominidae
Subfamilia: Homininae
Tribus: Hominini
Subtribus: Hominina
Genus: †Australopithecus
Species: Australopithecus afarensis

Name

Australopithecus afarensis Johanson & White, 1978

Vernacular names

中文: 阿法南方古猿

Simple English

Australopithecus afarensis ("Lucy")
Fossil range: Pliocene
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Hominidae
Subfamily: Homininae
Genus: Australopithecus
Species: A. afarensis
Binomial name
Australopithecus afarensis
Johanson & White, 1978

Australopithecus afarensis is an extinct hominid. It lived between 3.9 and 2.9 million years ago. A. afarensis is similar to the younger Australopithecus africanus. It had a thin build. It has been thought that A. afarensis was an ancestor to both the genus Australopithecus and the genus Homo. Homo includes the modern human species, Homo sapiens.[1][2].

References

  1. Johanson 1981, p. 283-297
  2. Jones, S. Martin; & R. Pilbeam (ed.) (2004). The Cambridge Encyclopedia of Human Evolution (8th ed.).Cambridge University Press. ISBN 0-521-46786-1

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