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| {{dablink|This article is about the biological superfamily. For a discussion of the inclusion of humans as apes, see ] below. For other uses, see ].}} | |||
| {{Taxobox | |||
| | name = Apes<ref name=MSW3>{{MSW3 Groves|pages=178-184}}</ref> | |||
| | fossil_range = Late ] - Recent | |||
| | image = Weisshandgibbon tierpark berlin.jpg | |||
| | image_width = 250px | |||
| | image_caption = ] (''Hylobates lar'') | |||
| | regnum = ]ia | |||
| | phylum = ] | |||
| | classis = ]ia | |||
| | ordo = ]s | |||
| | subordo = ] | |||
| | parvordo = ] | |||
| | superfamilia = '''Hominoidea''' | |||
| | superfamilia_authority = ], 1825 | |||
| | subdivision_ranks = ] | |||
| | subdivision = | |||
| ]<br> | |||
| ]<br> | |||
| †]<br> | |||
| †]<br> | |||
| †]</i> | |||
| }} | |||
| '''Apes''' are the members of the '''Hominoidea''' ] of ]s, which includes ]s. Under the current classification system there are two ] of '''hominoids''': | |||
| *the family ] consists of 4 genera and 13 species of ]s, including the ] and the ], collectively known as the "lesser apes" | |||
| *the family ] consisting of ]s, ]s, ]s, and ]s,<ref name=MSW3/><ref name=Goodman2>{{cite journal | journal = Journal of Molecular Evolution | year = 1990 | volume = 30 | pages = 260–266 | url = http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?cmd=Retrieve&db=PubMed&list_uids=2109087&dopt=Abstract | title = Primate evolution at the DNA level and a classification of hominoids | author = M. Goodman, D. A. Tagle, D. H. Fitch, W. Bailey, J. Czelusniak, B. F. Koop, P. Benson, J. L. Slightom}}</ref> collectively known as the great apes. | |||
| A few other primates, such as the ], have the word "ape" in their common names (usually to indicate lack of a tail), but they are not regarded as true apes. | |||
| Except for gorillas and most humans, all true apes are agile climbers of trees. They are best described as ], their diet consisting of fruit, grass seeds, and in most cases some quantities of meat and invertebrates—either hunted or scavenged—along with anything else available and easily digested. They are native to ] and ], although humans have spread to all parts of the world. | |||
| Most ape species are rare or ]. The chief threat to most of the endangered species is loss of tropical ] habitat, though some populations are further imperiled by hunting for ]. | |||
| ==Historical and modern terminology== | |||
| "Ape" (Old Eng. ''apa''; Dutch ''aap''; Old Ger. ''affo''; Ger. ''Affe''; Welsh ''epa''; Old Czech ''op'') is a word of uncertain origin and is possibly an ] imitation of animal chatter. The term has a history of rather imprecise usage. Its earliest meaning was a tailless (and therefore exceptionally human-like) non-human primate, but as ] knowledge developed it became clear that taillessness occurred in a number of different and otherwise unrelated ]. | |||
| The original usage of "ape" in ] may have referred to the ], an ]n ]. Two tailless species of ] are commonly named as apes, the ] of North Africa (introduced into ]), ''Macaca sylvanus'', and the Sulawesi Black Ape or ], ''M. nigra''. | |||
| Until a handful of decades ago, humans were thought to be distinctly set apart from the other apes (even from the other great apes), so much so that many people still don't think of the term "apes" to include humans at all. However, it is not considered accurate by many biologists to think of apes in a biological sense without considering humans to be included.{{Fact|date=February 2007}} The terms "non-human apes" or "non-human great apes" is used with increasing frequency to show the ] relationship of humans to the other apes while yet talking only about the non-human species.{{Fact|date=February 2007}} | |||
| A group of apes may be referred to as a ''troop'' or a ''shrewdness''. | |||
| ==Biology== | |||
| <!--]--> | |||
| The gibbon family, Hylobatidae, is composed of thirteen medium-sized species. Their major distinction is their long arms, which they use to ] through the trees. As an evolutionary adaptation to this ] lifestyle, their wrists are ball and socket joints. The largest of the gibbons, the ], weighs up to 23 kg (50 lb). In comparison, the smallest great ape is the ] at a modest 40 to 65 kg (88 to 143 lb). | |||
| The great ape family was previously referred to as ], and humans (and fossil hominids) were omitted from it, but there is no biological case for doing this. However, this definition is still used by many ]s and by ]. However, that definition makes Pongidae ], whereas most taxonomists nowadays encourage ] groups. Chimpanzees, gorillas, humans and orangutans are all more closely related to one another than any of these four genera are to the gibbons. However, the term "]" is still used with the specific meaning of extinct animals more closely related to humans than the other great apes (for example, ]s), even though "]" is now correct in that usage. It is now usual to use even finer divisions, such as ] and ]s to distinguish which hominoids are being discussed. Current evidence implies that humans share a common, extinct, ancestor with the chimpanzee line, from which we separated more recently than the gorilla line. | |||
| Both great apes and lesser apes fall within ], which also includes the ] of ] and ]. Within this group, both families of apes can be distinguished from these ]s by the number of ] on their ]s (apes have five—the "Y-5" molar pattern, Old World monkeys have only four in a ] pattern). Apes have more mobile shoulder joints and arms due to the dorsal position of the ], broad ribcages that are flatter front-to-back, and a shorter, less mobile spine compared to Old World monkeys (with caudal vertebrae greatly reduced, resulting in tail loss in most species). These are all anatomical adaptations to vertical hanging and swinging locomotion (brachiation) in the apes, as well as better balance in a ] pose. All living members of the Hylobatidae and Hominidae are tailless, and humans can therefore accurately be referred to as bipedal apes. However, there are also primates in other families that lack tails, and at least one (the ]) that has been known to walk significant distances bipedally. The front skull is characterised by its sinuses, fusion of the frontal bone and ]. | |||
| Although the hominoid fossil record is far from complete, and the evidence is often fragmentary, there is enough to give a good outline of the evolutionary history of humans. The time of the split between humans and living apes used to be thought to have occurred 15 to 20 million years ago, or even up to 30 or 40 million years ago. Some apes occurring within that time period, such as '']'', used to be considered as ]s, and possible ancestors of humans. Later fossil finds indicated that ''Ramapithecus'' was more closely related to the orangutan, and new biochemical evidence indicated that the last common ancestor of humans and other hominins occurred between 5 and 10 million years ago, and probably in the lower end of that range. | |||
| ==Cultural aspects of non-human apes== | |||
| The name "Hominoidea" can be loosely translated as "ape". However, although the superfamily of Hominoidea has always included ] such as humans, as well as the ], a different connotation of the word "ape" exists in the vernacular. The historical, common usage of the word often excludes humans when referring to apes. Humans are also often excluded from the larger classifications of "]" and "]" in common usage, despite belonging to both of these groups as well. The reason for this is that scientific nomenclature and everyday language abide by different rules. Other examples of this are "]" (not a member of ]), "]" (not a member of ]) and "]" (not a ]). Taxonomic labels can be redefined according to the latest scientific findings; as such, they may or may not overlap with their vernacular counterparts. | |||
| Often, non-human apes are said to be the result of a curse—a ] ] claims that one of the races who built the ] became apes as punishment, while ] lore says that the Jews of ] became non-human apes as punishment for ] on the ]. Some sects of ] have folklore that claims that these apes are a symbol of ] and were created by ] in response to ]'s creation of humans. It is uncertain whether any of these references are to any specific apes. All of these concepts date from a period when neither the distinction between apes and monkeys, nor the fact that humans are apes, was widely understood, or understood at all. | |||
| ==History of hominoid taxonomy== | |||
| The history of hominoid taxonomy is somewhat confusing and complex. The names of subgroups have changed their meaning over time as new evidence, from fossil discoveries and comparisons of anatomy and DNA sequences, has changed understanding of the relationships between hominoids. The story of the hominoid taxonomy is one of gradual demotion of humans from a special position in the taxonomy to being one branch among many. It also illustrates the growing influence of ] (the science of classifying living things by strict descent) on taxonomy. | |||
| ], there are eight extant ] of hominoids. They are the four great ape genera ('']'' (humans), '']'' (chimpanzees), '']'', and '']'' (orangutans)), and the four genera of ]s ('']'', '']'', '']'', and '']'').<ref name=MSW3/> (The genus for the ]s was recently changed from '']'' to ''Hoolock''.<ref name=Mootnick>{{cite journal | last = Mootnick | first = A. | coauthors = Groves, C. P. | year = 2005 | title = A new generic name for the hoolock gibbon (Hylobatidae) | journal = International Journal of Primatology | issue = 26 | pages = 971-976}}</ref>) | |||
| In 1758, ], relying on second- or third-hand accounts, placed a second species in ''Homo'' along with ''H. sapiens'': ''Homo troglodytes'' ("cave-dwelling man"). It is not clear to which animal this name refers, as Linnaeus had no specimen to refer to, hence no precise description. Linnaeus named the orangutan ''Simia satyrus'' ("satyr monkey"). He placed the three genera ''Homo'', '']'' and ''Lemur'' in the family of Primates. | |||
| The ''troglodytes'' name was used for the chimpanzee by ] in 1775 but moved to the genus ''Simia''. The orangutan was moved to the genus '']'' in 1799 by ]. | |||
| Linnaeus's inclusion of humans in the primates with monkeys and apes was troubling for people who denied a close relationship between humans and the rest of the animal kingdom. Linnaeus's Lutheran ] had accused him of "impiety." In a letter to ] dated ], ], Linnaeus wrote: | |||
| :It is not pleasing to me that I must place ] among the ]s, but man is intimately familiar with himself. Let's not quibble over words. It will be the same to me whatever name is applied. But I desperately seek from you and from the whole world a general difference between men and ]s from the principles of ]. I certainly know of none. If only someone might tell me one! If I called man a ] or vice versa I would bring together all the ] against me. Perhaps I ought to, in accordance with the law of Natural History.<ref>{{cite web | url = http://linnaeus.c18.net/Letters/display_txt.php?id_letter=L0783 | title = Letter, Carl Linnaeus to Johann Georg Gmelin. Uppsala, Sweden, ] 1747 | publisher = Swedish Linnaean Society}}</ref> | |||
| Accordingly, ] in the first edition of his ''Manual of Natural History'' (1779), proposed that the primates be divided into the ] (four-handed, i.e. apes and monkeys) and ] (two-handed, i.e. humans). This distinction was taken up by other naturalists, most notably ]. Some elevated the distinction to the level of ]. | |||
| However, the many affinities between humans and other primates — and especially the great apes — made it clear that the distinction made no scientific sense. ] wrote, in '']'': | |||
| :The greater number of naturalists who have taken into consideration the whole structure of man, including his mental faculties, have followed ] and ], and have placed man in a separate Order, under the title of the Bimana, and therefore on an equality with the orders of the Quadrumana, ], etc. Recently many of our best naturalists have recurred to the view first propounded by ], so remarkable for his sagacity, and have placed man in the same Order with the ], under the title of the ]s. The justice of this conclusion will be admitted: for in the first place, we must bear in mind the comparative insignificance for classification of the great development of the brain in man, and that the strongly-marked differences between the skulls of man and the Quadrumana (lately insisted upon by ], ], and others) apparently follow from their differently developed brains. In the second place, we must remember that nearly all the other and more important differences between man and the Quadrumana are manifestly adaptive in their nature, and relate chiefly to the erect position of man; such as the structure of his hand, foot, and pelvis, the curvature of his spine, and the position of his head.<ref>{{cite book | author = ]|title = ] | year = 1871}}</ref> | |||
| <!-- ...history from 1870 to 1960 goes here... --> | |||
| ===Changes in taxonomy over time=== | |||
| {| class="wikitable" | |||
| | Until about 1960, the hominoids were usually divided into two families: humans and their extinct relatives in ], the other apes in ].<ref>{{cite journal|author = ] | year = 1945 | title = The principles of classification and a classification of mammals | journal = Bull. Am. Mus. Nat. Hist. | volume = 85 | pages = 1–350}}</ref> | |||
| | ] | |||
| |- | |||
| | The 1960s saw the application of techniques from ] to primate taxonomy. Goodman used his 1964 immunological study of serum proteins to propose a division of the hominoids into three families, with the non-human great apes in Pongidae and the lesser apes (gibbons) in ].<ref name=Goodman>{{cite book | author = M. Goodman | year = 1964 | chapter = Man’s place in the phylogeny of the primates as reflected in serum proteins | editor = S. L. Washburn | title = Classification and human evolution | publisher = Aldine, Chicago|pages = 204–234}}</ref> The ] of hominoid families, however, prompted scientists to ask which family ] first from the common hominoid ancestor. | |||
| | ] | |||
| |- | |||
| | Within the superfamily Hominoidea, gibbons are the ]: this means that the rest of the hominoids are more closely related to each other than any of them are to gibbons. This led to the placing of the other great apes into the family Hominidae along with humans, by demoting the Pongidae to a subfamily; the Hominidae family now contained the subfamilies ] and ]. Again, the three-way split in Ponginae led scientists to ask which of the three genera is least related to the others. | |||
| | ] | |||
| |- | |||
| | Investigation showed ]s to be the outgroup, but comparing humans to all three other hominid genera showed that African apes (chimpanzees and gorillas) and humans are more closely related to each other than any of them are to orangutans. This led to the placing of the African apes in the subfamily Homininae, forming another three-way split. This classification was first proposed by M. Goodman in 1974.<ref name=Goodman1>{{cite journal | journal = Annual Review of Anthropology | volume = 3 | pages = 203–228 | year = 1974 | url = http://arjournals.annualreviews.org/doi/abs/10.1146/annurev.an.03.100174.001223 | title = Biochemical Evidence on Hominid Phylogeny | author = M. Goodman}}</ref> | |||
| | ] | |||
| |- | |||
| | To try to resolve the hominine trichotomy, some authors proposed the division of the subfamily Homininae into the tribes Gorillini (African apes) and Hominini (humans). | |||
| | ] | |||
| |- | |||
| | However, DNA comparisons provide convincing evidence that within the subfamily Homininae, gorillas are the outgroup. This suggests that chimpanzees should be in Hominini along with humans. This classification was first proposed (though one rank lower) by M. Goodman et al. in 1990.<ref name=Goodman2/> See ] for more information on the speciation of humans and great apes. | |||
| | ] | |||
| |- | |||
| | Later DNA comparisons split the gibbon genus ''Hylobates'' into four genera: '']'', '']'', '']'', and '']''.<ref name=MSW3/><ref name=Mootnick/> | |||
| | ] | |||
| |} | |||
| ==Classification and evolution== | |||
| As discussed above, hominoid taxonomy has undergone several changes. Current understanding is that the apes diverged from the ]s about 25 million years ago. The lesser and greater apes split about 18 mya, and the hominid splits happened 14 mya (''Pongo''), 7 mya (''Gorilla''), and 3-5 mya (''Homo'' & ''Pan''). | |||
| * '''Superfamily Hominoidea'''<ref name=MSW3/> | |||
| ** Family ]: gibbons | |||
| *** Genus '']'' | |||
| *** Genus '']'' | |||
| *** Genus '']'' | |||
| *** Genus '']'' | |||
| ** Family ]: great apes | |||
| *** Genus '']'': orangutans | |||
| *** Genus '']'': gorillas | |||
| *** Genus '']'': humans | |||
| *** Genus '']'': chimpanzees | |||
| ==Behaviour and cognition== | |||
| Although there had been earlier studies, the scientific investigation of behaviour and cognition in non-human apes expanded enormously during the latter half of the twentieth century. Major studies of behaviour in the field were completed on the three better-known great apes, for example by ], ] and ] (field work on gibbons and the ] is still relatively underdeveloped). These studies have shown that in their natural environments, the different apes show sharply varying social structure: gibbons are monogamous, territorial pair-bonders, orangutans are solitary, gorillas live in small troops dominated by a single adult male, while chimpanzees live in larger troops with Bonobos exhibiting promiscuous sexual behaviour. Their diets also vary; gorillas are ]s while the others are all primarily ]s, although the Common Chimpanzee does some hunting for meat. Foraging behaviour is correspondingly variable. | |||
| All the apes are generally thought of as highly intelligent, and scientific study has broadly confirmed that they perform outstandingly well on a wide range of cognitive tests - though again there is relatively little data on gibbon cognition. The early studies by ] demonstrated exceptional ] abilities in chimpanzees, which Köhler attributed to ]. The use of ]s has been repeatedly demonstrated; more recently, the manufacture of tools has been documented, both in the wild and in laboratory tests. ] is much more easily demonstrated in great apes than in other primate species. Almost all the studies in ] have been completed with great apes, and though there is continuing dispute as to whether they demonstrate real language abilities, there is no doubt that they involve significant feats of learning. Chimpanzees in different parts of Africa have developed tools that are used in food acquisition, demonstrating a form of animal culture.<ref>{{cite book | author = ]|title = Chimpanzee material culture: implications for human evolution | year = 1992}}</ref> | |||
| ==Legal status== | |||
| :''See ]'' | |||
| ==See also== | |||
| * ] | |||
| * ] | |||
| * ] (for notable non-fictional apes) | |||
| ==References== | |||
| {{reflist}} | |||
| ==External links == | |||
| {{sisterlinks|Ape}} | |||
| {{Wikispecies|Hominoidea|Ape}} | |||
| {{Wikibookspar|Dichotomous Key|Hominoidea}} | |||
| * at the ] | |||
| * | |||
| *{{1911}} | |||
| * by ] {{Entrez Pubmed|10995486}}. Agreement between cladograms based on molecular and anatomical data. | |||
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