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Hominoids
Apes
Temporal range: Miocene-Holocene
Male chimpanzee (Pan troglodytes)
Scientific classification Edit this classification
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 (/ˌhɒmɪˈnɔɪdi.ə/), are a superfamily 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.

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 extant 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

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"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] 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

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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

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Main article: List of hominoids

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

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Further information: Human taxonomy § History

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

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See also: Hominidae and Human taxonomy
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

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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

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See also: Monkey § Historical and modern terminology

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

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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

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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

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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

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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.[49]

See also

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Notes

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References

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Revisions and contributorsEdit on WikipediaRead on Wikipedia

Ape

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from Grokipedia
Apes are tailless catarrhine of the superfamily Hominoidea, divided into lesser apes (family Hylobatidae, including and siamangs) and great apes (family , excluding the genus Homo), with approximately 22 extant non-human species distributed across and . Distinguished from other by the absence of a , a broadened , flexible shoulder joints facilitating brachiation and suspensory locomotion, a shortened spine, and larger brain-to-body ratios enabling advanced problem-solving, apes demonstrate notable adaptations for arboreal lifestyles, though great apes like gorillas exhibit increased terrestriality. Exhibiting complex social structures, vocalizations, and tool use—particularly among chimpanzees, who modify sticks for fishing and stones for nut-cracking—apes display cognitive capacities rivaling those observed in early , yet face existential threats from , , and , rendering all great ape species and most gibbon taxa threatened with extinction per IUCN assessments.

Terminology and Etymology

Definition and Scope

Apes constitute the superfamily Hominoidea within the suborder Catarrhini of Old World primates, distinguished by the absence of an external tail, broad noses, and adaptations for brachiation and suspensory locomotion. This superfamily encompasses two families: Hylobatidae, comprising the lesser apes such as gibbons and siamangs, which are smaller-bodied and highly arboreal; and Hominidae, which includes the great apes—orangutans, gorillas, chimpanzees, bonobos—and humans. Lesser apes typically weigh 5–12 kg and exhibit lighter builds suited to agile swinging through forest canopies, whereas great apes range from 30–180 kg with more robust frames supporting knuckle-walking or upright postures in some species. Biologically, humans belong to and share a with other great apes approximately 5–7 million years ago, rendering the vernacular category "apes" paraphyletic when humans are excluded, as it omits a descendant without capturing the full monophyletic group. This exclusion persists in common usage to differentiate humans from non-human hominoids, despite phylogenetic evidence confirming humans as apes under a cladistic . The term thus scopes non-human members of Hominoidea, focusing on 20–25 extant species across these families, all endangered due to habitat loss and . Geographically, apes are restricted to tropical and subtropical forests: Hylobatidae and orangutans (Pongo spp.) inhabit , including , , and parts of and ; while ( spp.), chimpanzees (Pan troglodytes), and bonobos (Pan paniscus) occupy Central and , from equatorial rainforests to montane habitats up to 4,000 elevation. No apes are native to the or , reflecting their origins in and subsequent dispersal.

Historical and Linguistic Origins

The English word "ape" derives from apa, which traces back to Proto-Germanic *apô, denoting a tailless and appearing in cognates across such as api and affo. Its ultimate origin remains uncertain, potentially linked to an Indo-European root implying or , reflecting early observations of apes' behavioral similarities to humans, or possibly onomatopoeic associations with their vocalizations; by the 13th century, compounds like "Martin Halfape" appear in English records, suggesting derogatory connotations of ugliness or brutishness. This Germanic term contrasted with later borrowings for tailed primates, as "" entered English around the from or Dutch via Romance influences like monne or Italian monno, highlighting a linguistic distinction between tailless "apes" and tailed "monkeys" that persisted in European usage. In ancient Greek texts, the term pithēkos referred to apes or monkeys, often evoking notions of mockery or deformity; , in his (circa 350 BCE), described the ape (pithēkos) as resembling humans in face and posture but quadrupedal otherwise, positioning it as an intermediate form sharing traits with both humans and other animals, while distinguishing the tailed "" (kēbos) as a variant. Roman authors adopted similar views, with Latin simia (from simus, "snub-nosed") applied to apes for their flat faces and imitative habits, as noted by in (77 CE), where apes were portrayed as cunning mimics prone to vanity and theft. During the medieval period, European bestiaries depicted apes as tailless symbols of and the devil—lacking a tail like the fallen —and as grotesque imitators of human folly, reinforcing moral allegories in illuminated manuscripts from the onward. The 18th-century Linnaean system formalized nomenclature in (1758), classifying humans (Homo sapiens) alongside ape-like genera such as Simia satyrus (, evoking mythical satyrs) within the order , prompting Linnaeus to challenge contemporaries by questioning bodily distinctions between humans and apes, though the English "ape" term retained its pre-Linnaean focus on non-human forms. This classification influenced English scientific usage, narrowing "ape" to denote non-human hominoids like chimpanzees and gorillas by the 19th century, amid debates on human-ape continuity sparked by Darwin's (1859). In the , taxonomic refinements explicitly excluded humans from "ape" in popular and cladistic contexts to underscore distinctions, as pre-1960 divisions separated Hominoidea into families like (great apes excluding humans) from (humans), a convention persisting in vernacular despite molecular evidence later integrating humans phylogenetically. This shift avoided anthropocentric blurring, with "ape" standardizing as non-human tailless in encyclopedias and texts by mid-century, reflecting a deliberate terminological boundary amid evolutionary insights.

Taxonomy and Phylogeny

Current Classification

Apes constitute the superfamily Hominoidea, divided into two extant families: Hylobatidae (lesser apes) and (great apes, excluding humans). The family Hylobatidae comprises four genera—, Hoolock, , and Symphalangus—encompassing 19 recognized of as of taxonomic revisions accounting for genetic and vocalization data. These are primarily delineated by differences in morphology (e.g., body size, fur coloration), chromosomal variations, and geographic distribution across Southeast Asian forests. Within , the subfamily includes the genus Pongo with three species: the (P. pygmaeus), (P. abelii), and (P. tapanuliensis), distinguished by genetic divergence, cranial morphology, and habitat isolation on and . The subfamily Gorillinae features the genus with two species: the (G. gorilla, including subspecies G. g. gorilla and G. g. diehli) and (G. beringei, including G. b. graueri and G. b. beringei); these are separated by over 1 million years of divergence evidenced in , alongside morphological traits like skull shape and body size, and geographic barriers such as the . The subfamily contains the genus Pan with two species: the common (P. troglodytes) and (P. paniscus), differentiated by genetics (e.g., fixed chromosomal inversions), pelage patterns, and the as a vicariant barrier. Species and subspecies boundaries in apes are determined through integrated criteria including morphological traits (e.g., skeletal robusticity, dentition), genetic markers (e.g., mitochondrial and nuclear DNA sequences showing divergence thresholds of 1-2% for species), and allopatric geography, though debates continue on thresholds for gorillas, where some analyses suggest eastern-western splits merit full species status due to reproductive isolation implications, while others emphasize gene flow potential.

Phylogenetic Relationships

The superfamily Hominoidea diverged from the superfamily Cercopithecoidea ( monkeys) approximately 25–30 million years ago during the epoch, marking the basal split within primates based on analyses and calibrations. This divergence is supported by phylogenetic reconstructions integrating genomic data, which place the common ancestor of apes and monkeys in Afro-Arabia. Within Hominoidea, the family Hylobatidae ( and siamangs) separated from (great apes and humans) around 15–20 million years ago in the early , as estimated from relaxed models calibrated with fossil priors. then underwent further branching, with the genus Pongo (orangutans) diverging basally from the lineage leading to African apes and humans approximately 12–16 million years ago. The gorilla lineage () split next, around 8–10 million years ago, followed by the divergence between the genus Pan (chimpanzees and bonobos) and the genus approximately 6–7 million years ago. Cladistic analyses consistently recover Hominidae as monophyletic, with orangutans as the outgroup to a clade comprising gorillas, chimpanzees, bonobos, and humans; the latter three form the Homininae subfamily, underscoring closer genetic affinity among African apes and humans relative to Asian apes. Humans share about 98.5–98.8% DNA sequence identity with chimpanzees, their closest relatives, but these figures primarily reflect nucleotide substitutions while underemphasizing structural variants, insertions/deletions, and regulatory differences that drive profound functional divergences in morphology, cognition, and behavior.

Historical Developments in Taxonomy

In the 10th edition of Systema Naturae published in 1758, Carl Linnaeus classified apes within the order Primates, placing them under the genus Simia alongside monkeys, while grouping humans as the genus Homo in the same order; this arrangement reflected morphological similarities such as taillessness and upright posture but subordinated apes to humans without recognizing a distinct superfamily for tailless primates. Earlier editions had used the term Anthropomorpha for a broader group including humans, apes, sloths, and bats, emphasizing superficial resemblances to humans, though Linnaeus relied on limited, often second-hand descriptions of ape anatomy. This initial framework treated apes as a heterogeneous assemblage lacking precise delineation from monkeys, prioritizing descriptive traits over phylogenetic inference. By the 19th century, taxonomic separations emerged based on enhanced anatomical studies, with British zoologist establishing the family in 1840 to encompass great apes (Pongo, , and later Pan), distinct from the human-exclusive ; this reflected observations of shared arboreal adaptations but maintained humans in a separate family due to bipedalism and brain size differences. further refined classifications in the 1860s, proposing suborders for catarrhines including apes, yet debates persisted over whether morphological convergences, such as in African apes, warranted closer human-ape linkage or reinforced separation. Twentieth-century taxonomy grappled with human inclusion amid fossil discoveries like (1924 onward), prompting debates on whether great apes should join or remain in ; morphological favored exclusion until molecular data intervened. In 1967, Vincent Sarich and Allan Wilson's immunological comparisons of blood proteins introduced molecular clocks, estimating human-chimpanzee divergence at approximately 5 million years ago—closer than chimpanzee-gorilla splits—challenging morphology-based distances and supporting ape excluding . These findings fueled revisions, culminating in the 1980s-1990s consensus to dissolve and subsumed great apes into as subfamilies (e.g., for orangutans, Gorillinae, for humans and African apes), driven by DNA hybridization and sequence data over traditional metrics. Post-2000 genomic advancements refined ape , with formally recognized as the distinct family Hylobatidae since Gray's 1870 proposal but bolstered by molecular phylogenies confirming their early divergence around 15-18 million years ago. Haplotype-resolved assemblies in the 2020s, including complete telomere-to-telomere sequences for chimpanzees, bonobos, , and orangutans published in 2024, have validated boundaries—such as Pan troglodytes schweinfurthii versus P. t. troglodytes—through divergence timing (e.g., human-chimp split at 5.5-6.3 million years ago) and diversity, shifting emphasis from gross morphology to genetic coalescence and selection pressures. These refinements underscore ' superiority in resolving cryptic variation, though morphological data retains utility for integration.

Evolutionary History

Fossil Record and Origins

The earliest known hominoids, representing the basal radiation of the ape lineage, appeared in the early Miocene epoch, approximately 23 to 17 million years ago, primarily in East Africa. The genus Proconsul, discovered in sites such as those in Kenya and Uganda, exemplifies these primitive forms, with species like P. africanus and P. heslon exhibiting tailless bodies, a broad thoracic cage, and adaptations for arboreal quadrupedalism combined with suspensory locomotion, such as flexible shoulder joints and long forelimbs. These traits mark a departure from cercopithecoid monkeys, though Proconsul retained some primitive features like convergent incisors and lacked the specialized brachiation seen in later apes. By the middle , around 16 to 11 million years ago, hominoid diversity increased, with taxa dispersing into and showing more derived ape-like morphologies. In Europe, , known from sites in and dated to about 12.5 to 11.1 million years ago, displayed thin , a Y-5 molar pattern, and postcranial evidence of below-branch suspension, suggesting affinities to the great ape clade. Concurrently, in , Sivapithecus from the , approximately 12.5 to 8.6 million years old, featured a short face and thick molars akin to modern orangutans, supporting its role as an early pongine (Pongo lineage) ancestor. forms, such as Nakalipithecus nakayamai from around 10 million years ago, further illustrate this diversification, with robust jaws and thick-enameled teeth indicating a great ape-like dietary to tougher . The fossil record of non-human great apes becomes markedly sparse from the (5.3 to 2.6 million years ago) through the Pleistocene (2.6 million to 11,700 years ago), contrasting with the abundance of early hominins like in open habitats. This scarcity stems from the apes' persistence in environments, where acidic soils, high humidity, and rapid organic decay hinder fossilization, compounded by geological processes like lixiviation and erosion that destroy potential remains before preservation. Transitional fossils bridging hominoids to extant great ape genera remain limited, underscoring gaps in the record despite evidence of a last common ancestor with humans around 9 to 6.5 million years ago.

Molecular and Genetic Evidence

Molecular analyses employing and nuclear gene sequences, calibrated via molecular clocks, estimate the initial radiation of Hominoidea around 25 million years ago (mya), marking the divergence from cercopithecoids. These clocks account for rate variations across lineages, with nuclear DNA providing more precise calibrations than mitochondrial alone due to reduced saturation effects in deeper divergences. Whole-genome sequencing has further refined great ape splits: orangutans diverged from African apes approximately 12-16 mya, gorillas from the chimpanzee-bonobo-human around 8-10 mya, and chimpanzees/bonobos from humans at 5.5-6.3 mya. Advancements in 2025 produced haplotype-resolved, telomere-to-telomere genome assemblies for chimpanzees, bonobos, , Bornean and Sumatran orangutans, and siamangs, enabling detailed reconstruction of phased haplotypes without human contamination. These assemblies reveal mechanisms, including recurrent structural variations and recent turnover in loci like 17q21.31, which exhibit inverted haplotypes in chimpanzees relative to and orangutans. Comparative analyses highlight regulatory variations driving lineage-specific adaptations, such as elevated segmental duplications in chimpanzees, bonobos, and —exceeding those in humans—potentially linked to immune and neural traits. Genomic scans of in great apes identify footprints of in genes influencing locomotion and skeletal morphology, with African apes showing signatures in loci associated with via eccentric muscle contractions and wrist stabilization. exhibit distinct regulatory enhancements for brachiation, reflected in expanded gene families for strength and mobility, though independent of locomotor traits underscores over shared ancestry in some cases. Endangered populations, including mountain gorillas and Tapanuli orangutans, display critically low —comparable to inbred isolates—with coefficients exceeding 0.2 in some groups, elevating risks of deleterious mutations and reduced fitness. These patterns, quantified via heterozygosity metrics below 0.001 in bottlenecked lineages, inform conservation by prioritizing to mitigate load accumulation.

Physical Characteristics

Morphology and Anatomy

Apes, or members of the superfamily Hominoidea, are characterized by the absence of an external tail, a feature that sets them apart from monkeys and reflects adaptations to suspensory locomotion rather than . Their skeletons typically feature relatively short trunks, broad chests, elongated arms relative to legs, and long hands suited for grasping and swinging (brachiation in lesser apes) or knuckle-walking (in great apes). These proportions facilitate arboreal suspension and terrestrial , with arm lengths often exceeding leg lengths by 10-20% in species like chimpanzees and . Body sizes vary widely across ape taxa, from the smallest hylobatids (gibbons) weighing 5-12 kg to large great apes like male gorillas exceeding 170 kg, enabling diverse ecological niches from forest canopies to ground foraging. Sexual dimorphism is pronounced, particularly in great apes, where males average 1.5-2 times the body mass of females—e.g., adult male chimpanzees weigh 40-60 kg compared to 30-50 kg for females—linked to intrasexual competition. The follows the catarrhine formula of 2.1.2.3 (two incisors, one canine, two premolars, three molars per quadrant, totaling 32 teeth), with broad incisors and reduced canines relative to body size compared to earlier . Great apes possess robust jaws and larger molars adapted for processing tough, fibrous , though dental wear patterns vary by diet. Sensory anatomy emphasizes vision over olfaction: forward-facing eyes provide stereoscopic essential for navigating complex three-dimensional environments, while the is diminished, with fewer functional receptor genes than in strepsirrhine . This shift correlates with increased reliance on visual cues for foraging and social interactions.

Distinctions from Monkeys and Other

Apes differ from s in lacking an external , a feature present in most that aids in balance during quadrupedal locomotion. This absence in apes facilitates greater flexibility in suspensory behaviors, such as brachiation, where the would otherwise hinder swing. Prosimians, like lemurs and lorises, retain tails but exhibit more primitive grasping adaptations suited to vertical clinging and leaping rather than the sustained suspension seen in apes. In terms of locomotor anatomy, apes possess highly mobile joints with a broad, cranially oriented that enables extensive rotation and overhead arm positioning, contrasting with the narrower, more laterally positioned scapulae of monkeys adapted for pronograde . This configuration in apes supports weight suspension from above, reducing reliance on propulsion during arboreal travel, whereas monkeys' morphology prioritizes stable, ground-oriented . Prosimians display even less mobility, with scapulae geared toward leaping rather than prolonged . Apes exhibit larger brain-to-body size ratios compared to monkeys, with relative encephalization quotients higher in hominoids, reflecting adaptations for complex spatial problem-solving in varied arboreal niches. Monkeys, occupying more and folivorous roles, maintain smaller relative brain sizes suited to predictable patterns. Prosimians have the lowest ratios among , correlating with simpler sensory-motor demands in nocturnal, insectivorous lifestyles. Dentally, apes feature Y-5 molar patterns with five cusps arranged in a Y-shape, optimized for shearing fibrous fruits and leaves in canopy environments, distinct from the bilophodont molars of monkeys that form transverse ridges for grinding tougher vegetation. New World monkeys vary but lack this precise configuration, while prosimians retain more primitive, multi-cusped molars without the derived Y-5 shearing efficiency. Ecologically, apes lack specialized features like cheek pouches for , common in some monkeys for opportunistic caching during terrestrial , and ischial callosities for prolonged ground sitting, which monkeys use in habitats. Instead, apes' niches emphasize suspensory access to high-canopy resources, differing from monkeys' versatile across arboreal and terrestrial zones, and prosimians' niche in insectivory with wet noses for scent detection.

Behavior and Ecology

Social Structures and Group Dynamics

, the lesser apes, typically live in small, stable family units composed of a monogamous and their immature , ranging from 2 to 6 individuals. These units maintain exclusive territories defended through coordinated singing, primarily by adults, which serves to advertise pair bonds and deter intruders. Among the great apes, social structures diverge significantly by species. Chimpanzees (Pan troglodytes) form large communities of 20 to 150 individuals exhibiting fission-fusion dynamics, where the group splits into temporary parties of 3 to 10 members for and reconvenes at night. Male chimpanzees remain in their natal community, forming linear dominance hierarchies enforced through aggressive displays, coalitions, and occasional lethal violence, while females typically disperse at to avoid . Bonobos (Pan paniscus) also organize in multi-male, multi-female communities with fission-fusion patterns, but feature stronger female bonds and matrifocal structures where coalitions of related females hold higher status than males. Group sizes vary similarly to chimpanzees, with interactions emphasizing affiliation over aggression, though males form kin-based alliances for status. Females disperse from natal groups, promoting . Gorillas (Gorilla spp.) reside in cohesive, harem-like troops averaging 5 to 30 members, led by a dominant silverback male who mates with multiple females (typically 3 to 6) and their offspring. The silverback maintains cohesion through displays and protects against predators and rivals; females often transfer between groups, while young males may leave to form bachelor groups or challenge for leadership. occurs when a new silverback assumes control, killing unrelated infants to eliminate future competitors and resume female reproduction sooner. Orangutans (Pongo spp.) exhibit semi-solitary organization, with flanged adult males ranging independently over large territories and unflanged males or females associating transiently, mainly mothers with dependent young for 6 to 8 years. Social interactions are infrequent and opportunistic, lacking stable groups, though males may consort with estrous females briefly. Dominance in ape groups is generally established via physical displays, vocalizations, and rather than constant conflict, with coalitions enhancing status in chimpanzees and bonobos. by unrelated males is documented in chimpanzees and , accelerating interbirth intervals by ending amenorrhea, though rarer in bonobos and absent in orangutans. , where non-mothers assist in care, occurs in and chimpanzees, fostering group cohesion, while manifests in defense against predators or rivals, particularly in chimpanzees. Sex-biased dispersal predominates, with female transfer common across species to mitigate , except in where both sexes may disperse.

Diet, Foraging, and Habitat Adaptation

Apes exhibit primarily plant-based diets dominated by frugivory, with comprising 50-80% of intake in many , supplemented by foliage, , bark, , and occasionally other animal matter. This composition reflects to habitats where ripe patches provide high-energy resources, though dietary flexibility allows shifts to lower-quality fallback foods like mature leaves and herbs during seasonal scarcity to maintain energy balance. Such strategies prioritize nutrient-dense patches, minimizing travel costs while exploiting spatiotemporal availability, as evidenced by selective in high-quality arboreal sources. Chimpanzees (Pan troglodytes) display the most opportunistic diets among great apes, with fruit at 50-75% but including significant animal protein from insects (up to 10%) and hunted vertebrates like colobus monkeys, alongside leaves and pith. Foraging involves tool-assisted extraction, such as modifying sticks for termite fishing from epigeal mounds, where probes are inserted to withdraw adherent termites, enhancing caloric intake during lean periods. In African rainforests, this enables exploitation of understory insects inaccessible without tools, linking habitat structure—dense undergrowth and termite nests—to specialized behaviors. Gorillas (Gorilla spp.) are folivore-frugivores, with western lowland populations consuming up to 230 plant items but favoring herbaceous vegetation (40-60%) over (15-25%), contrasting mountain gorillas' lower frugivory (<5% ). They process fibrous foods via fermentation, adapting to central African rainforests rich in herbs and shoots as fallback staples when phenology declines, thus buffering against seasonal gaps in preferred ripe fruits. emphasizes ground-level in clearings, with selective intake of protein-rich stems and avoidance of , optimizing digestion in habitats with variable productivity. Orangutans (Pongo spp.), particularly Bornean populations, rely heavily on fruit (60-80%) in swamp forests, where low nutrients yield unpredictable mast fruiting events, prompting fallback to bark, leaves, and during inter-mast periods. These habitats, with acidic and sparse canopies, necessitate arboreal travel adjustments—shorter strides and compliance—to access dispersed resources, sustaining energy via prolonged suspension feeding. Gibbons (Hylobatidae), as lesser apes, maintain frugivorous diets with at ~55%, leaves ~25%, and flowers/seeds supplementing in Southeast Asian dipterocarp forests. Foraging centers on terminal branch feeding for pulpy , with daily patterns prioritizing fruit-rich breakfast trees to plan brachiation routes, adapting to canopy gaps by favoring energy-maximizing patches over uniform depletion. Seasonal fallback to leaves sustains amid shortages, aligning with habitats of emergent trees and lianas that support suspensory locomotion for efficient harvest.

Locomotion and Daily Activities

Gibbons, as lesser apes, exhibit highly specialized dominated by brachiation, a form of arm-swinging suspensory movement that constitutes more than 50% of their active time, enabling efficient travel through the forest canopy. This mode relies on elongated forelimbs and flexible shoulder joints adapted for continuous swinging between branches, with spending the vast majority of their time in trees. In contrast, great apes employ for terrestrial quadrupedal progression, supporting body weight on the dorsal surfaces of flexed fingers, a trait prominent in African species like chimpanzees, bonobos, and . Arboreally, great apes engage in clambering and , utilizing powerful upper body strength to navigate larger supports and vertical trunks, though less suspensory than . Apes are predominantly diurnal, with activity patterns structured around morning travel and foraging followed by midday resting to conserve energy in tropical environments, allocating roughly 30-50% of daylight hours to rest depending on species and resource availability. Daily travel distances vary by and diet but typically range from 3-5 km for most great apes, with chimpanzees occasionally covering up to 10 km in fruit-scarce periods to track dispersed resources. Nocturnal activity is rare among apes, though some crepuscular behaviors occur or dusk for group movement or predator avoidance in species like orangutans. Great apes routinely construct new nests each evening from branches and leaves for overnight sleeping, a that enhances by abandoning old sites and provides elevated from ground predators. This nightly rebuilding, observed across chimpanzees, , and orangutans, consumes 1-2 hours of late-afternoon activity and reflects adaptations for arboreal safety without permanent shelters. Low-activity periods, often midday siestas in shade, minimize metabolic costs in high-humidity forests where demands energy efficiency.

Cognition and Intelligence

Tool Use and Problem-Solving

Chimpanzees (Pan troglodytes) exhibit the most extensive and varied tool use among apes in the wild, including probe tools modified from twigs or sticks for extracting from mounds, a behavior first documented in 1960 at Gombe Stream National Park by . These tools often involve sequential use of a perforating stick followed by a fishing probe, with individuals selecting and modifying stems based on length, flexibility, and frayed tips for optimal efficacy, as observed in the Ndoki Forest. Nut-cracking with stone hammers and anvils is prevalent in West African populations, such as at , where chimpanzees select heavy stones (averaging 2-7 kg) suited to nut hardness and reuse sites over years, leaving archaeological traces of durable tools and fragments. Tool repertoires vary culturally across communities, with over 30 distinct behaviors transmitted socially rather than genetically, as evidenced by absence in some groups despite similar habitats. Orangutans (Pongo spp.) demonstrate tool use primarily in Sumatran populations at sites like Suaq Balimbing, where they fashion sticks to extract from tree holes or use leaves as gloves to handle irritant fruits and as impromptu umbrellas during rain. Wild individuals have been observed improvising shelters by weaving branches or using tools for seed extraction, with innovations like hanging tools for future reuse documented in 2018. These behaviors occur at higher frequencies in resource-rich swamp forests but are less common in Bornean orangutans, reflecting ecological opportunities rather than cognitive deficits. Tool use in (Gorilla spp.) is rarer in the wild, with the first verified instance in 2005 involving a western lowland female using a as a probe to gauge swamp water depth before crossing, followed by occasional ant-fishing or stick-testing in mountain . (family Hylobatidae), being strictly arboreal and adapted to fruit-abundant forests, show minimal tool use, limited to rare captive observations of manipulation for food extraction, attributable to reduced selective pressure in their where manual dexterity suffices. In captive settings, apes like chimpanzees innovate tool solutions to novel puzzles, such as modifying objects into metatools or overcoming traps, with early proficiency in simple manipulations but challenges in complex sequences mirroring wild constraints on elaboration. Wild tool complexity remains bounded by immediate ecological demands, with no sustained accumulation observed.

Learning, Memory, and Communication

Chimpanzees demonstrate superior for numerical sequences compared to adult s in tasks requiring rapid recall of briefly presented digits. In experiments conducted by Tetsuro Matsuzawa at University's Primate Research Institute, young chimpanzees such as Ayumu remembered the positions and order of nine numerals flashed for 200 milliseconds, outperforming human participants who required longer exposure times. This capability persists in trained individuals like Ai, the first chimpanzee to use symbolically, achieving errorless performance in sequencing up to nine items after extensive training starting in 1979. in chimpanzees also excels, with individuals forming long-term recollections of food cache locations after minimal exposure, as evidenced by field studies in Uganda's Budongo Forest where subjects relocated hidden rewards with high accuracy after delays of up to nine days. Social learning in apes relies heavily on imitation rather than individual trial-and-error, particularly in great apes raised in human-like environments. Enculturated chimpanzees and orangutans exhibit deferred imitation, reproducing observed actions after delays of up to 24 hours, a process distinct from asocial learning as it involves selective copying of demonstrator behaviors. Mirror self-recognition, a marker of self-awareness tested via the mark procedure, occurs reliably in great apes including chimpanzees, orangutans, and some gorillas, who touch marked body parts visible only in reflection, but fails in lesser apes like gibbons, indicating a cognitive threshold tied to encephalization rather than phylogenetic proximity to humans. Ape communication features intentional and vocalizations that convey context-specific meanings without syntactic structure. Great ape gestures, such as arm extensions for play invitations or ground slaps for copulation requests, exhibit first-order : producers adjust signals based on recipient attention and response, persisting or desisting accordingly across species like and bonobos. Vocal signals, including chimpanzee pant-hoots for group coordination or alarm calls differentiated by predator type, function referentially but lack recursive syntax or combinatorial rules observed in , relying instead on innate or learned repertoires limited to about 30-60 gesture types per individual. These systems prioritize immediate social goals over propositional content, with empirical playback studies confirming recipients respond appropriately to gesture without of displaced .

Comparative Assessments with Humans

Apes demonstrate episodic-like memory, as evidenced by chimpanzees recalling the location, content, and timing of past events, such as distinguishing food caches hidden at different intervals and revisiting them accordingly over periods exceeding a year. This capacity integrates "what," "where," and "when" elements, mirroring aspects of human but lacking the autonoetic awareness of re-experiencing subjective past states. Proxies for appear in apes through behaviors like gaze-following and tactical , where chimpanzees infer others' attentional states to hide resources or compete effectively. However, empirical tests reveal limitations; apes rarely pass stringent false-belief tasks requiring attribution of unobservable mental states, succeeding primarily on observable cues rather than full representational understanding, unlike consistent human performance from age four. Apes exhibit cultural transmission of behaviors, such as nut-cracking techniques in chimpanzees, yet lack cumulative culture, where innovations ratchet into increasingly complex forms across generations; experimental assessments show no progressive efficiency gains in tool use, with successes attributable to individual or simple rather than modification of prior techniques. Abstract reasoning remains constrained, with no evidence of symbolic manipulation or hypothetical beyond immediate contexts. Genetic differences underpin vocalization gaps; the gene in humans features two amino acid substitutions absent in chimpanzees, bonobos, and , correlating with enhanced fine for articulate speech and vocal learning, which apes do not exhibit despite shared core sequence conservation. prefrontal cortex shows disproportionate expansion relative to body size compared to apes, particularly in frontopolar regions supporting like planning and integration of abstract information, enabling capabilities beyond ape domain-specific adaptations. Ape prosocial behaviors, often labeled as , align with and mechanisms, prioritizing genetic relatives or exchange partners without evidence of impartial judgment or rule-based independent of immediate fitness benefits. Intelligence in apes is modular, excelling in spatial navigation or tool manipulation tasks but failing generalization across unrelated domains, contrasting fluid intelligence that transfers principles broadly.

Reproduction and Life History

Mating Systems and Parental Care

Ape mating systems exhibit significant variation across species, shaped by ecological pressures and in wild populations. Chimpanzees (Pan troglodytes) and bonobos (Pan paniscus) employ promiscuous multi-male/multi-female strategies, where females with multiple males during periodic estrus, signaled by conspicuous anogenital swellings that peak in size near and attract male attention, thereby facilitating female choice amid male . This promiscuity generates high paternity uncertainty, prompting males to pursue frequent copulations to maximize potential rather than targeted guarding. In contrast, (Gorilla spp.) feature polygynous harems dominated by a single silverback male who monopolizes access to females through coercion, including forced s and of unrelated infants upon group takeover to hasten female fertility, ensuring higher paternity certainty for the resident male. Orangutans (Pongo spp.) display semi-solitary opportunistic , with unflanged males often coercing dispersed females, while ( spp.) form long-term pair bonds approximating , minimizing overt male . Reproductive investment centers on extended maternal care, as ape gestation periods range from 7 to 9 months across great ape species, typically yielding a single due to the high energetic costs of large neonatal size and development. Infants are born altricial, clinging to the for transport and , with occurring between 3 and 5 years in chimpanzees and , extending to 6-8 years in orangutans, during which females forgo additional breeding to support survival in resource-variable habitats. Paternal remains minimal in most species, limited by paternity uncertainty in promiscuous systems or the absence of stable male-female associations, though gibbon pairs may involve biparental provisioning. Infanticide risks, particularly from incoming males in and chimpanzees, drive female counterstrategies like accelerated post-takeover to confuse paternity and induce male tolerance of existing young.

Growth, Development, and Longevity

Apes display extended ontogenetic trajectories relative to most mammals, featuring prolonged infancy, juvenility, and that support extended and behavioral learning. Weaning generally occurs between 3 and 5 years in chimpanzees and , while orangutans extend this to approximately 7.7 years, reflecting species-specific adaptations in maternal care duration. Following , a multiyear juvenile phase persists until onset, succeeded by marked by physical maturation and , with full adulthood delayed until skeletal and reproductive completion. Sexual maturity emerges between 7 and 15 years across great ape species, varying by sex and ecology; females typically mature earlier than males, with chimpanzees reaching reproductive viability around 10-13 years after a post-weaning interval exceeding a . This slow maturation rate, characterized by bimaturism in some species where males grow faster post-puberty, contrasts with faster-developing monkeys and underscores apes' investment in extended dependency for acquiring complex and . Brain development in apes involves protracted cerebral tissue maturation through prepuberty, with prefrontal volume increasing gradually beyond onset, facilitating akin to but less extended than in s. Prolonged juvenility correlates positively with neocortical expansion across , enabling extended learning periods that underpin social complexity without the extreme delays seen in . Lifespans in apes exceed those of most monkeys, with wild individuals entering after 30 years and rarely surpassing 50 due to predation, disease, and resource scarcity. Captive conditions extend through veterinary care and nutrition, though differences persist.
SpeciesWild Average Lifespan (years)Wild Maximum (years)Captive Lifespan (years)
33-38~5040-60+
35-40~5050+
35-45~5050-60+

Conservation and Threats

Population Status and Endangerment

All species of apes are threatened with , with great apes (family ) classified by the International Union for Conservation of Nature (IUCN) as either Endangered or Critically Endangered, reflecting severe population declines driven by historical and ongoing pressures. Lesser apes (family Hylobatidae, ) range from Vulnerable to Critically Endangered, with 19 of 20 species in the latter two categories. Population estimates derive primarily from field surveys using methods such as line transects, nest counts for density calculations, and camera traps, compiled in databases like the IUCN APES (Ape Populations, Environments, and Surveys) repository; however, significant data gaps persist in remote, unsurveyed forest regions, leading to conservative figures that likely underestimate totals. The 2023–2025 edition of in Peril, produced by the IUCN Species Survival Commission Specialist Group and partners, underscores drastic declines across ape taxa, listing multiple species—including the and —among the world's 25 most endangered . For great apes, global estimates indicate fewer than 500,000 individuals across all taxa, with subpopulations fragmented and isolated. Chimpanzees (Pan troglodytes) number 170,000–300,000 wild individuals and are listed as Endangered. Bonobos (Pan paniscus), also Endangered, persist at 15,000–20,000 individuals, confined to the Democratic Republic of Congo. Gorilla subspecies exhibit extreme variation: western lowland gorillas ( gorilla gorilla) at approximately 316,000 (Critically Endangered), eastern lowland or Grauer's gorillas ( beringei graueri) at about 3,800 (Critically Endangered), mountain gorillas ( beringei beringei) at 1,063 (Critically Endangered), and Cross River gorillas ( gorilla diehli) at fewer than 300 (Critically Endangered). Orangutan species are all Critically Endangered, with Bornean orangutans (Pongo pygmaeus) at around 104,700, Sumatran orangutans (Pongo abelii) at 13,846, and Tapanuli orangutans (Pongo tapanuliensis) at about 800.
Species/SubspeciesIUCN StatusEstimated Wild Population
Chimpanzee (Pan troglodytes)Endangered170,000–300,000
Bonobo (Pan paniscus)Endangered15,000–20,000
Western lowland gorilla (G. g. gorilla)Critically Endangered~316,000
Eastern lowland/Grauer's gorilla (G. b. graueri)Critically Endangered~3,800
Mountain gorilla (G. b. beringei)Critically Endangered1,063
Cross River gorilla (G. g. diehli)Critically Endangered<300
Bornean orangutan (P. pygmaeus)Critically Endangered~104,700
Sumatran orangutan (P. abelii)Critically Endangered~13,846
Tapanuli orangutan (P. tapanuliensis)Critically Endangered~800
Gibbon populations are less aggregated due to taxonomic diversity and across , but totals likely number in the low hundreds of thousands, with many species reduced to dozens or hundreds in isolated patches. Critically Endangered examples include the Hainan black-crested gibbon (Nomascus hainanus) at about 30 individuals and the Cao Vit or eastern black-crested gibbon (Nomascus nasutus) at 74. Density estimates from auditory surveys and direct observations reveal patchy distributions, with ongoing declines noted in the Primates in Peril report, which flags several gibbon taxa for their extreme rarity and limited survey coverage in border regions.

Primary Threats from Human Activities

Habitat destruction driven by agricultural expansion, commercial logging, and mining constitutes a leading anthropogenic threat to ape populations, affecting over 60% of primate species through deforestation. In great ape habitats across Africa and Southeast Asia, these activities fragment forests and reduce available range, with agriculture alone implicated in threats to 76% of primate species and logging to 60%. For instance, palm oil plantations and subsistence farming have accelerated forest clearance in orangutan habitats in Borneo and Sumatra, while mining operations in central Africa exacerbate gorilla and chimpanzee range contraction. Bushmeat hunting represents a severe direct threat, particularly in central and , where it surpasses habitat degradation as the dominant pressure on ape populations due to high demand for protein in growing human communities. Great apes, with their slow reproductive rates, suffer disproportionate impacts from sustained harvesting, as evidenced by widespread depletion in logged forests where access roads facilitate hunters. In , hunting pressure correlates strongly with proximity to human settlements and logging concessions, leading to local extirpations of chimpanzees and . Poaching for the illegal pet trade and live animal commerce further diminishes ape numbers, with an estimated 22,218 great apes lost to trafficking between 2005 and 2011 alone, equivalent to over 3,000 annually. Infants are often targeted after mothers are killed, disrupting social structures and reducing recruitment; this trade persists despite listings, fueled by demand in affluent markets for exotic pets. Chimpanzees and are particularly vulnerable, with seizures indicating underreported volumes nearly nine times higher than official records. Human encroachment heightens disease transmission risks to apes, as activities like and habitat invasion bring pathogens into contact with immunologically naive populations. virus outbreaks have decimated and groups, with human-mediated spillover suspected in some cases due to shared interfaces; apes serve as amplifying hosts rather than reservoirs, but proximity amplifies cross-species jumps. Respiratory viruses from habituated apes in tourism sites also illustrate bidirectional risks, though apes face higher mortality from human-borne strains. Climate change compounds these pressures by inducing habitat shifts and altering resource availability, projecting up to 94% range loss for African great apes by 2050 under combined scenarios. Rising temperatures and altered rainfall patterns may render current forests unsuitable, forcing potential elevational or latitudinal migrations that apes' limited dispersal capacity hinders. Synergistically, from human activities curtails , while correlations with human amplify encroachment, as denser settlements drive intensified resource extraction.

Conservation Strategies and Outcomes

Protected areas, such as national parks and reserves, form the cornerstone of great ape conservation, with patrols proving effective in reducing illegal hunting within these zones; for instance, ranger-based monitoring has curtailed poaching-related threats in African habitats. In , legal protections including designations and laws have supported localized efforts, though enforcement remains inconsistent outside formal boundaries. These interventions have yielded positive biological outcomes, such as stabilized local populations in well-patrolled sites, but broader empirical data indicate they insufficiently counter ongoing declines driven by external pressures. Rewilding and reintroduction trials for apes have produced mixed results, often hampered by high post-release mortality. releases, including attempts with Grauer's and mountain subspecies, recorded failure rates exceeding 66%, with four of six individuals dying shortly after translocation due to predation, , or failures. Soft-release techniques, involving pre-release acclimation, have improved translocation success by up to 77% in movement-based metrics compared to hard releases, yet ape-specific applications remain limited and empirically unscaled for population recovery. Advancements in genomic sequencing, including haplotype-resolved reference genomes for six ape species published in April 2025, enable better assessment of and inbreeding risks, potentially enhancing programs by guiding mate selection to bolster viability. Private sector collaborations, such as grants from foundations like Arcus, fund holistic initiatives integrating habitat protection with community incentives, contributing to incremental gains in site-specific ape survival rates as of 2024. Critiques highlight the limited efficacy of these strategies against , which fragments ape habitats and intensifies human- conflict; across African and Southeast Asian ranges, great apes coexist with approximately 97 million people, equating to one ape per 77-129 individuals, underscoring inherent trade-offs between conservation and human economic needs. Empirical analyses emphasize that without redirecting agricultural development away from high-priority ape areas, protected zones and patrols yield only marginal long-term outcomes, as loss persists and local communities prioritize over preservation. Overall, while interventions have averted total collapse in select locales, continental trends show persistent population reductions, with African down 76% over the past 50 years despite escalated efforts.

Human Interactions and Controversies

Role in Scientific Research

Chimpanzees have played a pivotal role in biomedical research, particularly in development for human diseases. Jonas Salk's cultivation of the inactivated in the 1950s relied on chimpanzee kidney cells, following experiments with over 100 s to identify non-neurovirulent strains safe for human use. Similarly, Albert Sabin's oral development incorporated chimpanzee testing to ensure intestinal replication without paralysis. For , chimpanzees served as the primary model due to their susceptibility mirroring humans, enabling the creation of plasma-derived vaccines that demonstrated protection against high-dose viral challenges, ultimately preventing millions of infections worldwide. Genomic sequencing of ape species has yielded insights into human disease mechanisms and . The chimpanzee , initially sequenced in 2005 with approximately 95% alignment to s, revealed mutations linked to traits like brain development and immunity. In April 2025, haplotype-resolved, telomere-to-telomere assemblies of six ape genomes—, , , and orangutans—enhanced comparative analyses, identifying segmental duplications associated with human-specific adaptations and disease susceptibilities, such as neurological disorders. These advancements facilitate modeling of genetic parallels, aiding research into conditions like Alzheimer's and infectious diseases without relying solely on invasive methods. Field studies of ape behavior have advanced understanding of , social structures, and tool use, informing evolutionary models. Jane Goodall's observations at Gombe Stream National Park from 1960 documented chimpanzees fashioning tools from twigs to extract termites, challenging prior assumptions of human uniqueness in this domain, and revealed predatory hunting and intergroup conflicts. Dian Fossey's long-term monitoring of mountain gorillas in Rwanda's Virunga region from 1967 provided foundational data on their , , and familial bonds, elucidating parallels to human systems. Contemporary non-invasive techniques, including camera traps, drones, and fecal genetic sampling, have sustained these behavioral insights while minimizing disturbance, supporting ongoing studies of wild populations. Regulatory shifts, such as the European Union's 2010 ban on great ape experimentation and the U.S. National Institutes of Health's 2015 retirement of most invasive protocols, have curtailed such uses, yet prior ape has yielded disproportionate benefits, including vaccines averting epidemics and genomic tools accelerating therapeutic development.

Ethical Debates and Research Restrictions

Ethical debates surrounding the use of apes in scientific research center on balancing potential benefits against concerns, with proponents arguing that apes' close genetic and physiological similarities to s provide irreplaceable models for studying complex neurological disorders and infectious diseases. In , great apes like s have facilitated insights into function and that smaller mammals cannot replicate due to differences in cortical structure and cognition, enabling translational advances such as understanding and relevant to conditions like . Similarly, historical studies contributed to early understandings of and potential models, where alternatives fail to mimic human-like immune responses. Advocates maintain that non-invasive alternatives, including computational models or organoids, remain incomplete for validating therapies requiring whole-organism testing, emphasizing that ethical oversight already minimizes harm through the "3Rs" principle of replacement, reduction, and refinement. Opponents highlight apes' demonstrated , , and emotional depth—evidenced by mirror self-recognition in chimpanzees and orangutans—as grounds for granting them moral consideration akin to s, arguing that invasive procedures inflict undue psychological and physical disproportionate to benefits, especially given high failure rates in translating findings to human trials. Welfare costs include from and experimentation, with critics asserting that apes' (up to 60 years) amplifies lifetime impacts, and that public sentiment, influenced by animal rights activism, views such research as outdated in an era of advanced and genetic tools. These positions often reflect broader ideological divides, where institutional biases in academia toward anthropomorphic interpretations may overstate ape "" at the expense of empirical human-centric priorities. Policy responses have trended toward restriction, with the enacting Directive 2010/63/EU in 2010, effectively banning great ape experiments except in unforeseen cases of human epidemic threats, a move solidified by 2013 and driven by ethical campaigns from groups like the . In the United States, the announced in 2015 the retirement of all federally owned chimpanzees from research, transferring approximately 50 remaining animals to sanctuaries after a 2011 Institute of Medicine review deemed most uses unnecessary, phasing out invasive biomedical studies amid similar activist pressures. These shifts, occurring between 2011 and the 2020s, have prompted critiques that they impede scientific progress by limiting models essential for unpredictable threats like emerging viruses, potentially delaying breakthroughs where alternatives prove insufficient. Controversies persist, including the 2025 scrutiny of Mauritius's export of thousands of long-tailed macaques for global research—though not great apes, this highlights ethical parallels in sourcing amid welfare allegations of and —underscoring tensions between supply demands and conservation. Historically, interpretations of ape dominance hierarchies and sexual behaviors have been accused of sexist biases, with mid-20th-century studies sometimes extrapolating coercive patterns to justify norms, reflecting researcher preconceptions rather than neutral observation. Such politicized restrictions, often amplified by media and activist narratives skeptical of industry motives, risk prioritizing symbolic over causal of net welfare gains from targeted research.

Cultural Representations and Misconceptions

In various global folklores, apes and closely related have been represented as tricksters or demonic figures, embodying cunning and rather than . For instance, in ancient Asian and African traditions, monkey-like entities often appear as deceptive intermediaries between humans and the divine, such as in African American oral tales or demons in Hindu epics. These depictions stem from observed behaviors like tool improvisation and social in wild populations, yet they exaggerate apes' agency into guile without evidence of intent beyond instinctual survival. Modern media has amplified dual stereotypes of apes as either brutish aggressors or anthropomorphized intellectuals, often detached from empirical observations. Iconic examples include the 1933 film King Kong, portraying a gigantic gorilla as a rampaging force symbolizing untamed primal fury, and the Planet of the Apes franchise starting in 1968, where apes evolve articulate societies overthrowing human dominance. Such narratives project human social conflicts onto apes, inflating their cognitive capacities—apes demonstrate problem-solving like nut-cracking with stones since at least 4,000 years ago in Côte d'Ivoire, but lack symbolic language or cumulative culture-building seen in humans. This anthropomorphism fosters misconceptions of apes as proto-humans, overlooking genetic divergences of 1-7% from Homo sapiens that preclude equivalent abstract reasoning. A persistent casts bonobos as utopian pacifists in female-led, sex-mediated harmony, contrasting "killer ape" , but field data reveal comparable or exceeding male-on-male violence, including lethal coalitions and . Studies from across multiple sites document bonobo males initiating fatal attacks at rates rivaling chimpanzees, driven by mating competition rather than resolved through purportedly peaceful rituals, debunking the "" ideal that ignores coercion in all great ape hierarchies. These errors arise from selective observations in , where reduced territorial pressures mask wild dynamics, and reflect broader cultural biases favoring egalitarian projections over causal drivers like resource scarcity and . Empirical tracking via camera traps and genetic paternity tests confirms as adaptive, not aberrant, underscoring in mitigating such impulses through institutions rather than innate benevolence.

References

  1. https://www.thoughtco.com/apes-hominoidea-profile-130639
  2. https://onlinelibrary.wiley.com/doi/10.1002/9781119179313.wbprim0150
  3. https://australian.museum/learn/science/human-evolution/humans-are-apes-great-apes/
  4. https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/hominoids
  5. https://pmc.ncbi.nlm.nih.gov/articles/PMC7521350/
  6. https://www.nature.com/articles/s41562-024-01836-5
  7. https://iucn.org/news/species/201609/four-out-six-great-apes-one-step-away-extinction-%25E2%2580%2593-iucn-red-list
  8. http://www.primate-sg.org/red_list_threat_status/
  9. https://www.biologyonline.com/dictionary/ape
  10. https://www.ncbi.nlm.nih.gov/books/NBK232185/
  11. https://study.com/learn/lesson/greater-lesser-apes-overview-differences.html
  12. https://evolution.berkeley.edu/what-are-evograms/the-emergence-of-humans/
  13. https://whyevolutionistrue.com/2012/03/22/were-all-apes-including-dawkins/
  14. https://a-z-animals.com/animals/ape/
  15. https://socialsci.libretexts.org/Courses/Fresno_City_College/ANTH_1%253A_Introduction_to_Biological_Anthropology_%28Taylor%29/05%253A_Meet_the_Living_Primates/5.01%253A_Meet_the_Living_Primates/5.1.03%253A_Primate_Diversity
  16. https://www.etymonline.com/word/ape
  17. https://www.johnhawks.net/p/how-did-english-come-to-have-different-words-for-ape-and-monkey
  18. https://www.reddit.com/r/history/comments/99poou/what_did_ancient_and_medieval_people_think_of_apes/
  19. https://brewminate.com/monkey-see-monkey-do-monkey-sin-the-devilish-ape-of-medieval-manuscripts/
  20. https://www.linnean.org/learning/who-was-linnaeus/linnaeus-and-race-easy-read
  21. https://www.realclearscience.com/articles/2018/01/08/how_did_we_realize_that_humans_are_just_one_of_the_great_apes_110510.html
  22. https://www.britannica.com/animal/Hominidae
  23. https://www.researchgate.net/publication/306304777_Phylogeny_and_Classification_of_Gibbons_Hylobatidae
  24. https://seaworld.org/animals/all-about/gorilla/classification/
  25. https://www.berggorilla.org/en/gorillas/species/taxonomy/articles-taxonomy/gorilla-diversity/
  26. https://www.nature.com/scitable/knowledge/library/primate-speciation-a-case-study-of-african-96682434/
  27. https://sites.lsa.umich.edu/tsebelis/wp-content/uploads/sites/382/2016/04/nature09094.pdf
  28. https://pmc.ncbi.nlm.nih.gov/articles/PMC4059600/
  29. https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1601-5223.2000.00217.x
  30. https://www.nature.com/articles/s41598-019-51685-w
  31. https://www.science.org/doi/10.1126/science.abb4363
  32. https://www.nature.com/scitable/knowledge/library/hominoid-origins-135874580/
  33. https://www.scientificamerican.com/article/what-does-the-fact-that-w/
  34. https://pl.khanacademy.org/partner-content/amnh/human-evolutio/human-evolution-the-evidence/a/understanding-our-past-dna
  35. https://bmcgenomics.biomedcentral.com/articles/10.1186/s12864-020-06962-8
  36. https://www.linnean.org/learning/who-was-linnaeus/linnaeus-and-race
  37. http://www.macroevolution.net/ape-to-human-evolution.html
  38. https://www.sciencedirect.com/topics/agricultural-and-biological-sciences/pongidae
  39. https://www.nature.com/scitable/knowledge/library/hominin-taxonomy-and-phylogeny-what-s-in-142102877/
  40. https://pmc.ncbi.nlm.nih.gov/articles/PMC223432/
  41. https://link.springer.com/chapter/10.1007/978-1-4684-8854-8_5
  42. https://bioone.org/journals/primate-conservation/volume-2006/issue-21/0898-6207.21.1.103/Gibbon-Hylobatidae-Species-Identification-Recommended-for-Rescue-or-Breeding-Centers/10.1896/0898-6207.21.1.103.full
  43. https://pmc.ncbi.nlm.nih.gov/articles/PMC11312596/
  44. https://www.pacb.com/publications/complete-sequencing-of-ape-genomes-2/
  45. https://www.donsmaps.com/proconsul.html
  46. https://socialsci.libretexts.org/Bookshelves/Anthropology/Biological_Anthropology/Explorations_Lab_and_Activities_Manual/08%253A_Primate_Evolution/8.01%253A_Fossil_Primates
  47. https://pmc.ncbi.nlm.nih.gov/articles/PMC1571308/
  48. https://evolution-outreach.biomedcentral.com/articles/10.1007/s12052-010-0251-z
  49. https://pmc.ncbi.nlm.nih.gov/articles/PMC2148271/
  50. https://www.tandfonline.com/doi/full/10.1080/08912963.2022.2057226
  51. https://bmcecolevol.biomedcentral.com/articles/10.1186/1471-2148-9-259
  52. https://www.pnas.org/doi/10.1073/pnas.1600374113
  53. https://www.nature.com/articles/s41586-025-08816-3
  54. https://www.biorxiv.org/content/10.1101/2025.08.15.670618v2.full.pdf
  55. https://phys.org/news/2025-04-ape-species-genomes-sequenced-telomere.html
  56. https://anatomypubs.onlinelibrary.wiley.com/doi/10.1002/ar.23743
  57. https://pmc.ncbi.nlm.nih.gov/articles/PMC2732797/
  58. https://pmc.ncbi.nlm.nih.gov/articles/PMC7770136/
  59. https://www.reuters.com/article/lifestyle/science/genetic-study-finds-severe-inbreeding-in-mountain-gorillas-idUSKBN0N021J/
  60. https://www.biorxiv.org/content/10.1101/2025.02.18.638799v1.full-text
  61. https://socialsci.libretexts.org/Courses/Fresno_City_College/ANTH_1%253A_Introduction_to_Biological_Anthropology_%28Taylor%29/09%253A_Early_Hominins/9.01%253A_Early_Hominins/9.1.03%253A_Derived_Adaptations-_Bipedalism
  62. https://animaldiversity.org/accounts/Hominidae/
  63. https://pubmed.ncbi.nlm.nih.gov/11786990/
  64. https://www.psu.edu/news/research/story/not-much-size-difference-between-male-and-female-australopithecines
  65. https://www.nature.com/scitable/knowledge/library/characteristics-of-crown-primates-105284416/
  66. https://pmc.ncbi.nlm.nih.gov/articles/PMC2817421/
  67. https://www.britannica.com/animal/primate-mammal/Sensory-reception-and-the-brain
  68. https://primatesinc.com/primate-differences/
  69. https://pmc.ncbi.nlm.nih.gov/articles/PMC10480693/
  70. https://bio.libretexts.org/Courses/Lumen_Learning/Fundamentals_of_Biology_I_%28Lumen%29/15%253A_Module_12-_Vertebrates/15.17%253A_Primates
  71. https://pmc.ncbi.nlm.nih.gov/articles/PMC2860095/
  72. https://www.researchgate.net/figure/Scapular-and-humeral-characteristics-of-an-ape-chimpanzee-compared-to-a-quadrupedal_fig1_316658418
  73. https://pubmed.ncbi.nlm.nih.gov/17170554/
  74. https://www.pnas.org/doi/10.1073/pnas.082105099
  75. https://www.npr.org/sections/thesalt/2017/03/27/521423216/what-gave-some-primates-bigger-brains-a-fruit-filled-diet
  76. https://quizlet.com/378314899/anthro-2200-unit-1112-flash-cards/
  77. https://pmc.ncbi.nlm.nih.gov/articles/PMC4804131/
  78. https://socialsci.libretexts.org/Courses/Lake_Tahoe_Community_College/ANT-103%253A_Physical_Biological_Anthropology/05%253A_Meet_the_Living_Primates/5.03%253A_Primate_Diversity
  79. https://www.aza.org/connect-stories/stories/navigating-chimpanzee-society-fission-fusion-maryland-zoo
  80. https://www.cambridge.org/core/books/wild-chimpanzees/chimpanzee-fissionfusion-social-organization-and-its-conservation-implications/58838D21EE07134F0526CF89D37F43FD
  81. https://pmc.ncbi.nlm.nih.gov/articles/PMC9515636/
  82. https://www.ab.mpg.de/377628/complex-sociality-lessons-from-bonobos
  83. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-bonobo/
  84. https://gorillafoundation.nl/social-structure/
  85. https://www.berggorilla.org/en/gorillas/general/behaviour/articles-behaviour/gorilla-group-formation-and-dynamics/
  86. https://onlinelibrary.wiley.com/doi/abs/10.1111/j.1439-0310.1989.tb00754.x
  87. https://orangutan.org/orangutan-facts/orangutan-behavior/
  88. https://ielc.libguides.com/sdzg/factsheets/orangutans/behavior
  89. https://pmc.ncbi.nlm.nih.gov/articles/PMC6971177/
  90. https://www.researchgate.net/publication/277666307_Primate_Allomaternal_Care
  91. https://nagonline.net/wp-content/uploads/2013/12/Chimpanzee-Nutrition.pdf
  92. https://pmc.ncbi.nlm.nih.gov/articles/PMC4495928/
  93. https://pmc.ncbi.nlm.nih.gov/articles/PMC3083508/
  94. https://journals.plos.org/plosone/article?id=10.1371/journal.pone.0129254
  95. https://pmc.ncbi.nlm.nih.gov/articles/PMC7816224/
  96. https://chimpsnw.org/2013/08/tool-use-and-the-termite-mound/
  97. https://ielc.libguides.com/sdzg/factsheets/westernlowlandgorilla/diet
  98. https://www.eva.mpg.de/documents/Wiley-Blackwell/Ostrofsky_Fruit-feeding_AmJPhysAnthr_2020_3221596.pdf
  99. https://link.springer.com/article/10.1023/A:1012021617737
  100. https://www.researchgate.net/publication/311006743_Nutritional_ecology_of_wild_Bornean_orangutans_Pongo_pygmaeus_wurmbii_in_a_peat_swamp_habitat_Effects_of_age_sex_and_season
  101. https://phys.org/news/2011-07-orangutans-movements-swamp-forest.html
  102. https://sites.lsa.umich.edu/ajmarsha/wp-content/uploads/sites/162/2021/11/Marshall-et-al.-2021-Biotic-and-abiotic-drivers-of-dispersion-dynamics-in-a-large%25E2%2580%2590bodied-tropical-vertebrate-the-Western-Bornean-orangutan.pdf
  103. https://royalsocietypublishing.org/doi/10.1098/rspb.2023.0430
  104. https://pubmed.ncbi.nlm.nih.gov/28431195/
  105. https://pmc.ncbi.nlm.nih.gov/articles/PMC2750765/
  106. https://www.researchgate.net/figure/Frequencies-of-support-size-use-during-locomotion-of-cao-vit-gibbons_tbl2_259351417
  107. https://www.nature.com/scitable/knowledge/library/primate-locomotion-105284696/
  108. https://www.sciencedirect.com/science/article/abs/pii/S0047248418301192
  109. https://www.cell.com/current-biology/pdf/S0960-9822%2822%2901474-9.pdf
  110. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-lar-gibbon/
  111. https://www.nature.com/articles/s41598-023-44192-6
  112. https://www.sciencedirect.com/science/article/pii/S0003347224000708
  113. https://www.sciencedirect.com/science/article/abs/pii/S0047248404000119
  114. https://www.masterclass.com/articles/jane-goodall-explains-chimpanzee-tool-use
  115. https://www.jstor.org/stable/4535262
  116. https://www.eva.mpg.de/fileadmin/content_files/staff/boesch/pdf/fol_prim_tool_use_making.pdf
  117. https://www.sciencedirect.com/science/article/pii/S0047248424000903
  118. https://pmc.ncbi.nlm.nih.gov/articles/PMC9593238/
  119. https://www.orangutanrepublik.org/learn/orangutan-specifics/intelligence-and-innovation/
  120. https://suaq.org/wp-content/uploads/2017/06/Dissertation_E.Meulman_2013.pdf
  121. https://www.nature.com/articles/s41598-018-31331-7
  122. https://www.discovermagazine.com/tooling-through-the-trees-11790
  123. https://pmc.ncbi.nlm.nih.gov/articles/PMC1236726/
  124. https://neprimateconservancy.org/white-handed-gibbon/
  125. https://primate.wisc.edu/primate-info-net/pin-factsheets/pin-factsheet-white-cheeked-gibbon/
  126. https://www.pnas.org/doi/10.1073/pnas.0901008106
  127. https://onlinelibrary.wiley.com/doi/10.1002/ajp.23311
  128. https://www.sciencedirect.com/science/article/pii/S0168159102002575
  129. https://www.sciencedirect.com/science/article/pii/S096098220702088X
  130. https://inference-review.com/article/primate-memory
  131. https://onlinelibrary.wiley.com/doi/10.1002/ajp.22248
  132. https://www.bps.org.uk/psychologist/deferred-imitation-children-and-apes
  133. https://pmc.ncbi.nlm.nih.gov/articles/PMC2660989/
  134. https://pmc.ncbi.nlm.nih.gov/articles/PMC5486474/
  135. https://pmc.ncbi.nlm.nih.gov/articles/PMC9358316/
  136. https://royalsocietypublishing.org/doi/10.1098/rstb.2018.0403
  137. https://www.nature.com/articles/s42003-024-07206-1
  138. https://www.mpi.nl/publications/item1170576/keeping-track-time-evidence-episodic-memory-great-apes
  139. https://www.sciencedirect.com/science/article/pii/S0960982213008397
  140. https://caplab.yale.edu/sites/default/files/files/2016b-MartinSantos.pdf
  141. https://pmc.ncbi.nlm.nih.gov/articles/PMC2346530/
  142. https://www.biorxiv.org/content/10.1101/2023.08.02.551487v1.full.pdf
  143. https://wires.onlinelibrary.wiley.com/doi/10.1002/wcs.1516
  144. https://www.eva.mpg.de/documents/Springer/Tennie_Island_Nature-of-culture_2016_2255850.pdf
  145. https://www.nature.com/articles/s41598-017-16844-x
  146. https://scholars.duke.edu/publication/1402109
  147. https://pubmed.ncbi.nlm.nih.gov/28162899/
  148. https://pmc.ncbi.nlm.nih.gov/articles/PMC10907097/
  149. https://files.commons.gc.cuny.edu/wp-content/blogs.dir/4133/files/2018/08/deWaal.altruism.pdf
  150. https://royalsocietypublishing.org/doi/abs/10.1098/rstb.2012.0191
  151. https://www.researchgate.net/publication/230749122_Are_there_geniuses_among_the_apes
  152. https://pmc.ncbi.nlm.nih.gov/articles/PMC4704114/
  153. https://www.cmnh.org/exhibits/g4-3-301
  154. https://carta.anthropogeny.org/moca/topics/control-paternity
  155. https://www.pnas.org/doi/10.1073/pnas.1307903110
  156. https://www.berggorilla.org/en/gorillas/general/behaviour/articles-behaviour/translate-to-englisch-mehrschichtige-gesellschaften/
  157. https://mytext.cnm.edu/lesson/chapter-7-primates-2/
  158. https://www.milkgenomics.org/?splash=evolving-motherhood-when-to-wean-part-i
  159. https://www.sciencedirect.com/science/article/abs/pii/S0047248404001605
  160. https://pmc.ncbi.nlm.nih.gov/articles/PMC9699942/
  161. https://www.science.org/doi/10.1126/sciadv.aaz5746
  162. https://pmc.ncbi.nlm.nih.gov/articles/PMC9094693/
  163. https://pmc.ncbi.nlm.nih.gov/articles/PMC3574345/
  164. https://www.nature.com/articles/npre.2010.4411.1.pdf
  165. https://www.researchgate.net/publication/6897135_Evolution_of_brain_size_and_juvenile_periods_in_primates
  166. https://pmc.ncbi.nlm.nih.gov/articles/PMC3448989/
  167. https://a-z-animals.com/animals/chimpanzee/chimpanzee-lifespan/
  168. https://chimpsnw.org/2013/03/how-long-do-chimpanzees-live/
  169. https://www.wildlifenomads.com/blog/how-long-do-apes-live-types-more-facts/
  170. https://davidshepherd.org/species/chimpanzees/
  171. https://www.ifaw.org/animals/chimpanzees
  172. https://www.awf.org/wildlife-conservation/bonobo
  173. https://ielc.libguides.com/sdzg/factsheets/bonobo/population
  174. https://www.berggorilla.org/en/gorillas/gorilla-numbers/gorilla-numbers/gorillas-on-the-iucn-red-list-of-threatened-species/
  175. https://acaciasafari.co.ug/faqs/how-many-mountain-gorillas-are-left-in-the-wild/
  176. https://gorillas.org/about-gorillas/cross-river-gorillas/
  177. https://www.worldwildlife.org/species/orangutan/
  178. https://socialsci.libretexts.org/Bookshelves/Anthropology/Biological_Anthropology/EXPLORATIONS%253A__An_Open_Invitation_to_Biological__Anthropology_1e/18%253A_Primate_Conversation/18.2%253A_Threats_to_Primates
  179. https://www.berggorilla.org/en/gorillas/threats-protection/threats/articles-threats/grasp-iucn-great-apes-status-report-2018-summary/
  180. https://p-wac.org/what-is-a-primate/threats/
  181. https://www.sciencedirect.com/science/article/abs/pii/S0006320710004714
  182. https://pmc.ncbi.nlm.nih.gov/articles/PMC7123567/
  183. https://www.un.org/en/chronicle/article/illegal-commercial-bushmeat-trade-central-and-west-africa
  184. https://www.lpzoo.org/primates-as-pets-by-the-numbers/
  185. https://www.science.org/content/article/illegal-ape-trade-nearly-nine-times-bigger-thought
  186. https://davidshepherd.org/species/chimpanzees/trade-statement/
  187. https://pmc.ncbi.nlm.nih.gov/articles/PMC7107455/
  188. https://pmc.ncbi.nlm.nih.gov/articles/PMC9323791/
  189. https://www.woah.org/en/disease/ebola-virus-disease/
  190. https://www.weforum.org/stories/2021/06/great-apes-could-lose-94-of-african-home-due-to-climate-crisis-and-other-human-actions-study-finds/
  191. https://onlinelibrary.wiley.com/doi/10.1111/ddi.13358
  192. https://www.scientificamerican.com/article/great-apes-biggest-threat-is-human-activity-not-habitat-loss/
  193. https://www.researchgate.net/publication/378324526_Future_coexistence_with_great_apes_will_require_major_changes_to_policy_and_practice
  194. https://pmc.ncbi.nlm.nih.gov/articles/PMC12139133/
  195. https://conbio.onlinelibrary.wiley.com/doi/full/10.1111/csp2.13205
  196. https://www.sciencedirect.com/science/article/pii/S000632072400082X?via%253Dihub
  197. https://besjournals.onlinelibrary.wiley.com/doi/full/10.1111/1365-2664.13873
  198. https://www.arcusfoundation.org/advancing-ape-conservation-from-many-angles/
  199. https://www.frontiersin.org/journals/conservation-science/articles/10.3389/fcosc.2023.1225911/full
  200. https://kar.kent.ac.uk/104044/1/E.%2520Meijaard%2520-%2520~Apes%2520and%2520Agriculture%2520-%2520PPDF.pdf
  201. https://news.[mongabay](/page/Mongabay).com/2024/10/wwf-report-offers-glimmer-of-conservation-hope-yet-warns-of-a-planet-in-peril/
  202. https://news.umich.edu/new-methods-of-conservation-needed-to-prevent-the-extinction-of-great-apes/
  203. https://www.pbs.org/wgbh/aso/databank/entries/dm52sa.html
  204. https://cjh.uchicago.edu/issues/fall13/2.4.pdf
  205. https://jamanetwork.com/journals/jama/fullarticle/346443
  206. https://academic.oup.com/ilarjournal/article/42/2/85/727168
  207. https://www.nature.com/articles/nrg981
  208. https://www.technologynetworks.com/genomics/news/ape-genomes-hold-new-clues-on-evolution-immunity-and-brain-development-398339
  209. https://janegoodall.org/our-story/about-jane/
  210. https://www.science.org/content/article/jane-goodall-famed-primatologist-changed-way-we-thought-about-apes
  211. https://inmersionafrica.com/dian-fossey-and-her-contribution-to-gorilla-conservation/
  212. https://link.springer.com/article/10.1007/s10764-021-00245-z
  213. https://projectchimps.org/2010s-our-legacy-for-research-chimps/
  214. https://pmc.ncbi.nlm.nih.gov/articles/PMC5636196/
  215. https://www.sciencedirect.com/science/article/abs/pii/S0140673608604894
  216. https://pmc.ncbi.nlm.nih.gov/articles/PMC5886318/
  217. https://pmc.ncbi.nlm.nih.gov/articles/PMC7888082/
  218. https://faunalytics.org/using-primates-in-research-scientific-and-ethical-considerations/
  219. https://bmcmedethics.biomedcentral.com/articles/10.1186/s12910-021-00580-z
  220. https://www.peta.org/issues/animals-used-for-experimentation/chimpanzees-laboratories/
  221. https://www.theguardian.com/us-news/2015/nov/19/national-institutes-health-end-chimpanzee-research
  222. https://www.theguardian.com/science/2008/nov/02/primate-monkey-animal-testing-drugs
  223. https://www.science.org/content/article/mauritius-research-monkeys-are-big-business-and-big-controversy
  224. https://www.scientificamerican.com/blog/primate-diaries/httpblogsscientificamericancomprimate-diaries20110720science-of-sexism/
  225. https://www.theatlantic.com/magazine/archive/2022/06/ape-gender-frans-de-waal-lucy-cooke/629637/
  226. https://www.luminarium.org/mythology/monkeygods.htm
  227. https://cybermunkiez.com/blogs/news/monkeys-in-ancient-mythology-tricksters-gods-and-demons
  228. https://primatefolklore.wordpress.com/
  229. https://www.complex.com/pop-culture/a/jamieiovine/top-10-most-influential-apes-popular-culture
  230. https://the-artifice.com/primate-representation-in-media/
  231. https://www.emory.edu/LIVING_LINKS/OurInnerApe/pdfs/anthropodenial.html
  232. https://www.theguardian.com/science/2024/apr/12/bonobos-not-the-peace-loving-primates-once-thought-study-reveals
  233. https://cosmosmagazine.com/nature/animals/bonobos-arent-as-peaceful-as-previously-thought/
  234. https://crev.info/2023/12/pat-a-pan-chimp-bonobo-myth-debunked/
  235. https://www.psychologytoday.com/us/blog/smarter-than-you-think/202311/bonobos-the-peaceful-apes
  236. https://dokumen.pub/aggression-in-humans-and-other-primates-biology-psychology-sociology-9783110291360-9783110291339.html
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