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エイプ, APE
APE
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Hominoids or Apes Temporal range: Late Oligocene–Holocene PreЄ
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Bornean orangutan (Pongo pygmaeus) | |
Scientific classification | |
Kingdom: | Animalia |
Phylum: | Chordata |
Class: | Mammalia |
Order: | Primates |
Suborder: | Haplorhini |
Infraorder: | Simiiformes |
Parvorder: | Catarrhini |
Superfamily: | Hominoidea Gray, 1825 |
Type species | |
Homo sapiens Linnaeus, 1758 |
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Families | |
†Proconsulidae |
Apes (Hominoidea) are a branch of Old World tailless anthropoid catarrhine primates native to Africa and Southeast Asia; they are distinguished from other primates by a wider degree of freedom of motion at the shoulder joint as evolved by the influence of brachiation. There are two extant branches of the superfamily Hominoidea: the gibbons, or lesser apes; and the hominids, or great apes.
Members of the superfamily Hominoidae are called hominoids—which term is not to be confused with hominids, the family of great apes; or with the hominins, the tribe of humans also known as the human clade; or with other very similar terms of primate taxa. (Compare terminology of primate names.)
Recent evidence has changed our understanding of the relationships between the hominoids, especially regarding the human lineage; and the traditionally used terms have become somewhat confused. Competing approaches re methodology and terminology are found among current scientific sources. See below, History of hominoid taxonomy and see Primate: Historical and modern terminology for discussions of the changes in scientific classification and terminology regarding hominoids.
Some and, recently, all, hominoids are also called "apes", but the term is used broadly and has several different senses within both popular and scientific settings. "Ape" has been used as a synonym for "monkey" or for naming any primate with a humanlike appearance, particularly those without a tail.[7] Thus the Barbary macaque, a kind of monkey, is popularly called the "Barbary ape". Biologists have traditionally used the term "ape" to mean a member of the superfamily Hominoidea other than humans,[1] but more recently to mean all members of Hominoidea. So "ape"—not to be confused with "great ape"—now becomes another word for hominoid including humans.[4][8]
Except for gorillas and humans, hominoids are agile climbers of trees. Their diet is best described as frugivorous and folivorous, consisting mainly of fruit, nuts, seeds, including grass seeds, leaves, and in some cases other animals, either hunted or scavenged, or (solely in the case of the humans) farmed—along with anything else available and easily digested.[9][10]
Most non-human hominoids are rare or endangered. The chief threat to most of the endangered species is loss of tropical rainforest habitat, though some populations are further imperiled by hunting for bushmeat. The great apes of Africa are also facing threat from the Ebola virus. Currently considered to be the greatest threat to survival of African apes, Ebola is responsible for the death of at least one third of the species since 1990.[11]
"Ape", from Old English apa, is a word of uncertain origin.[12] The term has a history of rather imprecise usage—and of comedic or punning usage in the vernacular. Its earliest meaning was generally of any non-human anthropoid primate,[7][13] as is still the case for its cognates in other Germanic languages.[14] Later, after the term "monkey" had been introduced into English, "ape" was specialized to refer to a tailless (therefore exceptionally human-like) primate.[15] Two tailless species of macaque still have common names using "ape": the Barbary ape of North Africa (introduced into Gibraltar), Macaca sylvanus, and the Sulawesi black ape or Celebes crested macaque, M. nigra. Thus, the term "ape" obtained two different meanings, as shown in the 1910 Encyclopædia Britannica entry: it could be used as a synonym for "monkey" and it could denote the tailless humanlike primate in particular.[7]
The primates called "apes" today became known to Europeans after the 18th century. As zoological knowledge developed, it became clear that taillessness occurred in a number of different and otherwise distantly related species. Sir Wilfrid Le Gros Clark was one of those primatologists who developed the idea that there were trends in primate evolution and that the extant members of the order could be arranged in an ".. ascending series", leading from "monkeys" to "apes" to humans. Within this tradition "ape" came to refer to all members of the superfamily Hominoidea except humans.[1] As such, this use of "apes" represented a paraphyletic grouping, meaning that even though all species of apes were descended from a common ancestor this grouping did not include all the descendant species, because humans were excluded from being among the apes.[16]
The cladogram of superfamily Hominoidae shows the descendant relationships of the hominoids that are broadly accepted today.[6] The grouping traditionally called apes is bracketed from the right:
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apes
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The traditional grouping is further divided into the great apes and the lesser apes:[17]
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great apes
lesser apes
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Thus, there are (at least) three common, or traditional, uses of the term "ape": non-biologists may not distinguish between "monkeys" and "apes", that is, they may use the two terms interchangeably; or they may use "ape" for any tailless monkey or non-human hominoid; whereas biologists have traditionally used the term "ape" to mean the non-human hominoids only, as above.
But, tradition is being changed; see immediately below.
In recent years when delineating classifications, biologists have generally preferred to use monophyletic groups only;[18] that is, they use only those groups that include all the descendants of a common ancestor.[19] The superfamily Hominoidea is such a group—also known as a clade. Some scientists now use the term "ape" to mean all members of the superfamily Hominoidea, including humans. For example, in his 2005 book, Benton wrote "The apes, Hominoidea, today include the gibbons and orang-utan ... the gorilla and chimpanzee ... and humans".[4] As such, the grouping traditionally called "apes" by biologists might now be referred to as the "non-human" apes. And today, to delineate humans-only, scientists broadly, other than paleoanthropologists, appear to be adopting the term "hominin" to identify the human clade, replacing the term "hominid" for this use. See terminology of primate names.
See below, History of hominoid taxonomy, for a discussion of changes in scientific classification and terminology regarding hominoids.
This section does not cite any references or sources. Please help improve this section by adding citations to reliable sources. Unsourced material may be challenged and removed. (July 2011) |
The lesser apes are the gibbon family, Hylobatidae, of sixteen species; all are native to Asia. Their major differentiating characteristic is their long arms, which they use to brachiate through trees. Their wrists are ball and socket joints as an evolutionary adaptation to their arboreal lifestyle. Generally smaller than the African apes, the largest gibbon, the siamang, weighs up to 14 kg (31 lb); in comparison, the smallest "great ape", the bonobo, is at 34 to 60 kg (75 to 132 lb).
Formerly, all the great apes except humans were classified as the family Pongidae, which conveniently provided for separating the human family from the apes; see The "great apes" in Pongidae. As noted above, such a definition would make a paraphyletic grouping of the Pongidae great apes. Current evidence indicates that humans share a common ancestor with the chimpanzee line—from which they separated more recently than from the gorilla line; see Gorillas the outgroup
The superfamily Hominoidea falls within the parvorder Catarrhini, which also includes the Old World monkeys of Africa and Eurasia. Within this grouping, the two families Hylobatidae and Hominidae can be distinguished from Old World monkeys by the number of cusps on their molars; hominoids have five—in the "Y-5" molar pattern, where Old World monkeys have only four in a bilophodont pattern.
Further in comparison with Old World monkeys, hominoids are noted for: more mobile shoulder joints and arms due to the dorsal position of the scapula; broader ribcages that are flatter front-to-back; and a shorter, less mobile spine, with greatly reduced caudal (tail) vertebrae—resulting in complete loss of the tail in living hominoid species. These are anatomical adaptations, first, to vertical hanging and swinging locomotion (brachiation) and, later, to developing balance in a bipedal pose. Note there are primates in other families that also lack tails, and at least one, the pig-tailed langur, is known to walk significant distances bipedally. The front of the ape skull is characterised by its sinuses, fusion of the frontal bone, and by post-orbital constriction.
Although the hominoid fossil record is still incomplete and fragmentary, there is enough evidence now to provide an outline of the evolutionary history of humans. Previously, the divergence between humans and other living hominoids was thought to have occurred 15 to 20 million years ago, and several species of that time period, such as Ramapithecus, were once thought to be hominins and possible ancestors of humans. But later fossil finds indicated that Ramapithecus was more closely related to the orangutan. And new biochemical evidence indicates that the last common ancestor of humans and non-hominins (that is, the chimpanzees) occurred between 5 and 10 million years ago, and probably nearer the lower end of that range; see Chimpanzee–human last common ancestor (CHLCA).
This section needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed. (July 2011) |
Although there had been earlier studies, the scientific investigation of behaviour and cognition in non-human members of the superfamily Hominoidea 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 Jane Goodall, Dian Fossey and Birute Galdikas. These studies have shown that in their natural environments, the non-human hominoids show sharply varying social structure: gibbons are monogamous, territorial pair-bonders, orangutans are solitary, gorillas live in small troops with a single adult male leader, while chimpanzees live in larger troops with bonobos exhibiting promiscuous sexual behaviour. Their diets also vary; gorillas are foliovores while the others are all primarily frugivores, although the common chimpanzee does some hunting for meat. Foraging behaviour is correspondingly variable.
All the non-human hominoids are generally thought of as highly intelligent, and scientific study has broadly confirmed that they perform very well on a wide range of cognitive tests—though there is relatively little data on gibbon cognition. The early studies by Wolfgang Köhler demonstrated exceptional problem-solving abilities in chimpanzees, which Köhler attributed to insight. The use of tools has been repeatedly demonstrated; more recently, the manufacture of tools has been documented, both in the wild and in laboratory tests. Imitation is much more easily demonstrated in "great apes" than in other primate species. Almost all the studies in animal language acquisition have been done 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.[20]
Apes do not possess a tail, unlike most monkeys. Monkeys are more likely to be in trees and use their tails for balance. While the great apes are considerably larger than monkeys, gibbons (lesser apes) are smaller than some monkeys. Apes are considered to be more intelligent than monkeys, which are considered to have more primitive brains.[21]
The history of hominoid taxonomy is complex and somewhat confusing. Over time, authorities have changed the names and the meanings of names of groups and subgroups as new evidence—that is, new discoveries of fossils and tools and of observations in the field, plus continual comparisons of anatomy and DNA sequences—has changed the understanding of relationships between hominoids. There has been a gradual demotion of humans from being 'special' in the taxonomy to being one branch among many. This recent turmoil (of history) illustrates the growing influence on all taxonomy of cladistics, the science of classifying living things strictly according to their lines of descent.
Today there are eight extant genera of hominoids. They are the four genera in the family Hominidae, namely Homo, Pan, Gorilla, and Pongo; plus four genera in the family Hylobatidae (gibbons): Hylobates, Hoolock, Nomascus and Symphalangus.[5] (The two subspecies hoolock gibbons were recently moved from the genus Bunopithecus to the new genus Hoolock).[22])
In 1758, Carolus Linnaeus, 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, Simia and Lemur in the order of Primates.
The troglodytes name was used for the chimpanzee by Blumenbach in 1775 but moved to the genus Simia. The orangutan was moved to the genus Pongo in 1799 by Lacépède.
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 archbishop had accused him of "impiety". In a letter to Johann Georg Gmelin dated 25 February 1747, Linnaeus wrote:
Accordingly, Johann Friedrich Blumenbach in the first edition of his Manual of Natural History (1779), proposed that the primates be divided into the Quadrumana (four-handed, i.e. apes and monkeys) and Bimana (two-handed, i.e. humans). This distinction was taken up by other naturalists, most notably Georges Cuvier. Some elevated the distinction to the level of order.
However, the many affinities between humans and other primates — and especially the "great apes" — made it clear that the distinction made no scientific sense. Charles Darwin wrote, in The Descent of Man:
Humans the non-apes: Until about 1960, taxonomists typically divided the superfamily Hominoidea into two families. Humans and their extinct relatives, considered as quite distant from kinship with apes, were classified as the family Hominidae and were known as the hominids. All other hominoids were known as apes and were referred to the family Pongidae.[25] | |
The "great apes" in Pongidae: The 1960s saw the methodologies of molecular biology applied to primate taxonomy. Goodman's 1964 immunological study of serum proteins led to re-classifying the hominoids into three families: the humans in Hominidae; the great apes in Pongidae; and the "lesser apes" (gibbons) in Hylobatidae.[26] However, a trichotomy—Pan, Gorilla, and Pongo—of the "great apes" in Pongidae presented a puzzle; scientists wanted to know which genus speciated first from the common hominoid ancestor. | |
Gibbons the outgroup: New studies indicated that gibbons are the outgroup within the superfamily Hominoidea, meaning: the rest of the hominoids are more closely related to each other than (any of them) to the gibbons. The splitting of the gibbons was isolated by moving the great apes into the same family as humans. Now the term "hominid" encompassed a larger collective taxa within family Hominidae. The trichotomy still required scientists to learn which genus is 'least' related to the others. | |
Orangutans the outgroup: Investigations comparing humans and the three other hominid genera disclosed that the African apes (chimpanzees and gorillas) and humans are more closely related to each other than any of them to orangutans (Pongo); that is, the orangutans and not humans, are the outgroup of the family Hominidae. This led to reassigning the African apes to the subfamily Homininae with humans—which presented a new three-way split (trichotomy), Homo, Pan, Gorilla.[27] | |
Hominins: To resolve the trichotomy, the subfamily Homininae was divided into two tribes: Gorillini, comprising genus Pan and genus Gorilla; and Hominini, comprising genus Homo (the humans). Humans and close relatives now began to be known as "hominins", that is, of tribe Hominini. Thus, the term "hominin" succeeded to the previous use of "hominid", which meaning had changed with changes in Hominidae (see above: 3rd graphic, Gibbons the outgroup). | |
Gorillas the outgroup: New DNA comparisons now provided evidence that gorillas, not humans, are the outgroup in the subfamily Homininae; this suggested that chimpanzees should be grouped with humans in tribe Hominini, but in separate subtribes.[6] Now the name "hominin" delineated Homo plus those earliest Homo relatives and ancestors that arose after the divergence from the chimpanzees. | |
Speciation of gibbons: Later DNA comparisons disclosed previously unknown speciation of genus Hylobates (gibbons) into four genera: Hylobates, Hoolock, Nomascus, and Symphalangus.[5][22] See Human evolutionary genetics re the speciation of humans and great apes. |
As discussed above, hominoid taxonomy has undergone several changes. Genetic analysis combined with fossil evidence indicates that hominoids diverged from the Old World monkeys about 25 million years ago, near the Oligocene-Miocene boundary.[28] The gibbons split from the rest about 18 mya, and the hominid splits happened 14 mya (Pongo), 7 mya (Gorilla), and 3–5 mya (Homo & Pan).[citation needed]
The families, and extant genera and species of hominoids are:
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Through careful study taxonomists today struggle to eliminate polyphyletic and paraphyletic groups and taxons, reclassifying their members into appropriate monophyletic taxa
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