World Library  
Flag as Inappropriate
Email this Article




Temporal range: Late Miocene - Early Pliocene, 5.6–4.4 Ma
Ardipithecus ramidus specimen, nicknamed Ardi
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Family: Hominidae
Subfamily: Homininae
Tribe: (debated[1] Hominini)
Genus: Ardipithecus
White et al., 1995

Ardipithecus kadabba
Ardipithecus ramidus

Ardipithecus is a genus of an extinct hominine that lived during Late Miocene and Early Pliocene in Afar Depression, Ethiopia. Originally described as one of the earliest ancestors of humans after they diverged from the main ape lineage, the relation of this genus to human ancestors and whether it is a hominin is now a matter of debate.[1] Two fossil species are described in the literature: A. ramidus, which lived about 4.4 million years ago[2] during the early Pliocene, and A. kadabba, dated to approximately 5.6 million years ago (late Miocene).[3] Behavioral analysis showed that Ardipithecus could be very similar to those of chimpanzees, indicating that the early human ancestors were very chimpanzee-like in behaviour.[4]


  • Ardipithecus ramidus 1
    • Ardi 1.1
  • Ardipithecus kadabba 2
  • Lifestyle 3
  • Alternative views and further studies 4
  • See also 5
  • References 6
  • External links 7

Ardipithecus ramidus

Map showing discovery locations.

A. ramidus was named in September 1994. The first fossil found was dated to 4.4 million years ago on the basis of its stratigraphic position between two volcanic strata: the basal Gaala Tuff Complex (GATC) and the Daam Aatu Basaltic Tuff (DABT).[5] The name Ardipithecus ramidus stems mostly from the Afar language, in which Ardi means "ground/floor" (borrowed from the Semitic root in either Amharic or Arabic) and ramid means "root". The pithecus portion of the name is from the Greek word for "ape".[6]

Like most hominids, but unlike all previously recognized hominins, it had a grasping hallux or big toe adapted for locomotion in the trees. It is not confirmed how much other features of its skeleton reflect adaptation to bipedalism on the ground as well. Like later hominins, Ardipithecus had reduced canine teeth.

In 1992–1993 a research team headed by [7] The fossils were dated to between 4.35 and 4.45 million years old.[8]

Ardipithecus ramidus had a small brain, measuring between 300 and 350 cm3. This is slightly smaller than a modern bonobo or female common chimpanzee brain, but much smaller than the brain of australopithecines like Lucy (~400 to 550 cm3) and roughly 20% the size of the modern Homo sapiens brain. Like common chimpanzees, A. ramidus was much more prognathic than modern humans.[9]

The teeth of A. ramidus lacked the specialization of other apes, and suggest that it was a generalized omnivore and frugivore (fruit eater) with a diet that did not depend heavily on foliage, fibrous plant material (roots, tubers, etc.), or hard and or abrasive food. The size of the upper canine tooth in A. ramidus males was not distinctly different from that of females. Their upper canines were less sharp than those of modern common chimpanzees in part because of this decreased upper canine size, as larger upper canines can be honed through wear against teeth in the lower mouth. The features of the upper canine in A. ramidus contrast with the sexual dimorphism observed in common chimpanzees, where males have significantly larger and sharper upper canine teeth than females.[10]

The less pronounced nature of the upper canine teeth in A. ramidus has been used to infer aspects of the social behavior of the species and more ancestral hominids. In particular, it has been used to suggest that the last common ancestor of hominids and African apes was characterized by relatively little aggression between males and between groups. This is markedly different from social patterns in common chimpanzees, among which intermale and intergroup aggression are typically high. Researchers in a 2009 study said that this condition "compromises the living chimpanzee as a behavioral model for the ancestral hominid condition."[10]

A. ramidus existed more recently than the most recent common ancestor of humans and chimpanzees (CLCA or Pan-Homo LCA) and thus is not fully representative of that common ancestor. Nevertheless, it is in some ways unlike chimpanzees, suggesting that the common ancestor differs from the modern chimpanzee. After the chimpanzee and human lineages diverged, both underwent substantial evolutionary change. Chimp feet are specialized for grasping trees; A. ramidus feet are better suited for walking. The canine teeth of A. ramidus are smaller, and equal in size between males and females, which suggests reduced male-to-male conflict, increased pair-bonding, and increased parental investment. "Thus, fundamental reproductive and social behavioral changes probably occurred in hominids long before they had enlarged brains and began to use stone tools," the research team concluded.[3]


On October 1, 2009, paleontologists formally announced the discovery of the relatively complete A. ramidus fossil skeleton first unearthed in 1994. The fossil is the remains of a small-brained 50-kilogram (110 lb) female, nicknamed "Ardi", and includes most of the skull and teeth, as well as the pelvis, hands, and feet.[11] It was discovered in Ethiopia's harsh Afar desert at a site called Aramis in the Middle Awash region. Radiometric dating of the layers of volcanic ash encasing the deposits suggest that Ardi lived about 4.4 million years ago. This date, however, has been questioned by others. Fleagle and Kappelman suggest that the region in which Ardi was found is difficult to date radiometrically, and they argue that Ardi should be dated at 3.9 million years.[12]

The fossil is regarded by its describers as shedding light on a stage of human evolution about which little was known, more than a million years before Lucy (Australopithecus afarensis), the iconic early human ancestor candidate who lived 3.2 million years ago, and was discovered in 1974 just 74 km (46 mi) away from Ardi's discovery site. However, because the "Ardi" skeleton is no more than 200,000 years older than the earliest fossils of Australopithecus, and may in fact be younger than they are,[12] some researchers doubt that it can represent a direct ancestor of Australopithecus.

Some researchers infer from the form of her pelvis and limbs and the presence of her abductable hallux, that "Ardi" was a facultative biped: bipedal when moving on the ground, but quadrupedal when moving about in tree branches.[3][13][14] A. ramidus had a more primitive walking ability than later hominids, and could not walk or run for long distances.[15] The teeth suggest omnivory, and are more generalised than those of modern apes.[3]

Ardipithecus kadabba

Ardipithecus kadabba fossils.

Ardipithecus kadabba is "known only from teeth and bits and pieces of skeletal bones",[11] and is dated to approximately 5.6 million years ago.[3] It has been described as a "probable chronospecies" (i.e. ancestor) of A. ramidus.[3] Although originally considered a subspecies of A. ramidus, in 2004 anthropologists Yohannes Haile-Selassie, Gen Suwa, and Tim D. White published an article elevating A. kadabba to species level on the basis of newly discovered teeth from Ethiopia. These teeth show "primitive morphology and wear pattern" which demonstrate that A. kadabba is a distinct species from A. ramidus.[16]

The specific name comes from the Afar word for "basal family ancestor".[17]


The toe and pelvic structure of A. ramidus suggest to some researchers that the organism walked upright.[7]

According to [7]

Alternative views and further studies

Due to several shared-characters with chimpanzees, its closeness to ape divergence period, and due to its fossil incompletenes, the exact position of Ardipithecus in the fossil record is a subject of controversy.[18] Independent researcher such as Esteban E. Sarmiento of the Human Evolution Foundation in New Jersey, had systematically compared in 2010 the identifying characters of apes and human ancestral fossils in relation to Ardipithecus, and concluded that the comparison data is not sufficient to support an exclusive human lineage. Sarmiento noted that Ardipithecus does not share any characters exclusive to humans and some of its characters (those in the wrist and basicranium) suggest it diverged from the common human/African ape stock prior to the human, chimpanzee and gorilla divergence [19] His comparative (narrow allometry) study in 2011 on the molar and body segment lengths (which included living primates of similar body size) noted that some dimensions including short upper limbs, and metacarpals are reminiscent of humans, but other dimensions such as long toes and relative molar surface area are great ape-like. Sarmiento concluded that such length measures can change back and forth during evolution and are not very good indicators of relatedness. The Ardipithecus length measures, however, are good indicators of function and together with dental isotope data and the fauna and flora from the fossil site indicate Ardipithecus was mainly a terrestrial quadruped collecting a large portion of its food on the ground. Its arboreal behaviors would have been limited and suspension from branches solely from the upper limbs rare.[20]

However, some later studies still argue for its classification in the human lineage. Comparative study in 2013 on carbon and oxygen stable isotopes within modern and fossil tympanic, and a shortened, trapezoidal basioccipital element) in Ardipithecus are also found only Australopithecus and Homo clade.[23] Comparison of the tooth root morphology with those of Sahelanthropus tchadensis also indicated strong resemblance,[24] implying its correct inclusion in human lineage.

In a study that assumes the hominin status of Ardithecus ramidus, it has been argued the species represents a heterochronic alteration of the more general great ape body plan.[25] In this study the resemblance of the species' craniofacial moprhology with that of subadult chimpanzees is attributed to dissociation of craniofacial growth from brain growth and associated life history trajectories such as eruption of the first molar and age of first birth. Consequently, it is argued the species represents a unique ontogeny unlike any extant ape. The reduced growth in the sub-nasal alveolar region of the face, which houses the projecting canine complex in chimpanzees, suggests the species had rates of growth and reproductive biology unlike any living primate species. In this sense the species may show the first trend towards human social, parenting and sexual psychology. Consequently, the authors argue it is no longer tenable to extrapolate from chimpanzees in reconstructions of early hominin social and mating behaviour, providing further evidence against the so-called 'chimpanzee referential model'.[26] As the authors write when discussing the species unusual pattern of cranio-dental growth and the light it may throw on the origins of human sociality:

'The contrast [of humans] with chimpanzees is instructive, for when humans start developing broader social bonds after the permanent dentition begins erupting, at the same developmental milestone, chimpanzee facial projection increases. In other words, humans seem to have replaced craniofacial growth with an extended and intensified period of socio-emotional development. As A. ramidus no longer has an ontogeny that results in the development of a prognathic jaw with a C/P3 complex (which is one of the most important means by which males vie for status within the mating hierarchies of other primate species), young and sub-adult members of the species must have pursued other avenues by which to become reproductively successful members of the social group. The implication of these interspecific differences is that A. ramidus would have most likely had a period of infant and juvenile socialisation different from that of chimpanzees. Consequently, it is possible that in A.ramidus we see the first, albeit incipient trend toward human forms of child socialisation and social organisation'.[27]

It should be noted that this view has yet to be corroborated by more detailed studies of the ontogeny of A.ramidus. The study also provides support for Stephen Jay Gould's theory in Ontogeny and Phylogeny that the paedomorphic form of early hominin craniofacial morphology results from heterochronic dissociation of growth trajectories.

See also


  1. ^ a b
  2. ^
  3. ^ a b c d e f
  4. ^
  5. ^
  6. ^
  7. ^ a b c
  8. ^
  9. ^
  10. ^ a b
  11. ^ a b
  12. ^ a b
  13. ^
  14. ^
  15. ^
  16. ^
  17. ^
  18. ^
  19. ^
  20. ^
  21. ^
  22. ^
  23. ^
  24. ^
  25. ^ Clark, G.; Henneberg, M. 'The life history of Ardipithecus ramidus: a heterochronic model of sexual and social maturation'. Anthropological Review, Volume 78, Issue 2 (June 2015).
  26. ^ Sayers, K et al. 'Human Evolution and the Chimpanzee referential Model' Annual Review of Anthropology, Vol. 41: 119-138.
  27. ^ Clark, G.; Henneberg, M. pp126-7.

External links

  • specialArdipithecusScience Magazine: (requires free registration)
  • The Smithsonian Institution's Human Origins Program:
    • Ardipithecus kadabba
    • Ardipithecus ramidus
  • Ardipithecus ramidus at Archaeology info
  • Explore Ardipithecus at
  • - Science Journal ArticleArdipithecus ramidus
  • Discovering Ardi - Discovery Channel
This article was sourced from Creative Commons Attribution-ShareAlike License; additional terms may apply. World Heritage Encyclopedia content is assembled from numerous content providers, Open Access Publishing, and in compliance with The Fair Access to Science and Technology Research Act (FASTR), Wikimedia Foundation, Inc., Public Library of Science, The Encyclopedia of Life, Open Book Publishers (OBP), PubMed, U.S. National Library of Medicine, National Center for Biotechnology Information, U.S. National Library of Medicine, National Institutes of Health (NIH), U.S. Department of Health & Human Services, and, which sources content from all federal, state, local, tribal, and territorial government publication portals (.gov, .mil, .edu). Funding for and content contributors is made possible from the U.S. Congress, E-Government Act of 2002.
Crowd sourced content that is contributed to World Heritage Encyclopedia is peer reviewed and edited by our editorial staff to ensure quality scholarly research articles.
By using this site, you agree to the Terms of Use and Privacy Policy. World Heritage Encyclopedia™ is a registered trademark of the World Public Library Association, a non-profit organization.

Copyright © World Library Foundation. All rights reserved. eBooks from Project Gutenberg are sponsored by the World Library Foundation,
a 501c(4) Member's Support Non-Profit Organization, and is NOT affiliated with any governmental agency or department.