Ape

Hominoids
Apes
Temporal range: Miocene-Holocene
Male chimpanzee (Pan troglodytes)
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Mammalia
Order: Primates
Suborder: Haplorhini
Infraorder: Simiiformes
Parvorder: Catarrhini
Superfamily: Hominoidea
Gray, 1825[1]
Type species
Homo sapiens
Families

Apes (collectively Hominoidea /hɒmɪˈnɔɪdi.ə/) are a clade of Old World simians native to sub-Saharan Africa and Southeast Asia (though they were more widespread in Africa, most of Asia, and Europe in prehistory, and counting humans are found globally). Apes are more closely related to Old World monkeys (family Cercopithecidae) than to the New World monkeys (Platyrrhini) with both Old World monkeys and apes placed in the clade Catarrhini. Apes do not have tails due to a mutation of the TBXT gene.[2][3] In traditional and non-scientific use, the term ape can include tailless primates taxonomically considered Cercopithecidae (such as the Barbary ape and black ape), and is thus not equivalent to the scientific taxon Hominoidea. There are two extant branches of the superfamily Hominoidea: the gibbons, or lesser apes; and the hominids, or great apes.

  • The family Hylobatidae, the lesser apes, include four genera and a total of 20 species of gibbon, including the lar gibbon and the siamang, all native to Asia. They are highly arboreal and bipedal on the ground. They have lighter bodies and smaller social groups than great apes.
  • The family Hominidae (hominids), the great apes, include four genera comprising three extant species of orangutans and their subspecies, two extant species of gorillas and their subspecies, two extant species of chimpanzees and their subspecies, and humans in a single extant subspecies.[a][4][5][6]

Except for gorillas and humans, hominoids are agile climbers of trees. Apes eat a variety of plant and animal foods, with the majority of food being plant foods, which can include fruits, leaves, stalks, roots and seeds, including nuts and grass seeds. Human diets are sometimes substantially different from that of other hominoids due in part to the development of technology and a wide range of habitation.

All non-human hominoids are rare and threatened with extinction. The main threat is habitat loss, though some populations are further imperiled by hunting. The great apes of Africa are also facing threat from the Ebola virus.[7]

Name and terminology

"Ape", from Old English apa, is a word of uncertain origin.[b] 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, as is still the case for its cognates in other Germanic languages.[c][8] Later, after the term "monkey" had been introduced into English, "ape" was specialized to refer to a tailless (therefore exceptionally human-like) primate.[9] Thus, the term "ape" obtained two different meanings, as shown in the 1911 Encyclopædia Britannica entry: it could be used as a synonym for "monkey" and it could denote the tailless human-like primate in particular.[10]

Some, or 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 human-like appearance, particularly those without a tail.[10] Biologists have traditionally used the term "ape" to mean a member of the superfamily Hominoidea other than humans,[4] 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.[6][d]

The taxonomic term hominoid is derived from, and intended as encompassing, the hominids, the family of great apes. Both terms were introduced by Gray (1825).[11] The term hominins is also due to Gray (1824), intended as including the human lineage (see also Hominidae#Terminology, Human taxonomy).

The distinction between apes and monkeys is complicated by the traditional paraphyly of monkeys: Apes emerged as a sister group of Old World Monkeys in the catarrhines, which are a sister group of New World Monkeys. Therefore, cladistically, apes, catarrhines and related contemporary extinct groups such as Parapithecidae are monkeys as well, for any consistent definition of "monkey". "Old World monkey" may also legitimately be taken to be meant to include all the catarrhines, including apes and extinct species such as Aegyptopithecus,[12][13][14][15][citation needed] in which case the apes, Cercopithecoidea and Aegyptopithecus emerged within the Old World monkeys.

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.[4] 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.[e]

Traditionally, the English-language vernacular name "apes" does not include humans, but phylogenetically, humans (Homo) form part of the family Hominidae within Hominoidea. Thus, there are at least three common, or traditional, uses of the term "ape": non-specialists 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; or they may use the term "ape" to just mean the non-human hominoids.

Modern taxonomy aims for the use of monophyletic groups for taxonomic classification;[16][f] Some literature may now use the common name "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".[6] Modern biologists and primatologists refer to apes that are not human as "non-human" apes. Scientists broadly, other than paleoanthropologists, may use the term "hominin" to identify the human clade, replacing the term "hominid". See terminology of primate names.

See below, History of hominoid taxonomy, for a discussion of changes in scientific classification and terminology regarding hominoids.

Evolution

Although the hominoid fossil record is still incomplete and fragmentary, there is now enough evidence to provide an outline of the evolutionary history of humans. Previously, the divergence between humans and other extant 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 (more recent); see Chimpanzee–human last common ancestor (CHLCA).

Taxonomic classification and phylogeny

Genetic analysis combined with fossil evidence indicates that hominoids diverged from the Old World monkeys about 25 million years ago (mya), near the Oligocene–Miocene boundary.[17][18][19] The gibbons split from the rest about 18 mya, and the hominid splits happened 14 mya (Pongo),[20] 7 mya (Gorilla), and 3–5 mya (Homo & Pan).[21] In 2015, a new genus and species were described, Pliobates cataloniae, which lived 11.6 mya, and appears to predate the split between Hominidae and Hylobatidae.[22][23][24][6][clarification needed]

Crown Catharrhini (31)
(29)
Hominoidea (30)

Dendropithecidae (†7 Mya)

Ekembo heseloni (†17 Mya)

Proconsulidae (†18 Mya)

Ekembo nyanzae (†17 Mya)

(29)

Equatorius (†16)

Pliobates (†11.6 Mya)

(29)
Afropithecidae (28)

Morotopithecus (†20)

Afropithecus (†16)

Crown Hominoidea (22)
Catarrhini (31.0 Mya)
Hominoidea/apes (20.4 Mya)
Hominidae/great apes (15.7 Mya)
Homininae (8.8 Mya)
Hominini (6.3 Mya)

humans (genus Homo)

chimpanzees (genus Pan)

gorillas (genus Gorilla)

orangutans (genus Pongo)

gibbons/lesser apes (family Hylobatidae)




Traditional apes

Cercopithecoidea  Old World monkeys

Skeletons of members of the ape superfamily, Hominoidea. There are two extant families: Hominidae, the "great apes"; and Hylobatidae, the gibbons, or "lesser apes".
From left: Comparison of size of gibbon, human, chimpanzee, gorilla and orangutan. Non-human hominoids do not stand upright as their normal posture.

The families, and extant genera and species of hominoids are:

History of hominoid taxonomy

The history of hominoid taxonomy is complex and somewhat confusing. 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 to methodology and terminology are found among current scientific sources. 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.[citation needed]

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.[25] (The two subspecies of hoolock gibbons were recently moved from the genus Bunopithecus to the new genus Hoolock and re-ranked as species; a third species was described in January 2017).[28]

In 1758, Carl Linnaeus, relying on second- or third-hand accounts, placed a second species in Homo along with H. sapiens: Homo troglodytes ("cave-dwelling man"). Although the term "Orang Outang" is listed as a variety – Homo sylvestris – under this species, it is nevertheless not clear to which animal this name refers, as Linnaeus had no specimen to refer to, hence no precise description. Linnaeus may have based Homo troglodytes on reports of mythical creatures, then-unidentified simians, or Asian natives dressed in animal skins.[29] 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:

It is not pleasing to me that I must place humans among the primates, 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 simians from the principles of Natural History. I certainly know of none. If only someone might tell me one! If I called man a simian or vice versa I would bring together all the theologians against me. Perhaps I ought to, in accordance with the law of Natural History.[30]

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. In his 1871 book The Descent of Man, and Selection in Relation to Sex, Charles Darwin wrote:

The greater number of naturalists who have taken into consideration the whole structure of man, including his mental faculties, have followed Blumenbach and Cuvier, 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, Carnivora, etc. Recently many of our best naturalists have recurred to the view first propounded by Linnaeus, so remarkable for his sagacity, and have placed man in the same Order with the Quadrumana, under the title of the Primates. 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 Bischoff, Aeby, 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.[31]

Changes in taxonomy and terminology

Humans the non-apes: Until about 1960, taxonomists typically divided the superfamily Hominoidea into two families. The science community treated humans and their extinct relatives as the outgroup within the superfamily; that is, humans were considered as quite distant from kinship with the "apes". Humans 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.[32]
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.[33] However, this arrangement had two trichotomies: Pan, Gorilla, and Pongo of the "great apes" in Pongidae, and Hominidae, Pongidae, and Hylobatidae in Hominoidea. These presented a puzzle; scientists wanted to know which genus speciated first from the common hominoid ancestor.
Gibbons the outgroup: New studies indicated that gibbons, not humans, are the outgroup within the superfamily Hominoidea, meaning: the rest of the hominoids are more closely related to each other than (any of them) are to the gibbons. With this splitting, the gibbons (Hylobates, et al.) were isolated after moving the great apes into the same family as humans. Now the term "hominid" encompassed a larger collective taxa within the family Hominidae. With the family trichotomy settled, scientists could now work to learn which genus is 'least' related to the others in the subfamily Ponginae.
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 are to the Asian orangutans (Pongo); that is, the orangutans, not humans, are the outgroup within the family Hominidae. This led to reassigning the African apes to the subfamily Homininae with humans—which presented a new three-way split: Homo, Pan, and Gorilla.[34]
Hominins: In an effort to resolve the trichotomy, while preserving the nostalgic "outgroup" status of humans, 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 the 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 the tribe Hominini, but in separate subtribes.[35] Now the name "hominin" delineated Homo plus those earliest Homo relatives and ancestors that arose after the divergence from the chimpanzees. (Humans are no longer recognized as an outgroup, but are a branch, deep in the tree of the pre-1960s ape group).
Speciation of gibbons: Later DNA comparisons disclosed previously unknown speciation of genus Hylobates (gibbons) into four genera: Hylobates, Hoolock, Nomascus, and Symphalangus.[25][28] The ordering of speciation of these four genera are being investigated as of 2022.

Characteristics

Like those of the orangutan, the shoulder joints of hominoids are adapted to brachiation, or movement by swinging in tree branches.

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 34 to 60 kg (75 to 132 lb).

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, whereas 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 extant 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.

Distinction from monkeys

Cladistically, apes, catarrhines, and extinct species such as Aegyptopithecus and Parapithecidaea, are monkeys,[citation needed] so one can only specify ape features not present in other monkeys.

Unlike most monkeys, apes do not possess a tail. 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.[36]

The enzyme urate oxidase has become inactive in all apes, its function having been lost in two primate lineages during the middle Miocene; first in the common ancestors of Hominidae, and later in the common ancestor of Hylobatidae. It has been hypothesized that in both incidents it was a mutation that occurred in apes living in Europe when the climate was getting colder, leading to starvation during winter. The mutation changed the biochemistry of the apes and made it easier to accumulate fat, which allowed the animals to survive longer periods of starvation. When they migrated to Asia and Africa, this genetic trait remained.[37][38]

Behaviour

Major studies of behaviour in the field were completed on the three better-known "great apes", for example by Jane Goodall, Dian Fossey and Birutė 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 hunts for meat. Foraging behaviour is correspondingly variable.

In November 2023, scientists reported, for the first time, evidence that groups of primates, including apes, and, particularly bonobos, are capable of cooperating with each other.[39][40]

Diet

Apart from humans and gorillas, apes eat a predominantly frugivorous diet, mostly fruit, but supplemented with a variety of other foods. Gorillas are predominantly folivorous, eating mostly stalks, shoots, roots and leaves with some fruit and other foods. Non-human apes usually eat a small amount of raw animal foods such as insects or eggs. In the case of humans, migration and the invention of hunting tools and cooking has led to an even wider variety of foods and diets, with many human diets including large amounts of cooked tubers (roots) or legumes.[41] Other food production and processing methods including animal husbandry and industrial refining and processing have further changed human diets.[42] Humans and other apes occasionally eat other primates.[43] Some of these primates are now close to extinction with habitat loss being the underlying cause.[44][45]

Cognition

This series of images shows a gorilla using a small tree trunk as a tool. The trunk is used to maintain balance while fishing for aquatic plants

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.[46]

Threats and conservation

All non-human hominoids are rare and threatened with extinction. The eastern hoolock gibbon is the least threatened, only being vulnerable to extinction. Five gibbon species are critically endangered, as are all species of orangutan and gorilla. The remaining species of gibbon, the bonobo, and all four subspecies of chimpanzees are 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.[47] Currently considered to be the greatest threat to survival of African apes, Ebola infection is responsible for the death of at least one third of all gorillas and chimpanzees since 1990.[48]

All the species of great apes in Africa, are considered endangered. Hunting, logging, agricultural expansion and mining are among the main threats. Recently mining has expanded due to the energy transition. According to researchers "This means that current climate solutions could lead to more industrialization in these places, which could worsen the climate crisis". The Sustainable Critical Minerals Alliance was created for solve problems like this.[49]

See also

Notes

  1. ^ Although Dawkins is clear that he uses "apes" for Hominoidea, he also uses "great apes" in ways which exclude humans. Thus in Dawkins 2005: "Long before people thought in terms of evolution ... great apes were often confused with humans" (p. 114); "gibbons are faithfully monogamous, unlike the great apes which are our closer relatives" (p. 126).
  2. ^ The hypothetical Proto-Germanic form is given as *apōn (F. Kluge, Etymologisches Wörterbuch der Deutschen Sprache (2002), online version, s.v. "Affe"; V. Orel, A handbook of Germanic etymology (2003), s.v. "*apōn" or as *apa(n) (Online Etymology Dictionary (2001–2014), s.v. "ape"; M. Philippa, F. Debrabandere, A. Quak, T. Schoonheim & N. van der Sijs, Etymologisch woordenboek van het Nederlands (2003–2009), s.v. "aap"). Perhaps ultimately derived from a non-Indo-European language, the word might be a direct borrowing from Celtic, or perhaps from Slavic, although in both cases it is also argued that the borrowing, if it took place, went in the opposite direction.
  3. ^ "Any simian known on the Mediterranean during the Middle Ages; monkey or ape"; cf. ape-ward: "a juggler who keeps a trained monkey for the amusement of the crowd." (Middle English Dictionary, s.v. "ape").
  4. ^ Dawkins 2005; for example "[a]ll apes except humans are hairy" (p. 99), "[a]mong the apes, gibbons are second only to humans" (p. 126).
  5. ^ Definitions of paraphyly vary; for the one used here see e.g. Stace 2010, pp. 106
  6. ^ Definitions of monophyly vary; for the one used here see e.g. Mishler 2009, pp. 114

References

  1. ^ Gray, J. E. "An outline of an attempt at the disposition of Mammalia into tribes and families, with a list of the genera apparently appertaining to each tribe". Annals of Philosophy. New Series. 10: 337–344. Archived from the original on 27 April 2022. Retrieved 27 April 2022.
  2. ^ Xia, Bo; Zhang, Weimin; Wudzinska, Aleksandra; Huang, Emily; Brosh, Ran; Pour, Maayan; Miller, Alexander; Dasen, Jeremy S.; Maurano, Matthew T.; Kim, Sang Y.; Boeke, Jef D. (16 September 2021). "The genetic basis of tail-loss evolution in humans and apes". bioRxiv 10.1101/2021.09.14.460388.
  3. ^ Weisberger, Mindy (23 March 2024). "Why don't humans have tails? Scientists find answers in an unlikely place". CNN. Archived from the original on 24 March 2024. Retrieved 24 March 2024.
  4. ^ a b c Dixson 1981, pp. 13.
  5. ^ Grehan, J. R. (2006). "Mona Lisa smile: the morphological enigma of human and great ape evolution". Anatomical Record. 289B (4): 139–157. doi:10.1002/ar.b.20107. PMID 16865704.
  6. ^ a b c d Benton, M. J. (2005). Vertebrate Palaeontology. Wiley-Blackwell. ISBN 978-0-632-05637-8. Archived from the original on 3 April 2023. Retrieved 10 July 2011., p. 371
  7. ^ Rush, J. (23 January 2015). "Ebola virus 'has killed a third of world's gorillas and chimpanzees' – and could pose greatest threat to their survival, conservationists warn". The Independent. Archived from the original on 30 March 2015. Retrieved 26 March 2015.
  8. ^ Terry 1977, pp. 3.
  9. ^ Terry 1977, pp. 3–4.
  10. ^ a b Chisholm, Hugh, ed. (1911). "Ape" . Encyclopædia Britannica. Vol. 2 (11th ed.). Cambridge University Press. p. 160.
  11. ^ Gray, JE. "An outline of an attempt at the disposition of Mammalia into tribes and families, with a list of the genera apparently appertaining to each tribe". Annals of Philosophy. New Series. 10: 337–344. Archived from the original on 27 April 2022. Retrieved 27 April 2022.
  12. ^ Osman Hill, W. C. (1953). Primates Comparative Anatomy and Taxonomy I—Strepsirhini. Edinburgh Univ Pubs Science & Maths, No 3. Edinburgh University Press. p. 53. OCLC 500576914.
  13. ^ Martin, W. C. L. (1841). A General Introduction to the Natural History of Mammiferous Animals, With a Particular View of the Physical History of man, and the More Closely Allied Genera of the Order Quadrumana, or Monkeys. London: Wright and Co. printers. pp. 340, 361.
  14. ^ Geoffroy Saint-Hilaire, M. É. (1812). "Tableau des quadrumanes, ou des animaux composant le premier ordre de la classe des Mammifères". Annales du Muséum d'Histoire Naturelle. 19. Paris: 85–122. Archived from the original on 27 March 2019. Retrieved 16 July 2019.
  15. ^ Bugge, J. (1974). "Chapter 4". Cells Tissues Organs. 87 (Suppl. 62): 32–43. doi:10.1159/000144209. ISSN 1422-6405.
  16. ^ Springer; D. H. (1 July 2011). An Introduction to Zoology: Investigating the Animal World. Jones & Bartlett Publishers. p. 536. ISBN 978-0-7637-5286-6. Through careful study taxonomists today struggle to eliminate polyphyletic and paraphyletic groups and taxons, reclassifying their members into appropriate monophyletic taxa
  17. ^ "Fossils may pinpoint critical split between apes and monkeys". 15 May 2013. Archived from the original on 16 December 2022. Retrieved 30 June 2022.
  18. ^ Rossie, J. B.; Hill, A. (2018). "A new species of Simiolus from the middle Miocene of the Tugen Hills, Kenya". Journal of Human Evolution. 125: 50–58. Bibcode:2018JHumE.125...50R. doi:10.1016/j.jhevol.2018.09.002. PMID 30502897. S2CID 54625375.
  19. ^ Rasmussen, D. T.; Friscia, A. R.; Gutierrez, M.; et al. (2019). "Primitive Old World monkey from the earliest Miocene of Kenya and the evolution of cercopithecoid bilophodonty". Proceedings of the National Academy of Sciences. 116 (13): 6051–6056. Bibcode:2019PNAS..116.6051R. doi:10.1073/pnas.1815423116. PMC 6442627. PMID 30858323.
  20. ^ Alba, D. M.; Fortuny, J.; Moyà-Solà, S. (2010). "Enamel thickness in the Middle Miocene great apes Anoiapithecus, Pierolapithecus and Dryopithecus". Proceedings of the Royal Society of London B: Biological Sciences. 277 (1691): 2237–2245. doi:10.1098/rspb.2010.0218. ISSN 0962-8452. PMC 2880156. PMID 20335211.
  21. ^ Grabowski, M.; Jungers, W. L. (2017). "Evidence of a chimpanzee-sized ancestor of humans but a gibbon-sized ancestor of apes". Nature Communications. 8 (1): 880. Bibcode:2017NatCo...8..880G. doi:10.1038/s41467-017-00997-4. ISSN 2041-1723. PMC 5638852. PMID 29026075.
  22. ^ "A new primate species at the root of the tree of extant hominoids". 29 October 2015. Archived from the original on 29 October 2015. Retrieved 29 October 2015.
  23. ^ Nengo, I.; Tafforeau, P.; Gilbert, C. C.; et al. (2017). "New infant cranium from the African Miocene sheds light on ape evolution" (PDF). Nature. 548 (7666): 169–174. Bibcode:2017Natur.548..169N. doi:10.1038/nature23456. PMID 28796200. S2CID 4397839. Archived from the original (PDF) on 22 July 2018. Retrieved 15 July 2019.
  24. ^ Dixson 1981, p. 16.
  25. ^ a b c Groves, C. P. (2005). Wilson, D. E.; Reeder, D. M. (eds.). Mammal Species of the World: A Taxonomic and Geographic Reference (3rd ed.). Baltimore: Johns Hopkins University Press. pp. 178–184. ISBN 0-801-88221-4. OCLC 62265494.
  26. ^ Cochrane, J. (2 November 2017). "New Orangutan Species Could Be the Most Endangered Great Ape". The New York Times. Archived from the original on 17 April 2018. Retrieved 3 November 2017.
  27. ^ a b Wilson, Don E.; Cavallini, Paolo (2013). Handbook of the Mammals of the World. Barcelona: Lynx Edicions. ISBN 978-84-96553-89-7. OCLC 1222638259.
  28. ^ a b Mootnick, A.; Groves, C. P. (2005). "A new generic name for the hoolock gibbon (Hylobatidae)". International Journal of Primatology. 26 (4): 971–976. doi:10.1007/s10764-005-5332-4. S2CID 8394136.
  29. ^ Frängsmyr, T.; Lindroth, S.; Eriksson, G.; Broberg, G. (1983). Linnaeus, the man and his work. Berkeley and Los Angeles: University of California Press. ISBN 978-0-7112-1841-3., p. 166
  30. ^ "Letter, Carl Linnaeus to Johann Georg Gmelin. Uppsala, Sweden, 25 February 1747". Swedish Linnaean Society. Archived from the original on 27 February 2009. Retrieved 4 February 2009.
  31. ^ Darwin, C. (1871). The Descent of Man. Barnes & Noble. ISBN 978-0-7607-7814-2.
  32. ^ G. G., Simpson (1945). "The principles of classification and a classification of mammals". Bulletin of the American Museum of Natural History. 85: 1–350.
  33. ^ Goodman, M. (1964). "Man's place in the phylogeny of the primates as reflected in serum proteins". In Washburn, S. L. (ed.). Classification and Human Evolution. Chicago: Aldine. pp. 204–234.
  34. ^ Goodman, M. (1974). "Biochemical evidence on hominid phylogeny". Annual Review of Anthropology. 3 (1): 203–228. doi:10.1146/annurev.an.03.100174.001223.
  35. ^ Goodman, M.; Tagle, D. A.; Fitch, D. H.; et al. (1990). "Primate evolution at the DNA level and a classification of hominoids". Journal of Molecular Evolution. 30 (3): 260–266. Bibcode:1990JMolE..30..260G. doi:10.1007/BF02099995. PMID 2109087. S2CID 2112935.
  36. ^ Call, J.; Tomasello, M. (2007). The Gestural Communication of Apes and Monkeys. Taylor & Francis Group/Lawrence Erlbaum Associates.
  37. ^ Johnson, R. J.; Lanaspa, M. A.; Gaucher, E. A. (2011). "Uric acid: A Danger Signal from the RNA World that may have a role in the Epidemic of Obesity, Metabolic Syndrome and CardioRenal Disease: Evolutionary Considerations". Seminars in Nephrology. 31 (5): 394–399. doi:10.1016/j.semnephrol.2011.08.002. PMC 3203212. PMID 22000645.
  38. ^ Johnson, Richard J.; Andrews, Peter (2015). "The Fat Gene". Scientific American. 313 (4): 64–69. Bibcode:2015SciAm.313d..64J. doi:10.1038/scientificamerican1015-64.
  39. ^ Zimmer, Carl (16 November 2023). "Scientists Find First Evidence That Groups of Apes Cooperate - Some bonobos are challenging the notion that humans are the only primates capable of group-to-group alliances". The New York Times. Archived from the original on 16 November 2023. Retrieved 17 November 2023.
  40. ^ Samuni, Liran; et al. (16 November 2023). "Cooperation across social borders in bonobos". Science. 382 (6672): 805–809. Bibcode:2023Sci...382..805S. doi:10.1126/science.adg0844. PMID 37972165. Archived from the original on 17 November 2023. Retrieved 17 November 2023.
  41. ^ Lawton, G. (2 November 2016). "Every human culture includes cooking – this is how it began". New Scientist. Archived from the original on 29 July 2021. Retrieved 27 August 2021.
  42. ^ Hoag, Hannah (2 December 2013). "Humans are becoming more carnivorous". Nature. doi:10.1038/nature.2013.14282. S2CID 183143537. Archived from the original on 29 November 2014. Retrieved 26 November 2014.
  43. ^ Callaway, E. (13 October 2006). "Loving bonobos have a carnivorous dark side". New Scientist. Archived from the original on 29 October 2014. Retrieved 26 November 2014.
  44. ^ M., Michael. "Chimpanzees over-hunt monkey prey almost to extinction". BBC Earth. Archived from the original on 6 June 2018. Retrieved 28 May 2018.
  45. ^ "Extinction threat to monkeys and other primates due to habitat loss, hunting". Science Daily. Archived from the original on 28 May 2018. Retrieved 28 May 2018.
  46. ^ McGrew, W. (1992). Chimpanzee Material Culture: Implications for Human Evolution.
  47. ^ Rush, J. (23 January 2015). "Ebola virus 'has killed a third of world's gorillas and chimpanzees' – and could pose greatest threat to their survival, conservationists warn". The Independent. Archived from the original on 30 March 2015. Retrieved 26 March 2015.
  48. ^ Rush, J. (23 January 2015). "Ebola virus 'has killed a third of world's gorillas and chimpanzees' – and could pose greatest threat to their survival, conservationists warn". The Independent. Archived from the original on 30 March 2015. Retrieved 26 March 2015.
  49. ^ Junker, Jessica; Quoss, Luise; Valdez, Jose; Arandjelovic, Mimi; Barrie, Abdulai; Campbell, Geneviève; Heinicke, Stefanie; Humle, Tatyana; Kouakou, Célestin Y.; Kühl, Hjalmar S.; Ordaz-Németh, Isabel; Pereira, Henrique M.; Rainer, Helga; Refisch, Johannes; Sonter, Laura; Sop, Tenekwetche (3 April 2024). "Threat of mining to African great apes". Science Advances. 10 (14). Bibcode:2024SciA...10L.335J. doi:10.1126/sciadv.adl0335. PMID 38569032. Retrieved 19 September 2024.

Literature cited

Read other articles:

Bagian dari seriGereja Katolik menurut negara Afrika Afrika Selatan Afrika Tengah Aljazair Angola Benin Botswana Burkina Faso Burundi Chad Eritrea Eswatini Etiopia Gabon Gambia Ghana Guinea Guinea-Bissau Guinea Khatulistiwa Jibuti Kamerun Kenya Komoro Lesotho Liberia Libya Madagaskar Malawi Mali Maroko Mauritania Mauritius Mesir Mozambik Namibia Niger Nigeria Pantai Gading Republik Demokratik Kongo Republik Kongo Rwanda Sao Tome dan Principe Senegal Seychelles Sierra Leone Somalia Somaliland ...

 

Sharafuddin Idris Shah شرف الدين إدريس شاه الحاجDKM DK(Selangor) DK(Johor) DK(Kelantan) DK(Kedah) DK(Perlis) DK(Pahang) DK(Perak) DK(Terengganu) DKNS(Negeri Sembilan) LHSultan SelangorBerkuasa21 November 2001 – sekarangPenobatan9 Maret 2003PendahuluSalahuddin Abdul Aziz ShahPutra MahkotaTengku Amir ShahInformasi pribadiKelahiran11 Desember 1945 (umur 78)Klang, SelangorAyahSultan Salahuddin Abdul Aziz ShahIbuRaja Saidatul Ihsan Tengku Badar ShahPasanganRaja Zarina R...

 

MeritamenPermaisuri MesirIstri Kerajaan AgungNyonya Dua TanahNyonya Hulu dan Hilir Mesir Putri Raja, kekasihnya, dll.Patung MeritamenKematianThebes?Pemakamanmakam QV68, Lembah Para Ratu, ThebesAyahRamses IIIbuNefertariPasanganFiraun Ramses IIAgamaagama Mesir Kuno Patung Meritamen di depan patung kolosal ayahandanya Ramses II Meritamen Era: Kerajaan Baru(1550–1069 BC) Hieroglif Mesir Meritamen (juga dieja Meritamun, Merytamen, Merytamun, Meryt-Amen; Bahasa Mesir Kuno: Kekasih Amun) meru...

Men's senior national football team of South Vietnam This article includes a list of general references, but it lacks sufficient corresponding inline citations. Please help to improve this article by introducing more precise citations. (March 2011) (Learn how and when to remove this template message) Republic of Vietnam1947–1976Nickname(s)Rồng vàng(Golden Dragon)AssociationVietnam Football AssociationConfederationAFC (Asia)Top scorerDucCu Sinh Le (6)Home stadiumCộng Hòa StadiumFIFA co...

 

Division of Air India Limited This article is about a defunct airline. For other airlines of India, see List of airlines of India. This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed.Find sources: Indian Airlines – news · newspapers · books · scholar · JSTOR (July 2012) (Learn how and when to remove this message) Indian Airli...

 

A wrap-style sandwich For other uses, see Roti (disambiguation). RotiA roti filled with curried goat and potatoesRegion or stateCaribbeanCreated bySackina Karamath in the mid-1940s in San Fernando, Trinidad and Tobago[1][2]Main ingredientsCurried meats or vegetables wrapped in a paratha, dhalpuri, or dosti roti  Media: Roti Karamath Roti Shop a.k.a. D' Humming Bird Roti Shop at Coffee Street in San Fernando, Trinidad and Tobago where the roti was invented Originating ...

House elections for the 60th U.S. Congress This article needs additional citations for verification. Please help improve this article by adding citations to reliable sources. Unsourced material may be challenged and removed.Find sources: 1906 United States House of Representatives elections – news · newspapers · books · scholar · JSTOR (January 2015) (Learn how and when to remove this message) 1906 United States House of Representatives elections ͛...

 

American actor For other people with the same name, see Jonathan Katz (disambiguation). Jonathan KatzKatz in 2006BornJonathan Paul Katz (1946-12-01) December 1, 1946 (age 77)New York City, New York, U.S.OccupationsActorcomedianWebsitewww.jonathankatz.com Jonathan Paul Katz (born December 1, 1946)[1] is an American actor and comedian best known for his starring role in the animated sitcom Dr. Katz, Professional Therapist as Dr. Katz.[2] He also is known for voicing Erik Ro...

 

This template does not require a rating on Wikipedia's content assessment scale.It is of interest to the following WikiProjects:African diaspora This template is within the scope of WikiProject African diaspora, a collaborative effort to improve the coverage of African diaspora on Wikipedia. If you would like to participate, please visit the project page, where you can join the discussion and see a list of open tasks.African diasporaWikipedia:WikiProject African diasporaTemplate:WikiProject A...

Северный морской котик Самец Научная классификация Домен:ЭукариотыЦарство:ЖивотныеПодцарство:ЭуметазоиБез ранга:Двусторонне-симметричныеБез ранга:ВторичноротыеТип:ХордовыеПодтип:ПозвоночныеИнфратип:ЧелюстноротыеНадкласс:ЧетвероногиеКлада:АмниотыКлада:Синапси...

 

Argentine diplomat and electrical engineer Susana MalcorraMalcorra in 2016Minister of Foreign AffairsIn office10 December 2015 – 12 June 2017PresidentMauricio MacriPreceded byHéctor TimermanSucceeded byJorge FaurieUnder Secretary-General of the United Nations for Field SupportIn office14 March 2008 – 1 March 2012Secretary-GeneralBan Ki-moonPreceded byJane LuteSucceeded byAnthony Banbury Personal detailsBorn (1954-11-18) 18 November 1954 (age 69)Rosario, ArgentinaPo...

 

American high school in Missouri St. Louis University High SchoolAddress4970 Oakland AvenueSt. Louis, Missouri 63110United StatesCoordinates38°37′41″N 90°16′01″W / 38.6281°N 90.2669°W / 38.6281; -90.2669InformationSchool typePrivate secondaryMottoReligioni et Bonis ArtibusReligion and the Fine ArtsAd Majorem Dei GloriamFor The Greater Glory of GodMen for OthersReligious affiliation(s)Roman Catholic,JesuitEstablished1818; 206 years ago (181...

John Sanborn PhillipsBorn(1861-07-02)July 2, 1861Council Bluffs, IowaDiedFebruary 28, 1949(1949-02-28) (aged 87)Goshen, New YorkEducation Knox College Harvard College OccupationPublisherSpouse Emma Delia West ​(m. 1885)​Children5RelativesSamuel Huntington (grandson) John Sanborn Phillips (1861–1949) attended Knox College in Illinois, where he worked on the student newspaper and met S. S. McClure. After earning an associate's degree, he entered Harvard Coll...

 

Pegunungan Himalaya, rangkaian jajaran gunung dilihat dari angkasa. Pegunungan Jayawijaya di Papua Pegunungan atau disebut juga banjaran dan barisan merupakan suatu area geografis berupa rentetan dari beberapa gunung-gunung atau bukit yang terkait secara geologis yang membentuk suatu deret atau bentangan.[1] Pegunungan yang dibatasi oleh dataran tinggi atau terpisah dari pegunungan lain dengan melewati punggung gunung atau lembah. Di Bumi, pegunungan biasanya terbentuk dari pergerakan...

 

Fort de la MontagneMontreal, Quebec Image of fort in front of Collège de MontréalCoordinates45°29′38″N 73°35′05″W / 45.4938°N 73.5846°W / 45.4938; -73.5846TypeFortSite informationControlled byNew France: QuebecSite historyBuilt1685In use1685-1854 National Historic Site of CanadaOfficial nameSulpician Towers / Fort de la Montagne National Historic Site of CanadaDesignated1970 Patrimoine culturel du QuébecTypeHistoric monumentDesignated1974 The ...

Gereja di Chelyabinsk Eparki Chelyabinsk adalah sebuah eparki Gereja Ortodoks Rusia yang terletak di Chelyabinsk, Federasi Rusia. Eparki tersebut didirikan pada tahun 1918.[1] Referensi ^ http://www.patriarchia.ru/db/text/136527.html lbsKeuskupan Gereja Ortodoks RusiaPatriark MoskwaEparki di Rusia Abakan dan Khakassia Akhtubinsk Alapayevsk Alatyr Alexdanrov Almetyevsk Amur Anadyr Ardatov Arkhangelsk Armavir Arsenyev Astrakhan Balashov Barnaul Barysh Belgorod Belyov Bezhetsk Birobidzha...

 

Romano FogliFogli al Catania nella stagione 1973-1974Nazionalità Italia Altezza172 cm Peso69 kg Calcio RuoloAllenatore (centrocampista) Termine carriera1974 - giocatore2000 - allenatore CarrieraGiovanili 1948-1955 Torino Squadre di club1 1955-1958 Torino49 (1)1958-1968 Bologna285 (6)1968-1970 Milan43 (5)1970-1974 Catania113 (3) Nazionale 1958-1967 Italia13 (0) Carriera da allenatore 1977-1979 BolognaVice[1]1980-1983 Reggiana1983-1984 Foggi...

 

Pour les articles homonymes, voir Académie impériale des Beaux-Arts, Académie des Beaux-Arts et Académie des sciences de Saint-Pétersbourg. Académie russe des beaux-artsHistoireFondation 1757StatutType AcadémieFondateur Ivan ChouvalovSite web en.rah.ruLocalisationPays RussieVille Saint-Pétersbourg Géolocalisation sur la carte : Saint-Pétersbourg (centre) modifier - modifier le code - modifier Wikidata L'Académie russe des beaux-arts, anciennement Académie impériale des beau...

Cet article est une ébauche concernant une localité de l'Alaska. Vous pouvez partager vos connaissances en l’améliorant (comment ?) selon les recommandations des projets correspondants. Kachemak SeloGéographiePays  États-UnisÉtat AlaskaCoordonnées 59° 47′ 28″ N, 151° 04′ 30″ OFonctionnementStatut Communauté non incorporée aux États-Unis (en)modifier - modifier le code - modifier Wikidata Kachemak Selo est une localité d'Alaska au...

 

ستار البامبو عام 1959. يعبر عن الستار باللون الأسود مع ملاحظة أن في ذلك الوقت كانت لاوس حليفةً للولايات المتحدة حيث لم يسيطر الباثيت لاو على البلاد حتى 1975. أيضًا لم يتوحد الشمال والجنوب الفيتنامي بعد. كذلك تعرض حدود الدول السوفيتية غير المستقلة وقتها للتوضيح. ستار البامبو هو �...