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Hominidae

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(Redirected fromGreat apes)

For an explanation of very similar terms, seeHomininaeandHominini.
Not to be confused withHominoidea.
"Great apes" and "Hominid" redirect here. For other uses, seeGreat apes (disambiguation)andHominid (disambiguation).
"Ape-men" redirects here. For other uses, seeApeman (disambiguation).

Hominidae[1]
Temporal range:Miocene–present,17–0Ma
The eight extant hominid species, one row per genus (humans,chimpanzees,gorillas,orangutans)
Scientific classificationEdit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Parvorder: Catarrhini
Superfamily: Hominoidea
Family: Hominidae
Gray,1825[2]
Type genus
Homo
Linnaeus, 1758
Subfamilies

sister:Hylobatidae

Distribution of great ape species
Synonyms
  • PongidaeElliot, 1913
  • GorillidaeFrechkop, 1943
  • PanidaeCiochon, 1983

TheHominidae(/hɒˈmɪnɪdiː/), whose members are known as thegreat apes[note 1]orhominids(/ˈhɒmɪnɪdz/), are a taxonomicfamilyofprimatesthat includes eightextantspecies in fourgenera:Pongo(theBornean,SumatranandTapanuli orangutan);Gorilla(theeasternandwestern gorilla);Pan(thechimpanzeeand thebonobo); andHomo,of which onlymodern humans (Homo sapiens)remain.[1]

Numerous revisions in classifying the great apes have caused the use of the termhominidto change over time. The original meaning of "hominid" referred only to humans (Homo) and their closest extinct relatives. However, by the 1990s humans, apes, and their ancestors were considered to be "hominids".

The earlier restrictive meaning has now been largely assumed by the termhominin,which comprises all members of the human clade after the split from the chimpanzees (Pan). The current meaning of "hominid" includes all the great apes including humans. Usage still varies, however, and some scientists and laypersons still use "hominid" in the original restrictive sense; the scholarly literature generally shows the traditional usage until the turn of the 21st century.[5]

Within the taxon Hominidae, a number of extant and extinct genera are grouped with the humans, chimpanzees, and gorillas in the subfamilyHomininae;others with orangutans in the subfamilyPonginae(seeclassification graphicbelow). Themost recent common ancestorof all Hominidae lived roughly 14 million years ago,[6]when the ancestors of the orangutans speciated from the ancestral line of the other three genera.[7]Those ancestors of the family Hominidae had already speciated from the familyHylobatidae(thegibbons), perhaps 15 to 20 million years ago.[7][8]

Due to the close genetic relationship between humans and the other great apes, certainanimal rightsorganizations, such as theGreat Ape Project,argue that nonhumangreat apes are personsand should be given basichuman rights.Twenty-nine countries have institutedresearch bansto protect great apes from any kind of scientific testing.[9]

Evolution

See also:Human evolution
Sumatran orangutan (Pongo abelii)

In the earlyMiocene,about 22 million years ago, there were many species oftree-adaptedprimitivecatarrhinesfrom East Africa; the variety suggests a long history of prior diversification. Fossils from 20 million years ago include fragments attributed toVictoriapithecus,the earliest Old World monkey. Among the genera thought to be in the ape lineage leading up to 13 million years ago areProconsul,Rangwapithecus,Dendropithecus,Limnopithecus,Nacholapithecus,Equatorius,Nyanzapithecus,Afropithecus,Heliopithecus,andKenyapithecus,all from East Africa.

At sites far distant from East Africa, the presence of other generalized non-cercopithecids,that is, non-monkey primates, of middle Miocene age—Otavipithecusfrom cave deposits in Namibia, andPierolapithecusandDryopithecusfrom France, Spain and Austria—is further evidence of a wide diversity of ancestral ape forms across Africa and the Mediterranean basin during the relatively warm and equable climatic regimes of the early and middle Miocene. The most recent of these far-flung Miocene apes (hominoids) isOreopithecus,from the fossil-rich coal beds in northern Italy and dated to 9 million years ago.

Molecular evidence indicates that the lineage of gibbons (family Hylobatidae), the "lesser apes", diverged from that of the great apes some 18–12 million years ago, and that of orangutans (subfamily Ponginae) diverged from the other great apes at about 12 million years. There are no fossils that clearly document the ancestry of gibbons, which may have originated in a still-unknown South East Asian hominoid population; but fossil proto-orangutans, dated to around 10 million years ago, may be represented bySivapithecusfrom India andGriphopithecusfrom Turkey.[10]Species close to the last common ancestor of gorillas, chimpanzees and humans may be represented byNakalipithecusfossils found in Kenya andOuranopithecusfossils found inGreece.Molecular evidence suggests that between 8 and 4 million years ago, first the gorillas (genusGorilla), and then the chimpanzees (genusPan) split off from the line leading to humans. HumanDNAis approximately 98.4% identical to that of chimpanzees when comparing single nucleotide polymorphisms (seehuman evolutionary genetics).[11]The fossil record, however, of gorillas and chimpanzees is limited; both poor preservation—rain forest soils tend to be acidic and dissolve bone—andsampling biasprobably contribute most to this problem.

Otherhomininsprobably adapted to the drier environments outside the African equatorial belt; and there they encountered antelope, hyenas, elephants and other forms becoming adapted to surviving in the East Africansavannas,particularly the regions of theSaheland theSerengeti.The wet equatorial belt contracted after about 8 million years ago, and there is very little fossil evidence for the divergence of the hominin lineage from that of gorillas and chimpanzees—which split was thought to have occurred around that time. The earliest fossils argued by some to belong to the human lineage areSahelanthropus tchadensis(7 Ma) andOrrorin tugenensis(6 Ma), followed byArdipithecus(5.5–4.4 Ma), with speciesAr. kadabbaandAr. ramidus.

Taxonomy

Further information:Human taxonomy

Terminology

Humans are one of the fourextanthominid genera.

The classification of the great apes has beenrevised several timesin the last few decades; these revisions have led to a varied use of the word "hominid" over time. The original meaning of the term referred to only humans and their closest relatives—what is now the modern meaning of the term "hominin".The meaning of thetaxonHominidae changed gradually, leading to a modern usage of "hominid" that includes all the great apes including humans.

A number of very similar words apply to related classifications:

  • Ahominoid,sometimes called anape,is a member of the superfamilyHominoidea:extant members are the gibbons (lesser apes,family Hylobatidae) and the hominids.
  • Ahominidis a member of the family Hominidae, the great apes: orangutans, gorillas, chimpanzees and humans.
  • Ahominineis a member of the subfamilyHomininae:gorillas, chimpanzees, and humans (excludes orangutans).
  • Ahomininis a member of the tribeHominini:chimpanzees and humans.[12]
  • Ahomininan,following a suggestion by Wood and Richmond (2000), would be a member of the subtribeHomininaof the tribeHominini:that is, modern humans and their closest relatives, includingAustralopithecina,but excluding chimpanzees.[13][14]
  • Ahumanis a member of the genusHomo,of whichHomo sapiensis the only extant species, and within thatHomo sapiens sapiensis the only survivingsubspecies.

A cladogram indicating common names (cf.more detailed cladogram below):

Hominoidea

Hylobatidae
gibbons

Hominidae
hominids, great apes
hominoids, apes

Extant and fossil relatives of humans

Afossilhominid exhibit atThe Museum of Osteology,Oklahoma City,Oklahoma

Hominidae was originally the name given to the family of humans and their (extinct) close relatives, with the other great apes (that is, the orangutans, gorillas and chimpanzees) all being placed in a separate family, thePongidae.However, that definition eventually made Pongidaeparaphyleticbecause at least one great ape species (the chimpanzees) proved to be more closely related to humans than to other great apes. Most taxonomists today encouragemonophyleticgroups—this would require, in this case, the use of Pongidae to be restricted to just one closely related grouping. Thus, manybiologistsnow assignPongo(as thesubfamilyPonginae) to the family Hominidae. The taxonomy shown here follows the monophyletic groupings according to the modern understanding of human and great ape relationships.

Humans and close relatives including the tribesHomininiandGorilliniform the subfamilyHomininae(see classification graphic below). (A few researchers go so far as to refer the chimpanzees and the gorillas to the genusHomoalong with humans.)[15][16][17]But, those fossil relatives more closely related to humans than the chimpanzees represent the especially close members of the human family, and without necessarily assigning subfamily or tribal categories.[18]

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 includeGigantopithecus,Orrorin,Ardipithecus,Kenyanthropus,and theaustralopithecinesAustralopithecusandParanthropus.[19]

The exact criteria for membership in the tribe Hominini under the current understanding of human origins are not clear, but the taxon generally includes thosespeciesthat share more than 97% of theirDNAwith the modern humangenome,and exhibit a capacity forlanguageor for simpleculturesbeyond their 'local family' or band. Thetheory of mindconcept—including such faculties as empathy, attribution of mental state, and even empathetic deception—is a controversial criterion; it distinguishes the adult human alone among the hominids. Humans acquire this capacity after about four years of age, whereas it has not been proven (nor has it been disproven) that gorillas or chimpanzees ever develop a theory of mind.[20]This is also the case for someNew World monkeysoutside the family of great apes, as, for example, thecapuchin monkeys.

However, even without the ability to test whether early members of the Hominini (such asHomo 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 shown the development of culture comparable to that of chimpanzees,[21]and some[who?]say the orangutan may also satisfy those criteria for thetheory of mindconcept. These scientific debates take on political significance for advocates ofgreat ape personhood.

Phylogeny

Below is a cladogram with extinct species.[22][23][24][failed verification]It is indicated approximately how many million years ago (Mya) the clades diverged into newer clades.[25]

Hominidae (18)


Taxonomy ofHominoidea(emphasis on family Hominidae): After an initial separation from the main line by the Hylobatidae (gibbons) some 18 million years ago, the line ofPonginaebroke away, leading to the orangutan; later, the Homininae split into the tribesHominini(led to humans and chimpanzees) andGorillini(led to gorillas).

Extant

See also:List of hominoids

There are eight living species of great ape which are classified in fourgenera.The following classification is commonly accepted:[1]

Fossil

Replica ofthe skull sometimes known as "Nutcracker Man",found byMary Leakey

In addition to the extant species and subspecies,archaeologists,paleontologists,andanthropologistshave discovered and classified numerous extinct great ape species as below, based on the taxonomy shown.[27]

Family Hominidae

Description

Gorilla

The great apes are tailless primates, with the smallest living species being the bonobo at 30 to 40 kilograms (66 to 88 lb) in weight, and the largest being the eastern gorillas, with males weighing 140 to 180 kilograms (310 to 400 lb). In all great apes, the males are, on average, larger and stronger than the females, although the degree ofsexual dimorphismvaries greatly among species. Hominid teeth are similar to those of theOld World monkeysand gibbons, although they are especially large in gorillas. Thedental formulais2.1.2.32.1.2.3.Human teeth and jaws are markedly smaller for their size than those of other apes, which may be an adaptation to not only having supplanted with extensive tool use the role of jaws in hunting and fighting, but also eating cooked food since the end of thePleistocene.[32][33]

Behavior

Although most living species are predominantlyquadrupedal,they are all able to use their hands for gathering food or nesting materials, and, in some cases, for tool use.[34]They build complex sleeping platforms, also called nests, in trees to sleep in at night, but chimpanzees and gorillas also build terrestrial nests, and gorillas can also sleep on the bare ground.[35]

All species areomnivorous,[36]although 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 ofbamboo,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. Humans, since theNeolithic revolution,have consumed mostlycerealsand otherstarchyfoods, including increasingly highlyprocessed foods,as well as many otherdomesticated plants(including fruits) andmeat.

Gestationin great apes lasts 8–9 months, and results in the birth of a single offspring, or, rarely, twins. The young are born helpless, and require care for long periods of time. Compared with most other mammals, great apes have a remarkably long adolescence, not beingweanedfor several years,[37]and not becoming fully mature for eight to thirteen years in most species (longer in orangutans and humans). As a result, females typically give birth only once every few years. There is no distinct breeding season.[34]

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 males can no longer control them and the females often mate with other subordinate males. 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.

This fact is related to gorillas' greater sexual dimorphism relative to that of chimpanzees; that is, the difference in size between male and female gorillas is much greater than that between male and female chimpanzees. This enables gorilla males to physically dominate female gorillas more easily. In both chimpanzees and gorillas, the groups include at least one dominant male, and young males leave the group at maturity.

Legal status

Main articles:Great ape personhood,Great Ape Project,andCountries banning non-human ape experimentation

Due to the close genetic relationship between humans and the other great apes, certainanimal rightsorganizations, such as theGreat Ape Project,argue that nonhumangreat apes are personsand, per theDeclaration on Great Apes,should be given basichuman rights.In 1999, New Zealand was the first country to ban any great ape experimentation, and now 29 countries have currently instituted aresearch banto 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.[38]On 8 September 2010, theEuropean Unionbanned the testing of great apes.[39]

Conservation

The following table lists the estimated number of great ape individuals living outside zoos.

Species Estimated
number 		Conservation
status 		Refs
Bornean orangutan 61,234 Critically endangered [40]
Sumatran orangutan 6,667 Critically endangered [41]
Tapanuli orangutan 800 Critically endangered [42]
Western gorilla 200,000 Critically endangered [43]
Eastern gorilla 6,000 Critically endangered [43]
Chimpanzee 200,000 Endangered [44][45]
Bonobo 10,000 Endangered [44]
Human 8,040,640,000 N/A [46]

See also

Notes

  1. ^"Great ape" is a common name rather than a taxonomic label, and there are differences in usage, even by the same author. The term may or may not include humans, as when Dawkins writes "Long before people thought in terms of evolution... great apes were often confused with humans"[3]and "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives."[4]

References

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External links

Wikimedia Commons has media related toHominidae.
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The WikibookDichotomous Keyhas a page on the topic of:Hominidae


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