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]
Hominidae[1] | |
---|---|
The eight extant hominid species, one row per genus (humans,chimpanzees,gorillas,orangutans) | |
Scientific 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 | |
|
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 and other apes 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
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
Terminology
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 | |
hominoids, apes |
Extant and fossil relatives of humans
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.[clarification needed][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.
Description
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 relative to body size compared to those of other apes. This may be an adaptation not only to the extensive use of tools, which has supplanted the role of jaws in hunting and fighting, but also to eating cooked food since the end of thePleistocene.[22][23]
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.[24]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.[25]
All species areomnivorous,[26]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,[27]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.[24]
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
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.[28]On 8 September 2010, theEuropean Unionbanned the testing of great apes.[29]
Conservation
The following table lists the estimated number of great ape individuals living outside zoos.
Species | Estimated number |
Conservation status |
Refs |
---|---|---|---|
Bornean orangutan | 104,700 | Critically endangered | [30] |
Sumatran orangutan | 6,667 | Critically endangered | [31] |
Tapanuli orangutan | 800 | Critically endangered | [32] |
Western gorilla | 200,000 | Critically endangered | [33] |
Eastern gorilla | <5,000 | Critically endangered | [34] |
Chimpanzee | 200,000 | Endangered | [35][36] |
Bonobo | 10,000 | Endangered | [35] |
Human | 8,040,640,000 | N/A | [37] |
Phylogeny
This sectionneeds additional citations forverification.(July 2019) |
Below is a cladogram with extinct species.[38][39][40][failed verification]It is indicated approximately how many million years ago (Mya) the clades diverged into newer clades.[41]
Hominidae (18) |
| |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Extant
There are eight living species of great ape which are classified in fourgenera.The following classification is commonly accepted:[1]
- Family Hominidae:humans and other great apes; extinct genera and species excluded[1]
- SubfamilyPonginae
- TribePongini
- GenusPongo
- Bornean orangutan,Pongo pygmaeus
- Northwest Bornean orangutan,Pongo pygmaeus pygmaeus
- Northeast Bornean orangutan,Pongo pygmaeus morio
- Central Bornean orangutan,Pongo pygmaeus wurmbii
- Sumatran orangutan,Pongo abelii
- Tapanuli orangutan,Pongo tapanuliensis[42]
- Bornean orangutan,Pongo pygmaeus
- GenusPongo
- TribePongini
- SubfamilyHomininae
- TribeGorillini
- GenusGorilla
- Western gorilla,Gorilla gorilla
- Western lowland gorilla,Gorilla gorilla gorilla
- Cross River gorilla,Gorilla gorilla diehli
- Eastern gorilla,Gorilla beringei
- Mountain gorilla,Gorilla beringei beringei
- Eastern lowland gorilla,Gorilla beringei graueri
- Western gorilla,Gorilla gorilla
- GenusGorilla
- TribeHominini
- SubtribePanina
- GenusPan
- Chimpanzee,Pan troglodytes
- Central chimpanzee,Pan troglodytes troglodytes
- Western chimpanzee,Pan troglodytes verus
- Nigeria-Cameroon chimpanzee,Pan troglodytes ellioti
- Eastern chimpanzee,Pan troglodytes schweinfurthii
- Bonobo,Pan paniscus
- Chimpanzee,Pan troglodytes
- GenusPan
- SubtribeHominina
- GenusHomo
- Human,Homo sapiens
- Anatomically modern human,Homo sapiens sapiens
- Human,Homo sapiens
- GenusHomo
- SubtribePanina
- TribeGorillini
- SubfamilyPonginae
Fossil
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.[43]
Family Hominidae
- SubfamilyPonginae[44]
- TribeLufengpithecini†
- Lufengpithecus
- Lufengpithecus lufengensis
- Lufengpithecus keiyuanensis
- Lufengpithecus hudienensis
- Meganthropus
- Meganthropus palaeojavanicus
- Lufengpithecus
- Tribe Sivapithecini†
- Ankarapithecus
- Ankarapithecus meteai
- Sivapithecus
- Sivapithecus brevirostris
- Sivapithecus punjabicus
- Sivapithecus parvada
- Sivapithecus sivalensis
- Sivapithecus indicus
- Gigantopithecus
- Gigantopithecus bilaspurensis
- Gigantopithecus blacki
- Gigantopithecus giganteus
- Ankarapithecus
- Tribe Pongini
- TribeLufengpithecini†
- SubfamilyHomininae[45][46]
- TribeDryopithecini†
- Kenyapithecus(placement disputed)
- Kenyapithecus wickeri
- Danuvius
- Danuvius guggenmosi
- Pierolapithecus(placement disputed)
- Pierolapithecus catalaunicus
- Ouranopithecus
- Ouranopithecus macedoniensis
- Otavipithecus
- Otavipithecus namibiensis
- Morotopithecus(placement disputed)
- Morotopithecus bishopi
- Oreopithecus(placement disputed)
- Oreopithecus bambolii
- Nakalipithecus
- Nakalipithecus nakayamai
- Anoiapithecus
- Anoiapithecus brevirostris
- Hispanopithecus(placement disputed)
- Hispanopithecus laietanus
- Hispanopithecus crusafonti
- Dryopithecus
- Rudapithecus(placement disputed)
- Rudapithecus hungaricus
- Samburupithecus
- Samburupithecus kiptalami
- Graecopithecus†[47]
- Graecopithecus freybergi
- Kenyapithecus(placement disputed)
- TribeGorillini
- Chororapithecus†(placement debated)
- Chororapithecus abyssinicus
- Chororapithecus†(placement debated)
- TribeHominini
- SubtribePanina
- SubtribeHominina
- Sahelanthropus†
- Sahelanthropus tchadensis
- Orrorin†
- Orrorin tugenensis
- Orrorin praegens
- Ardipithecus†
- Kenyanthropus†
- Kenyanthropus platyops
- Australopithecus†
- Paranthropus†
- Homo– close relatives of modern humans
- Homo gautengensis†(probableH. habilisspecimens)
- Homo rudolfensis†(membership inHomouncertain)
- Homo habilis†(membership inHomouncertain)
- Homo naledi†(membership inHomouncertain)
- Dmanisi Man,Homo georgicus†(probable early subspecies ofHomo erectus)
- Homo ergaster†(AfricanHomo erectus)
- Homo erectus†
- Homo erectus bilzingslebenensis†
- Java Man,Homo erectus erectus†
- Lantian Man,Homo erectus lantianensis†
- Nanjing Man,Homo erectus nankinensis†
- Peking Man,Homo erectus pekinensis†
- Solo Man,Homo erectus soloensis†(possible separate species)
- Tautavel Man,Homo erectus tautavelensis†
- Yuanmou Man,Homo erectus yuanmouensis†
- Flores Manor Hobbit,Homo floresiensis†(membership inHomouncertain)
- Homo luzonensis†(membership inHomouncertain)
- Homo antecessor†
- Homo heidelbergensis†
- Homo cepranensis†(probableH. heidelbergensisspecimens)
- Homo helmei†(probable earlyH. sapiensspecimens)
- Homo tsaichangensis†(thought by some to be a subspecies ofH. erectusor a Denisovan; unlikely to be separate species)
- Denisovans(scientific name not yet assigned)†
- Neanderthal,Homo neanderthalensis†
- Homo rhodesiensis†(probable lateH. heidelbergensisspecimens)
- Modern human,Homo sapiens(sometimes calledHomo sapiens sapiens)
- Sahelanthropus†
- TribeDryopithecini†
See also
- Bili ape
- Dawn of Humanity(2015 PBS film)
- Great ape language
- Planet of the Apes franchise
- Great Ape Project
- Great ape research ban
- Great Apes Survival Partnership
- International Primate Day
- Kinshasa Declaration on Great Apes
- List of human evolution fossils
- List of individual apes
- Monkeys and apes in space
- Oldest hominids
- Prehistoric Autopsy(2012 BBC documentary)
- Primate cognition
- The Mind of an Ape
- Timeline of human evolution
Notes
- ^"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][better source needed]and "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives."[4][better source needed]
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External links
- The Animal Legal and Historical Center at Michigan State University College of Law,Great Apes and the Law (archived 13 April 2011)
- NPR News: Toumaï the Human Ancestor
- Hominid SpeciesatTalkOrigins Archive
- For more details on Hominid species, including photos of fossil hominids(archived 30 April 2013)
- Scientific Americanmagazine (April 2006 Issue) Why Are Some Animals So Smart?(archived 14 October 2007)
- A new mediterranean hominoid-hominid link discovered,Anoiapithecus brevirostris,"Lluc":A unique Middle Miocene European hominoid and the origins of the great ape and human cladeLink to graphical reconstruction
- Human Timeline (Interactive)–Smithsonian,National Museum of Natural History(August 2016).