The Hominidae (anglicized hominids, also known as great apes[notes 1]) form a taxonomic family, including four extant genera: chimpanzees, gorillas, humans, and orangutans.
A number of known extinct genera are grouped with humans in the Homininae subfamily, others with orangutans in the Ponginae subfamily. The most recent common ancestor of the Hominidae lived roughly 14 million years ago, when the ancestors of the orangutans speciated from the ancestors of the other three genera. The ancestors of the Hominidae family had already speciated from those of the Hylobatidae family, perhaps 15-20 million years ago.
On July 19, 2001, a 7-million-year-old fossil skull, classified as Sahelanthropus tchadensis and nicknamed "ToumaΓ―", was discovered in Chad, Africa. It is possibly the earliest hominine fossil ever found. In addition to its age, ToumaΓ―, unlike the three- to four-million-year younger gracile australopithecine dubbed "Lucy", has a relatively flat face without the prominent snout seen on other pre-Homo hominids. There is some dispute over the importance of Sahelanthropus tchadensis. Professor Daniel E. Lieberman, has suggested this previously unknown species may be a direct ancestor of modern humans (or at least closely related to a direct ancestor). Others such as Professor Bernard Wood argue that one fossil is not enough to make such a claim because appearance is an unreliable method of determining evolutionary relations.
Researchers such as David Reich, Eric Lander, et al argue that the species associated with the divergence between chimpanzees and proto-humans swapped genes over time through interbreeding, before finally separating. It is generally believed that the Pan/Homo separation occurred about 6 â 8 million years ago, but the molecular clock (a method of calculating evolution based on the rate at which genes mutate) suggests the genera division 4.6 â 6.2 million years ago. Previous studies looked at average genetic differences between human and chimp. The new study compares the ages of key sequences of genes of modern humans and modern chimps. Some sequences are younger than others, indicating that chimps and humans gradually separated over a period of 4 million years. The youngest human chromosome is the X chromosome which is about 1.2 million years more recent than the 22 autosomes. The X sex chromosome is known to be vulnerable to selective pressure. Its age suggests there was an initial division between the two species, followed by gradual divergence and interbreeding that resulted in younger genes, and then a final separation.
An alternative minority viewpoint is that Homo diverged from a common ancestor with Pongo perhaps as early as 13 million years ago while Pan is more closely related to Gorilla. This alternative is supported by characteristics uniquely shared between humans and orangutans such as dental structure, thick enamel, shoulder blade structure, thick posterior palate, single incisive foramen, high estriol production, and beard and mustache. There are at least 28 well corroborated such features compared with perhaps as little as one unique feature shared between humans and chimpanzees. It is widely believed that these physical features are misleading, but an alternative possibility is that orangutans have undergone more genetic change than humans and African apes have since their divergence from the common ancestor. If this had happened, then the apparent genetic similarity between humans and chimpanzees would not necessarily be due to a close evolutionary relationship. This theory has been proposed as an explanation as to why early hominids such as the australopiths not only look more like orangutans than either African ape, but also share characters unique to orangutans and their close fossil relatives such as a thickened posteror palate and anterior zygomatic roots.
 Taxonomic history
Humans, as depicted on a Pioneer plaque
, are one of the four extant hominid genera
The classification of the great apes has been revised several times in the last few decades. Originally, Hominidae was the name given to humans and their extinct relatives, with the other great apes being placed in a separate family, the Pongidae. However, that definition makes Pongidae paraphyletic because at least one great ape species appears to be more closely related to humans than other great apes. Most taxonomists nowadays encourage monophyletic groups so this would require the use of Pongidae to be restricted to one of the great ape groups only. Thus many biologists consider Hominidae to include Pongidae as the subfamily Ponginae, or restrict the latter to the orangutans and their extinct relatives like Gigantopithecus. The taxonomy shown here follows the monophyletic groupings according to the two theories of human and great ape relationships.
Especially close human relatives form a subfamily, the Homininae. Some researchers go so far as to include chimpanzees and gorillas in the genus Homo along with humans, but it is more commonly accepted to describe the relationships as shown here. Alternatively, those fossil relatives that are more closely related to humans than the nearest living great ape species represent members of Hominidae without necessarily assigning subfamily or tribal categories. If the orangutan is the closest living relative of humans, there would be a sister group relationship between Hominidae and Pongidae, with the African apes comprising a separate family (Panidae) according to the morphological evidence.
Many extinct hominids have been studied to help understand the relationship between modern humans and the other extant hominids. Some of the extinct members of this family (as defined to encompass humans and chimpanzees) include Gigantopithecus, Orrorin, Ardipithecus, Kenyanthropus, and the australopithecines Australopithecus and Paranthropus. In the orangutan model of human origin the Hominidae would include the australopiths, and possibly Orrorin and Kenyanthropus, but not Ardipithecus and Sahelanthropus that lack features necessary to provide strong corroboration as hominids.
The exact criteria for membership in the Homininae under the chimpanzee theory of human origins are not clear, but the subfamily generally includes those species that share more than 97% of their DNA with the modern human genome, and exhibit a capacity for language or for simple cultures beyond the family or band. The theory of mind including such faculties as mental state attribution, empathy and even empathetic deception is a controversial criterion distinguishing the adult human alone among the hominids. Humans acquire this capacity at about four and a half years of age, whereas it has neither been proven nor disproven that gorillas and chimpanzees develop a theory of mind. This is also the case for some New World monkeys outside the family of great apes, as, for example, the capuchin monkeys.
However, without the ability to test whether early members of the Homininae (such as Homo erectus, Homo neanderthalensis, or even the australopithecines) had a theory of mind, it is difficult to ignore similarities seen in their living cousins. Orangutans have also been shown to have culture comparable to that of chimpanzees and some say that the orangutan may also satisfy these criteria. These scientific debates take on political significance for advocates of Great Ape personhood.
Skulls of an orangutan and a gorilla
The seven living species of great ape are classified in four genera. The following classification is commonly accepted:
- Family Hominidae: humans and other great apes; extinct genera and species excluded.
- Subfamily Ponginae
- Subfamily Homininae
- Tribe Gorillini
- Tribe Hominini
- Genus Pan
- Common Chimpanzee, Pan troglodytes
- Central Chimpanzee, Pan troglodytes troglodytes
- West African Chimpanzee, Pan troglodytes verus
- Nigerian Chimpanzee, Pan troglodytes vellerosus
- Eastern Chimpanzee, Pan troglodytes schweinfurthii
- Bonobo (Pygmy Chimpanzee), Pan paniscus
- Genus Homo
- Human, Homo sapiens sapiens
In addition to the extant species and subspecies above, archaeologists, paleontologists, and anthropologists have discovered numerous extinct great ape species. The list below are some of the genera of those discoveries.
 Physical description
The great apes are large, tailless primates, with the smallest living species being the Bonobo at 30 â 40 kilograms in weight, and the largest being the gorillas, with males weighing 140 â 180 kilograms. In all great apes, the males are, on average, larger and stronger than the females, although the degree of sexual dimorphism varies greatly among species. Although most living species are predominantly quadrupedal, they are all able to use their hands for gathering food or nesting materials, and, in some cases, for tool use.
Most species are omnivorous, but fruit is the preferred food among all but humans. Chimpanzees and orangutans primarily eat fruit. When gorillas run short of fruit at certain times of the year or in certain regions, they resort to eating shoots and leaves, often of bamboo, a type of grass. Gorillas have extreme adaptations for chewing and digesting such low-quality forage, but they still prefer fruit when it is available, often going miles out of their way to find especially preferred fruits. In contrast, humans consume a large proportion of highly processed, low fiber foods, unusual proportions of grains (only very recently) and vertebrate flesh, as well as a wide variety of other foodstuffs. The teeth are similar to those of the Old World monkeys and gibbons, although they are especially large in gorillas. The dental formula is: Human teeth and jaws are markedly smaller for their size than those of other apes. Some scholars have speculated that humans' small teeth and jaws are adaptations to eating cooked food for more than a million years, but this idea remains controversial.
Gestation in great apes lasts 8â9 months, and results in the birth of a single offspring, or, rarely, twins. The young are born helpless, and the mother must care for them for long periods of time. Compared with most other mammals, great apes have a remarkably long adolescence, not being weaned for several years, and not becoming fully mature for 8â13 years in most species (longer in humans). As a result, females typically give birth only once every few years. There is no distinct breeding season.
Gorillas and chimpanzees live in family groups of around five to ten individuals, although much larger groups are sometimes noted. Chimpanzees live in larger groups that break up into smaller groups when fruit becomes less available. When small groups of female chimpanzees go off in separate directions to forage for fruit, the dominant male(s) can no longer control them and the females often mate with other subordinate males, whether by choice or not. In contrast, groups of gorillas stay together regardless of the availability of fruit. When fruit is hard to find, they resort to eating leaves and shoots. Because gorilla groups stay together, the male is able to monopolize the females in his group. This fact is related to gorillas' greater sexual dimorphism than chimpanzees'. In both chimpanzees and gorillas, the groups include at least one dominant male, and females leave the group at maturity. By contrast, orangutans are generally solitary. The social structure of humans is complex and highly variable. DNA studies suggest that, as in other apes, female humans are the sex that leaves the group at maturity, or at least has done so over evolutionary time.
 Legal status
Due to the close genetic relationship between humans and other great apes, certain animal rights organizations, such as the Great Ape Project, argue that non-human great apes are persons and should be given basic human rights. Some countries have instituted a research ban to protect great apes from any kind of scientific testing.
On 25 June 2008, the Spanish parliament supported a new law that would make "keeping apes for circuses, television commercials or filming" illegal.
 See also
- ^ "Great ape" is a common name rather than a taxonomic label and there are differences in usage. Subtly, it may seem to exclude human beings ("humans and the great apes") or to include them ("humans and non-human great apes").
- ^ a b c d e Groves, C. (2005). Wilson, D. E., & Reeder, D. M, eds. ed. Mammal Species of the World (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 181â184. ISBN 0-801-88221-4. http://www.bucknell.edu/msw3/browse.asp?id=12100786.
- ^ Andrew Hill & Steven Ward (1988). "Origin of the Hominidae: The Record of African Large Hominoid Evolution Between 14 My and 4 My". Yearbook of Physical Anthropology 31 (59): 49â83. doi:10.1002/ajpa.1330310505
- ^ a b Dawkins R (2004) The Ancestor's Tale.
- ^ http://www.timetree.org/time_e_query.php?taxon_a=Hominidae&taxon_b=Hylobatidae
- ^ Brunet, Michel; Guy, Pilbeam, Taisso, et al (11 July 2002). "A new hominid from the Upper Miocene of Chad, Central Africa". Nature (Nature Publishing Group) 418 (418): 145â151. doi:10.1038/nature00879. ISSN 0028-0836. PMID 12110880. http://www.nature.com/nature/journal/v418/n6894/full/nature00879.html. Retrieved 3 May 2010.
- ^ Whitfield, John (11 July 2002). "Oldest member of human family found". Nature. http://www.nature.com/news/2002/020711/full/news020708-12.html. Retrieved 3 May 2010.
- ^ a b Patterson, Nick; Richter, Daniel J.; Gnerre, Sante; Lander, Eric S.; Reich, David (2006). "Genetic evidence for complex speciation of humans and chimpanzees". Nature 441 (7097): 1103â1108. doi:10.1038/nature04789. PMID 16710306. http://www.nature.com/nature/journal/v441/n7097/abs/nature04789.html
- ^ Jay Kelley (1994). "Evolution of apes". in Steve Jones, Robert Martin & David Pilbeam. Cambridge Encyclopedia of Human Evolution. Cambridge, U.K: Cambridge University Press. pp. 223â230. ISBN 0-521-46786-1 (see pp229-230 for summary of DNA dating)
- ^ Purvis, Andy (29 June 1995). "A composite estimate of primate phylogeny.". Philosophical Transactions of the Royal Society B: Biological Sciences 348 (1326): 405â421. doi:10.1098/rstb.1995.0078. PMID 7480112
- ^ Goodman, M., Porter, C.A., Czelusniak, J., Page, S.L., Schneider, H., Shoshani, J., Gunnell, G., & Groves, C.P (June 1998). "Toward a phylogenetic classification of primates based on DNA evidence complemented by fossil evidence". Molecular Phylogenetics and Evolution 9 (3): 585â598. doi:10.1006/mpev.1998.0495. PMID 9668008. http://homopan.wayne.edu/Research/MPE_toward_(1998).pdf
- ^ Feng-Chi Chen1, & Wen-Hsiung Li (February 2004). "Genomic Divergences between Humans and Other Hominoids and the Effective Population Size of the Common Ancestor of Humans and Chimpanzees". The American Journal of Human Genetics 68 (2): 444â456. doi:10.1086/318206. PMID 11170892. PMC 1235277. https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1235277/
- ^ Schaffner, Stephen F (January 2004). "The X Chromosome in Population Genetics". Nature Reviews Genetics 5 (1): 43â54. doi:10.1038/nrg1247. PMID 14708015. http://www.broadinstitute.org/~sfs/nrg_Xchrom.pdf
- ^ Schwartz, J.H. (2005) The Red Ape. Westview Press.
- ^ a b Grehan, J.R. (2006) Mona Lisa Smile: The morphological enigma of human and great ape evolution. Anatomical Record 289B: 139-157.
- ^ Schwartz, J.H. (2004) Barking up the wrong ape - australopiths and the quest for chimpanzee characters in hominid fossils. Colloquim Antropologicum Supplement 28: 87-100.
- ^ Pickrell, John (2003-05-20). "Chimps Belong on Human Branch of Family Tree, Study Says". National Geographic. http://news.nationalgeographic.com/news/2003/05/0520_030520_chimpanzees.html. Retrieved 2007-08-04.
- ^ Relationship Humans-Gorillas.
- ^ Watson, E. E. et al. (2001). "Homo genus: a review of the classification of humans and the great apes". in eds. Tobias, P. V. et al.. Humanity from African Naissance to Coming Millennia. Florence: Firenze Univ. Press. pp. Pp. 311â323.
- ^ Schwartz, J.H. (1986) Primate systematics and a classification of the order. Comparative primate biology volume 1: Systematics, evolution, and anatomy (ed. by D.R. Swindler, and J. Erwin), pp. 1-41, Alan R. Liss, New York.
- ^ Schwartz, J.H. (2004b) Issues in hominid systematics. Zona ArqueologΓa 4, 360-371.
- ^ Heyes, C. M. (1998). "Theory of Mind in Nonhuman Primates". Behavioral and Brain Sciences 21. doi:10.1017/S0140525X98000703. bbs00000546. http://www.bbsonline.org/documents/a/00/00/05/46/index.html.
- ^ Van Schaik, C.P. et al. (2003) Orangutan cultures and the evolution of material culture. Science 299: 102-105.
- ^ a b Harcourt, A.H., MacKinnon, J. & Wrangham, R.W. (1984). Macdonald, D.. ed. The Encyclopedia of Mammals. New York: Facts on File. pp. 422â439. ISBN 0-87196-871-1.
- ^ Brace, C. Loring; Mahler, Paul Emil (1971). "Post-Pleistocene changes in the human dentition". American Journal of Physical Anthropology 34 (2): 191â203. doi:10.1002/ajpa.1330340205. PMID 5572603. http://deepblue.lib.umich.edu/handle/2027.42/37509
- ^ Richard Wrangham (2007). "Chapter 12: The Cooking Enigma". in Charles Pasternak (ed.). What Makes Us Human?. Oxford: Oneworld Press. ISBN 978-1851-68519-6.
- ^ "Spanish parliament to extend rights to apes". Reuters. 2008-06-25. http://www.reuters.com/article/scienceNews/idUSL256586320080625. Retrieved 2008-07-11.
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